00

?r
OU 162034 >m
 QSMANIA UNIVERSITY LIBRARY
Call No. fiftft/ ft S&" Accession Na

J, C.I
Title Hi * i^ ...... 4

This book should be r&urncti' on or before the datfe last marke

 KEY TO TISSUES
STEMS AND PETIOLES

SCLERENCHYMA XYU.E

CHLORENCHYMA COLLENCHYMA

CORK PHLOEM

WOOD
VESSELS

RAYS PARENCHYMA

SEPTATE FIBRES PHL.OEM

TRACHEIDS SCLERENCHYMA

which have
The above types of shading have been used for the tissues in the drawings
been specially prepared for this book. They do not apply to drawings where an author's
name is given. Septate fibres not shaded except where forming patches in the ground-
tissue of the wood. Transverse sections of Wood x 50, Rays x 90.
 ANATOMY OF THE
DICOTYLEDONS
ANATOMY OF THE
DICOTYLEDONS
LEAVES, STEM, AND WOOD
IN RELATION TO TAXONOMY
WITH NOTES ON ECONOMIC USES
BY
C. R. METCALFE
MJV., PH.D.
JODRELL LABORATORY, ROYAL BOTANIC GARDENS, KEW
AND
L. CHALK
M.A., D.PHIL.
IMPERIAL FORESTRY INSTITUTE, OXFORD

WITH THE ASSISTANCE OF

M. M. CHATTAWAY
HARE
C. L.
F. R. RICHARDSON
E. M. SLATTER

VOLUME I

OXFORD
AT THE CLARENDON PRESS
Oxford University Press, Amen House, London E.C. 4
GLASGOW NEW YORK TORONTO MFLBOURNF WELLINGTON
BOMBAY CALCUTTA MADRAS KARACHI
CAPE TOWN IBADAN NAIROBI ACCRA SINGAPORE

FIRST EDITION I95O

REPRINTED LITHOGRAPHICALLY IN GREAT BRITAIN
AT THE UNIVERSITY PRESS, OXFORD
BY CHARLES BATEY, PRINTER TO THE UNIVERSITY
FROM CORRECTED SHEETS OF FIRST EDITION
1957
 FOREWORD
IT is more than forty years since L. A. Boodle, then in charge of the Jodrell
Laboratory, and F. E. Fritsch, then also working at Kew, collaborated in the
translation of Solereder's work on the systematic anatomy of the Dicotyledons.
Boodle brought to this task a profound knowledge of the subject, acquired in
the course of his official duties, which gave to the translation an even greater
value than that of the German original. During the past forty years the range
of material available for study has been greatly augmented, and a large mass
of data has been accumulated at Kew, which has enabled generalizations to be
based on a surer foundation and provided the means of filling many of the
gaps in Solereder's work. In this tisfe Dr. Metcalfe, who succeeded Boodle
in the charge of the Jodrell Laboratory, has ably carried forward the work
which Boodle began, to such good purpose that the present publication must
be regarded as a new work incorporating and correcting the old, and in a far
more real sense as a
comprehensive survey of the anatomical features of
dicotyledonous plants considered as taxonomic characteristics.
In no aspect of plant anatomy has more extensive progress been made
during the past half-century than in the study of the secondary wood, in no
small degree because of the growing appreciation of the importance of
anatomical characters for the identification of commercial timbers. It is
therefore a pleasure to record the debt to Dr. L. Chalk who has collaborated
in the production of this work and to whom the original descriptions of the
secondary woods are entirely due.
The whole constitutes a monument to the assiduous labours of the authors
and remain the standard reference work on the subject and indis-
will long

pensable to every Botanical Library of any importance.

The living collections at Kew, now numbering some 45,000 species, have
provided an invaluable source of material, but in addition specimens have
been sought from all parts of the world, and there are but few larger groups
of all the vast assemblage of dicotyledonous plants that have not been examined
in respect to one or more representatives. The large collection of wood
specimens and microscope slides at the Imperial Forestry Institute, Oxford,
formed the basis of the work on wood anatomy, and this was supplemented
by data specially collected by Dr. M. M. Chattaway for all the genera in the
Yale Collection not available in Oxford. Thus the picture of the range of
structure evolved by the various taxonomic groups is presented with a com-
pleteness that was never before possible and with a simplification of treatment
that a mastery of detail can alone achieve.
E. J. SALISBURY
ROYAL BOTANIC GARDENS
KEW
 PREFACE
THIS reference book aims at providing a summary of our present knowledge
of the anatomy of the vegetative organs of the Dicotyledons. It has been
written mainly so as to emphasize the taxonomic and phylogenetic value of
histology, thus perpetuating and extending Solereder's chief aim when he
wrote his Systematic Anatomy of the Dicotyledons, of which the English transla-
tion was published in 1908. At the same time the authors hope that botanists
who are interested in other aspects of the anatomy of the Dicotyledons will
find the book useful as a work of reference. Some attention has accordingly
been devoted to ecological and developmental anatomy, and references have
been included in the bibliography which should enable workers to find more
detailed information concerning these subjects. In addition an attempt has
been made to appeal to a wider range of readers by including, under many
of the families, a section headed 'Economic Uses'. In particular it is hoped
that this will be of interest to pharmacognosists, foresters, wood technologists,
public analysts, and all who have to deal with plant products of economic
importance and their microscopical structure. Since the book is concerned
solely with the anatomy of the vegetative organs, the microscopy of economic
products derived from flowers or fruits has been omitted.
Although British taxonomists frequently recognize the value of attempt-
ing to investigate taxonomic problems along anatomical lines, this method
is seldom practised in the herbarium. This is partly because herbarium
botanists are so few that they are very hard pressed to complete their everyday
work of identification and description of new species. Furthermore it is very
seldom that an herbarium botanist receives more than a very elementary
training in plant structure, so that he lacks the confidence and knowledge
which are necessary to assess the value of anatomical characters. In Britain,
the relationship between plant structure and physiological function or en viron-
ment has commanded more attention than that which exists between struc-
ture and taxonomic affinity. Herbarium botanists on the European continent
have devoted more attention to systematic anatomy, while it is well known
that the Englerian system of classification is partly based on anatomical
characters observed by continental taxonomists. In recent years there has
been a world-wide interest in the systematic anatomy of the secondary xylem,
but this subject has been investigated largely by specialists whose basic train-
ing has been in forestry or wood technology rather than, in taxonomy. This
has had two consequences: (i) that herbarium botanists find the need of
some authoritative guidance in interpreting the anatomical evidence (ii) that
;

anatomists tend to be segregated into those with a knowledge of wood struc-

ture and those who are familiar with the structure of other parts of the plant
body. In order to meet these two defects a book is required in which the
combined knowledge of the general systematic anatomist together with that
of the specialist in wood structure is summarized and presented in a form
which can be generally understood by anatomists, and its taxonomic signi-
ficance made clear to the herbarium botanist.
We undertook to write this book in the hope that a modern counterpart to
viii PREFACE
Solereder's well-known treatise on the same subject would lead towards a
more natural classification of the Dicotyledons and make some contribution
to our knowledge of phylogeny. Although we had felt for many years that a
reference book of the kind we have written was badly needed, and had indeed
been recording facts and making reference collections of microscope slides
with some such purpose in mind, it was not until after a meeting of the
Systematics Association in 1938 that the late Sir Arthur Hill, then Director
of Kew, raised the question of publication with the Delegates of the Clarendon
Press. The task of writing the book was started in 1939.
In writing the descriptions of the 'Leaf and 'Stem' which constitute a large
part of the present book, Solereder's previous treatise has provided the
starting point, and, indeed, were it not for the assistance derived from this
source, our task would not have been completed for many years. The facts
obtained from Solereder's book have been combined with those obtained
from work published in journals and books since 1908 and further expanded
by a large number of original observations. The information thus accumulated
ispresented in as concise a form as possible. Solereder's somewhat ponderous
style has undoubtedly deterred many botanists from using his treatise or fully
appreciating the value of its contents. The more condensed mode of presenta-
tion now adopted has necessitated the omission of some of the exceedingly
detailed information included in Solereder's book and in the subsequent
literature. It seemed to the authors, however, that it is preferable for specialists

studying certain families or groups to consult the original sources for such
details. For this reason a very full bibliography to literature that has
appeared since 1908 has been included, whilst references to the older litera-
ture that may be found in Solereder's book have been omitted.
Solereder fully realized and stressed the taxonornic importance of wood
structure. A
justifiable criticism of his book, however, is that most of his
remarks on this subject refer to the wood of small twigs such as occur in
herbarium specimens rather than to samples which may be described as
'timber'. The microscopical features of the wood from young twigs frequently
differ considerably from those visible in wood specimens of the type which
are the special concern of the forester or wood technologist. In the present
work we have, as far as possible, drawn a clear distinction between the xylem
of young twigs, which is described under 'Stem*, and the wood of the forester's
sample, which is described under 'Wood'.
The illustrations include some of those which appeared in Solereder's book,
but as his treatise has been frequently criticized because they were so few,
very many new ones have been added. The new drawings are intended to
show diagrammatically the structure of young stems and petioles in transverse
sections, of the wood in transverse sections ( x 50), and of the medullary
rays of the wood in tangential sections ( x 95). The species illustrated have
been selected in order to indicate the range of structure in each family, so far
as the material available to us has made this possible. This has frequently
meant that little-known species have been illustrated in preference to those
which are more generally familiar or of greater economic importance. The
authors believe that this course will be found more valuable, from the
scientific point of view, in a book which is basically taxonornic. Almost all of
the new illustrations are original, and have been drawn from slides in our
 PREFACE ix

respective reference collections. The authors are particularly indebted to

Dr. C. L. Hare for his painstaking assistance in preparing and arranging a
large part of these drawings, many of which are solely his own work. Thanks
are also due to Mr. J. S. Shaw who prepared for the press many of the draw-
ings of the wood made by Dr. Chalk. More recently considerable assistance in
the arrangement of the drawings has been received from Miss E. M. Slatter
and Mr. F. R. Richardson.
We are greatly indebted to our numerous friends and colleagues for their
encouragement, helpful suggestions, and criticisms throughout our long task.
When the work began, many of the preliminary descriptions of the wood
structure were ably drawn up by Dr. M. M. Chattaway. Later Dr. C. L.
Hare worked systematically through a large proportion of the reference collec-
tion of slides at Kew, recording his observations in the form of notes and
tables which have been of the greatest value when writing the text. We are
especially grateful to these two workers for their collaboration. As neither of
the authors is an expert in systematic botany it has been necessary to make
frequent calls on our taxonomic colleagues for their assistance. In the early
stages of our work we were helped particularly by Dr. T. A. Sprague and
Dr. W. B. Turrill, and later on by Dr. J. Ilutchinson, F.R.S. Taxonomic
advice concerning certain groups has also been given by Mr. H. K. Airy-Shaw,
Mr. N. Y. Sandwith, and other members of the staff of the Kew herbarium.
The work at the Jodrell Laboratory could not have been completed so soon
without the energy, patience, and skill which Mr. F. R. Richardson devoted to
preparing many thousands of microscope slides of material cultivated in the
gardens or housed in the reference collections in the Kew museums and
herbarium. We are also indebted to the gardens staff at Kew for the large
quantities of living material which they were able to supply, to Major A. A.
Dorrien Smith, D.S.O., for living specimens from his gardens at Tresco in
the Isles of Scilly, and to Mr. E. Nelmes for his assistance in preparing the
index.
The work on wood anatomy has been made possible by the large collection
of microscope slides at the Imperial Forestry Institute, Oxford, and, though
it is
impossible to list all those who have helped to build up this collection,
mention must be made of the Forest Products Research Laboratory, Princes
Risborough (Mr. B. J. Rendle), the Yale School of Forestry (Prof. S. J.
Record), the Forest Research Institute, Kepong, Malaya (Dr. H. E. Desch),
and the Forest Department, Gold Coast Colony (Mr. C. Vigne) for their
exceptionally valuable contributions of material, and of Mr. A. A. Shaw,
Mr. C. W. Bond and Mr. P. Franklin for their assistance in preparing the
slides.

January 1950. C.R.M.

L.C.
 CONTENTS
VOLUME I

INTRODUCTION
ANATOMY AND TAXONOMY
ANATOMY AND PHYLOGENY .....
. . . . xi

xxix

SELECTION AND PREPARATION OF MATERIAL

ARRANGEMENT OF THE FAMILIES
THE DESCRIPTIONS OF THE FAMILIES
.....
....
. . . liii

Iv

Ixii

DESCRIPTIONS OF THE FAMILIES

i. RANUNCULACEAE 156. UMBELLIFERAE

VOLUME II
... i

DESCRIPTIONS OF THE FAMILIES

157. ARALIACEAE 263. CERATOPHYLLACEAE . . .
725

LISTS OF FAMILIES IN WHICH CERTAIN DIAGNOSTIC

FEATURES OCCUR . . . . ,
.1326
APPENDIX I

WOODS

BIBLIOGRAPHY
.......
DISTRIBUTION OF CELL DIMENSIONS,

. .
ETC., IN

.
DICOTYLEDONOUS

. .
.1363
1360

INDEX 1459

KEY TO TISSUES IN THE DRAWINGS front endpaper

in each volume

PLATES
A. APOTRACHEAL PARENCHYMA . . . . \

between
B. PARATRACHEAL PARENCHYMA . . . .1
I
pp. xlviii-xlix
c. RAY TYPES . . . . .1
 INTRODUCTION
ANATOMY AND TAXONOMY
THE historical development of botany has been such that physiological and
anatomical investigations of plants have been unnecessarily separated from
studies of systematic arrangement, and workers in these artificially divided
fields have developed their own special techniques and outlooks. From time
to time anatomists, though in general more interested in structure in relation to
function than to classification, have made excursions into the realms of taxo-
nomy, and have added some solid contributions to the knowledge of systematy.
In general, however, the work of anatomists has tended to be overlooked or
mistrusted by their taxonomic colleagues. The chief reason for this is that
anatomists have not always realized the limitations of their mode of investiga-
tion and have sometimes drawn conclusions that, to a taxonomist, are
obviously highly improbable. Conversely many highly skilled taxonomists
have sometimes been unable to assess the value of anatomical investigations.
There have been signs in recent years, however, that as taxonomists have
learned the value of co-operation with cytologists and geneticists, so they are
coming to appreciate the contribution which anatomists can make to their
investigations. At the same time anatomists are ready to admit that anatomical
characters cannot by themselves form the basis of classification, but are to be
regarded rather as an extension of, and supplement to, those external morpho-
logical characters on which classifications have been built.

Practical Reasons for Using Anatomical Characters

The taxonomic value of cytological and genetical studies lies chiefly in the
interpretation of species and groups of lower rank. Anatomical structure, on
the other hand, is most likely to provide evidence concerning the inter-
relationships of larger groups such as families, or in helping to establish the
real affinities of genera of uncertain taxonomic status. Anatomy is of restricted
value for distinguishing species or groups of less than specific rank, because
the differences between them are usually quantitative rather than qualitative,
and often demand statistical treatment based on a greater amount of material
than is frequently available.
Anatomy sometimes proves very helpful for individual identifications. For
example, microscopical methods are of great value in establishing the identity
of herbarium specimens which are not accompanied by flowers or fruits.
That this can frequently be done has repeatedly been proved at Kew, when,
by microscopical examination, it has been possible to assign sterile specimens
to a family or genus, thus greatly simplifying the task of the herbarium
botanist. Microscopical methods are also often necessary to establish the
botanical identity of commercial samples of medicinal plants, timbers, fibres,
&c., and may play an important part in checking adulteration, substitution,
and fraud, and have on occasion been instrumental in helping to establish the
guilt or innocence of suspected criminals. These practical applications alone
provide sufficient cause to justify the use of anatomical methods in taxonomic
xii INTRODUCTION
investigations. Finally may be pointed out that a complete anatomical
it

survey is necessary as a preliminary to the interpretation of palaeobotanical

remains.

Characters of Taxonomic Importance

It will be readily appreciated that the taxonomic value of anatomical
characters is inversely proportional to their plasticity, those which are particu-
larly liable to become modified in response to environmental change being
most unsatisfactory. On the other hand no characters are strictly immutable.
In practice, however, many characters are more fixed than others, and it is on
those that are less plastic that the systematic anatomist must rely. It should
also be realized that conclusions supported or suggested by combinations of
characters are more likely to be correct than those which rest on one character
alone.^The feature or features to be employed will, however, depend on the
nature of the material to be examined as well as on the taxonomic problem to
be settled. For instance, an entirely different method of approach would be
necessary if the problem were to determine to what family a leafy twig belonged,
compared with the procedure a pharmacognosist would adopt when distin-
guishing between the leaves of two species of Digitalis. In solving the second
problem differences in the exact type and structure of the trichomes would be
of importance whilst one could also make use of measurable differences such
as those of the vein-islet number. In attempting to deal with the first problem
qualitative differences would be more important. Dimensions are generally
more liable to vary with conditions of growth than are differences in kind, and
should therefore be used with great caution when attempting to separate
species on anatomical grounds. ^ For example, a distinction between two
species based on the density of the hairs on the leaves or on stomatal size and
frequency is not likely to be satisfactory unless the differences are very marked,
as these features vary with environment or even in different parts of an
individual plant. Differences in the type of hair, on the other hand, usually
provide much more reliable grounds for separation. Failure to appreciate
the variability of dimensional characters has sometimes led to erroneous con-
clusions that have tended to bring the anatomical method into disrepute.
Plants which are not closely related to one another sometimes develop
similar habit forms or structural features. For example, there are certain
members of the Euphorbiaceae, Cactaceae, Geraniaceae, Compositae, and
Apocynaceae in which a stem succulent habit has been evolved, presumably
as a direct response to the dry ecological habitat of the plants in which they
are exemplified. Corresponding anatomical similarities, such as enlargement
of parenchymatous cells, have also arisen in the same plants. Although these
anatomical resemblances possess diagnostic value because they occur in but
few families, the greatest care must be taken to ensure that they are not by
themselves taken as evidence of taxonomic affinity between the plants in
which they are exhibited.
Many other examples of anatomical responses to ecological conditions could
be cited. All of these are well known to those who are interested in ecological
anatomy, but it is necessary to draw attention to a few of them, if only to
emphasize that they must be carefully distinguished from characters possess-
ing taxonomic value. Thus it may be noted that well- developed systems of
 INTRODUCTION xiii

intercellular spaces are characteristic of aquatic plants. The structure of the

mesophyll is also dependent on the nature of the habitat. For example,
some of the plants in dry localities may possess small leaves of the ericoid
type with inrolled margins, others such as some of the acacias have leaves in
which the mesophyll is isobilateral, and in plants with cylindrical leaves the
mesophyll is centric. Another variation sometimes occurs in species with
adpressed leaves in which spongy tissue is developed towards the adaxial
surface and palisade tissue towards the abaxial surface, thus reversing the
arrangement which is characteristic of most leaves with a dorsiventral meso-
phyll. Ecological specializations that occur in stems include the specially
large vessels in the xylem of climbing plants, and the ring-porosity of many
woods from temperate regions.
Another difficulty, which the systematic anatomist must face, is that some
very useful diagnostic characters, which have no apparent ecological signi-
ficance, have arisen quite independently in unrelated families. The following
examples will serve to illustrate this point.
Laticiferous elements are of restricted distribution and therefore of
diagnostic value, but the possession of these structures does not necessarily
imply affinity. They are to be found in such diverse families as Papaveraceae,
Eucommiaceae, Compositae, and Euphorbiaceae. Intraxylary phloem is
common to the closely related Apocynaceae and Asclepiadaceae and in this
instance provides confirmatory evidence of affinity. It nevertheless occurs
also in certain of the Lythraceae, Combretaceae, and Gentianaceae, as well as
in many other families that are not closely related to one another. Here again
it can be seen that intraxylary phloem is of taxonomic value because of its

restricted occurrence, but it may or may not be indicative of affinity. A

third
example is afforded by the nature of the petiolar vascular system. If trans-
verse sections be cut through corresponding regions (preferably immediately
below the lamina) of the leaves of Dicotyledons, it will be found that the
vascular systems can be arranged in a series of intergrading types; closely
similar forms frequently occur, however, in quite unrelated families. Never-
theless, the different types of petiolar structure are of diagnostic value because
of their restricted occurrence and relative stability.
It is important to realize that an anatomical feature which is of considerable

diagnostic value in one family is not necessarily of equivalent value in another.

This is not very remarkable when one recalls that the same principle applies
when using exomorphic characters for taxonomic purposes. All families are
not of equal status. Some consist of only one or a few genera, but many
species, others of many genera, but few species. Besides this, families vary-
considerably in size, and the lines of demarcation between them are largely a
matter of personal opinion, since no absolute criteria for their separation exist.
It is sometimes averred that the use of anatomical characters is undesirable
because this practice will lead to taxonomic conclusions completely at variance
with those that are now generally accepted. Experience shows, however, that
this is by no means wholly true. Many anatomical characters indicate relation-
ships in harmony with those suggested by exomorphic features including
those of the flower. Where exomorphic and anatomical characters are alleged
to point to different conclusions it will frequently be found that the discrepancy
is due to an incorrect
interpretation of the available facts. In cases of this kind
xiv INTRODUCTION
a joint reinvestigation by an anatomist and a systematist may quite well lead to
readjustments more in line with those suggested by the anatomical characters.
It is, however, most important to rely on anatomical characters of which the
taxonomic value has become well established. These will now be enumerated.

1. HAIRS
The diversity of external hairs is familiar to all taxonomists. Firstly there
are the glandular and non-glandular categories, each of which may be sub-
divided according to the number of component cells, degree of branching,
a,nd so forth. Whole families may frequently be recognized by the occurrence
of one or more distinctive types of hair. In other instances hairs are of more
value for the determination of species or genera. The length, size, and
density of hairs are more liable to vary with environment than is the occurrence
of different kinds, so that the former features are of more restricted taxonomic
value. There are families, e.g. the Theaceae, in which hairs are infrequent
and others, e.g. Cistaceae, in which they are abundant. Differences such as
these are of value. Smaller variations in size and density should, however, be
accepted as a basis for the separation of closely related genera and species only
after exhaustive investigation of a wide range of material.

2. STOMATA
It is known that stomatal frequency not only varies from point to point in
a single leaf but depends also on the level of insertion of the leaf on the stem,
whilst it is also strongly influenced by the conditions prevailing in the habitat.
It is scarcely surprising to find, therefore, that the number of stomata per
unit area, and the extent to which they are raised above or sunk below the
epidermis, is of very restricted importance compared with their appearance
in surface view, especially the nature and orientation of the subsidiary cells in
relation to the guard cells. The principal types of stoma are described in
detail since they are of undoubted importance.
The term stoma is here taken to mean the pair of guard cells together with
the aperture between them. When the cells immediately surrounding a stoma
clearly differ from the remaining epidermal cells they are described as sub-
sidiary cells.
Thefour main types of stoma which occur amongst the Dicotyledons are
generally known respectively as 'ranunculaceous', 'cruciferous', 'caryo-
phyllaceous', and 'rubiaceous'. These names are taken from the families in
which they are well exemplified or in which they were first observed. It is
now well known, however, that stomata of each of the above types occur in
many other families besides those after which they were originally named.
Another difficulty about the terms is that they have not always been used in
exactly the same sense. They were originally applied to the mode of develop-
ment rather than to the appearance of the stomata in a fully grown leaf, but
since it is seldom possible, when identifying a small sample, to follow the
development of stomata in detail, it has become usual to apply the terms to
stomata as they appear when mature. For these reasons it seems very desirable
to replace the present terms by new ones devoid of any taxonomic or onto-
genetic implications. After consultation with Mr. H. K. Airy- Shaw the
following new terms were devised and are put, forward below for the first
 INTRODUCTION xv
time. If they prove to be generally acceptable, it is to be hoped that they will
come into general use. Meanwhile the older and more familiar terms have
been retained book to allow time for the new ones to become assimilated.
in this
Some additional descriptive terms for stomata will probably be found neces-
sary in the future. For instance the so-called 'ranunculaceous' or anomocytic
stoma includes a variety of types for which future investigation will probably
provide a more satisfactory classification. Then again there are stomata sur-
rounded by a circle of radiating cells which might appropriately be described
as actinocytic.

Type A. Stoma surrounded by a limited number of cells that are indistin-

guishable in size, shape, or form from those of the remainder of the
epidermis the 'ranunculaceous' or anomocytic (irregular-celled) type.
Type B. Stoma surrounded by three cells of which one is distinctly smaller
*
than the other two the cruciferous* or anisocytic (unequal-celled) type.

Type C. Stoma accompanied on either side by one or more subsidiary cells

parallel to the long axis of the pore and guard cells the 'rubiaceous' or
paracytic (parallel-celled) type.
Type D. Stoma enclosed by a pair of subsidiary cells whose common wall
is at right angles to the guard cells the 'caryophyllaceous' or diacytic
(cross-celled) type.

Reference must also be made to the so-called 'gramineous' stoma. This

type does not really concern us, however, since it is believed to be confined to
the Monocotyledons where it is especially characteristic of the Gramineae and
Cyperaceae. The 'gramineous' stoma possesses guard cells of which the
middle portions are much narrower than the ends so that the cells appear, in
surface view, to be shaped like dumb-bells. The walls of the narrow portions
of the guard cells are also strongly thickened, so that a transverse section
passing through this region shows a narrow, slit-shaped lumen. There are
frequently subsidiary cells as well lying parallel to the long axis of the pore.
Whilst the above classification of stomata is useful for diagnostic purposes,
it should be realized that the various types tend to merge into one another.

For example, it would be difficult to decide whether a stoma surrounded by

three cells of equal or nearly equal size should be classed as ranunculaceous
or cruciferous. It should also be remembered that stomata of more than one
kind sometimes occur together on the same leaf surface, or stomata on the
upper and lower surfaces respectively may not be alike.

3. EPIDERMAL CELLS AND HYPODERM

Epidermal cells differ considerably in size, shape, and outline in different
plants, but external factors such as light intensity and atmospheric humidity
often have a marked effect on these features. Their diagnostic value is further
restricted by the fact that similar types of epidermal cell are often to be found
in quite unrelated families. Nevertheless the shape of the epidermal cells
provides useful confirmatory evidence, if the identity of a leaf is already
suspected on other grounds. Characters such as a partly or wholly crystalli-
ferous epidermis or the inclusion of cells with specific chemical contents, are
of more importance because of their restricted occurrence. They must not,
xvi INTRODUCTION
however, be taken as indicators of affinity. Epidermal cells with vertical or
horizontal partitions are likewise of specific or at most of generic /aluc,
since these features are of restricted and sporadic occurrence. A horizontally
divided epidermis is not always easy to distinguish from a hypoderm, because
the only difference between the two is that a true hypoderm is derived from
the mesophyll in the course of development, whereas the divided epidermis,
as its nameimplies, arises by partition of the epidermal cells themselves.
When the cells of the outer and inner layers are opposite each other it is
probably correct in most instances to assume that one is dealing with a divided
epidermis, and, when this is not so, that the inner layer of cells is hypoderm.
In a mature leaf the two features cannot be readily distinguished, but this is
of no great practical importance, since, for taxonomic purposes, the occur-
rence of a hypoderm and a divided epidermis are of about equal value.

4. VEINS
In comparing the veins of two leaves, and in particular the structure of the
vascular bundles and their relationship to the surrounding tissues, it is impor-
tant to ensure that veins of the same order are being examined. Larger and
smaller veins in an individual leaf often differ from each other to a consider-
able extent. Nevertheless certain characters of the veins are of confirmatory
value in the identification of species and genera. For instance the vascular
bundles of the veins may be wholly immersed in the mesophyll or raised above
the general level of the upper or lower surface of the lamina. Alternatively
they may be surrounded by a distinct parenchymatous sheath, or be partly
or wholly enclosed by sclerenchyma. Veins in which the vascular bundles
are accompanied on either side by parenchyma devoid of chlorophyll or by
sclerenchymatous or collenchymatous tissue occupying the whole of the space
between the bundle and the upper and lower epidermis are described as
vertically transcurrent. The possession of this character is likewise of specific
or generic value.

5. PETIOLE
The petiole is of considerable taxonomic importance, since its structure
appears to be but little affected by environmental change. In comparing
petioles by means of transverse sections, however, it is essential that sections
of strictly comparable portions of the petioles in question are examined.
The vascular system, in its passage through a petiole, frequently displays a
complex and highly characteristic series of changes, so that sections taken at
different levels in a single petiole may be very unlike one another. To obtain
a complete picture of the vascular system of the petiole a series of sections
must be cut, but since this operation requires more time than is frequently
justified by the results, it has become customary to examine sections through
the distal end immediately below the lamina, and to compare them with sec-
tions taken from the same position in other leaves. Supplementary sections
from the base or proximal end of the petiole are sometimes used. As might
be expected, the vascular structure is less complex in simple than in compound
leaves, but this direct relationship between elaboration of leaf form and com-
plexity of vascular structure is by no means universal. v^
The principal types of vascular system which can be seen in transverse
 INTRODUCTION XVH

sections through the distal ends of dicotyledonous petioles are shown in

Fig. i A-I. Each type may consist of a single vascular strand or be dissected
into a number of bundles according to the species. Types of structure inter-
mediate between those shown in the figures also occur. It should also be
remembered that in certain families, such as the Dipterocarpaceae, the petiolar
vascular structure is more complex (Fig. 53 A-D). There is also a type of

FIG. i A-I, Types of vascular structure visible in transverse sections through the distal ends of
.

dicotyledonous petioles. Each type may appear as a single vascular strand or be dissected into
a number of bundles in different genera and species.

petiole with scattered bundles, e.g. in Anemone vitifolia (Fig. 4 j), and some
special types of vascular structure that occur in some species of Populus are
illustrated (Fig. 314 D and F). It should also be emphasized that the structure
of the petiole is very imperfectly known, but such facts as have been recorded
suggest very strongly that this field would repay further investigation.

6. MICROCHEMISTRY. (See also under 13 (i) on p. xxviii)

In the course of a. broad
anatomical survey one cannot fail to be impressed
by the many types of chemical deposits which occur in plant tissues. Some-
times these occur in cells which do not differ in appearance from their neigh-
bours or, alternatively, they may be in special secretory structures or elements.
The diverse nature of the substances secreted suggests that there must be
correspondingly great physiological differences between the plants in which
they are exemplified. The author feels that, because the nature of secretory
xviii INTRODUCTION
structures and their contents are intimately bound up with the basic physio-
logical processes of the plants in which they occur, they might be expected to
be particularly valuable as indicators of taxonomic affinity. In practice there
is good evidence in favour of this suggestion. For example, taking the genus

Semcio in a broad sense, we find similar secretory structures throughout the

various species quite irrespective of their range of habit forms which includes
annual herbs, stem- and leaf-succulents, shrubs, and even trees. In spite
of this diversity of habit, the secretory system of Senecio at once enables
the stem-succulent species to be distinguished from similar stem-succulent
plants belonging to other families, whilst the task of identifying the herbaceous
and arboreal members of the genus is greatly simplified by the restricted
number of families in which a similar secretory system is known to occur.
The chief types of secretory structures and secretions will now be considered.

(a) Crystals. (See also under 13 (i)

on p. xxviii)
The most common crystals consist of variously shaped deposits of calcium
oxalate. Some types, such as solitary prisms or clusters of various descrip-
tions, are relatively widespread. Nevertheless, even with these relatively
widely occurring types, the kind or kinds secreted by any one species appear
to be fixed. Where more than one kind is secreted in a particular species the
relative proportions of each are frequently found to vary, generally in accord-
ance with the types of nutriment available or with the time of year. Other
types of crystalline secretion such as raphides and crystal-sand are more
restricted in distribution and therefore of still greater taxonomic interest.
Sometimes a whole family is characterized by the secretion of a highly
distinctive material. This applies for instance to the Cruciferae where
myrosin is very common. Likewise berberin is secreted in the Berberidaceae
and to a certain extent in related families such as the Menispermaceae and in
certain Ranunculaceae. A
word of caution is necessary, however, as there are
instances where similar chemical substances are synthesized in quite unrelated
coumarin in certain grasses and a few Leguminosae, or rubber
families, e.g. in

Compositae, Euphorbiaceae, Apocynaceae, and other unrelated families.

(b) Starch
The
size, shape, and other characters of starch grains, such as their appear-
ance in polarized light, are highly distinctive and consequently of considerable
taxonomic value. The botanical origin of commercial starches can be estab-
lished by their microscopical characters.

(c) Cystoliths
. Thesecretion of silica, calcium carbonate, and other chemical substances in
the form of relatively large concretions known as cystoliths is highly charac-
teristic of certain families, e.g. Acanthaceae, Urticaceae, &c.

(d) Laticiferous System

In certain families there is a well-developed system of tubes or cells in
which variously coloured colloidal fluids known as latex are secreted. Latici-
ferous tubes and cells sometimes occur together in the same plant. The term
latex, although convenient, covers a series of fluids of diverse chemical com-
 INTRODUCTION xix

position. Thus the white milky latex of the Para Rubber Tree (Hevea
brasiliensis) isthe chief commercial source of rubber, while the well-known
yellow latex of Chelidonium majus contains very little. Nevertheless laticiferous
elements are of diagnostic value, whatever the precise chemical nature of their
contents may be, because of their restricted occurrence and the ease with
which they may be seen. Since latex is secreted in quite unrelated families
its presence must not in itself be taken as indicative of taxonomic affinity
unless this is also supported by other characters.

(e) Secretory elements

Besides the laticiferous system, other secretory elements in which oils,
resins, mucilage, tanniniferous substances, &c., are deposited are of con-
siderable taxonomic value. These substances are deposited in cells, inter-
cellular cavities, elongated sacs, and canals, the last often being lined with a
definite epithelium. The morphology of the secretory elements, their distribu-
tion within the plant, and the chemical nature of their contents are often highly
distinctive. The nature of the contents has sometimes been assumed rather
than demonstrated by actual tests, so that many terms such as 'oil cells',
4
resin canals', &c,, have frequently been used in a rather loose sense in the
literature of plant anatomy. Another reason for this lies in the fact that the

deposits in secretory elements are seldom pure substances, so that tests applied
by different authors to the same material may lead to various conclusions.
Thus we find in the Lauraceae that the familiar, large secretory cells have
sometimes been described as 'oil cells', though in some species these contain
mucilage and not oil. There appears to be considerable scope for more
intensive work on the chemistry of secreted material as an index of taxonomic
relationship. If circumstances do not permit a sufficiently comprehensive
chemical investigation to be made, the investigator would be well advised
to apply descriptive terms to the contents of these elements, rather than to
assume their chemical nature and so perpetuate inaccuracies. To describe
deposits as being 'solid, red, amorphous* is less misleading than to describe
them as 'gum' when they may be chiefly 'resinous' or 'tanniniferous'.

7. CORK
Solereder attached considerable diagnostic importance to the position in
which the cork originates in a young stem, and this character is undoubtedly
of value within limits. It must be borne in mind that, even in an individual
species, the first cork to be formed is often more superficial than that which
arises later on. Then again, within a single family there are frequently species
with either superficial or deep-seated cork respectively, although in other
families the position of origin of the cork appears to be more constant.
Solereder further stressed the importance of the exact position in which the
cork arises even within the superficial and deep-seated categories. This
character, however, does not appear to the author to merit such detailed
investigation.

8. ENDODERMIS
The presence of a well-marked endodermis in stems is of diagnostic
value because of its restricted occurrence. In most dicotyledonous stems
xx INTRODUCTION
the endodermis is by no means obvious in some it consists of a more or less
;

clearly defined layer of cells which differ from their neighbours in containing
starch; in a third group the endodermis, in the same way as is normal for the
root, consists of cells with well-marked casparian thickenings. In a few species
the endodermis becomes wholly suberized.

9. THRICYCLIC' SCLERENCHYMA
In the stems of
many plants the 'pericycle' is not clearly defined, nor, in
fact, isthere any general agreement concerning the precise meaning of this
term. There is, nevertheless, a portion of the stem between the inner boundary
of the primary cortex and the outer part of the primary phloem in which
mechanical elements usually arise. These mechanical elements are generally
referred to as 'pericyclic*.
Thepresence or absence and nature of the 'pericyclic' sclerenchyma is
easily determined and of taxonomic value. The most common types in trans-
verse sections through an internode are: (i) An interrupted ring of fibres,
(ii)
A continuous ring of fibres, (iii) An interrupted ring of mixed fibres and
stone cells, (iv) A continuous ring of mixed fibres and stone cells, which is
commonly described as 'a composite, continuous ring of sclerenchyma'.
(v) Stone cells present, but no fibres, (vi) Sclerenchymatous elements entirely
lacking. More than one of these arrangements is frequently to be seen in a
single species, especially in stems of different ages. In particular a scler-
enchymatous ring may be continuous or interrupted at different stages of its
development. For this reason it is desirable when comparing the 'pericyclic'
sclerenchyma in two specimens to ensure that the sections are taken through
internodes of the same age.
The amount of sclerenchyma in the 'pericycle' does not necessarily depend
on whether a plant is woody or herbaceous. For example, a well-defined ring
of fibres occurs in herbaceous species of Linum, whereas there are only isolated
strands of fibres in certain shrubs, such as species of Forsythia. On the other
,

hand, 'pericyclic' sclerenchyma is particularly well developed in the Beech

(Fagus sylvaticd), but very little is developed in the herbaceous Lamium album
and it rarely occurs at all in the Umbelliferae.
The nature of 'pericyclic' sclerenchyma is helpful in separating species or
genera in some families, but in a few instances the arrangement may be typical
of a whole family, e.g. 'pericyclic' sclerenchyma is typically absent from all
investigated members of the Pittosporaceae, whilst a sclerenchymatous ring
of a very characteristic type occurs throughout the Geraniaceae.

10. WIDTH OF MEDULLARY RAYS

In the internodes of young stems of many plants the primary vascular
bundles appear to be widely separated from one another by parenchymatous
tissues which may be regarded as constituting very broad medullary rays,

e.g. in the herbaceous Ranunculaceae. In extreme cases the

vascular bundles
are scattered, or show a tendency to be scattered, in the same way as those of
Monocotyledons, e.g. in Piperaceae, herbaceous Berberidaceae, &c. In trans-
verse sections through the internodes of other plants, however, the xylem
appears as a closed ring, traversed by very narrow medullary rays. In these
plants the primary bundles are not individually distinct, e.g. in the Theaceae.
 INTRODUCTION xxi

Other plants exhibit an intermediate structure. These different kinds of

arrangement have not received, from the taxonomist, all the attention they
appear to merit, presumably in the mistaken belief that they are chiefly
expressions of habit differences, a continuous ring of xylem having been
regarded as typical of woody plants, and separate bundles as typical of herbs.
If this were entirely true little or no taxonomic significance could be attached
to these differences. As will be seen from the descriptions of the families in
this book, however, there are woody plants like Clematis and Menispermum in
which the primary bundles are separated by broad medullary rays; on the
other hand, the herbaceous Linaceae and the tribe Galieae amongst the
Rubiaceae have a continuous ring of xylem, whilst in Veronica even the most
slender herbs are devoid of medullary rays. Solereder states that the diversity
of arrangement of the vascular system of herbs is so varied that it is of little
practical taxonomic value, and he pays but little attention to the subject.
Experience suggests, however, that the primary vascular structure is probably
of greater taxonomic significance than many of the features that Solereder
emphasized. Much new information on this subject is incorporated in this
book.

11. BICOLLATERAL BUNDLES

The habitual occurrence of bicollateral vascular bundles in the axis of a
family, e.g. the Cucurbitaceae,is often a useful diagnostic character.

12. CORTICAL AND MEDULLARY BUNDLES, AND ANOMALOUS SECONDARY

THICKENING
All of these features are of diagnostic value on account of their restricted
occurrence, but the families in which they are to be found are seldom closely
related to one another.

13.WOOD. (See also pp. xl-li)

The structure of the xylem in the young twigs of shrubs and trees often
from that of the mature trunk. Solereder's remarks
differs in certain respects

concerning the wood were often based on features seen in twigs taken from
herbarium sheets and, being, therefore, not typical of the mature wood, are of
doubtful value for the identification of commercial timbers. In a young twig
the dimensions of the cells, both diameter and length, are markedly smaller
than in the mature wood, there often being an increase of 100 per cent, over
those of the first ring in addition the distribution of the vessels, the occurrence
;

and arrangement of the parenchyma, and even the nature of the fibres that
form the ground-tissue of the xylem may be unlike those of the mature wood.
It is not unusual to find vessels with scalariform perforation plates in the

young wood of a species which has exclusively simple perforations in the

mature wood, or heterogeneous rays in the first rings changing later to homo-
geneous rays.
On the whole the wood structure appears to possess features that are more
conservative than those of the external morphology, and in consequence
specific differences are often less clearly expressed in the wood than in the
external features. microscopical characters of the wood are also less
Many
plastic than both the exomorphic features and many anatomical characters of
 xxii INTRODUCTION
other parts of the plant body such as the leaf; this, combined with its more
conservative nature, adds to its value for the study of larger groups. There
are, however, some characters, such as the number and size of the vessels and
the amount of parenchyma present, that may be markedly influenced by the
environment, and these features are relatively unimportant for taxonomic
purposes.
The characters of the wood that have been found by experience to have the
greatest taxonomic value are discussed below (see also p. xl).
100 n 200 u
SMALL MEDIUM LARGE

30

20

40 SO 120 I6O 20O 24-O 28O 32O 56C 4OO

VESSEL DIAMETER IN MICRONS
DIAGRAM I. Distribution of Mean Vessel Diameter in the Dicotyledons.

(a) Vessels

The most useful characters are distribution, pattern, diameter, and fre-
quency as seen in transverse sections, type of perforation, pitting between one
vessel and another (intervascular) and between vessels and wood- or ray-
parenchyma cells, and the occurrence of spiral thickening. The presence of

tyloses may be of value, but it is sometimes doubtful if their formation is a

constant feature and they are normally lacking from sapwood. Where tyloses
are pitted, contain crystals, or have a characteristic shape, as in the Sapotaceae,
they have a greater diagnostic value. Vessel diameter is best recorded as a
mean figure; though varying according to position in the tree and conditions
of growth, often of considerable value for identification, as it may differ
it is

markedly in closely related woods. It is not practicable to use mean vessel

diameter for a ring-porous wood. The range of mean diameters in the
Dicotyledons is shown in the above histogram, which is based on 1,500
species (see also Appendix I, Table i, on p. 1360). Vessels are described as
'small' if their mean diameter is less than IOO/LI and 'large' if it is more than
200 jit.
The number of vessels per sq. mm., though useful, is difficult to determine.

In many cases it is extremely difficult to distinguish the smallest vessels of a

 INTRODUCTION xxiii

group as seen in transverse section from tracheids or fibre-traeheids, and a

convention has been commonly adopted to treat all the vessels of a group as a
single unit when making counts. The figures obtained in this way (see
Appendix I, do not conform to any normal distribution curve. The
Table 3)
range is very great, from fewer than i per sq. mm. to over 300. Very low or
very high figures tend to be characteristic of some families. For example,
most of the woods of the Caesalpiniaceae, Papilionaceae, and Rosaceae-
Chrysobalanoideae have fewer than 5 vessels per sq. mm., whereas the woods
of the other Rosaceae typically have more than 40 per sq. mm.
Vessel member length is seldom used for identifying timbers. It can only
be measured satisfactorily in macerated material and should consist of the
total length (359). It must be emphasized that vessel member length, in
common with other dimensions, varies considerably within any species
cell
and even in different parts of the same tree. Bailey and Tupper (100), for
example, have shown that the first-formed members are shorter than those
developed later, that slow-growing plants often have shorter members than
fast-growing and that, within the same species, dwarf plants tend to have
shorter members than normal individuals. Nevertheless, vessel member
length is of very great importance for phylogenetic studies, and the distribution
of mean length throughout the Dicotyledons is shown in the following histo-
gram, which is based on 600 species (see also Appendix I, Table 2, on p. 1360).
Vessel members whose mean length is less than 350/4 are described as short,
and those over 8oo/z as long.

(b)Wood Parenchyma
The distinction between the maintypes depends on the distribution as seen
There has been, and still is, considerable difference of
in transverse sections.
opinion concerning both the types to be distinguished and the terms that
should be applied to them. It is therefore desirable to describe here the
meaning of the terms used later in the descriptions of the families.
The scheme adopted is a combination of the terms given by the International
Association of Wood Anatomists in its Glossary (1119) and the subsequent
modifications suggested by Kribs (1287), Bailey and Howard (83), and
Chalk (354). For the benefit of readers who are not specialists in wood
anatomy, the parenchyma has usually been described in the text as well as
labelled, as, for example, 'Parenchyma scattered as isolated cells among the
fibres (diffuse)'.
A basic distinction is drawn between which the
'apotracheal' types, in
distribution is
fundamentally independent of the vessels, and 'paratracheaF
types, whose distribution is determined primarily by the vessels. Apotracheal
types may often touch some or all of the vessels, particularly where either the
vessels or the parenchyma are abundant, but such contacts are, so to speak,
accidental. Three main types of apotracheal parenchyma are distinguished
terminal, diffuse, and metatracheal. Terminal parenchyma occurs between
the end of one growth ring and the beginning of another, either as scattered
cells or, more commonly, as a continuous band (Plate A, i). The name
'terminal' has been shown by Chowdhury (412, 413) to be in some instances a
misnomer, as the parenchyma band may be the first-formed tissue of the new
ring and not the last-formed tissue of the old. For this type he has proposed
xxiv INTRODUCTION
the term 'initial', but, as not practicable to make this distinction on the
it is

ordinary wood sample, the term has not been used in this work, except where
it is recorded in the literature.
Of the other apotracheal forms there is complete series from 'diffuse',
a
with isolated cells scattered among the fibres (Plate A, 2), to regular, continuous
'mctatracheal' bands (Plate A, 5 7). Bailey suggests that the term 'meta-
tracheal' should be dropped, owing to the different senses in which it has
been used, in favour of 'banded'. The intermediate forms present some
difficulty, particularly where the cells tend to be grouped into short lines from

350* 800*
40
SHORT '
MEDIUM LONG

2 -4 -6 -8 1-0 1-2 1-4- 1-6 1-8 2-O 2-2 2-4 26

MEAN VESSEL MEMBER LENGTH IN MILLIMETRES
DIAGRAM II. Distribution of Vessel Member Length.

ray to ray (Plate A, 3 and 4). These are sometimes classed with the isolated
cells under the term diffuse, but there is much to recommend the use of a
separate term, such as 'diffuse-aggregate' or 'diffuse-in-aggregates', as sug-
gested by Kribs and Bailey respectively. In the text this type has usually been
described in some such terms as 'scattered among the fibres as isolated cells
and short uniseriate lines'.
The term 'reticulate* is applied to banded types in which the parenchyma
bands are similar to the rays in width and spacing, so that the two together
give the appearance of a net, as in Plate A, 3 and 6.
Various forms of paratracheal parenchyma are illustrated in Plate B. The
three main types are 'vasicentric' (B, 3), a sheath concentric with the vessel,
'aliform' (B, 4), a sheath projecting from the sides, and 'confluent' (B, 6), the

parenchyma round some of the vessels linking up to form bands. There has
been some difference of opinion concerning whether a few cells round the
vessels (not forming a complete sheath) should be included under the term
vasicentric. Kribs suggested the phrase 'scanty vasicentric', which Bailey
amended to 'scanty paratracheal', and the latter has been used in the text (see
 INTRODUCTION xxv

Plate B, i). Another useful term suggested by Bailey and adopted here is

'abaxial', to describe forms in which the parenchyma is limited to the side of

the vessel farthest away from the pith and often extends from the vessel as
lateralwings (Plate B, 2). Kribs has also used the term 'banded paratracheal'.
An
entirely different parenchyma feature, which has been described in the
text, and which may need a brief explanation, is the number of cells in a
parenchyma strand or longitudinal file of cells formed from a single cambial
initial. Most commonly of four cells, the number may be up to as many as
sixteen in woods with long cambial initials, and in many short strands there
may often be a single cross-wall or none. The latter strands, each consisting of a
single cell, are now known as fusiform parenchyma cells in the older literature
;

they are referred to as substitute or intermediate wood fibres.

(c) Kays
The most widely used character of the rays is width, either in actual dimen-
sions or in number of cells height is also of value, but usually only in extremes,
;

when very high or very low. 'Rays exclusively uniseriate' is another feature
of great value for identification, particularly as it represents a type of structure
that occurs rather sporadically, and may thus offer a distinction between
closely related genera or species, e.g. between particular species of Guarea
and Terminalia. Other useful characters are the various types of heterogeneous
and homogeneous rays (Plate C), and oil or mucilage cells. The
tile 1 cells

frequency of rays is often a useful character. It is usually measured by

counting the rays along a line at right angles to the rays on either a transverse
or tangential section, and is expressed as the number per millimetre. The
number is sometimes characteristically high or low in particular families,
numerous rays being, for example, typical of the Myrtaceae and Rubiaceae.
The distribution of ray number in the Dicotyledons is shown in the following

histogram, which is based on 1,800 species (see also Appendix I, Table 6).
Ray width, though theoretically best expressed as a mean value, is so much
more easily measured as a maximum that it is usually given in this form. The
distribution of maximum ray width in the Dicotyledons is shown in the
histogram, which is based on 650 species, in Diagram IV (see also Appendix I,
Table 5). Rays whose maximum width is less than 50 are described as fine
JJL

and those over 100 /z as broad.

(d) Fibres
It isusual to distinguish between 'libriform fibres' with simple pits, and
'fibre-tracheids'with bordered pits, but no exact definition of these two types
appears to be universally acceptable. .Intermediates are so numerous that it
was found more satisfactory in the text to group all types under the single
heading 'Fibres'. The most important characters of fibres are the nature of
the pitting and the presence or absence of septa. Of particular interest are
instances where septate fibres are grouped together in a manner suggestive of
parenchyma, as in many of the species of the Celastraceae. Fibre length,
1
Tile cells have been defined (1119) as a 'special type of apparently empty upright or
square cells of approximately the same height as the procumbent cells and occurring in
indeterminate horizontal series usually interspersed among the procumbent cells. (Common
in certain of the Tiliales and Malvales)', and are described and illustrated by Chattaway (371).
 20-

-
18

>- 12
U
Z
3"
a

4 6 8 10 12 14 16 IS 20 22 24 26 28 30
NUMBER OF RAYS PER MILLIMETRE
DIAGRAM III. Distribution of Ray Number.

50 u 100*
40
FINE MEDIUM BROAD
36
-

32
~

> 2S -

3 20
O
ui

20 40 6O 80 IOO 120 I4O I6O I8O 20O 22O 24O 260 280 30O
MAXIMUM RAY WIDTH IN MICRONS
DIAGRAM IV. Distribution of Maximum Ray Width.
 INTRODUCTION xxvii

though used for diagnostic purposes, is of some practical interest and

little

has been included in the text. The distribution of mean fibre length in the
Dicotyledons is given in the following histogram, which is based on 534
species (see also Appendix I, Table 4). Fibres whose mean length is less than
900 fi are described as short, and those over 1,600 p as long.

(e) Storied Structure

This
is a most useful character. It consists of the
arrangement of the cells
or tissues in horizontal series as seen in tangential section
(see Fig. 1170).

900 u 1600M
r
SHORT MEDIUM LONG

20

3 -6 8 12 1-5 1-8 2-1 2-4 2-7 3-0 3-3 36

MEAN FIBRE LENGTH IN MILLIMETRES
DIAGRAM V. Distribution of Mean Fibre Length.

In some cases only the rays are storied, or grouped in horizontal series like
thewindows of a house; in others all the elements are storied; in others, again,
only the wood parenchyma. Storied structure has been described in the text
under each element or tissue in which it occurs. A list of the families in which
the feature occurs is given on p. 1352.

(/) Growth rings

Although this character is a standard feature of most descriptions of species,

it was omitted from the present work because it was found that so little
useful information could be added to a family description under this
heading.
The heading has, however, been retained to record any investigations on
cambial activity and the development of the tissues. Any special points con-
nected with the growth ring, such as terminal parenchyma, markedly thicker-
walled fibres at the end of the ring, &c., have been recorded under the type of
cell concerned.

(g) Included (Interxylary) Phloem

This form of anomalous structure has been recorded, where appropriate,
under the description of the wood. A list of the families in which it occurs is
given on p. 1352. The terminology adopted is that given by Pfeiffer (1712) and
used also by Chalk and Chattaway (362).
xxviii INTRODUCTION
(//) Intercellular Canals (see also under 6 (e) on p. xix)
Both the vertical and radial types are of great diagnostic value. Lists of the
families in which they occur are given on pp. 13534. Latex tubes, which
sometimes occur in the rays, are also of value, but are rare.

(?) Deposits and Chemical Reactions (see also under 6 (e) on p. xix)
The presence of deposits of unusual substances may be of great value where
facilities are available for identifying them, and such facts as are recorded in
the literature and appear likely to be of taxonomic use have been included in
the text, Otherwise, observations have been limited to the visible aspects of
deposits, such as colour and whether crystalline or gum-like, and the terms
'gum-like' and 'gummy' have been used without any chemical significance,
'['he degree of ligniiication of the cell wall, as indicated by stains, has some-
times been referred to, particularly the occurrence of a mucilaginous layer in
the wall of the fibre or areas of tissue with unlignified cell walls.

GENERAL CONCLUSIONS
If one surveys the Dicotyledons as a whole, the anatomical evidence con-
firms that some families are relatively homogeneous and therefore probably
natural, while others are seen to be heterogeneous and therefore in need of
revision or at least reinvest igation. So far as the constitution of families is
concerned, anatomy points to the desirability of local amendments rather than
complete reorganization. For example, a genus may be shown to need trans-
ferring to another family or possibly to form the basis of a new one, or the
dividing line betvveen closely related families may need modification. Examples
will also be found in this book of groups of families that have sufficient
anatomical similarity to justify the conclusion that they constitute larger
natural groups or orders, e.g. the Malvales. Other orders seem to be more
artificial, with little in common between their constituent members, in which
case it is better to recognize that the families are isolated than to try to force
them into a group. Every family must obviously be included somewhere in
a flora, but, where affinities are doubtful, this should be clearly indicated.
Modern angiosperms represent only the existing end-points in an evolutionary
process of which the early course is almost unknown, and previously existing
groups may have become extinct without leaving any known record for our
guidance. When, in addition, one must acknowledge that the very origin of
the angiosperms is a matter of conjecture, the difficulties in the way of making
a truly natural classification will be seen to be very great. It is therefore
important that any attempt to do so should neglect no evidence and should be
based on as many attributes as possible tw o of the most important of these
;
r

are external morphology and microscopic anatomy.

woody stems obviously reduces the general taxonomic value
Limitation to
of wood anatomy, but that it can be of great value in particular groups is now
generally recognized. It must be emphasized again, however, that the evidence
from wood anatomy needs careful interpretation, particularly in order to
distinguish between characters, such as type of perforation and intervascular
pitting, that are associated primarily with level of specialization and those that
are likely to indicate affinity.
 INTRODUCTION xxix

This section may be appropriately ended by a quotation from T. A, Sprague

(2172):
'It should be clear that future progress in the classification of the groups of higher
rank will be by synthetic methods not by attempting to make a complete scheme
embracing every family, but by placing such families together as show maximum
correlation of characters, and by building up larger groups from them. A natural
classification built up in this way will for many years be very incomplete, but it will
have the supreme merit of being based on established facts instead of individual
opinions/

Selected Literature. (See also p. Ixii)

Anderson 26, Bailey 71, Bandulska 1259-32, Bartlctf 145, Camp 332, CariipbHI 335,
Fritsch 723, Greenman 813, Gundcrson 846, HeinKch and Wctmore 939, Hill 970,
Hutchinson 1113, Metcalfe 1500-2, Netolitzky 1588, Odell 1628, Rushy 1968, Scwurd
2o8iA, Sherif 2088, Sprague 2172, 2I72A, Slopes 2208, Slopes and Fujii 2209, Tippo
2263, Turrill 2303, 2304, Watson 2368, Weiss 2392, Wettstein 2416.

-*
ANATOMY AND PHYLOGENY
IT a familiar fact that very little is known about the ancestry of the (lowering
is

plants, owing to the imperfect geological record. For this reason most of the
views which have been expressed concerning angiosperrn phytogeny rest
mainly on facts of comparative morphology tempered by a generous seasoning
of speculation. Taxonomists and morphologists, stimulated by the classical
writings on evolution during the last century, have sought for characters
which may respectively be regarded as primitive or advanced from the
evolutionary standpoint. Series of characters which are believed to possess
these attributes in varying degrees have been found, and, by making use of
them, the families of flowering plants have been arranged by systematists in
branched, linear series that are commonly treated as if they represent phylo-
genetic trends along which evolution may be assumed to have progressed.
The result has been the various systems of classification which are well known
to taxonomists. These systems are all much alike so far as the composition of
families and taxonomic units of lower rank are concerned. It is true that there
are differences concerning the lines separating closely related families, that
there is lack of agreement concerning the advisability of raising certain genera
or groups of genera to the status of families, and that opinions differ concern-
ing the true affinities of certain genera. Further investigation is, however,
gradually leading to a more general agreement concerning those matters which
are still unsettled. Nevertheless the fact that there is such a large measure of
agreement common to all reliable taxonomic systems is in itself very remark-
able, and tends to show that modern taxonomy rests on a firm basis of well-
established facts.
When we turn to taxonomic units above the rank of families, it is at once
apparent that there is no general agreement concerning their composition.
The differences of opinion are still greater when we pass from the purely
taxonomic realm into that of phytogeny. One of the chief reasons for this lies
in the fact that there has been much speculation concerning the phylogeny of
the existing plant families. The reliability of a phylogenetic system is largely
 xxx INTRODUCTION
a measure of the knowledge and wisdom of its author, and all views which
have been expressed concerning the phylogenetic interrelationships of plant
families are largely a matter of personal opinion. To the author, as indeed to
many other botanists, it appears highly improbable that the families of flower-
ing plants as we know them to-day have been evolved from one another.
Those that agree with each other taxonomically in many respects may well
have had a common ancestry, but it seems fundamentally misleading to
arrange them in a single, linear, phylogenetic series even if this has the form
of a branched tree. It is surely more profitable to recognize that the families
and taxonomic groups of lower rank represent the present levels which have
been reached by plants that have evolved along lines of descent between which
there has been no direct connexions except possibly in a remote geological
age. Even then there can have been a direct connexion between them only if
all of the modern flowering plants have been derived from a common
pre-
angiospermic stock. That this was so has yet to be proved by irrefutable
evidence.
i
It is well known to students of palaeobotany that one of the greatest

evolutionary problems is how the flowering plants originated, and from what
pre-angiospermic stock they made their sudden appearance in the Cretaceous
era. The palaeobotanical record from the Cretaceous deposits of Greenland
affords evidence of the contemporary existence of forests of plants resembling
modern plane trees, as well as of species which can be identified as having
affinities with such wholly unrelated modern genera as Artocarpus, Cinnamo-

mum, Dalbergia, Magnolia, Menispermum, and Quercus. In the Cretaceous of

Colorado and New Mexico the remains of palm-like plants, and of species,
which modern genera Ftcus, Juglans, and Quercus, have been found.
recall the
In the lateCretaceous, in addition to the plants already mentioned, genera
allied to Salix, Myrica, Bauhinia, Sterculia, and Eucalyptus are known to have
occurred. In view of the coexistence of such diverse types amongst the earliest
known fossil angiosperms one cannot, in the absence of a more exact geo-
is more
logical record, decide which of them primitive than others. There is
no fundamental reason why they should not have been evolved from different
pre-angiospermic stocks. In fact this seems no more improbable than that
the origin of the angiosperms was a solitary evolutionary event.
Because of the paucity of geological evidence, taxonomists make use of
comparative morphology when attempting to elucidate phylogenetic sequences.
In this connexion series of morphogenetic progressions have been proposed
which demonstrate possible lines along which relatively advanced plants may
have been evolved from simpler forms. Floral examples are afforded by the
changes from polypetaly to gamopetaly, from radial to bilateral symmetry,
from the superior to the inferior gynoecium, or from the atropous to anatropous
position of the ovule. Others are also recognized, and further details may be
found in Sprague's (ZIJZA) paper. Morphogenetic progressions such as these
have not yet been proved to represent actual evolutionary sequences, but it is
more reasonable to assume that changes have taken place in the directions
indicated rather than in the reverse. If it be accepted that this line of reasoning
is legitimate, it can at once be seen that there are families of
Dicotyledons, such
as the Papilionaceae and Compositae, which may be regarded as relatively
advanced and others, such as the Ranunculaceae, which are relatively primitive.
 INTRODUCTION xxxi

In the same way it is possible within a family to recognize relatively primitive

and advanced genera and species. IVe thus obtain a glimpse of possible modes
^f^evolution within the angiosperms, hut the mere fact that such sequences
probably occurred must not be taken to mean that any existing families have
necessarily been evolved from one another directly.
During the past fifty years, the problem of angiosperm phylogcny has also
been approached by anatomists working along several distinct lines, The
early investigations were concerned with stelar structure, followed by a period
devoted to studies of the relationship between the arboreal and herbaceous
habits, whilst more recent phylogenetic work has dealt with nodal anatomy
and wood structure. Each of these lines of inquiry will now be considered ia
more detail. ^

Stelar structure
At the beginning of the present century, speculation concerning the origin
of the angiosperms led to frequent comparison being made between their
vascular anatomy and that of plants from which they might have been derived.
It was commonly held that the primitive vascular system of the axis was a proto-

stele, in which a solid central cylinder of xylern was surrounded by phloem

and an endodermis (Fig. 2 A). This type of structure persists in modern
plants amongst some of the ferns and lycopods, whilst other Pteridophytes,
which possess a more complex vascular system when mature, nevertheless
exhibit a protostelic structure when the sporophyte is still young.
The protostele is still generally held to be the ancestral to other types of
vascular structure, but there has been speculation concerning the manner in
which the transitional stages were effecte^l. From the point of view of the
prehistory of the angiosperms the most essential conception is that the central
portion of the protostele became medullated or converted to parenchyma..
According to one view this change may have occurred by the intrusion of the
parenchyma of the cortex through gaps in the protostele associated with the
departure of vascular strands to supply the leaf. By others it was held that
the conversion of the central tissues of the protostele to parenchyma was a
direct process in which intrusion of parenchyma through leaf gaps was of no
significance. Whichever of these alternatives represents the actual course of
events, the important point is that a tubular vascular system was formed as a
result, and this was termed a medullated protostele or siphonostele (Fig. 2 B)
and represents the basic structure of many angiosperms at the present day.
A variant of the continuous siphonostele was one in which the xylem and
phloem became dissected so as to give a ring of separate vascular strands each
consisting of xylem and phloem (Fig. 2 c). This polyfascicular type of stele
occurs in many angiosperms at the present day, and, as indicated below, there
has been considerable controversy concerning the mode of transition from
the continuous to the dissected type. Some botanists have held that this
change was effected by the devascularization of tissues opposite leaf-traces,
whilst others believe that it took place by the reduction or suppression of

cambial activity accompanied by enlargement of primary rays, without the

m

leaf- traces having taken any important part in the process. Another variant of
the siphonostele is to be seen in Fig. 2 F. This differs from the structure
shown in Fig. 2 B in having a layer of phloem on the inside as well as on the
XXX11 INTRODUCTION
outside of the xylem. This type of structure is to be seen amongst the
Ptcridophyta such plants as A-h'trsileg. and Loxsoma, whilst amongst the
in

flngiosperms it is represented by families possessing internal phloem, e.g.

c or Asclcpiadaccac. Other variations of the medullated protostele
and siphonostele also occur depending on the extent to which the phloem and
cndodennis extend into the interior of the xylem cylinder through the leaf
gaps and also on the extent to which the external phloem has the form of a

FIG. 2. A, Protostele.B, Medullated protostele or siphonostele. Type I. C, Medullated protested.

Type II. Astelic, e.g. as in Nymphaea. E, Polystelic, e.g. as in Pteridium aquilinum (L.) Kuhn and
D,
certain species of Primula. F, Medullated monostele \vith internal phloem and endodermis, e.g. as in
Marsilea or Loxsoma. The thick black lines adjoining the phloem represent endodermis. For further
details see p. xxxi.

continuous or dissected cylinder. These are not, however, important in the

present discussion, so it will suffice merely to note their existence. If, when
the siphonostele is dissected, each vascular strand becomes completely
enclosed by an endodermis, the resulting structure is a polystelejTig. 2 D, E)
which occurs amongst advanced ferns such as the common Bracken (Pteridium
aquilinum) and in certain of the Auricula section of the genus PrimulcL These
various types of stelar structure and their interrelationships have been more
fully described by T. G. Hill (974).
^
Herbs and Trees

During the period from 1911 to 1922 there were many investigations and
much speculation concerning Ihe phylogenetic relationship of the arboreal
and herbaceous habits in the angiosperms. The botanists mainly concerned
were Bailey, Eames, Jeffrey, Sinnott, Thompson, and Torrey, whilst the
leading American and British botanical journals of that period abound with
articles on the subject. After a lapse of several years Arber (30) took up the
 INTRODUCTION xxxiii

subject again, but even at the present time it seems improbable that the last
word has been said.
At the beginning of this period it was generally believed that, in the inter-
nodes of typical herbaceous stems, there is a ring of collateral vascular bundles
separated from one another by relatively broad areas of parenchymatous
tissue. This arrangement was also thought to exist in the young stems of
shrubs and trees, whilst, in the more woody plants, the cambium, instead of
remaining confined to the vascular bundles as in the herbs, became converted
to a continuous ring by the development of a zone of meristematic tissue across
the parenchyma between the bundles. There was thus a distinction between
the cambium of the primary bundles which was termed fascicular and that
which developed between the bundles and was described as interfascicular.
By the activity of the interfascicular cambium the original bundles became
united to form a continuous ring interruptedjonly by medullary faysjwhose
width varied considerably ifl different kinds of plants. This mode of develop-
ment, which had become familiar through the teachings of Sachs, de Bary, and
other botanists in the previous century, doubtless occurs in many plants. It
is now known, however, that in other and very numerous species a continuous
zone of cambium arises at a very short distance behind the stem apex, so that
the distinction between fascicular and interfascicular cambium breaks down.
All of the botanists who were interested in the phylogenetic relationship of
herbs to trees during the IQII to IQ22 period probably believed the arboreal
habit to have beer^ primitive and the herbaceous habit derived ^r advanced.
This commonly accepted view depends mainly on the well-known fact that
the immediate precursors of the angiosperms, together with the early angio-
sperms ^themselves revealed in the geological record, were known to have
been trees or at least woody plants. It was often forgotten, however, that the
geological record is certainly very imperfect, and, while we have no definite
knowledge of them, it is at least possible that herbs may have coexisted with
the arboreal types. The fact that they have not yet been discovered may be
because they were less readily preserved than trees or they may have been
comparatively rare. The question of climate was also taken into consideration,
especially by Bailey and Sinnott, who pointed out that, in the present era,
trees constitute the dominant vegetation of the tropics whilst Jierbs become
progressively more numerous in cooler climes. Since it is generally believed
that a warmer climate occurred over a much greater part of the world at the
time when the angiosperms first made their appearance than now, it seems
reasonable to suppose that arboreal plants may then have been more widely
distributed than at present. Bailey and Sinnott regarded the development of
a colder climate in the temperate zones, particularly in the Tertiary era, as
having played a fundamental part in producing herbs by the reduction of more
woody forms. An additional argument, which has frequently been put forward
in favour of the arboreal ancestry of the angiosperms, is the familiar one that
in other groups of plants below the angiosperms, arboreal forms, now extinct,
have been succeeded by herbs which persist until the present day. Here again
herbaceous forms may have previously existed beside their arboreal relatives,
without their remains having yet been discovered. *
In order to give an anatomical explanation of the phylogenetic transforma-
tion of trees to herbs a hypothesis was put forward by Eames (620), and later
xxxiv INTRODUCTION
elaborated by Jeffrey and his co-workers. This assumed the primitive cauline
vascular system of the angiospcrms to have had the form of a cylinder of
jtyleinsurrounding a central pith, that it was in fact a medullated protostele.
This cylinder became interrupted by parenchymatous tissue above the point
of attachment of the leaf-traces to the stem, the necessity for this parenchyma
being attributed to the need for storing food near the leaves in which it is
synthesized. By the upward and downward extension of the parenchymatous
tissue above and below the leaf insertions, and by the simultaneous thinning
of the woody cylinder the latter became dissected. If, as the holders of this
view believed, the leaf paps Qverlapped it is clear that the primitive vascular
|

cylinder would become converted into a series of distinct fibro-vascular

bundles as seen in transverse sections, and in this way the so-called typical
herbaceous structure was stated to have arisen. In support of this view the
results of anatomical comparisons between herbaceous and woody species
were presented in which the facts appeared to be consistent with the theory.
It is important to note, however, that most of these comparative studies were
limited to a few families, particularly the Compositae.
Eames's and Jeffrey's view that the primitive cauline vascular system had
the form of a closed cylinder was strongly disputed by Petersen (1706).
Petersen pointed out that there is no evidence to show that plants with a vas-
cular system which appears in transverse sections as a ring of individually
distinct vascular bundles is less primitive than the continuous xylem cylinder.
She emphasized that plants with widely spaced bundles may have existed
beside those with a closed xylem cylinder from the earliest times, so that there
is no necessity for regarding the one type of structure as derived from the
other. She also observed that even plants like herbaceous species of Linum,
where a continuous xylem ring is formed whilst the stem is still very young,
nevertheless pass through a very brief phase in which independent vascular
bundles can be recognized.
Jeffrey and his associates also used another line of argument in support of
their views. This is based on what they believed to have been the sequence of
morphological changes through which the medullary rays passed during their
phylogeny. Jeffrey and his followers regarded the narrow, uniseriate ray as
primitive, whilst the so-called 'aggregate* rays in Carpinus and other amenti-
ferous trees were believed to represent congeries of narrow or uniseriate
primitive rays which became closely associated in the region of leaf-traces.
Since many botanists then held, and it is still commonly believed, that the
Amentiferae are primitive plants, it was argued that the aggregate ray was a
primitive character which still persists. By the subsequent fusion of the com-
ponents of the aggregate ray and their complete conversion to parenchyma it
was believed that the broad Compound' ray such as occurs in the oaks was
formed. As an argument in favour of this view it was pointed out that in
certain seedling oaks the broad compound ray does not exist, whilst by
traumatic disturbance of mature oaks reversion to the aggregate ray could be
induced. The validity of this argument rests on the supposition that charac-
ters in a seedling which do not persist in the adult are relics of the type of
structure which occurred in mature members in the phylogenetic precursors
of the modern species. Furthermore that traumatically induced anatomical
changes necessarily represent reversions to an ancestral type. Whilst there
 INTRODUCTION xxxv

may be some support for both of these ideas, it is improbable that either
would command the universal support of present-day botanists. But
thesis
the difficulties do not end at this point, for it is still necessary to explain the
occurrence of narrow rays in the amentiferous Salicaceae as well as in a great
range of other angiosperms including many which are generally regarded as
advanced types. Jeffrey and his followers appear to have appreciated this
difficulty, but they attempted to overcome it by making the rather improbable
suggestion that the aggregate ray, besides developing phylogenetically into
the broad compound ray in the manner just described, also became split up
into progressively narrower components.
Thompson (2253) was one of the first to record evidence which he believed
to demonstrate that ancestral rays of the compound type have become
narrower by disintegration in certain of the Ericaceae, Casuarinaceae,
Betulaceae, and Fagaceae, and this theme was developed by Bailey and
Sinnott (97). The disintegration of rays in advanced members of the Betu-
laceae was also described by Hoar (981). More recent research (138, 1286)
suggests that the aggregate ray is a highly specialized structure which occurs
sporadically throughout the Dicotyledons, and it is now generally believed
that the most primitive ray type is a combination of multiseriate and uni-
seriate rays, and that woods with wholly uniseriate rays have been derived

phylogenetically by elimination of the multiseriates (p. xliii). Ontogenetic

dissection of large rays commonly leads to an increase, not a decrease, in their
number owing to the subsequent growth of the smaller units.
Bailey (67) appears at first to have been not unfavourably disposed to the
concept of the primitive continuous stele having been dissected by paren-
chyma associated with outgoing leaf-traces. A year later (68), when referring
to the modern Fagales, he speaks of the primitive aggregate ray having been
reduced and disintegrated in some species and compounded to form the broad
multiseriate type in others. Thus Castanea and Castanopsis with uniseriate
rays are regarded as reduced members of the oak family since Quercus is
characterized by broad rays. In the same way species of Alnus which possess
only uniseriate and no aggregate rays were regarded as derived from species
in which the aggregate rays previously existed. The broad rays of Quercus
were thought to have been formed by the compounding of previously existing
aggregate rays.
By 1914, however, Bailey had been collaborating with Sinnott (97), and in
the joint publications of these authors in that year a somewhat different point
of view is presented. Bailey and Sinnott agree with Jeffrey and his followers
in regarding the herbs as being derived from trees, but differ in their views

concerning the method by which this has been accomplished. They rightly
regard the fundamental difference between herbaceous and woody plants as
consisting^ of a reduction in cambia^activrty, accompanied, in sojne but by no
means all instances; by an expansion of interfascicular parenchyma or primary
medullary rays. These changes they believe to have been in no way dependent
on the formation of leaf gaps. They very rightly draw attention to the fact
that herbs are by no means always characterized by multifasciculate stems as
implied by Jeffrey and demanded by his hypothesis concerning the origin of
the herbaceous habit. In many herbs there is a cylindrical xylem interrupted
only by minute gaps associated with the departure of vascular traces to the
 XXXVI INTRODUCTION
appendages, whilst in others the structure is multifascicular as has already
been mentioned. Comparison of woody and herbaceous plants, in which each
of these two types of structure occurs, shows that the multifasciculate arrange-
ment persists in old steins of plants in whose young extremities it is at first

FIG. 3. A, Transverse section of one-year-old stem of arborescent or fruticose Dicotyledon, which is

devoid of wide, high multiseriate rays. B, Transverse section of one-year-old stem of arborescent or
fruticose Dicotyledon whose woody cylinder is dissected into separate strands by wide, high multi-
seriate rays. C, Transverse section of stem of woody or 'transitional* herb showing 'confronting' and
'flanking' parenchyma. This condition is emphasized by Jeffrey and Torrey in their theory of the
origin of the herbaceous type. D, Transverse section of slender, herbaceous stem which is devoid of
'foliar storage rays'. E, Transverse section of slender, herbaceous stem whose stele is dissected into
a series of discrete woody strands. (After Sinnott and Bailey 2113.) For further details see p. xxxvii.

established, whilst it is equally true that the closed cylinder of xylem persists
in the older parts of the axis of those plants in which it is laid down when

young. It has already been pointed out that Jeffrey and his co-workers based
many of their conclusions on investigations of woody and herbaceous members
of the Compositae which are multifasciculate when both young and old. Had
 INTRODUCTION xxxvii

they extended their researches to a wider range of herbs it is probable that, like
Bailey and Sinnott, they would have appreciated the difficulty of accepting
the hypothesis that the progressive conversion of vascular tissue to paren-
chyma in the region of leaf insertions provided the sole explanation of the
method by which herbs may have been evolved from trees. In 1914 Bailey
and Sinnott also took exception to the foliar ray hypothesis on the following
grounds. Firstly, if the hypothesis is correct, it is a necessary corollary that
certain of the Amentiferae must be amongst the most primitive living angio-
spernis, and this they believed to be doubtful, Secondly, because multiseriate
rays are known to have occurred in certain Dicotyledons from the middle and
upper Cretaceous, i.e. at a period before the cooler climate of the Tertiary
period began. It is difficult to understand why aggregation and subsequent
compounding of rays should have occurred by then, especially as upholders
of the theory stressed the importance of a cooler climate in inducing these
alleged changes. Moreover the occurrence of plants with broad rays in
tropical regions at the present time does not seem to indicate that low tempera-
tures have been necessary for their formation. Thirdly, the evidence from
seedling structure advanced in favour of the hypothesis is not always reliable,
since the seedlings of certain oaks are known to include broad rays, and it is
not true that the seedlings of Quercus differ universally from the adult in
lacking broad rays. Fourthly, the interfascicular parenchyma of a multifasci-
culate axis is not always subtended by a tiny leaf-trace bundle as demanded
by the hypothesis, but abuts directly on the pith tissue between strands of
primary wood. By 1914 Bailey and Sinnott regarded the broad ray as having
been evolved from a narrow type by simple enlargement which took place
quite independently of the insertion of leaf-traces. To the present author it
also seems that, if the foliar ray hypothesis were generally applicable, one
would expect to find a very much dissected stele in plants in which leaves are
particularly congested on the axis, and, conversely, a less dissected vascular
system in plants with more widely spaced leaves. This is not in accordance
with the facts, however, for if we examine a species of Ranunculus with widely

spaced leaves we find a multifasciculate cauline vascular system. In woody

members of the Ranunculaceae such as Clematis this is also true. In the
woody species Veronica traversii or indeed in any of the woody members of
this genus, in spite of the close approximation of the leaves on the axis, the
xylem cylinder is interrupted only by small gaps which do not extend upwards
and downwards in the axis to any appreciable extent, so that in transverse
sections through an internode the closed cylindrical form of the xylem is
maintained. The same applies in herbaceous species of Veronica. We thus
have a striking example, and it is only one of the many that must occur, which
strongly indicates that the degree of dissection of the cauline vascular system
isnot directly governed by the leaf-trace strands.
The essential difference between the attitude of Bailey and Sinnott on the
one hand and of Jeffrey and Torrey and those with similar views on the other
may perhaps be made clearer by the above series of diagrams (Fig. 3 A-E),
after those in one of Sinnott and Bailey's (2113) papers. In A we have the
primitive, arboreal angiosperm where wide rays flanking the leaf-traces are
absent. In c the traditional structure of a herb is shown in which the foliar
vascular strands are subtended on the outside and partly flanked by the
xxxviii INTRODUCTION
parenchyma of the foliar ray which is shown in black. By elimination of the
subtending and retention of the flanking portions of the foliar ray tissue the
polyfascicular type of stem (E) which is characteristic of advanced herbs is
obtained. In the view of Bailey and Sinnott, the type of structure shown in
H may, in some instances, have been derived from A through c as pointed out

by Jeffrey and Torrcy, but unlike the last two authors they also believe that
the structure in E may have been derived directly from an arboreal ancestor
with a structure like that shown in B merely by a reduction in cambial activity.
In B there are well-marked flanking rays by which the xylem is broken up into
separate bundles, In D is shown the structure of a herb in which the xylem
forms a continuous cylinder, and this Bailey and Sinnott regard as having
been derived from A, again by less prolonged cambial activity. To the present
author the reasoning of Bailey and Sinnott appears simpler and more generally
applicable than docs that of Jeffrey and Torrey, although the two conceptions
do not appear to be mutually exclusive in all cases. Whilst Bailey and Sinnott
have clearly demonstrated the probable anatomical relationship between
woody and herbaceous species which are closely related, their belief that
arboreal forms were the sole ancestors of the modern angiosperms needs
additional proof.
Thoday (2250), after making an intensive study of the stem of the sun-
flower (Helianthus annuus), concluded that there is no evidence of the deriva-
tion of herbs from trees by the conversion to parenchyma of segments of an
originally continuous stele. His views are highly critical of the respective
positions taken up both by Jeffrey on the one hand and Bailey and Sinnott on
the other. Thoday does not believe that the ontogenetic development of the
sunflower stem recapitulates its phylogenetic development. At the same time
he does not appear to take exception to Bailey and Sinnott's idea that a herba-
ceous stem in all its essentials is like the first annual ring of its woody relatives.
Smith (2147), who has studied the nodal anatomy of Acer, Platanus, and
Quercus, concurs with the view of Kostychev (1271, 1272) that a continuous
procambium ring is the most common arrangement in the Dicotyledons, the
dissected stele having been evolved by the conversion of segments to paren-
chyma. Smith believes, however, that far more ontogenetic work is needed
before the phylogeny of the angiosperm stele can be profitably considered.
It would be as well, before leaving the subject of rays, to recall that Stone

(2204) expressed the opinion that these structures are to be regarded as 'stop-
gap-tissue* of the same nature as callus, and that they occupy slits formed in
the cambium cylinder caused by the increase in the periphery of the stem.
At this point it may be appropriate to consider the views expressed by
Arber (30) who, unlike Bailey and Sinnott and many other botanists, believes
the herbaceous habit to be primitive and the tree habit derived. She agrees
that the floral characters of trees are often more primitive than those of
related herbs, e.g. in the Leguminosae, Violaceae, &c., but points out that this
may be due to the evolutionary lag in trees caused by the very fact that they
are unable to pass through so many generations in a given space of time as is
possible with herbs. The herbs therefore possess an evolutionary advantage,
which enables genetical changes and consequently the formation of new
species to proceed more quickly. Quoting Church's (424) Thalassiophyta she
draws attention to the suggestion that when plants first emerged from a
 INTRODUCTION xxxix

marine to a terrestrial environment they were subjected to more intense

insolation, resulting in an increase in photosynthesis and the incorporation of
excess carbohydrate in the plant body. One result was a tendency for herbs
to become transformed which may be kept in bounds
into trees, a tendency
in the youthful phases of a race, but one which may assume pathological pro-
portions when the race becomes senile. She therefore regards the inert matter
of a tree as in some respects comparable with the unbalanced development of
useless growths which are known to have occurred in animals which are now
extinct.
While, to the present writer, it seems somewhat distressing to have to regard
the graceful giants of the forest as in any way pathological, it is as well to bear
in mind, when discussing the phylogeny of herbs and trees, that much depends
on precisely what is meant by the term 'origin'. Assuming that life began in
the sea and migrated to land, it is difficult to imagine that this transmigration
was effected by plants in any way comparable with modern arboreal forms.
Plants of the migrationary period were probably more like the modern herbs,
so that on these purely speculative grounds the original land vegetation seems
likely to have been herbaceous and the arboreal habit derived. In the modern
era trees constitute the dominant element of tropical vegetation, and the
geological record points to woody plants other than angiosperms as having
made up a large proportion of warm climate vegetation of past ages. It there-
fore seems reasonable to imagine a primitively herbaceous vegetation having
given rise to a mixture of herbs and trees quite early in the history of terrestrial
vegetation. At this time climatic factors may have determined which of these
two types of habit was dominant in a given region, just as they seem to have
done in the present era.
It is true that a large proportion of fossilized plants which are known from
the Devonian period until after the angiosperms first appeared in the early
Cretaceous were arboreal, but there is no reason to suppose that herbaceous
forms may not have existed simultaneously. May it not be true that in
geological time herbaceous angiosperms existed in periods much preceding
those from which we have any definite fossilized remains ?
To the present author it seems quite possible that much of the difference
of opinion concerning the origin of herbs and trees would disappear if it could
be recognized that the terms 'herb' and 'tree' are commonly used in a very
loose sense. Herbs of the Linum, Veronica, and Galium types, in which a con-
tinuous ring of xylem is differentiated almost at the growing apex of the stem,
may be species which have evolved from arboreal ancestors by reduction of
cambial activity. A herb of the Ranunculus type, with a cauline vascular
system of widely spaced bundles, may, on the other hand, have been evolved
from herbaceous ancestors which never included any arboreal or woody forms.
We can thus see that 'herbs' may be of more than one type. By examining
the structure of stems with this possibility in view, there seems a reasonable
prospect that further information of considerable phylogenetic interest might
be obtained.

Structure of the Node

Sinnott, and Sinnott and Bailey (2111, 2114) have put forward views con-
cerning the phylogeny of the angiosperms which are based on a consideration
xl INTRODUCTION
of the numberof interruptions in the cauline vascular system which occur at
a node, in connexion with the insertion of a single leaf. An extensive survey
of fossil and modern plants showed that in some species the number of
interruptions is three, in others only one, whilst in a third group the number
isgreater than three. Each separate interruption is caused by the attachment
of an individual leaf-trace. The survey originally made by Sinnott led him to
believe the trilacunar node to be primitive, whilst the unilacunar type as well
as the nodes with more than three lacunae were regarded as more advanced.
The phylogenetic development of the unilacunar from the trilacunar type is
believed to have been caused in some cases by the elimination of two of the
gaps and in others by closing together of three which were originally separate.
The type of node was shown to be independent of the external morphology of
the leaf. For example the number of gaps is not constant for all leaves possess-
ing a sheathing base. Sinnott also found the number of gaps to be character-
istic for a single family or even for a whole order, as in the Centrospennae,
which were found to be unilacunar in all the material he examined. In other
families such as the Ranunculaceae there was found to be far more variation.
J

Looking through the table in Sinnott s first paper in which he gives the
distribution of each of the nodal types in some detail, it is striking to note the
very clear demonstration that the trilacunar node occurs in many wholly
unrelated families. It requires rather a stretch of the imagination to regard
all of these families as primitive, and even Sinnott himself admits in his paper

that there are numerous exceptional plants that do not fit in with his main
thesis. It would also be interesting to have more information concerning the

constancy of the type of node in different species of a large genus.

Sinnott and Bailey (2112) seem rather less certain than Sinnott was in his
earlier paper in the same year that the type of node is independent of the
external form of the leaf. In their joint paper the authors point out that a
majority of plants with trilacunar nodes possess stipules, whereas, with certain
exceptions, stipules are absent from those with unilacunar nodes. The same
authors emphasize that leaves with a multilacunar node possess a sheath-
ing base, and also express the view that the growth of the lateral leaf-traces
stimulates the development of stipules since the stipule was invariably found
to occur directly opposite the point of departure of the trace. At the same
time they point out that stipules are generally absent from families in which
the leaf margin is entire even when the node is trilacunar. Bailey and Sinnott
were somewhat uncertain about the morphology of stipules but inclined to the
idea that they are homologous with leaf teeth. In fact stipules, sheaths, ligules,
and similar structures at the base of the petiole are, in their opinion, all homo-
logous 'since they are dependent in position and character on the anatomy of
the node'.
Sinnott and Bailey (2114) further expressed the opinion that the palmate is
phylogenetically more primitive than the pinnate type of leaf, and again based
their conclusions on evidence derived from nodal anatomy, palaeobotany, and
comparative morphology. ^

PHYLOGENETIC EVIDENCE FROM WOOD ANATOMY

The controversy about the foliar ray hypothesis and its bearing on the
possible derivation of herbs from trees has been described above (p. xxxiv).
 INTRODUCTION xli

Another subject that was highly controversial during the same period con-
cerned the origin of the vessel in the Gnetales. It was thought that this group,
owing to its possession of vessels and the occurrence of broad, angiosperm-
like rays in some species, might prove to be a link between other gymnosperms
and the angiosperms. The controversy centred round the problem of whether
the simple perforations in the end-walls of the vessel members ('elements')
of the Gnetales and angiosperms respectively have been developed phylo-
genetically along the same lines. From the outset it appeared unlikely that the
vessel had had precisely the same origin in both groups, because the less
advanced perforation plates in the Gnetales are typically 'Ephedroid' or
'foraminate*, that is pierced by irregularly arranged, circular holes, whereas
the corresponding primitive multiperforate plates of the Dicotyledons are
scalariform, with regularly arranged, elongated holes.
It is generally believed that the ancestral tracheal element was the tracheid,
a long cell with bordered pitting, and that vessels have been derived from
tracheids by the disappearance of the pit-membrane from some of the pitting
in the overlapping walls, thus forming a vertical series of cells that gives an
uninterrupted passage for water. Thompson (2255) in 1918 postulated that
the simple perforation of the Gnetales must have been derived from tracheids
with circular bordered pits, whereas those of the angiosperms must have
been derived from tracheids with scalariform pitting, and that therefore there
can be no genetic connexion between the vessels of these two groups. He
considered, rather, that the vessels of the two groups furnish a remarkable
instance of the independent development of similar structures.
This view was strongly opposed at the time, particularly by Bliss (207),
Jeffrey, and MacDuffie (1408), but has been supported by the more recent
studies of the development of the vessel that are discussed later, and is now
generally accepted. The principal arguments against Thompson's views were
the occurrence of a few scalariform perforation plates in Gneium and of
occasional foraminate plates, similar to those of Ephedra, in some genera of
the Rosaceae and in Paeonia and Tropaeolum. Bliss also held that scalariform
pitting had its origin in the fusion of multiseriate circular pits. This latter
view has not been upheld by subsequent research and it seems highly probable
that the occasional foraminate perforation plates of the Rosaceae represent a
sporadic development from existing circular intervascular pitting in the side
walls, in much the same way as occasional reticulate perforation plates in
the highly specialized vessels of the Bignoniaceae appear to be related to the
hexagonal, alternate intervascular pitting.
Bailey (75) has recently returned to this problem as the result of ontogenetic
studies of the primary and secondary xylem. He considers that vessels arose
first in the secondary xylem and later in the last-formed part of the primary

xylem, i.e. where the secondary walls of the tracheary elements are pitted,
and that specialization in the primary xylem thus lags behind that of the
secondary xylem. He points out that Dicotyledons whose vessels in the
secondary wood are very unspecialized often possess vessels in the primary
xylem that are barely distinguishable from scalariformly pitted tracheids. He
states that 'the fact that structurally primitive angiosperms had scalariformly
pitted tracheids, from which vessels originated, rules out
any possibility of
deriving the angiosperms from the Gnetales or other representatives of the
xlii INTRODUCTION
higher gymnosperms. The Ginkgoales, Coniferalcs and Gnetales are charac-
terized by having a highly specialized and peculiar type of primary xylem
which is entirely unlike that of other known vascular plants, with the possible
exception of the Ophioglossales.' The development of the vessels in the
Gnetales from circular pitted tracheids he concludes to be unique and 'entirely
unlike the derivation of vessel members from scalariformly bordered-pitted
tracheids in Selaginella, Pteridium, monocotyledons and dicotyledons*. Bailey,
indeed, considers that vessels have originated independently not only in the
Gnetales and angiosperms, but in five distinct categories of the Tracheophyta,
namely the Selnginelnles, Filicales, Gnetales, Monocotyledons, and Dicoty-
ledons.
in 1918 Bailey and Tupper published a paper (100) on size variation in
tracheary cells, which marked the beginning of a new period of phylogenetic
wood anatomy. The significance of this work does not seem to have been
immediately appreciated, and it was not till Frost (728-30) applied this work
to the development of the vessel member that it was realized that there was
to hand a new tool of the greatest importance. The main conclusion reached

by Bailey and Tupper can be stated very simply that the length of the
tracheary elements and of the cambial initial is reduced as specialization
increases. The special value of this investigation lies in the fact that it was
based on observations over such a wide range of plants from the Cycadales
to the angiosperms, and including various fossil plants that, as Tippo (2261)
has said, it can be fairly claimed that the conception of reduction in length as
an index of specialization is not based on any system of angiosperm classifica-
tion and is independent of any preconceived notion that this or that group of
the gymnosperms has given rise to the angiosperms; or any idea that the
Ranales or the Amentiferae are primitive. Consequently it offers a means of
breaking the vicious circle caused by a taxonomic group being regarded as
primitive or advanced because its floral characters are primitive or advanced,
the floral characters themselves being judged as primitive or advanced because
they occur in primitive or advanced groups.
Frost correlated vessel member length with various other characters, of
which the most important was the nature of the end- wall. He concluded that
the scalariform perforation plate with many bars set in a very oblique end-
wall is the most primitive type because it is associated with the longest vessel
members he also concluded that subsequent specialization has taken the form
;

of a progressive reduction in the number of bars and perforations and in the

steepness of the end-wall to a single, simple perforation in an almost horizontal
end-wall. The relation between this series and vessel member length is shown
in the following table, taken from Frost.

TABLE i

Vessel Member Length and Type of Perforation Plate

 INTRODUCTION xitii

Frost applied a similar technique to intervascular pitting and demonstrated

a series with scalariform pitting as the most primitive, progressing through
transitional types to opposite and finally to alternate. The relation between
these types and vessel member length, as demonstrated by Frost, is shown in
Table 2.

TABLE 2
Vessel Member Length and Intervascular Pitting

The validity of this work is now generally accepted and the same technique,
applied to other characters in wood, has helped to demonstrate other phylo-
genetic series. The results make possible many interesting comparisons of
the relative specialization of different groups of the angiosperms. Before
discussing this later work in more detail, it is perhaps advisable to point out
that individual measurements of vessel member length based on single samples
must not be interpreted too exactly, as cell dimensions vary considerably
under different conditions and in different parts of the stem (see p. xxiii).
Further, most of this work is based on the assumption, probably quite
legitimate when based on large numbers of observations, that characters
constantly or predominantly associated with long tracheary elements are
themselves primitive.
Using the series of vessel characters established by Frost (729) as phylo-
genetic indicators and a similar statistical technique, Kribs (1286) has defined
certain types of ray, which are believed to exhibit degrees of specialization.
His conclusions may be summarized as follows. The most primitive ray type
consists of a combination of multiseriate rays with high uniseriate 'wings',
and numerous and high uniseriate rays composed of high upright cells.
(Plate C, Heterogeneous I). Specialization takes the form of reduction either
of the uniseriate rays, leading to a type with multiseriate rays only (Plate C,
Homogeneous II), or of the multiseriate rays, leading to types with wholly
uniseriate rays (Plate C, Heterogeneous III and Homogeneous III). Parallel
with reduction of the uniseriate or of the multiseriate rays, individual rays
become less heterogeneous and the component cells smaller. Wholly uni-
seriate rays are never primitive, being derived from multiseriate types, those
derived from less advanced types tending to be heterogeneous with large cells.
Kribs's views on the most primitive type appear to be generally acceptable,
but his assumption that reduction of either the multiseriate or the uniseriate
rays is always accompanied by a decrease in heterogeneity cannot be com-
pletely justified and leads to an over-simplification of the problem. It has
been objected by Barghoorn (138) that reduced cambial activity, such as
occurs in shrubs, is often accompanied by the retention or exaggeration of
heterogeneity. Such plants may retain a primitive type of ray, e.g. high-
celled uniseriate rays, or there may be a tendency to axial elongation of the
ray cells, converting procumbent cells to upright cells. These woods may
xliv INTRODUCTION
thus he specialized in so far as their rays are wholly multiseriate or wholly
uniseriate, yet the rays may be composed entirely of high upright cells. There
is no place in Kribs's classification for such rays. The tendency to axial
elongation of the cells may in extreme cases convert all the ray initials to
fusiform initials and result in a wood with no rays, e.g. in Bocconia, Santolina,
and Sesamum.
of perforation plate is a much more convenient index of specialization
Type
than vessel member length, as the latter can be satisfactorily measured only
in slides of macerated material; the relation between the two characters
appears to be sufficiently close to justify the use of type of perforation alone.
One shortcoming of this method, however, is that more than half the species
lie most advanced group obtainable with this feature perforations
in the
simple and end-walls transverse. This large group, however, can be sub-
divided by the use of storied structure to distinguish the most highly specialized
woods.
It has been shown by Beijer (168) and others that storied structure (see

p. xxvii) is related to the length cambial initial. The number of cells in

of the
the cambium constantly being increased to keep pace with its increasing
is

perimeter as the stem grows in girth. In most woods this increase in the
number of cambial cells is obtained rather indirectly, through the occasional
division of an initial into two halves by the formation of a horizontal cross-
wall; the two halves subsequently elongate till they overlap for a large part of
their length and so constitute two cells approximately side by side. This is

typical of woods with long cambial initials. In woods in which the mean
length of the initials is less than about 0*4 mm. the plane of division changes
and, instead of a small transverse wall, the cell forms a long radial wall. This
gives rise to two cells almost exactly side by side and, when repeated, to a
horizontal series of initials lying side by side, and so to storied structure in
the wood. Storied structure, therefore, being associated with very short
cambial initials, can be used as an index of a very high level of specialization.
Data from about 1,800 woods have been classified by the author into the
1

following groups of increasing specialization :

I. Perforation plates wholly scalariform (206 woods).

II. Perforation plates scalariform and simple (91 woods).
III. Perforations all simple, elements not storied (1,261 woods).
IV. Perforations all simple, rays or parenchyma storied (243 woods).

The percentages of the woods of each group exhibiting particular features

are given in Tables 3-5. In these tables each figure represents the percentage
of all the woods in one Group with a particular feature. For example, 132 out
of the 206 woods in Group I, or 65 per cent., have scalariform to opposite
pitting; the 35 per cent, of this Group that have other types of pitting are not
referred to in this table.
1
Many of the data accumulated during the writing of this book were recorded on 'punched
cards', and correlations between different features could then be simply obtained by counting
the numbers of cards punched for particular features. In general one card was used for each
species, but where a species was sufficiently variable it was necessary to use more than one
card for it, or, conversely, where species could not be distinguished, more than one species
may be represented by a single card. Such exceptions, however, are not sufficiently numerous
to mask any clearly marked trend and in any case tend to cancel each other out. More species
were grouped than were given more than one card per species.
 INTRODUCTION xlv

TABLE 3
Features associated with a Low Level of Specialization

The proportion of woods with scalariform and opposite intervascular

pitting shows the progressive decrease from Group I to Group IV that would
be expected from Kribs's work and calls for no special comment. The next
feature, exclusively solitary vessels, has been claimed, e.g. by Tippo (2261), to
represent an imspecialized type. This is confirmed by the above figures, the
percentage dropping from 35 in Group I to 5 in Group IV. The figures in
this table also confirm that both fibres with
distinctly bordered pits and
diffuse parenchyma are unspecialized. Specialization in the fibres
appears to
have proceeded from the fibre-tracheid, with large bordered pits in all walls
touching other fibre-tracheids and numerous pits to vessels, towards the
libriform wood fibre, with small, simple to slightly bordered pits on the radial
walls between fibres and with few or no pits to vessels.
Specialization appears
also to have been accompanied by a greater
proportional elongation of the
developing fibre, so that, while fibre-tracheids are seldom more than one and
a half times as long as the cambial initial from which
they were derived,
libriform fibres are commonly two or three times and may be up to eight or
nine times (374) as long.
The figures for diffuse parenchyma differ in some respects from those given
by Kribs in his study of the specialization of parenchyma
(1287), particularly
the persistence of diffuse parenchyma in the highest group (23
per cent, in
Group IV). In Kribs's highest group, woods with transverse simple perfora-
tions, the percentages are o for 'diffuse* and 2-7 for 'diffuse-aggregate'. This
discrepancy may perhaps be accounted for if Kribs included only genera in
which these types are dominant, as the author's figures include all species in
which diffuse parenchyma occurs, irrespective of other types in the same
wood. It seems probable that there is a more advanced type of diffuse paren-
chyma, possibly Kribs's 'diffuse aggregate', which should, if possible, be
distinguished from the unspecialized type.
The figures given in the table below fully bear out the view that homo-
geneity in rays is an advanced character. It is also apparent that paratracheal
parenchyma in some form or other is more commonly associated with advanced
than with primitive woods, the percentage of woods with predominantly
paratracheal parenchyma rising steadily from 14 in Group I to 57 in Group IV.
Of these paratracheal forms aliform and confluent appear to be the most
xlvi INTRODUCTION
TABLE 4
Features associated with a High Level of Specialization

advanced. This is in general agreement with the conclusions of Kribs (1287),

who, using a similar technique based entirely on type of perforation and slope
of end-wall, placed his parenchyma types in the following sequence and corre-
lated them with vessel member lengths (given in brackets) diffuse (0*92 mm.),
:

absent (0-78 mm.), diffuse aggregate (0*65 mm.), vasicentric scanty (0-60 mm.),
metatracheal narrow (0*51 mm.), terminal (0*44 mm.), metatracheal wide
(0-42 mm.), vasicentric abundant (0-31 mm.). Other conclusions drawn by
Kribs were that the absence of wood parenchyma indicates a primitive con-
dition, that terminal parenchyma is a specialization due to reduction and that,
as wood becomes more highly specialized, the individual wood parenchyma
cells become shorter and wider.
The number of cells in a single strand of parenchyma can also be used as
an index of level of specialization. Long cambial initials that give rise to
strands of 8 or more cells are characteristic of unspecialized woods. Strands
of 1-2 cells and numerous fusiform parenchyma cells are most frequent in
highly specialized woods. The tendency for the number of cells to decrease
as the strands become shorter in length is, however, partially offset by the
parallel tendency, mentioned above, of the cells to become shorter as they
become more highly specialized.
TABLE 5
Miscellaneous Features

Groups in order of increasing specialization

Minute, alternate, intervascular pitting might logically be assumed to

represent the end of the series scalariform opposite alternate and so be a
very advanced character. From Table 5 it will be seen that though it is less
common in Groups I and II, as would be expected from the mere fact that it
 INTRODUCTION xlvii

is a alternate pitting, there is no significant difference between the

form of
proportion of woods with this feature in Groups III and IV. It is suggested
that minute pitting represents a special development peculiar to particular
groups rather than a form of specialization towards which the vessels of all
moderately advanced woods tend. This feature is likely, therefore, to prove
of value as an index of affinity rather than of level of specialization.
Septate fibres tend to be of common occurrence throughout large natural
groups, such as that which includes the Meliaceae and Burseraceae, and are
a most useful index of affinity. Opinion is divided as to whether they repre-
sent an advanced or a primitive type of fibre. Chalk (354) in 1937 suggested
that they should be regarded as more primitive than the non-septate Hbriform
fibre, but Tippo (2261) a year later reached the opposite conclusion from his
study of the woods of the Moraceae and their presumed allies. The figures
given in Table 5 show that septate fibres occur just as commonly in primitive
as in moderately advanced woods (Groups I-I1I), but that they are far less
frequent in the most advanced woods (Group IV). A list of the families in
which septate fibres occur is given on p. 1351.
Ring-porousness, or the development of a marked zone of larger vessels at
the beginning of the growth ring, appears to be accompanied by an increase
in the length of the complete vessels in the pore zone, and has been shown by
Huber (1102) to increase the rate of flow of water through the wood. It is
associated with cold winters or alternating very dry and wet seasons and has
been generally assumed to be a specialized development of either an ecological
or an evolutionary nature, though the actual evidence adduced has been
rather slender. Recently Gilbert (765) has investigated this problem afresh,
using both the older approach of Jeffrey, based on recapitulation and sup-
posedly conservative parts of plants, and the newer technique introduced by
Frost. Gilbert has concluded that 'there is abundant evidence to indicate
that the ring-porous type of vessel arrangement represents an evolutionary
advance from the diffuse distribution*. He states that in any genus the ring-
porous species have other advanced characters, compared with the diffuse-
porous species, and that, within the north temperate regions, ring-porousness
is correlated with other specialized characters, such as simple perforations,

paratracheal parenchyma, and simple pits in the fibres, but that it is not
correlated with ray types as defined by Kribs (1286). He considers that this
specialization took place early in angiosperm history and affords no proof of
parallel lines of development.
Gilbert emphasizes strongly that morphological comparisons are valid only
with species from the limited region of the world in which ring-porousness
seems to have developed, and that the feature is absent from the temperate
regions of the southern hemisphere. This may account for the fact that the
regression shown in Table 5 between ring-porousness and degree of specializa-
tion is not significant statistically, as judged by the x z test, 1 as the data were

1
The relevance of this test is somewhat doubtful owing to the tendency of related woods
to give similar records of occurrence or non -occurrence of a particular character. This applies
to all the features measured, but those given in Tables 3 and 4 have such exceedingly large
2
X values that the significance can scarcely be in question. For the features, other than ring-
porousness, in Table 5, the differences are not significant according to the x* test, but the
comparatively steady trend of the percentages suggests that there is a real trend from Group I
to Group IV, though this cannot be so firmly established as for the other features.
xlviii INTRODUCTION
drawn from all over the world. It is to be noted, however, that the population
on which Gilbert bases his conclusions has some peculiar features, and neither
his ring-porous nor his diffuse-porous group corresponds with material from
the wider geographical range investigated by the author. For example, 85 per
cent, of the species in his ring-porous group have paratracheal parenchyma
and 27 per cent, in his diffuse-porous group, compared with 50 and 42 per
cent, for the author's material. The proportion of woods with scalariform
perforation plates in Gilbert's whole group of about 60 species is rather higher
(22'5 per cent.) than for the author's 1,800 woods, but, whereas Gilbert's
woods did not include a single ring-porous species that had scalariform per-
foration plates, the author's data included several.
The relation between the features discussed by Gilbert and the occurrence
of ring-porousness in the material examined by the author is shown in the
following table.
TABLE 6

Comparison of 128 Ring-porous with 2904 Diffuse-porous Woods

Percentage of Percentage of
j

ring-porous diffuse-porous

......
Feature woods woods
' '

With diffuse parenchyma 32 35

With predominantly paratracheal parenchyma . .
53 47
With fibres with distinctly bordered pits . . .
32 30
With homogeneous rays . . . . . . 22 18

For each feature the difference between the numbers of woods with and with-
out the feature in the ring-porous and the diffuse-porous groups was subjected
to the x 2 test, and in no case was the difference found to be significant.
It would appear unwise, therefore, to rule out the possibility that ring-

porousness may represent an ecological specialization, which occurs in wood

at very different levels of general specialization and in widely separated
taxonomic groups (for list of families see p. 1350).
Spiral thickening of the secondary wall of the vessel is regarded by Frost
(730) as an advanced character. In the material examined by the author,
however, spiral thickening was relatively more common in unspecialized than
in specialized woods it will be seen that in Table 5 the percentage of woods
;

in which spiral thickening occurs drops from 15 per cent, of Group I to 8 per
cent, of Group IV. Spiral thickening is much more closely linked with ring-

porousness. Of the diffuse-porous woods tabulated only 140 out of 2,780 woods
(6*5 per cent.) possessed spiral thickening, while among the ring-porous
and semi ring-porous woods 125 out of 289 (44 per cent.) had this feature.
It is possible that it is the ecological factors associated with ring-porousness
that are primarily responsible for this correlation. Spiral thickening occurs
sporadically in many widely separated families (for list see p. 1349).
So wood anatomy has been discussed mainly as an index of general
far,

specialization, and it remains to consider how it may be used to demonstrate

affinity. The fact, so important for the former purpose, that the number of
characters in wood is small and that each of the elements has tended to follow
one general trend of specialization, to some extent limits the use of wood
anatomy for this purpose. Paucity of characters also implies the probability
 PLATE A APOTRACHEAL PARENCHYMA

TERMINAL

Scattered Diffyse-in-aggtegates
DIFFUSE

METATRACHEAL OR BANDED
 U
2

E
x
o
a

PM

< X,

X O
o
CQ

% "V

L.J
 qjpS^SSgBsggyuaytaiieq

00
N
.~rttr.:"~**^
Btro^sai
U

^!I!^^ r/5

^ ;t***t*ww; si
ii
i
e
^ u:
P
t?/ j i,
^^.^^i^^jrfi.j.m,!

X O
tii
w
O X
w B w
a:
O
h S
O
B

O
W
 INTRODUCTION xKx
of many cases of parallel development. If two or more taxonomic groups
have but one important character in common, this should he regarded as
suggestive rather than as proof of affinity, unless supported hy other evidence,
There are some special lines of development limited to well-defined groups,
such as tile cells in the Bombacaceae and Sterculiaceae. It is, however, not
only unusual characters that are of special taxonomic interest, but also unusual
combinations of common characters. This is, perhaps, particularly true of
characters that indicate specialization, where one feature is at a markedly
different level from the others. For example, Betula is characterized by vessels
with exclusively scalariform perforation plates, but minute, alternate inter-
vascular pitting, the former being an unspecialized feature and the latter an
advanced one. Similarly, in the Zygophyllaceae there is an unusual combina-
tion of storied structure (specialized) with distinctly bordered pits in the fibres
(unspecialized).
The significance of the more important characters of the wood, not already
described on pp. xxi-xxvii, is discussed briefly below.

Vessels
The most valuable taxonomic vessel characters are the distribution and
pattern of the pores as seen in transverse sections. The majority of woods
have a preponderance of solitary vessels, mixed with some multiples of two
or three cells. Divergence in either direction to exclusively solitary vessels
or to more numerous or larger groups and multiples is useful for identifica-
tion, but the possession of exclusively solitary vessels, being a common un-
specialized type, cannot be used alone as indicating affinity. In some groups
there is a marked tendency for multiples to be more common or for the vessels
tobe arranged in radial, oblique, or tangential lines, and these characters can
be used positively as indicating affinity. They cannot, however, be safely
used negatively as evidence of lack of affinity with woods without any such
pattern. The marked radial or oblique pattern known as 'dendritic' (see
Fig. 175 A), however, seems to lack any such significance, as it occurs
sporadi-
cally in isolated genera in many widely separated families (for list see p. 1351).
It has already been mentioned that minute, alternate, intervascular pitting
tends to be characteristic of large natural groups; the same may possibly be
true of very large alternate pitting. The nature of the pitting between vessels
and ray or wood parenchyma cells also appears to have some taxonomic
significance, particularly the distinction between pitting that is essentially
similar in size and shape to the intervascular pitting, and that which includes
some much larger, and often irregularly shaped pits, one or other of these
types commonly persisting throughout a family or larger group. The feature
known as a Vestured pit* (formerly 'cribriform membrane') has been shown
by Bailey (78) to be characteristic of a limited number of families (for list see
p. 1350). Vasicentric tracheids tend to be associated with particular taxonomic
groups, but are characteristic of tribes rather than families (for list of families
seep. 1351).
Parenchyma
This is possibly the most important tissue indicating relationship, but, in
the present state of knowledge, is also the most difficult to interpret. Probably
the most fundamental distinction is that between apotracheal and paratracheal
1 INTRODUCTION
parenchyma (sec Plates A
and B), and in most families the parenchyma
is wholly or predominantly one or the other. In a few groups, e.g. in the
Malvalcs and in some of the Dipterocarpaceae, both types may be common
in the same wood, but this is sufficiently rare for it to constitute a distinctive
character. Some
of the banded types are difficult to interpret as definitely
apotracheal or paratracheal. Parenchyma also tends to be rather variable in
type, particularly under conditions that cause the amount to vary. This is
true not only of different specimens, but even of different parts of the same
ring, it being common, for example, for parenchyma to be more banded in
the outer part of the ring. Such variation, however, seldom affects the funda-
mental difference between apotracheal and paratracheal and appears to repre-
sent merely a step up or down in a series, e.g. from aliform to confluent or
from scattered diffuse cells to short apotracheal lines (diffuse-in- aggregates).
Use may be made of this variation for the interpretation of banded types;
where parenchyma is least abundant in a species, either in particular rings or
parts of a ring or in different specimens, 'it will usually be found that, if the
bands arc essentially paratracheal, the parenchyma will be broken down to
aliform-confluent types, whereas if the bands are apotracheal, there will be
some vessels without any parenchyma round them. Very little appears to be
known about the influence of external factors on such variation.
Single crystals of calcium oxalate scattered in the ordinary cells of wood-
or ray-parenchyma are too common to be of much value, but chambered

crystalliferous cells and forms of crystal, such as raphides and druses,

special
are of significance, as is the occurrence of silica and of less common inclusions
such as lapachol (1789) and aluminium (2515).

Rays
Most ray characters, such as heterogeneity and the number of uniseriate
than affinity. Size, parti-
rays, indicate general level of specialization rather
cularly the maximum number of cells wide, is sometimes characteristic of
taxonomic groups, e.g. the large rays of the Proteaceae and Rhizophoraceae
and the small rays of the Salicaceae. On the other hand, there is no reason to
suppose that ray size in itself indicates affinity between groups that are widely
separated on other grounds. Further, it must be remembered that exclusively
uniseriate rays are a form of specialization that has occurred independently
in occasional genera of many families.
One of the standard features used in wood descriptions is 'rays of two sizes*.
In some woods, e.g. in Quercus, the rays are either uniseriate or many cells
wide, with no intermediate sizes. On the other hand, in Fagus, the rays of
which are also often described as being of two sizes, all the intermediate
widths between uniseriate and the maximum number of cells can be found.
Without more adequate definition the feature has little significance. It seems
probable that it is associated with the relatively uncommon method of
increasing the ray tissue, to keep pace with increasing girth, by the splitting up
of large primary rays and the subsequent growth of the parts split off to
the maximum size.

Fibres
It has already been mentioned that septate fibres afford a valuable indica-
tion of affinity, and that they occur commonly throughout large and small
 INTRODUCTION li

groups that are considered to be closely related on other grounds (for list of
families see p. 1351).

Intercellular Canals

Normal canals are of very limited occurrence and tend to be characteristic

of particular groups. For example, the Dipterocarpaceae typically have
vertical canals and the Anacardiaceae and Burseraceac commonly have radial
canals. A tendency to form canals in response to injury also tends to be
characteristic of taxonomic groups, e.g. the Meliaceae (for list of families
with intercellular canals see p. 1353).

Vestal (2329), using a technique similar to that of Frost, concluded that the
phylogenetic value of anatomical studies rests mainly on the fact that the
characters employed are quite different from the cxomorphic ones that arc
normally employed by taxonomists. It is thus possible to obtain an entirely
independent check on the conclusions of the orthodox taxonomist. Vestal
(2330) also points out other strengths and weaknesses of the anatomical
method of testing phylogeny, and rightly emphasizes that anatomical charac-
ters can be successfully used only to supplement exomorphic features.
Heimsch and Wetmore (939) have shown how, by using those characters
of the secondary xylem which they describe as being generally accepted in-
dicators of phylogenetic development, it is possible to show specialization
tendencies within a family. It was found possible to arrange the constituent
genera of the Juglandaceae in a phylogenetic sequence that agrees in nearly
all details with one already prepared by Manning (1432), working with the

exomorphic features.
Chalk (354) has drawn attention to the fact that parallelism between the
wood structure and the accepted systems of classification tends to break down
in groups larger than the family. He has compared the evidence for a parti-
cular phylogenetic sequence derived from wood structure with the arrange-
ment in each of the taxonomic systems of Bentham and Hooker, Engler, and
Hutchinson. Hutchinson's arrangement of the Archichlamydeae was found
to agree more closely with the evidence from wood structure than does that
of Engler. Other conclusions drawn were that :

'The Monochlamydeae of Bentham and Hooker appears to be slightly less highly

specialized than the Polypetalae, but neither group differs very markedly from the
average for all the Dicotyledons examined. The Metachlamydeae (Sympetalae)
includes a mixture of specialized and unspecialized woods, but these can be separated
by the type of parenchyma present. The woods of Engler's Primulales, Phimba-
ginales, Contortae (part), Tubiflorae and Campanulatae have paratracheal paren-
chyma and constitute a highly specialized group/

CONCLUSIONS
The main emphasis in recent years has been on the anatomy of the secondary
xylem and in particular on the evolutionary trends of its elements. At present
direct comparison between woody and non-woody plants is not possible, but
work on anatomical specialization in secondary wood suggests that the
application of this knowledge and of similar techniques to the xylem and
Hi INTRODUCTION
other tissues of the non-woody plants may ultimately make such a comparison
feasible. Tippo (^64) has said:

'There is every reason to suppose that the trends already defined will he found
to prevail in the secondary wood of the herbs, and there is also some basis for the
belief that similar sequences will be unearthed in the primary xylem. Bailey (75),
for t-xample, has recently shown that the evolutionary development of the vessel
elements in the primary xylem parallels the development of these structures in the
secondary xylem/
The recent work of Cheadlc (382 -8) on the systematic anatomy of the Mono-
con ledons indicates other lines along which further advances are likely to
be made.
There seems to be a considerable body of evidence in favour of regarding
the Magnoliaceae and their allies as retaining many primitive characters.
This is supported especially by the lack of vessels in their allies, the homo-
\ylous angiosperms Drimys, Trochodendron, and Tetracentron. This, in all

probability, is to be regarded as the retention of a truly primitive character

rather than as a reduction., Modern views as to the origin and specialization
of the vessel, particularly those concerned with the derivation of perforations
from bordered pits, appear to rule out any possibility of deriving the angio-
sperms from the Gnetales (75).
It thus seems probable that one line of descent among the angiosperms
started with forms, possibly now extinct, which were in some respects similar
to the modern Winteraceae, Trochodendraceae, and their allies. This con-
clusion is in accord with the views of many taxonomists who work with floral
and other exomorphic characters, and also with the views expressed by
Wieland (2421).
The main general conclusions to be drawn from studies of the vessel have
been summarized by Bailey (75) as follows:
'The independent origins and specializations of vessels in monocotyledons and
dicotyledons clearly indicate that, if the angiosperms are monophyletic, the mono-
cotyledons must have diverged from the dicotyledons before the acquisition of
vessels by their common ancestors. This renders untenable all suggestions for
deriving monocotyledons from vessel-bearing dicotyledons or vice versa. Further-
more, the highly specialized structure of the xylem throughout both stems and
roots of herbaceous dicotyledons, not only affords conclusive supplementary
evidence of the derivation of herbaceous from arboreal or fruticose dicotyledons,
but also is an insuperable barrier to the derivation of monocotyledons from hej-
baceous dicotyledons.'
There seems to be some danger in assuming too readily that all modern
angiosperms have been derived from Magnolian or allied ancestors. It is
possible that a Magnolian stock may have provided the sole starting-point for
the evolution of the angiosperms, but, if this be so, it is somewhat remarkable
to find evidence of the co-existence of wholly unrelated flowering plants so
long ago as the early Cretaceous. That the angiosperms may be polyphyletic
is no new and Wieland implies that geological evidence tends to support
idea,
this view. Campbell (335) discusses this problem, and concludes that the
weight of evidence appears to favour the view of Engler and Wettstein that
the angiosperms are polyphyletic. He states that both comparative morphology
and the geological record indicate that the existing angiosperms represent a
 INTRODUCTION liii

number of distinct phyla which cannot be traced back to a single ancestral

type. The anatomical evidence appears to support this view. ~y

SELECTED LITERATURE. (See also p. xxix)

Arber 30, Bailey 67, 68, 71, 75, Bailey and Sinnott 97, Bailey and Tupper 100, Bancroft
121, Barghoorn 137-9, Bews 193, Bliss 207, Bouvrain 247, Bremerkamp 267, Brown,
F. B. H. 280, Campbell 335, Chalk 354, 357, Cheadle 382-8, Church 424, Coulter 483,
Dormer 604, Eames 620, Frost 728-30, Gilbert 765, Hallier 874, Hcimscli and Wetmore
939, Hill, T. G. 974, Hoar 981, Jeffrey 1162, 1163, 1168, 1169, Kostycht-v 1271, 1272,
Kribs 1286, MacDuffie 1408, Majumdar 1426, Manning 1432, McNair 1473-8, Metealfe
1500-2, Petersen 1706, Sinnott 2107, 2111, Sinnott and Bailey 2112, 2114, 2115, 3x16,
Smith, E. P. 2147, Stone 2204, Stopes 2208, Stopes and Fujii 2209, Tansley 2235, Thoday
2250, Thompson 2253-5, Thompson and Bailey 2256, Tippo 2261, 2263, 2264, Vestal
2329, 2330, Weiss 2392, Wieland 2421.

SELECTION AND PREPARATION OF

MATERIAL
FOR all investigations in systematic anatomy it is essential to work with
material that is accurately named. The ideal at which to aim is that all investi-
gated material should be accompanied by herbarium specimens from the
same plant. Since this ideal cannot always be attained, it is frequently neces-
sary to compare specimens to be identified with museum samples. If the
samples in the museum collections all agree with one another and have been
obtained from reliable sources, identifications which are sufficiently accurate
for practical purposes can usually be made, but it is desirable to compare
material of which the identity has been established beyond question whenever
possible. The
difficulty of obtaining flowering specimens of commercial
material often very great, especially when it is customary for the flowers to
is

be removed before the material is regarded as a commercial article. In con-

sequence there are, even now, economic products of vegetable origin whose
botanical identity has never been finally established in spite of their having
been known in commerce for many years. It can thus be seen that the
building up of collections of authentically named material for microscopical
examination is of fundamental importance.
For general purposes, herbaceous material for microscopical examination
is best fixed and preserved in formalin-acetic-alcohol or a similar solution
until slides can be made. The taxonomist, when collecting specimens to be
dried and mounted for the herbarium, would be doing a valuable service if he
could also preserve material for anatomical work at the same time, since the
authenticity of the material could then always be checked from the corre-
sponding herbarium sheets. This would entail considerable additional labour
on collecting expeditions, and might present difficulties in inaccessible
localities, but modern transport should make this easier as time goes
facilities
on. Collections of floral material in fluid preserving media are now assuming
increasing importance in herbaria, so it would be a logical development to
include portions of the vegetative organs for anatomical investigation at the
same time. Woody specimens can usually be preserved without any special
treatment.
Herbarium specimens can frequently be revived sufficiently for their
anatomical structure to be studied, but the ease with which this can be done
 liv INTRODUCTION
varies in different families. For example, revived specimens of the Solanaceae
are usually most unsatisfactory. The usual procedure is to place the material
in methylated spirit under reduced pressure to remove air from the tissues,
after which it is transferred to hot water, which helps to soften it ; then it is
treated in cold dilute solutions of caustic soda or eau de Javelle, which fre-
quently cause the cells to swell up and assume their original dimensions. The
duration of treatment in each of the reagents, the gradual stages in passing
from one to the other, and the strength of the caustic soda solution all need
modification to suit the nature of the material. After the caustic soda treat-
ment, the excess of alkali is neutralized with dilute acetic acid, the acid being
subsequently removed by prolonged washing in running water. This treat-
ment, or modifications of it, frequently enables material to be embedded in
wax and sectioned on a microtome in the usual way, or good sections of
relatively woody or rigid specimens may be cut on a sliding microtome with-
out any preliminary embedding. Great care is necessary, however, when
examining revived material, since the cells are not always fully restored to
their original sizeand shape. This danger is apt to be especially great when
revived material compared with better-preserved specimens of the same
is

species. Herbarium specimens are nearly always worth examining anatomic-

ally, but preference should be given to material preserved in a fluid medium.
Relatively hard material, such as wood, often needs no treatment other than
boiling in water to remove the air. Very hard wood, however, may need
softening. The most common methods of softening are by treatment with
hydrofluoric acid, glacial-acetic acid with hydrogen peroxide, phenol under
pressure, or a steam jet. Wood containing very thin-walled tissues, such as
included phloem, or wood that is attacked by fungus, may need embedding in
celloidin. A comprehensive account of the techniques of sectioning wood is

given by J. Kisser (Z. wiss. Mikr. 48, 320-42; 1932), and a briefer account of
the most commonly used methods is given in the For. Res. Leafl. No. 40;
1946. A critical comparison of different softening treatments is given by
M. Harlow (Tech. Publ. N.Y.S. Col. For. 63, 1944), and a more general
description of sectioning, staining, &c., by C. J. Chamberlain (Methods in
Plant Histology, Chicago, 1932). The acetic acid-hydrogen peroxide softening
technique has been described by G. L. Franklin (Trop. Woods, 88, 35-6;
1946).
The most common method of macerating wood is probably still that of
Schultze, and consists of placing chips of wood in a test-tube with a few
crystals of potassium chlorate and covering them with nitric acid diluted,with
its own volume of water. Another well-known method is to use a 5 per cent,
solution of chromic acid. A
newer technique, using glacial-acetic acid and
hydrogen peroxide, has been described by G. L. Franklin (Trop. Woods 49, ,

21-2; 1937).
Another important requirement is to make reference collections of per-
manent microscope slides. These collections should be housed at institutions
where taxonomic investigations are undertaken, especially at national herbaria.
Permanent slide collections are as important for investigations in systematic
anatomy as an ordinary herbarium collection is for taxonomic work based on

external morphology. The lack of slide collections has done much to delay
the development of systematic anatomy.
 INTRODUCTION Iv

Arrangements for the exchange and loan of slides could be made with other
institutions to the mutual advantage of all concerned. Work on these lines
has already been accomplished in the restricted field of wood anatomy, exten-
sive collections of authentically named timber slides having been assembled
at a number of institutions in different parts of the world, including Great
Britain. It is now desirable to extend these collections so as to include slides
of other parts of the plant. In view of the magnitude of the task, attention
should first be given to plants of economic importance or to those which are
likely to assume economic importance. Sections of other plants could then
be added gradually until a reasonably complete record of the anatomy of the
higher plants has been secured. The slides should be filed in a logical sequence
to facilitate easy reference, using a system which permits the collection to be
expanded indefinitely without having to undertake its complete reorganiza-
tion. This can easily be done in ordinary card index cabinets if the slides are

arranged vertically in standard metal slide holders such as are now on the
market. A filing system of this kind has considerable advantages over one in
which the slides are arranged in boxes or cabinets of the usual type since far
less space is required, and it is possible to add slides to the collection without

having to rearrange all of them. At Kew the nucleus of a slide collection on

these lines has been made during the last eighteen years, and has been found
in practice to work efficiently and to save time when material is received for
identification.
Examination under the microscope should not be confined to permanent
slides. Certain features, such as bordered pits, are more clearly visible in
sections mounted in dilute glycerine, whilst others, such as oily cell contents,
are revealed by microchemical reagents whose effect is only temporary.
Measurements of cells can best be made on macerated material, and clearing
reagents often enable the course of vascular bundles to be seen, or sclerenchy-
matous idioblasts to be observed.
Samples of wood should always be selected from the main stem and not
from branches. The very bottom of the stem should be avoided, even if there
is no obvious swelling or buttressing, as the wood there is apt to be abnormal.

Owing to the changes in cell dimensions that occur from the pith outwards
at any given level, specimens should be located at some distance from the

pith; a suitable position is often at the edge of the heartwood, with the
specimen including both heartwood and sapwood, and it is useful to have
the longest side of the specimen radial rather than tangential so as to include
several growth rings. A convenient size is 4! x i X 6 inches, or 8 X 3 X 15 cm.

ARRANGEMENT OF THE FAMILIES

IT was a matter of some difficulty for the authors to decide which system of
should be used in the present book. On the advice of Sir Arthur
classification
Hill, who was the Director of Kew when the work was started, it was decided
to arrange the families primarily in the sequence first introduced by Bentham
and Hooker in their Genera Plantarum. It is generally recognized, however,
that some of the families, as understood by Bentham and Hooker, do not
constitute homogeneous units, for which reason many systematic botanists
Ivi INTRODUCTION
have sought to improve the Bentham and Hooker classification either by
regrouping genera so as to constitute new families or by elevating certain
tribes or individual genera to family status. Many of the new families thus
created have become generally accepted by qualified taxonomists and have
been treated here as separate units. This procedure has greatly simplified the
task of description.
In the following list the families in the left-hand column are those which
were described by Bentham and Hooker in their Genera Plantarum. Those
in the right-hand column are families that have since come to be generally

accepted. In most instances the latter have been arranged in alphabetical

sequence, following the Bentham and Hooker family in which the constituent
genera of the new families were originally included. Thus, for example, the
Ternstroemiaceae of Bentham and Hooker have become the Theaceae in a
more restricted sense, together with nine families whose members were
previously included in the Ternstroemiaceae as originally understood. A very
few families, such as the Diclidantheraceae and Julianiaceae, have been
removed from their positions in the Bentham and Hooker sequence, since this
seems to be fully justified by the anatomical evidence. No claim is made that
the families which follow one another closely in this sequence are nearly
related in the true taxonomic sense. Some of them undoubtedly are, e.g.
the families which constitute the Malvales. On the other hand, this is not
true of others. For example, it is doubtful if the Hydrangeaceae (family 124)
have any very close affinities with the Crassulaceae (family 125). In those
families where the anatomical facts give some evidence of the true affinities
of the families concerned, this has been indicated in the 'Taxonomic Notes'
included in many of the family descriptions throughout the book. Until the
taxonomic and phylogenetic interrelationships of the families are more fully
understood, no arrangement can be entirely satisfactory. It is hoped, however,
that the sequence of families that has been adopted will, with the aid of the
index, facilitate easy reference, and at the same time avoid injuring the feelings
of those taxonomists who have strong predilections for any other system.

I. POLYPETALAE
A. THALAMIFLORAE
RANALES i. Ranunculaceae
2. Dilleniaceae
3. Crossosomataceae
4. Calycanthaceae
5. Magnoliaceae
6. Schisandraceae
7. Winteraceae
8. Cercidiphyllaceae
9. Eucommiaceae
10. Eupteleaceae
11. Himantandraceae
12. Lactoridaceae
13. Trochodendraceae
14. Tetracentraceae
15. Annonaceae
1 6. Eupomatiaceae
17. Menispermaceae
 INTRODUCTION Ivii

RANALES (cont.) 1 8. Berberidaceae

19. Circaeasteraceae
20* Lardizabalaceae
21. Sargentodoxaceae
x
22. Nymphaeaceae

PARIETALES 23. Sarraceniaceae

24. Papaveraceae
25. Fumariaceae
26. Cruciferae
27. Capparidaceae
28. Resedaceae
29. Cistaceae
30. Violaceae
31. Canellaceae
32. Bixaceae
33. Cochlospermaceae
34. Flacourtiaceae (including Samydaceae)

POLYGALALINAE 35. Pittosporaceae

36. Tremandraceae
37. Polygalaceae
38. Diclidantheraceae
39. Vochysiaceae
40. Trigoniaceae

CARYOPHYLLINAE 41. Frankeniaceae

42. Caryophyllaceae (including Illecebraceae)
43. Portulacaceae
44. Tamaricaceae
45. Fouquieraceae

GUTTIFERALES 46. Elatinaceae

47. Hypericaceae
48. Guttiferae
49. Quiinaceae
50. Theaceae (including the anomalous genera Clematoclethera^
Sladenia, and Trematanthera)
51. Actinidiaceae
52* Bonnetiaceae
53. Caryocaraceae
54. Marcgraviaceae
55. Medusagynaceae
56. Pellicieraceae
57. Pentaphylacaceae
58. Saurauiaceae
59. Stachyuraceae
60. Dipterocarpaceae
61. Ancistrocladaceae
62. Chlaenaceae

MALVALES 63. Malvaceae

64. Bombacaceae
65. Sterculiaceae (including the Buettneriaceae)
66. Tiliaceae
67. Elaeocarpaceae
68. Scytopetalaceae
Iviii INTRODUCTION
B. DlSCIFLORAE
GERANIALES 69. L,inaceae
70. Erythroxylaceae
71. Humiriaceae
72. Malpighiaceae
73. Zygophyllaceae
74. Geraniaceae
75. Balsaminaceae
76. Limnanthaceae
77. Oxalidaceae
78. Tropaeolaceae
79. Rutaceae
So. Simarubaceae
81. Brunelliaceae
82. Cneoraceae
83. Koeberliniaceae
84. Ochnaoeae (including Strasburgeria)
85. Tetrameristaceae
86. Burseraceae
87. Meliaceae
88. Dichapetalaceae (Chailletiaceae)

OLACALES 89. Olacaceae

90. Icacinaceae
91. Octoknemaceae
92. Opiliaceae
93. Aquifoliaceae (Ilicaceae)
94. Cyrillaceae

CKLASTRALES 95. Celastraceae

96. Goupiaceae
97. Hippocrateaceae
98. Stackhousiaceae
99. Rhamnaceae
100. Ampelidaceae (Vitaceae)

SAPINDALES ioi . 3 apindaceae

1 02. Aceraceae
103. Akaniaceae
104. Hippacastanaceae
105. Melianthaceae
1 06. Staphyleaceae
107. Didiereaceae
1 08. Sabiaceae
109. Anacardiaceae
no. Corynocarpaceae
in. Julianiaceae
112. Coriariaceae
113. Moringaceae

C. CALYCIFLORAE
ROSALES 114. Connaraceae
115. Leguminosae
115 A, Mimosaceae
1158. Caesalpinia
 INTRODUCTION lix

ROSALES (cant.) 1150. Papilionaceae

1 1 6. Krameriaceae
117. Rosaceae
H7A. Rosaceae-Chrysobalanoideae
Saxifragaceae
119. Cephalotaceae
1 20. Cunoniaceae
121. Escalloniaceae
122. Eucryphiaceae
123. Grossulariaceae
124. Hydrangeaceae
125- Crassulaceae
126. Droseraceae
127. Byblidaceae (Roridulaceae)
128. Hamamelidaceae
129. Myrothamnaceae
130. Bruniaceae
131. Haloragaceae
132. Callitrichaceae
133. Hippuridaceae

MYRTALES 134- Rhizophoraceae

135- Combretaceae
136. Myrtaceae
137. Lecythidaceae
138. Melastomaceae
139- Lythraceae
140. Crypteroniaceae
141. Oliniaceae
142. Punicaceae
143. Sonneratiaceae
144- Onagraceae (Oenotheraceae)
(Samydaceae incorporated in Flacourtiaceae)

PASSIFLORALES 145- Loasaceae

146. Turneraceae
147- Passifloraceae
148. Achariaceae
149. Caricaceae
150. Malesherbiaceae
151. Cucurbitaceae
152. Begoniaceae
153- Datiscaceae

FICOIDALES 154- Cactaceae

155- Ficoidaceae (Aizoaceae) (including Molluginaceae)

UMBELLALES 156. Umbelliferae

157- Araliaceae
158. Cornaceae
159. Alangiaceae
1 60. Garryaceae

161. Nyssaceae

II. GAMOPETALAE
A. INFERAE
RUBIALES 162. Caprifoliaceae
163. Adoxaceae
164. Rubiaceae
Ix INTROD UCTION
ASTERALES 165. Valerianaceae
1 66. Dipsacaceae
167. CaJyceraceae
1 68. Compositae

CAMPANULALES 169 . Sty lidiaceae

170. Goodeniaceae (including Brunoniaceae)
171. Campanulaceae
172. Lobeliaceae

B. HETEROMERAE
ERICALES 173. Vacciniaceae
174. Ericaceae
175. Clethraceae
176. Monotropaceae
177. Epacridaceae
178. E> iapensiaceae
179. Lennoaceae

PRIMULALES i So. Plumbaginaceae

181. Primulaceae
182. Myrsinaceae (including Theophrastaceae)

EBENALES 183. Sapotaceae

184. Ebenaceae
185. Styracaceae
1 86. Symplocaceae

C. BlCARPELLATAE
GENTIANALES 187. Oleaceae
1 88. Salvadoraceae
189. Apocynaceae
190. Asclepiadaceae
191. Loganiaceae
192. Gentianaceae

POLEMONIALES 193. Polernoniaceae

194. Hydrophyllaceae
195. Boraginaceae
196. Convolvulaceae (including Nolanaceae)
197. Solanaceae

PERSONALES 198. Scrophulariaceae

199. Orobanchaceae
200. Lentibulariaceae
201. Columelliaceae
202. Gesneriaceae
203. Bignoniaceae
204. Pedaliaceae (including Martyniaceae)
205. Acanthaceae

LAMIALES 206. Myoporaceae

207. Selaginaceae (including Globulaiiaceae)
208. Verbenaceae (including Phrymaceae)
209. Labiatae
 INTRODUCTION bd
III. MONOCHLAMYDEAE OR INCOMPLETAE
A. CURVEMBRYAE
210* Plantaginaceae
(Illecebraceae included in Caryophyllaceae)
211. Nyctaginaceae
212. Amaranthaceae
213. Chenopodiaceae
214. Basellaceae
215. Phytolaccaceae (including Achatocarpaceae and Gyrostemonaceae)
216. Batidaceae
217. Polygonaceae

B. MULTIOVULATAE AQUATICAE
2 1 8. Podostemaceae
219. Hydrostachyaceae

C. MULTIOVULATAE TERRESTRES
220, Nepenthaceae
221. Cytinaceae (RafResiaceae)
222. Hydnoraceae
223. Aristolochiaceae

D. MlCEMBRYAE
224, Piperaceae
225. S a uru raceae
226. Chloranthaceae
227. Myristicaceae
228. Monimiaceae

E. DAPHNALES
229. Lauraceae
230. Gomortegaceae
231. Hernandiaceae
232. Proteaceae
233. Thymelaeaceae
234. Gonystylaceae
235. Penaeaceae
236. Geissolomataceae
237. Elaeagnaceae

F. ACHLAMYDOSPOREAE
238. Loranthaceae
239. Santalaceae
.
240. Grubbiaceae
241. Myzodendraceae
242. Balanophoraceae
Ixii INTRODUCTION
G, UNISEXUALES
243. Euphorbiaceae
244. Buxaceae
245. Daphniphyllaceae
246. Balanopsidaceae
247. Urticaceae
248. Cannabinaceae
249. Cynocrambaceae
250. Moraceae
*

251. Ulmaceae
252. Platanaceae
253. Leitneriaceae
254. Juglandaceae
255. Myricaceae
256. Casuarinaceae
(Julianiaceae placed after Anacardiaceae)
257. Betulaceae
258. Corylaceae
259. Fagaceae

H. ANOMALOUS FAMILIES
260. Salicaceae
261. Lacistemaceae
262. Empetraceae
263. Ceratophyllaceae

THE DESCRIPTIONS OF THE FAMILIES

THE descriptions of the individual families have, for the most part, been
arranged according to the following plan. The system has been slightly
modified for some of the families.

SUMMARY
(i)
GENERAL \ A summary of the information given more fully under 'Leaf
(ii)
WOOD / and 'Axis'.

LEAF
Axis
(i) STEM
For woody plants this refers to stems of the first few seasons' growth or such
as are to be found on herbarium sheets. For herbaceous plants, stems of all
ages are dealt with in this section.

(ii)
WOOD
This xylem of mature stems large enough
refers typically to the secondary
to produce timber, but the wood of small trees, shrubs, and lianes has also
been described here if including the equivalent of several growth rings. The
immature xylem, such as occurs in the first few seasons' growth, has been
described separately under the section headed 'STEM*. For a complete picture
of the secondary xylem, therefore, it is usually necessary to consult both of
these sections.
 INTRODUCTION kiii

(iii)
ROOT
It has often been necessary to omit this section owing to the lack of informa-
tion concerning the root structure of many families.

(iv) ANOMALOUS STRUCTURE

This section usually includes all the information about anomalous struc-
ture, but, where the anomaly known as included or interxylary phloem is
characteristic of the secondary xylem of woody stems, this feature has
generally been dealt with separately in the description of the wood, with a
cross-reference in this section.

TAXONOMIC NOTES
The main object of this book has been to make anatomical data of taxonomic
interest generally available, and no attempt has been made to provide solutions
for all of the many taxonomic problems which have come to our notice. It
was, however, felt that it would be useful to include in this section some
notes on taxonomic conclusions drawn from anatomy by various investigators
and recorded in the literature. This has been done, in most instances, without
any additional comment from the authors. The authors have, however,
expressed opinions of their own wherever original observations have made
this appear desirable.

ECONOMIC USES
Since systematic anatomy can be and is generally applied in the identifica-
tion of economic products, it seemed desirable to indicate the nature of the
chief economic products derived from each family. It is hoped that the
information included here, though necessarily far from complete, will prove
useful to all who are concerned with the use of timber, as well as to students
of economic botany, public analysts, pharmacognoscists, and all who are con-
cerned with the identification of fragmentary material of vegetable origin.
For the wood, no attempt has been made to do more than show whether or not
the family is important as a source of timber and indicate the most important
species and genera.

GENERA DESCRIBED
At the end of most of the family descriptions there are two lists of the
genera that have been included, one for the stem, root, and leaf, the other for
the wood. Where the family is mainly or entirely herbaceous the second list
has been omitted. In the first list an asterisk has been used to indicate that
the genus is represented in the reference collection of microscope slides at
Kew. In the list for the wood, brackets round a name indicate that no material
of this genus was examined by the authors, but that information concerning
at least some characters was available in the literature.
The inclusion of a genus in these lists does not necessarily mean that either
the information or material available was fully representative, particularly in
the case of genera with a large number of species, or that information was
available all of the characters included in the family description. The lists,
on
however, should provide the reader with some indication of the extent to
Ixiv INTRODUCTION
which any particular family has been covered, and it is hoped that further
research may be stimulated on those groups about which our knowledge is
still rather meagre.

LITERATURE
Two separate lists are given at the end of most of the family descriptions,
one for general anatomy and one for wood. Where the family is mainly or
entirely herbaceous there is only one list. The numbers refer to the biblio-
graphy at the end of the book.
 DESCRIPTIONS OF THE FAMILIES
RANUNCULACEAE
1.
(FiG. 4 on p. 2 ; FIG. 7 on p. 14)

SUMMARY
(i) GENERAL
Mainly herbs, but including aquatic plants and some woody climbers. The
family occurs mainly in the Northern Hemisphere, but extends into the
tropics. The vascular bundles, in transverse sections, appear widely spaced,
collateral, typically with the xylem concave on the side towards the phloem,
so that the latter is often partly surrounded by xylem. Caps or arcs of fibres
are present immediately on the outside of the phloem. ring of pericyclic A
sclerenchyma is frequent in some genera, and in certain species the fibre
groups adjoining the phloem are bounded externally by, or partly embedded
in, the sclerenchymatous ring of the pericycle. In some genera, notably the
Japanese species of Ranunculus, the vascular strands are entirely surrounded
by sclerenchyma. Xylem vessels of herbaceous species have simple per-
forations, except in Paeonia, where scalariform plates have been recorded.
Medullary bundles are usual in certain genera. Petiole, in transverse sec-
tions through the distal end, variable in outline, but always supplied by widely
spaced vascular bundles arranged in an arc or circle or sometimes scattered.
Arm-palisade cells are common in the leaf.

(ii) WOOD
Clematis
Vessels in tangential or irregular groups, ring-porous, with spiral thicken-
ing, perforation plates simple, intervascular pitting alternate, members of
medium length to moderately short. Parenchyma paratracheal, storied.
Rays all large, up to 12 or more cells wide and very high. Fibres with simple
or bordered pits, storied, extremely short.

Paeonia
Vessels mostly solitary, ring-porous, perforation plates scalariform, inter-
vascular pitting transitional and opposite, members of medium length. Par-
enchyma diffuse, very sparse. Rays up to 3 cells wide, with numerous
uniseriates. Fibres with distinctly bordered pits, very short.

LEAF
Generally dorsiventral, rarely centric.Hairs including glandular and non-
glandular types. Non-glandular trichomes mostly unicellular, of varied
length. Simple, unicellular hairs recorded in Thalictrum. Small, unicellular,
thin-walled, glandular hairs occur in Adonis, Anemone, Aquilegia, Caltha,
Clematis, Delphinium, Eranthis, Helleborus, Nigella, Ranunculus, Trollius;
glandular hairs with rather longer stalks observed by Solereder in Helleborus
and Ranunculus. Cells of the lower epidermis with sinuous anticlinal walls;
those of the upper epidermis generally straight* Stomata ranunculaceous;
4504 B
 FIG. 4. RANUNCULACEAE
A, Nigella integrifolia Regel. Petiole X 19. B, Thalictrum 'rugosum*. Petiole x 8. C, Adonis vemalis
Linn. Petiole x 28. D, Clematis afoliata Buch. Stem x 10. E, Ranunculus acris Linn. Stem x 10.
F, Anemone vitifolia Buch-Ham. Stem X 10. G, Aconitum 'gracile'. Petiole X 19. H, Zanthorixa
apiifolia L'H^rit. Stem X i^. I, Helleborus 'corsicus*. Petiole X 8. J, Anemone vitifolia Buch-Ham.
Petiole X 8. K, Clematis vttalba Linn. Petiole X 19. L, Aquilegia canadensis Linn. Petiole X 19,
M, Aconitum napellus Linn. Stem x 10. N, Ranunculus acris Linn. Petiole X 19.
 RANUNCULACEAE 3

confined to the lower surface in some species, but present on both in others.
(The list of genera in which these two types of stomatal distribution occur as

given by Solereder probably needs revision and amplification according to

Maue (1460), but the taxonomic value of the variations in this character
appears to be somewhat limited.) Stomata ofHelleborus niger Linn, described
by Miihldorf (1568) as slightly below the general level of the epidermis, the
guard cells being provided with cuticular teeth which become interlocked
across the pores on closure. Hydathodes sometimes occur in association
with the crenate teeth at the margins of the leaves. Petiole (Fig. 4 A-C, G,
I-L, and N), in transverse sections through the distal end, exhibiting a con-
siderable range of variations in outline, but nearly always provided with a
number of widely spaced vascular bundles which may or may not be accom-
panied by pericyclic fibres according to the species. Petiolar vascular bundles
arranged in a simple arc, e.g. in species of Adonis, Helleborus, Nigella; forming
an almost or quite complete, sometimes adaxially flattened circle, e.g. in
species of Aconitum, Aquilegia, Clematis, and Ranunculus including a propor-
;

tion of medullary petiolar bundles, e.g. in species of Aquilegia and Thalictrum.

The petiolar structure might well afford characters of considerable diagnostic
value if more completely investigated. Petiolar bundles of Clematis flammula
Linn, not separate. (A study of 34 species and races of Pulsatilla by Zamels
and Paegle (2502) showed the following leaf characters to be of taxonomic value.
The presence or absence of a large ventral bundle in the petiole, and of a
parenchymatous sheath around the individual bundles. Individual races and
species are distinguishable by the amount and distribution of mechanical
tissue and the number of bundles. On the basis of the first of these characters
the genus may be divided into 4 groups.)

Axis
STEM (Fig. 4 D-F, H, and M)
General topography of the aerial stem as indicated above under 'Summary*.
Cork rarely formed, but known to be deep-seated in origin in a few woody
species. Vascular bundles, in most genera and species, provided with
xylem strands which are concave where adjoining the phloem, but outer
boundary of the xylem flattened in Paeonia. Vessels of herbaceous species
usually with simple perforations, but scalariform plates also occur in Paeonia.
The primary vascular bundles usually remain widely spaced as seen in trans-
verse sections, but the xylem of adjacent bundles becomes connected by
fibres in the mature stem of species of Paeonia. Interfascicular cambium
often absent from the woody genera Clematis and Naravelia, the bundles
remaining separated by the primary rays, although secondary rays arise
within the bundles. Secondary bundles originate from an interfascicular
cambium in other Species of Clematis. Additional rigidity is secured by
thickening of the walls of the pith and cortex.
Vascular bundles frequently to be seen in transverse sections arranged in
several circles, or more or less irregularly scattered; several circles present in
Actaea, Cimicifuga, and Thalictrum; medullary bundles present in certain
species of Anemone, Anemonopsis, Delphinium, Glaucidium, and 'Ranunculus
4 RANUNCULACEAE
chinensis*. A circle of medullary bundles extends from the base to the floral

receptacle in the erect stem of Glaucidium (cf. Berberidaceae). The course of

the medullary strands has been specially investigated by Kumazawa (1299)
who recognized the following distinct arrangements, (i) Large leaf-trace
strands becoming medullary at the nodes and gradually extending to the
periphery; perianth strands not entering the pith. This type observed in
species of Anemonopsis, Delphinium, Thalictrum, and in Cimidfuga foetida
Linn, and 'Ranunculus chinensis*. (ii) As (i) but perianth strands also entering
the pith (Anemone japonica Sieb, & Zucc., A. vitifolia Buch-Ham., and
Glaucidium). (iii) Leaf strands remaining at the periphery of the stem, but
cauline bundles, and bundles from the lateral branches tending to be medul-
9
'

lary in Cimidfuga japonica The taxonomic value of vascular arrangements

.

i-iii is unknown.
Pith generally parenchymatous, often with abundant intercellular spaces;
becoming hollow in many herbs.
Clematis afoliata Buch, with reduced leaves, is provided with alternate
bands of fibres and chlorenchyma in the cortex (Fig, 4 D).
Smith's (2146) examination of the stem anatomy in 138 species of
Clematis has shown the general constructional scheme to be remarkably
constant. Purely cauline bundles are absent throughout the genus, a
starch sheath is generally present, and the periderm intrastelar in origin.
Smith (2145) also found that vegetative propagation of Clematis by cuttings
is facilitated by previous etiolation, because this causes mechanical softening
of the tissues and restores the carbon-nitrogen ratio to the level that is neces-
sary for meristematic activity. Fibres reduced in aquatic species of Ranun-
culus; schizogenous air passages being present on the inner side of the
protoxylem. The nature of the interfascicular tissue and pericyclic scleren-
chyma may be of specific diagnostic value. This has recently been confirmed
for the species of Ranunculus which occur in Switzerland in a very detailed
investigation by Bergman (185), who found that the amount and distri-
bution of sclerenchyma in the pericycle and of fibre sheaths on the inner side
of the vascular bundles afford valuable characters for the identification of
species. Sheaths of sclerenchyma entirely surround the bundles in Japanese
species of Ranunculus.

WOOD
Clematis (Fig. 7 A)
Vessels of the pore-zone large, those of the late wood small and in tan-
gential or irregular groups ring- porous with spiral thickening (spiral thicken-
; ;

ing reported also (1851) in Naravelia). Perforations simple, horizontal in the

larger vessels; MacDuffie (1408) describes some rare, irregular, multiperforate
plates imperfect vessels common. Intervascular pitting moderate-sized and
;

alternate; pits to ray cells similar. Storied. Mean member length 0*25-
0*5 mm. Parenchyma paratracheal, associated with the vessel groups.
Fusiform strands common, strands otherwise of 2 cells. Storied. Rays all
multiseriate, commonly up to 12 cells wide and sometimes many more; very
high (more than 2 cm.); 1-2 per mm.; heterogeneous and with some sheath
cells. Fibres with numerous, small, simple or bordered pits in both radial
and tangential walls walls moderately thin storied mean length O'37-O'5 mm.
; ; ;
 RANUNCULACEAE 5
Paeonia
Vessels small and mostly solitary; ring-porous or semi-ring-porous; per-
foration plates scalariform, usually with 2-5 rather thick bars. Intervascular
pitting transitional to opposite, pits to ray cells similar. Mean member length
about 0*4 mm. Parenchyma limited to rare cells among the fibres. Rays
up to 3 cells wide and less than i mm.
high; with numerous uniseriates;
about 12 rays per mm. composed entirely of square to upright cells. Fibres
;

with numerous, conspicuously bordered pits in both radial and tangential

walls. Walls thick. Mean length about 0-55 mm.

ROOTS AND RHIZOMES

Fleshy and fibrous roots occur in Ranunculus, cambium being scanty in or
absent from* the thin ones. Tuberous roots present in some species, e.g.
Aconitum and Ficaria verna Huds. Ground tissue of the tubers chiefly con-
sisting of the amyliferous parenchyma of the cortex and sometimes the pith.
According to Holm (1015, 1028), in Aconitum uncinatum Linn, and Cimicifuga
americana Michx. the tuberous roots become thickened owing to the develop-
ment of secondary bark.
In the tuberous roots of certain Indian species of Aconitum there are,
according to Solereder, concentric vascular strands in the parenchymatous
ground tissue of the central cylinder, each strand being provided with a pith,
and growing in thickness by means of a partial cambium. These abnormal
vascular strands arise from circular, secondary meristems situated in the
pith, where they produce parenchyma and phloem towards the inside, and
parenchyma and sometimes a few vessels on the outside. The secondary
cambia subsequently become connected with the normal cambium. The
rhizome of the xerophytic Delphinium scaposum Greene splits up into separate
members during development. The accepted methods for distinguishing
the rhizomes of Helleborus niger Linn, from those of H. viridis Linn, in use
before 1924 are stated by Wallis and Saunders (2352) to be unreliable, but
no alternative distinctive characters recorded. Pith of the mature root of
Ranunculus bullatus Linn, described by Nicolas (1591) as becoming scleren-
chymatous; the limited number of secondary xylem vessels arising by
differentiation of cells of the conjunctive parenchyma. Contractile exoder-
mis recorded in the roots of cultivated forms of Anemone hepatica Linn.
Berberin present in the rhizome of Coptis trifolia Salisb. Fungal hyphae
stated by Holm (1062) to be normally present in the roots of Ranunculus
bulbosus Linn.

TAXONOMIC NOTES
The most interesting features of the herbaceous Ranunculaceae include the
typical closed bundles with V-shaped xylem, and the frequent occurrence of
several rings of or scattered bundles. These features suggest that the Ranun-
culaceae have close affinities with the Monocotyledons, especially the Alisma-
taceae. In this connexion the monocotylous condition of Ficaria verna Huds.
is significant. These and other facts indicate that the division of the angio-
sperms into Dicotyledons and Monocotyledons is not so absolute as has at
times been supposed or implied. The vascular system of the Ranunculaceae
6 RANUNCULACEAE
alsoshows affinities with that of some of the Berberidaceae. It has, in fact,
been suggested on anatomical grounds by Kumazawa (1297) that certain
genera of Berberidaceae and Ranunculaceae should be removed from their
respective families and merged into a single family, the Podophyllaceae.
Worsdell (2469) has also drawn attention to the supposed anatomical
affinities between Paeonia and the Magnoliaceae.
WorsdelFs views were, however, based on rather slender evidence, and it is
interesting to note that the resemblance between the secondary xylem of
Paeonia and that of the Magnoliaceae is not very close. The most similar
features are the scalariform perforation plates, but these indicate a similar
level of specialization rather than affinity. In connexion with the low phylo-

genetic status which is commonly assigned to the Ranunculaceae, it is of

interest that the wood of Clematis is of a moderately highly specialized type.
It is also noteworthy that the wood of Clematis is not very similar to that of
Paeonia.

ECONOMIC USES
The family includes well-known ornamental plants such as species of
Aconitum, Anemone, Clematis, Delphinium, Paeonia, and Ranunculus. The
Christmas Rose is Helleborus niger Linn.
Various members of the family are or have been used in medicine. One of
the best known is Aconite Root, consisting of the dried tuberous roots of
Aconitum napellus Linn. The conical roots are 4-10 cm. long when dried, and
bear fibrous lateral roots or scars from which they have become detached.
The pith appears stellate in transverse sections, and is bounded externally by
a line of cambium which is more conspicuous than the xylem and phloem,
the latter extending outwards to the endodermis, the cells of which have
casparian thickenings. Other microscopical characters of diagnostic value
include irregular, brown, tabular, suberized cells derived from the cortex
which form a protective external layer; thick- walled, pitted sclereids in the
cortex; 2-5 compound starch grains, the individual components being up to
30 (mostly 12-15) p in diameter; absence of crystals and fibres. The roots of
various other species of Aconitum are sometimes met with as substitutes or
adulterants for those of A. napellus.
Cimicifuga or Black Snakeroot, also used in medicine, consists of the dried
rhizomes and roots of Cimicifuga racemosa Nutt. obtained from North America.
The horizontal rhizome, about 15 cm. long and 1-2 cm. in diameter, bearing,
on the upper side, the bases or scars of stems and leaves, and brittle, often
broken roots attached to the lower surface, the positions of detached roots
being marked by scars. Microscopical features of the rhizome, visible in
transverse sections, include numerous, narrow, triangular xylem groups
separated by broad medullary rays a cambium ring between the xylem and
;

the thin, dark-coloured, horny bark; a dark-coloured pith; starch in all

tissues except the xylem. In the root the 3-6 triangular xylem groups are
united by their apices.
Black Hellebore, which was at one time used in medicine, consists of
tortuous pieces of the rhizome of Helleborus niger Linn., characterized by
cup-shaped scars of aerial stems, and root scars and roots 1-2 mm. thick
attached to the lower surface. Transverse sections of the rhizome show a
 RANUNCULACEAE 7
wide cortex, surrounding a circle of 3-10 triangular xylem groups separated
by wide medullary rays, the centre of the rhizome being occupied by a small
brown pith. In each root there is a pale-yellow stele.
Seeds of Nigella and Paeonia, &c., have also been used in medicine.
The anatomy of the mildly poisonous Ranunculus thora Linn, has been
described in some detail by Goris (797).

GEN.ERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aconitum,* Actaea,* Adonis,* Anemone,* Anemonella, Anemonopsis,
Aquilegia,* Cirnicifuga, Clematis,* Coptis, Delphinium,* Glaucidium, Helle-
borus,* Hepatica, Isopyrum, Myosurus,* Naravelia, Nigella,* Paeonia,*
Pulsatilla, Ranunculus,* Thalictrum,* Trollius,* Zanthoriza.*
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Clematis, (Naravelia), Paeonia.

LITERATURE
(i) On General Anatomy
Betts 187, Bird 199, Bretin 268, Crockart 5025, Giersch 764, Goris 797,
Bergman 185,
Holm 996, 997, 1015, 1029, 1050, 1062, 1068, Kingsley 1237, Kumazawa 1297, 1299,
1300, Maue 1460, Munch and Crosbye 1574, Muhldorf 1568, Nicolas 1591, Rabbas 1769,
Salisbury 1984, Smith, E. P. 2145, 2146, Wallis and Saunders 2352, Worsdell 2469,
Zamels and Paegle 2502.

(ii) On Wood Structure

Dadswell and Record 533, MacDuffie 1408, Pfeiffer, H. 1712, Record 1843, 1851.

2. DILLENIACEAE
(Fic. 5 on p. 8 ;
FIG. 6 on p. 12; FIG. 8 on p. 20)

SUMMARY
(i) GENERAL
A family of trees, shrubs, climbers, or, more rarely, herbs, which occurs in
tropical and subtropical regions. The hairs are simple, unicellular trichomes
or stellate; glandular types absent. The stomata are usually ranunculaceous,
except in Tetracera. Crystals of calcium oxalate occur chiefly in the form of
raphides, but transitional forms between these and crystal-sand have been
recorded in Curatella, Davilla, DiUenia, and Doliocarpus. Canals containing
crystal-sand have also been observed in certain members of the family, and
tetrahedral crystals are present in the phloem of the leaf veins and axis as
well as in the mesophyll of Hibbertia scandens (Willd.) Gilg. These are absent
from other members of the genus.

(ii)
WOOD
solitary, perforation plates scalariform, or simple and scalariform,
Vessels
intervascular pitting opposite to scalariform, members moderately to extremely
long. Parenchyma predominantly apotracheal, diffuse or in uniseriate lines,
with a few cells about the vessels; sometimes containing raphides. Rays
 8 DILLENIACEAE
8-10 wide or more and high, heterogeneous, sometimes containing
cells
Fibres with distinctly bordered pits, moderately to very long.
raphides.
Anomalous structure of the banded type present in Doliocarpus.
LEAF
Mostly dorsiventral with 1-4 layers of palisade tissue, but centric or some-
times ericoid, with furrows on the lower side, in certain members of the

FIG. 5. DILLENIACEAE
A, of Hibbertia furfumcea (R. Br.) Benth. B-C, spiny hair of Tetracera oblongata DC.:
stellate hair
B, transverse section through the spiny hair and through a piece of leaf-tissue. C, surface view of the
spiny hair with the surrounding cells of the upper side of the leaf. D~E, peltate hair of Hibbertia
lepidota R. Br.: D, in surface view; E, in section. By Solereder.

Hibbertieae from dry habitats in Australia. Expansion of the leaf is con-

trolled by specialized motor tissue at the margin in Hibbertia rhadinopoda
F. v. Muell. The leaves of other species of Hibbertia from more humid
regions exhibit typical dorsiventral structure. Hairs (Fig, 5 A-E) in the form
of simple, unicellular trichomes, or stellate; the stellate hairs and scales most
numerous in the xeromorphic Hibbertieae. Peltate scales also recorded in
Hibbertia lepidota R. Br. Epidermis often silicified. Stomata usually
ranunculaceous, but rubiaceous in Tetracera oblQnga DC.; generally level
with the surface of the leaf, although sunken in xeromorphic Hibbertieae,
especially in the phylloclades of Pachynema. Hypoderm recorded in certain
species of Wormia and in Dillenia indica Linn. Sclerenchymatous idioblasts
said to be present in Hibbertia grossulariaefolia Salisb. and H. linearis R. Br.
Midrib containing a closed ring of vascular bundles in Curatella, Davilla>
 DILLENIACEAE 9

Doliocarpus, and Tetracera, but with 2-7 or more concentric bundles in

Acrotrema and Dillenia or a circle of collateral bundles in the midrib of
Schumacheria. Petiole of Dillenia indica (Fig. 6 A), in transverse sections
through the distal end, exhibiting a closed, cylindrical vascular strand accom-
panied by subsidiary bundles in the wings. A wide crescent-shaped strand
with a smaller dorsal one between the arms present in the corresponding
position in Hibbertia volubilis Andr. (Fig. 6 c). All leaf tissues turn brown
when dried owing to the presence of tanniniferous substances. Crystals.
Raphides situated in elongated, often thick-walled, sacs or tubes, present in
the mesophyll and in the phloem of the veins. Similar sacs or tubes sometimes
filled with crystal-sand, or types of crystal intermediate between raphides and

crystal-sand.

Axis
YOUNG STEM (Fig. 6 B)
Cork deep-seated in origin in species of Davilla, Doliocarpus, Hibbertia,
and Tetracera; sub-epidermal in Dillenia. Cork cells of Dillenia pentagyna
Roxb. with thin walls, but in other species and genera usually with thicker
walls. Pericycle generally including a continuous ring of sclerenchyma
except in Acrotrema, species of Hibbertia and Tetracera scandens (Forst.)
Gilg et Werd. At first continuous, but becoming broken in Dillenia indica
Linn. Xylem traversed by broad rays in Dillenia indica and by less frequent
broad rays in Hibbertia volubilis Andr. containing vessels of very variable
;

diameter depending on the habit of the plant. Vessels mostly with scalari-
form perforation plates, although these sometimes become simple owing to
the destruction of the bars, notably in Davilla and Tetracera (see also Wood
below). Pith generally containing elongated, sclerenchymatous idioblasts
with white inclusions in Davilla, Doliocarpus, and Tetracera generally septate
;

in Wormia\ including groups of stone cells in Curatella, Davilla, Dillenia,

Doliocarpus, and Tetracera. Raphides, crystal-sand, and transitional forms
of crystals present in elongated sacs or canals in all unlignified parts of the
stem, notably in the cortex, phloem, and pith.

WOOD (Fig. 8 i, j, L)
Vessels usually medium-sized (mean tangential diameter 100-200 ^),
large (more than 200 p) in the lianes, e.g. in Davilla and Doliocarpus\
exclusively solitary, except for apparent tangential pairs due to overlapping
ends; usually i~8 per sq. mm., but, according to Vestal (2329), sometimes
up to 50 per sq. mm. Perforation plates typically scalariform, with few to
numerous bars (up to 130), though sometimes accompanied by simple per-
forations; perforations mostly simple in Davilla, Doliocarpus, and Tetracera
(2329), but with some scalariform plates in the smaller vessels. Inter-
vascular pitting rare- except in the overlapping end- walls, opposite to scalari-
form. Often filled with yellow to white chalky contents; tyloses reported
(2329) in Dillenia] Gonggrijp (794) has observed silica in Dillenia and Wormia.
Mean member length O'7-2*o mm. Parenchyma predominantly apotracheal,
as scattered cells (diffuse) or short, uniseriate tangential lines (Fig. 8 L), the
linesmore continuous in Curatella; often with a tendency to form incomplete
io DILLENIACEAE
sheaths round the vessels (Fig. 8 j) cells often filled with gum-like substance.
;

Enlarged cells containing raphides present in some species of Dillenia and in

the conjunctive tissue of Doliocarpus. Strands usually of 8 cells. Rays usually
of two distinct sizes, the larger commonly up to 8~io cells wide and up to
20 or more cells wide in Curatella and Davilla, but all narrow, according to
Tupper, in Acrotrema, Hibbertia, and Pachynema\ commonly more than
2 mm. high; uniseriates numerous and composed of upright cells; 4-12 rays
per mm. heterogeneous (Kribs's Types I and II A), with 4-10 (occasionally
;

more) marginal rows of square to upright cells with square and procumbent
;

cells intermingled in Curatella and Davilla, and, to a less extent, in Wormia.

Raphides present in enlarged mucilage cells (Fig. 8 i) in Curatella^ Dillenia

(lacking from D. aurea Smith (1154)), Tetracera (959), and Wormia. Scleroid
cells reported (2329) in Davilla. Fibres with distinctly bordered pits on both
radial and tangential walls, the pits very large in Doliocarpus. Walls thin
to moderately thick. Mean length 1-8-3 mm
Included (interxylary)
-

phloem of the concentric type (C. L circumvallatum) present in Doliocarpus,

the short-lived cambium being periodically replaced by new meristematic
tissue, which repeats the structure of the young stem.

ANOMALOUS STRUCTURE, see under 'WOOD'

TAXONOMIC NOTES
(i)
FROM GENERAL ANATOMY
Actmidia, Clematoclethra, Saurauia, Sladenia, Trematanthera resemble both
the Dilleniaceae and the Theaceae. This is of interest because, as Hutchinson
(1113) has pointed out, there are affinities between the Dilleniales and Theales.
For this reason it has been difficult to decide on the most appropriate position
for the inclusion of the above genera in this book. Since Actinidiaceae and
Saurauiaceae are commonly recognized as distinct families, the single genus
included in each has been described under these family names. Clematoclethra,
Sladenia, and Trematanthera, whose position is less well established, have
been provisionally described as anomalous genera under Theaceae. It is
interesting to note that Beauvisage (163) expressed the view that the anatomical
characters in general, and in particular those of the petiole, combined with
the presence of raphides, suggest that the affinities of all of the above genera
lie closer to the Dilleniaceae than to the Theaceae.

(ii) FROM WOOD STRUCTURE

Vestal (2329) notes that there is little difference in anatomy between mem-
bers of this family and those of the Actinidiaceae, Saurauiaceae, and Theaceae.
Bausch (154) found that the anatomy of the wood shows no very close relation
to that of the Eucryphiaceae.

ECONOMIC USES
The silicified cells on the surface enable the leaves of Tetracera to be used
in the same way as sand-paper for polishing wood.
Some
of the species of Dillenia provide timbers that are used locally for
general construction and boat-building, and occasionally for furniture.
 DILLENIACEAE n
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acrotrema, Curatella, Davilla, Dillenia,* Doliocarpus, Hibbertia,* Pachy-
nema, Schumacheria, Tetracera, Wormia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Acrotrema), Curatella, Davilla, Dillenia, Doliocarpus, (Hibbertia),
(Pachynema), (Schumacheria), (Tetracera), Wormia.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gilg and Werdemann 781, Hutchinson 1113, Kastory and Namylowsky
1223, Lechner 1328, Shelton 2086, Solereder 2165, Thompson 2254.
(ii) On Wood Structure
Bausch 154, den Berger 179, 182, Chalk and Chattaway 362, Dadswell and Record 533,
Desch 574, Foxworthy 704, Gonggrijp 794, Hess 959, Howard 1088, Janssonius 1154,
Kanehira 1206, Lechner 1328, Lecomte 1334, Pearson and Brown 1679, Pfeiffer, H. 1712,
Record 1843, I 8si Record and Hess 1886, Record and Mell, 1894, Tupper 2295, Vestal
2329, Williams 2430.

3. CROSSOSOMATACEAE
(FiG. 6 on p. 12)

SUMMARY
The
sole representatives of this family are the Californian shrubs belonging
to the genus Crossosoma. The following description of the leaf and stem is
based chiefly on fresh material grown at Kew. The wood exhibits the follow-
ing features: Vessels very small, ring-porous, perforations simple, inter-
vascular pitting alternate. Parenchyma sparse, paratracheal. Rays up to
6 cells wide, heterogeneous. Fibres with large bordered pits.

LEAF
Isobilateral, thecomponent cells containing small, dark-coloured bodies of
unknown nature. Epidermis composed of cells with sinuous anticlinal walls.
Stomata present on both surfaces but most numerous on the lower side;
ranunculaceous. Petiole (Fig. 6 E), in transverse sections through the distal
end, exhibiting an open arc of separate collateral bundles, not accompanied
by fibres in the pericyclic region. Vessels of the petiolar bundles in radial
rows. No raphides, crystalliferous sacs, or tubes present, but small rosettes or
dendritic groups of minute acicular crystals present in some of the epidermal
cells on both surfaces.

Axis
YOUNG STEM (Fig. 6 H)
Cork arising at a very early stage in the sub-epidermal region. Primary
cortex parenchymatous. Pericycle containing small, very narrow strands
of thick-walled fibres. Phloem devoid of sclerenchymatous elements.
Xylem forming an almost continuous cylinder, traversed by fairly narrow
but lignified, primary medullary rays; including vessels up to about 20 p, in
12 CROSSOSOMATACEAE
diameter, arranged in radial rows, and provided with minute bordered pits
and simple perforations. Pith composed of abundantly pitted parenchyma-
tous cells; occasionally hollow. Minute, yellow, acicular crystals closely

FIG. 6. DILLENIACEAE, A-C; CALYCANTHACEAEi D, F, and G;

CROSSOSOMATACEAE, E and H
A, Dillenia indica Linn. Petiole X 8. B, Hibbertia volubilis Andr. Stem X 13. C, H. volubilis Andr.
Petiole x8. D, Chimonanthus fragrans Lindl. Petiole x 19. E, Crossosoma californicum Nutt. Petiole
X27- F, Calycanthus floridus Linn. Petiole x 19. G, C. occidentalis Hook, et Am. Stem X 19.
H, Crossosoma californicum Nutt. Stem X 27.

packed in clusters of cells in the phloem. (These crystals are entirely unlike
the raphides and crystal-sand which occur in the phloem throughout the
Dilleniaceae.)
WOOD 1

Vessels small to minute (15-55 A6 ) an ^ angular, those of the late wood

solitary, but grouped into radial rows by the rays ring-porous. Perforations
;

1
Based on the description by Record and Hess (1886).
 CROSSOSOMA TACEAE 13

simple. Intervascular pitting alternate and small, pits to ray cells similar.
Many very small vessels with 'fibriform members*. Parenchyma sparse,
paratracheal. Rays up to 6 cells wide heterogeneous, with most of the cells
;

square or upright and the procumbent cells short. Fibres with numerous,
large-bordered pits and thick walls, scarcely distinguishable from the fibri-
form vessel members.

TAXONOMIC NOTES
The absence from Crossosoma of raphides as well as of tubes or sacs filled
with raphides or crystal-sand is a clear indication that the genus has no close
affinities with the Dilleniaceae.

GENUS DESCRIBED
(Crossosoma*)
* Kew
Represented in the slide collection.

LITERATURE
On Wood Structure. Record 1843, 1851, Record and Hess 1886.

4. CALYCANTHACEAE
(Fic. 6 on p. 12 ;
FIG. 7 on p. 14)

SUMMARY
(i) GENERAL
Shrubs from North America and the Far East. The most distinctive
anatomical feature of the young stem of this family is the presence of 4 inversely
orientated vascular bundles in the pericycle (Fig. 6 G). Secretory cells are
present. Hairs unicellular; surrounded by silicified cells. The stomata are
rubiaceous. The cork in the axis arises in the sub-epidermis, and the first-
formed periderm cells are radially elongated. The bark cells are stated by
Quinlan (1767) to be pitted.

(ii) WOOD
Vessels very small, with a marked oblique or flame-like pattern, ring-
porous, with spiral thickening, perforations simple, intervascular pitting
alternate and large; members moderately short to medium-sized. Paren-
chyma limited to a few cells round the vessels. Rays up to 4 cells wide and
composed almost entirely of square and upright cells. Fibres with simple or
small-bordered pits, very to moderately short.

LEAF
Dorsiventral. Bases of the unicellular hairs surrounded by a rosette of
cells with silicified walls, having the appearance of dark or translucent dots in
dried specimens. Stomata rubiaceous; confined to the lower surface.
Mesophyll. Palisade tissue indistinct in Calycanthus floridus Linn. Petiole
of Chimonanthus fragrans LindL (Fig. 6 D), in transverse sections through the
distal end, exhibiting a single, slightly arc-shaped vascular strand; that of

Calycanthus floridus Linn. (Fig. 6 F) provided with a larger and rather more
invaginated median vascular arc, accompanied by small accessory strands in
 '
B-D; TROCHQDENDRACEAE, E-F-
,

HIMANTANDRACEAE, G-H; CERCIDIPHYLLACEAE,!-]; EUCOMMIACEAE,K-i

S B Pa/^ n teJ"M* Li""- ^y. C, Chimonanthus
' '
vrfBr V ni fragram
T S ^ Trochodendron aralioides
a
^G 'Mmantandra
V,.
- -
Sieb. et Zucc.^ S
1

- ^' belgraveana Diels. Ray. H, H. bdgra

J' c -
 CALYCANTHACEAE 15
the wings. Secretory elements. Secretory cells present in the mesophyll
or lower epidermis. Oil cells, chiefly present in the veins, but to a smaller
extent in the mesophyll in Calycanthus floridus, according to Lehmann (1343)*

Axis
YOUNG STEM (Fig. 6 G)
Four pericyclic bundles with external xylem and internal
(leaf traces),
phloem, present in Calycanthusand Chimonanthus. Pericycle including a
continuous ring of sclerenchyma in the lower nodes of Chimonanthus.
Isolated strands of fibres situated in the corresponding position in Calycanthus.
Pericyclic fibres and stone cells showing U-shaped thickenings in longitudinal
sections. Xylem, in transverse sections, appearing as a continuous cylinder
traversed by narrow rays. Pith consisting of thin-walled parenchyma. Oil
cells present in the parenchymatous portions of the axis.
(The distinction recorded by Solereder that the external bundles of Caly-
canthus consist of undivided vascular strands in the cortex, whereas the
corresponding bundles in Chimonanthus are double strands in the pericycle,
was not substantiated by direct observation. In both genera the pericyclic
bundles are bounded externally by groups of fibres which appear to be con-
tinuous with the pericyclic sclerenchyma. Nor could the 'double* nature of
the strands in Chimonanthus be confirmed.)

WOOD (Fig. 7 B-D)

Vessels very small (25-50 p, mean tangential diameter), often little wider
than the fibres and angular in cross-section; in clusters and radial pore-
multiples that are grouped to give a distinct oblique or 'flame-like* pattern,
very numerous; ring-porous, or semi-ring-porous; with spiral thickening.
Perforations simple, oblique; imperfect vessel members common. Inter-
vascular pitting alternate and large, the apertures round in Calycanthus. Pits
to ray cells usually similar to the intervascular pitting in size and shape, but
occasionally larger and irregular in shape; simple. Mean member length
about 0-3 to 0-4 mm. Parenchyma very scanty; paratracheal, consisting of
a few cells around the vessels. Garratt (747 A) refers also to diffuse parenchyma
in Calycanthus, but this was not observed by the author. Rays small and
low, except for some wider and higher rays in Chimonanthus that appear to
have been dissected out from large primary rays. 1-2 (occasionally 3) cells
wide in Calycanthus^ 1-3 (occasionally 4) cells wide in Chimonanthus com- ;

posed almost entirely of square or upright cells with few truly procumbent
cells; with about 4 uniseriate marginal rows in Calycanthus and up to 8 rows
in Chimonanthus. Perforated ray cells very common (358). Uniseriate rays
numerous, often only 1-2 cells high; of mixed square and upright cells, as in
the multiseriate rays. Fibres with small pits mostly on the radial walls, with
small borders in Calycanthus walls thin; mean length about 0*7 mm. Vasi-
\

centric tracheids with spiral thickening present in both genera.

TAXONOMIC NOTES
The Calycanthaceae in the Bentham and Hooker system were placed in
the Ranales between the Dilleniaceae and Magnoliaceae, similarities to the
Rosaceae, Combretaceae, and Monimiaceae also being mentioned. In the
16 CALYCANTHACEAE
Engler system the family is likewise placed near the Magnoliaceae, whereas
Hutchinson (1113), includes them in the Resales, thus recognizing their
affinities with the Rosaceae and Dichapetalaceae. In this connexion it is inter-

esting to record Tippo's (2261) somewhat guarded view based on the

anatomical characters of the wood, 'that the derivation of the Calycanthaceae
from the more primitive members of the Rosaceae would be inconsistent with
the anatomical facts'. He also draws attention to features in which the
Calycanthaceae resemble the Himantandraceae and Lactoridaceae which are
included in the Magnoliales.
Garratt (747 A) says: 'The Calycanthaceae have very few anatomical
features in common with the great majority of Monimiaceous genera and the
points of distinction are so outstanding as to indicate that their relationship,
ifany exists, is remote. Only with Bracteanthus, Peumus, and Siparuna is
there sufficient similarity to the Calycanthaceae to indicate possible, although
somewhat indefinite, affinity/
According to Record and Hess (1886) the 'Structure does not suggest that
of the order Resales'.

ECONOMIC USES
Carolina Allspice (Calycanthus floridus Linn.) has an aromatic bark.

GENERA DESCRIBED
Calycanthus,* Chimonanthus.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Hutchinson 1113, Lehmann 1343, Quinlan 1767, Tison 2265, Worsdell 2469.

(ii) On Wood Structure

Chalk and Chattaway 358, Garratt 747A, Record 1843, 1851, Record and Hess 1886,
Tippo 2261.

5. MAGNOLIACEAE
(FiG. 8 on p. 20; FIG. 10 on p. 32)

SUMMARY
(i) GENERAL
Trees and shrubs, now with a comparatively restricted distribution in
parts of North America, the West Indies, Brazil, the Far East, India, and
Malaya, &c. The geological record shows that the family was at one time
much more widely distributed in the Northern Hemisphere. Stomata are
usually confined to the lower surface of the leaf. In the stem the cork arises
in the epidermis, hypodermis, or outer part of the cortex. Complete or in-
complete diaphragms of stone cells are common in the broad pith. Secretory
cells with mucilaginous or oily contents are very frequent, especially in
parenchymatous tissues. Calcium oxalate, when present, occurs as small
octahedral or prismatic crystals.
 MAGNOLIACEAE 17

(ii)
WOOD
Vessels solitary and in small groups, sometimes with spiral thickening,
perforation plates typically scalariform with few, widely spaced bars, but
simple perforations often present and sometimes predominant, intervascular
pitting scalariform to opposite, members moderately long. Parenchyma
terminal only. Rays usually up to 3 or 4 cells wide, with few uniseriates,
heterogeneous to homogeneous, sometimes with oil cells. Fibres with
bordered pits, of medium length to very long.

LEAF
Usually dorsiventral. Hairs mostly uniseriate, multicellular, but some-
times unicellular in Magnolia and Michelia. Tufted hairs with i- or 2-celled
rays recorded in Michelia spp. Epidermis composed of cells with straight or
sinuous anticlinal walls; outer walls sometimes striated or covered with
granular deposits of wax. Epidermis silicified, e.g. in Magnolia grandiflora
Linn, and M. virginiana Linn.; papillose in Liriodendron tulipifera Linn.,
Magnolia ferruginea Farm., and Michelia punduana Hook. f. et Thorns.
Hypoderm present below the upper epidermis in Liriodendron tulipifera,
Magnolia spp., Manglietia insignis BL, Michelia spp., and Talauma ovata
St. Hil. A second layer of cells, similar to those of the hypoderm, present
beneath the palisade layers in Liriodendron tulipifera according to Holm
(1036). Stomata usually confined to the lower surface; mostly rubiaceous,
but sometimes ranunculaceous types present as well, e.g. in Liriodendron.
Petiole (Fig. 10 A, c, G, and i), in transverse sections through the distal end,
exhibiting a circle of separate collateral bundles in Liriodendron, Magnolia,
Michelia, and Talauma. One of the bundles in the base of the petiole of
Liriodendron tending to become medullary. Bundles scattered in the base of
the petiole of Talauma hodgsoni Hook. f. et Thorns, according to Worsdell
(2469). Ozenda (1645, 1647) has recently drawn attention to the numerous
vascular strands in the base of the petiole of various members of the Magno-
liaceae. (See also ANNONACEAE under 'Petiole' for similar structure in Uvaria.)

Secretory cells, sometimes with suberized walls, common in parenchy-

matous tissues. The mode of development of the oil cells in Liriodendron
and Magnolia has been described by Lehman (1343).

Axis
YOUNG STEM (Fig. 10 B)
Cork superficial, generally arising in the sub-epidermis. Primary cortex
including groups of stone cells and/or branched sclerenchymatous idioblasts
in Magnolia, Manglietia, and, according to Holm (1036), in Liriodendron.
Phloem strands in Liriodendron tulipifera Linn, each bounded externally by
a cap of fibres; fibres also occurring but forming less definite caps at the
periphery of the phloem in species of Magnolia examined at Kew. Secondary
phloem sometimes including concentrically arranged strands of fibres.
Xylem and phloem of adjacent vascular strands in Liriodendron and Mag-
nolia usually sufficiently separated by conspicuous, primary medullary rays
to appear as distinct vascular bun dies in transverse sections. Vessels frequently
4594 o
i8 MAGNOLIACEAE
with scalariform perforationplates. Pith generally broad; often divided by
transverse septa composed of sclerenchymatous cells. Secretory cells com-
mon in parenchymatous tissues.
WOOD (Fig. 8 A-B, E-G, and K)
Vessels usually medium-sized (mean tangential diameter 100-200 /x),
but
small (less than 100 JJL) in Alcimandra, Liriodendron, Magnolia, Manglietia,
and Michelia p.p. solitary, in short radial multiples and in occasional clusters
; ;

radial multiples moderately pronounced in Elmerillia and Liriodendron very ;

variable in number, 5-100 per sq. mm.; most numerous in Liriodendron and
Magnolia, and fewest in Aromadendron and Elmerrillia p.p., e.g. E. mollis
Dandy; spiral thickening observed or reported (1467, 1851) in Alcimandra,
Aromadendron, Magnolia, Michelia, and Talauma (1851). Perforation plates
typically scalariform with few, widely spaced bars, but varying from all
scalariform to almost all simple, e.g. in Magnolia acuminata Linn. the number;

of simple perforations varying markedly in different species, even of the same

genus, being greatest in some species of Elmerrillia and Magnolia the number ;

of bars in the scalariform perforation plates typically fewer than 10, but,
according to Garratt (746), occasionally up to 20 in Magnolia and up to 25 in
Talauma. Intervascular pitting typically large and scalariform, but opposite
in Liriodendron. Pits to ray and wood parenchyma often large and simple or
with very narrow borders; often unilaterally compound, one large pit in a
vessel subtending several small pits in a ray cell, the latter separated by
vertical, rod-like partitions; similar to the intervascular pitting in Liriodendron
or occasionally unilaterally compound. Tyloses often present. Mean member
length o-8-i-i mm. Parenchyma in bands, 2-12 cells wide, that usually
appear to be obviously 'terminal' in some tropical species, however, there
;

may be more than one band at the end of what appears to be a single growth
ring, the inner band or bands often being discontinuous and sometimes
anastomosing; Chowdhury (415) states that the position of the bands in the
growth rings of Michelia champaca Linn, is uncertain; rare cells can occa-
sionally be found scattered among the fibres or touching the vessels. Cells
often markedly disjunctive. Silica sometimes present in Michelia (794).
Strands usually either of 8-12 moderately short cells or of 4 tall cells. Rays
usually up to 3 or 4 cells wide Solereder gives a maximum of 7 for Manglietia
; ;

slightly less than i mm. high; uniseriates usually very few, though occasionally
moderately numerous, e.g. in Magnolia pterocarpa Roxb. and Talauma minor
Urb., and composed of both procumbent and upright cells; 3-7 rays per mm. ;

heterogeneous (Kribs's Type II A and B), usually with 1-4 marginal rows of
square to upright cells; homogeneous in a few species of Magnolia, e.g. M.
acuminata Linn., and seldom with more than 2 marginal rows and with the
cells square rather than upright in Liriodendron. Enlarged oil cells present on
the margins of the rays in Aromadendron, Elmerrillia (Fig. 8 E), Michelia, and
Talauma. Gonggrijp (794) states that silica is 'present in some species of
Aromadendron, Magnolia, and Michelia, Fibres with small- to moderately
large-bordered pits, which occur mostly on the radial walls and are typically
rather few; apertures usually exserted. Occasional thin cross-walls have been
noted by Janssonius (1154) near the ends of the fibres in Magnolia javanica
Koord. et Valet, and Michelia montana BL; thin membranes (septa?) are
 MAGNOLIACEAE 19
sometimes moderately abundant in Manglietia glauca Bl. and M. hooheri
Cubitt et W. W. Sm. Walls moderately thin. Mean length 1-2-2-3 mm -

ROOT
The following characters recorded by Holm (1036) for Liriodendron
tuKpifera Linn. Resin cells present in the primary cortex, bark, and
parenchymatous rays. Scattered sclerenchyma strands situated in the
secondary phloem. Pith broad in lateral roots.

ECOLOGICAL ANATOMY
A 'dune'form of Liriodendron tulipifera Linn, described by Starr (2188)
with smaller epidermal cells and a deeper palisade layer than in 'mesophytic*
forms. Vessels more numerous in the midrib of the 'dune' form and fibres
and collenchyma more abundant.

ECONOMIC USES
Numerous species and horticultural Varieties' of Magnolia are cultivated
for the sake of their ornamental flowers. According to Holm (1036) the bark
of Liriodendron tulipifera Linn, was at one time used medicinally because it
contains 'liriodendrin' and 'tulipiferin*.
The woods of this family are typically rather soft and easy to work and are
suitable for general carpentry and joinery. That of Liriodendron tulipifera
Linn, is of considerable commercial importance and is known under a variety
of names such as Yellow Poplar (America), White wood (Britain), and Canary
Whitewood. The timber of Magnolia acuminata Linn, is very similar and is
also often exported from America. Various species of Magnolia, Manglietia^
Michelia, and Talauma produce similar timber in the East and are of local
importance, particularly Michelia champaca Linn., the source of Champak in
India,

TAXONOMIC NOTES
The occurrence of scalariform perforation plates and scalariform inter-
vascular pitting is generally accepted as corroborative evidence that this is a
very primitive group. It should, however, be borne in mind that the vessel
members are only moderately long and that the perforations appear to be in
a transitional condition between wholly scalariform and wholly simple. This
would indicate a moderately rather than a very low level of specialization;
further, the ray type, withfew uniseriates and a tendency to be homogeneous,
suggests an even higher level.
In this connexion it is interesting to note that Ozenda (1645, 1647) has
drawn attention to the somewhat complex nodal anatomy of Liriodendron and
certain other members of the Magnoliaceae, and has pointed out that this is
very different from the simpler node structure of the Annonaceae, Win-
and certain of the Rosaceae. The same author also
teraceae, Dilleniaceae,
found evidence of syncarpy in the Magnoliaceae. This evidence, like that of
the wood anatomy, indicates that the Magnoliaceae, although a primitive
group, may be less so than has sometimes been supposed.
The genera, other than Liriodendron, are not easily distinguished from one
another by their wood anatomy. McLaughlin (1467) was unable to find any
 FIG. 8. MAGKQLIACEAE, A-B, E-G, and K; DILLENIACEAE, I-J and L;
SCHISANDRACEAE, C and H; WINTERACEAE, D
A, Michelia champaca Linn. Ray. B, Manglietia virginiana Linn. Ray. C, Illicium cambodianus
var. crassifolium Ridl. Ray. D, Drimys dipetala F. Muell. Rays. E, Elmerillia mollis Dandy. Ray.
F, Liriodendron tulipifera Linn. Ray. G, L. tulipifera Linn. T.S. H, Illicium cambodianus Ridl. Ray.
I, Dillenia reticulata King. Rays. J t PToriwa excelsa Jack. T.S. K, Michelia champaca Linn. T.S.
L, Dillenia meliosmifolia Hook. f. et Thorns. T.S. C, cell containing raphides.
 MAGNOLIACEAE 21

marked characteristics to distinguish Aromadendron and Manglietia from

Talauma, or Elmerrillia from Michelia] he confirmed Dandy's statement that
Talauma spongocarpa King is a synonym for Michelia baillonii F. & G. but
opposed Dandy's transfer of Michelia kachirachira Kanehira et Yamamota
and Talauma sp. (Toroconte') to Magnolia.

GENERA DESCRIBED
(i) GENERAL
Liriodendron,* Magnolia,* Manglietia,* Michelia,* Talauma.*
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Alcimandra Dandy, Aromadendron BL, Elmerrillia Dandy, Liriodendron
Linn., Magnolia Linn., Manglietia BL, Michelia Linn., Talauma Juss.

LITERATURE
(i) On General Anatomy
Holm 1036, Lehmann 1343, Ozenda 1645, 1647, Starr 2188, Worsdell 2469.

(ii) On Wood Structure

Bailey 99, Beekman 167, den Berger 179, 182, 183, Brown, F. B. H. 280, Brown, H. P.
288, 289, Chowdhury 415, Coster 481, Desch 574, Garratt 746, 747, 747A, Gonggrijp
794, Hale 870, Howard 1088, Janssonius 1154, Jeffrey 1167, Jones 1191, Kanehira 1206,
1209, Knight 1250, Lecomte 1334, Lemesle 1351, McLaughlin 1466, 1467, Pearson and
Brown 1679, Record 1790, 1818, 1843, 1851, 1883, Record and Hess 1886, Record and
Meli, 1894, Stone 2203, Tang 2231, Thompson 2256, Tortorelli 2272, Tapper 2295,
Yamabayashi 2478.

6. SCHISANDRACEAE
(Fic. 8 on p. 20; FIG. 9 on p. 22; FIG. 10 on p. 32)

A. AUSTROBAILEYA, KABSURA, SCHISANDRA

SUMMARY
A family of shrubs some of which have a trailing habit. Its members occur
in China, the Malayan region, Australia, and in the south-eastern parts of the
U.S.A., its distribution thus recalling that of the Magnoliaceae. The wood
exhibits the following features. Vessels solitary, with spiral thickening, per-
foration plates scalariform or scalariform and simple, intervascular pitting
scalariform to opposite, members moderately to very long. Parenchyma
terminal only. Rays up to 3 cells wide, heterogeneous, with oil cells. Fibres
with conspicuous bordered pits, of medium length to moderately long.

LEAF
Epidermis composed of mucilaginous cells in Schisandra axillaris Hook, f .

et Thorns, and 5. elongata Hook. f. et Thorns. Cuticle ridged, especially on

the lower surface of the leaf, in Austrobaileya scandens BailL, and to a smaller
extent in Schisandra chinensis (Turcz.) BailL Stomata present on both sur-
faces in Kadsura japonica Juss., more numerous on the lower than on the
upper surface in Austrobaileya scandens^ confined to the lower surface in
Schisandra chinensis provided with specially large, oval guard cells in species
;
22 SCHISANDRACEAE
of Austrobaileya, Kadsura, and Schisandra. No definite subsidiary cells noted
in Austrobaileya scandens, but a tendency to form subsidiary cells parallel to
the pore observed in Schisandra chinensis. Petiole, in transverse sections
through the distal end of material examined at Kew, exhibiting an arc of
vascular bundles in Kadsura japonica\ an interrupted, crescentic vascular
strand in Schisandra chinensis (Fig. 10 j) and S.propinqua Hook. f. et Thorns.
(Fig. 10 L); an almost continuous crescentic vascular strand in Austrobaileya
scandens (Fig. 10 E). Ozenda (1646) found 3 vascular bundles in the base and
middle part and 5 at the distal end of the petiole of Kadsura and Schisandra.
The 3 strands are close together at the base of the petiole, the nodal structure
being unilacunar. Secretory elements. Mucilage cells present around the

FIG. 9. SCHISANDRACEAE
Crystal-bearing fibrous cells from the pith of Kadsura roxburghiana Arn. :

A, in transverse section; B, in longitudinal section. The crystals are shaded.

By Solereder.

petiolar vascular bundles in Schisandra, and mucilage cavities noted in the

corresponding position in Austrobaileya and Kadsura. 'Resin* cells noted in
the mesophyll and petiolar ground tissue in Austrobaileya and Kadsura.

Axis
YOUNG STEM (Figs. 9 A-B, 10 H)
Cuticle ridged in Austrobaileya scandens Baill. and Schisandra chinensis
(Turcz.) Baill. Cork arising superficially in material examined at Kew. Peri-
cycle including a somewhat interrupted to almost continuous ring of
sclerenchyma in Schisandra', pericyclic sclerenchyma in the form of a com-
posite and continuous ring in Austrobaileya scandens and of a similar but less
conspicuous and less lignified ring in Schisandra chinensis. Phloem forming
an almost closed cylinder in Schisandra. Xylem appearing, in transverse
sections, as an almost or quite continuous cylinder in species of Schisandra.
A continuous ring of xylem traversed by narrow rays, also noted in Austro-
baileya and Kadsura. Vessels angular in Austrobaileya and Kadsura mostly ;

solitary but also in tangential or oblique pairs in Austrobaileya scandens mostly ;

solitary, but tangential groups usually of more than 2 in Schisandra chinensis \

perforation plates scalariform in both of the last 2 species (see also under
'Wood'). Lateral pits of the vessels circular with slit-shaped oblique apertures
in Austrobaileya scandens; scalariform lateral pitting or very elongated hori-
zontal pits noted in Schisandra chinensis. Groundwork of wood, in both of
the same 2 species, composed of radial rows of rather thick-walled fibres, the
 SCHISANDRACEAE 23
latterhaving conspicuously bordered pits with slit-shaped frequently crossed
apertures. Pith consisting of oval, relatively thin-walled cells with granular
contents in Schisandra chinensis\ composed of fairly thick- walled, polygonal,
pitted cells, in Austrobaileya scandens. Crystalliferous fibrous cells occur in
the pith of Kadsura roxburghiana Arn. (Fig. 9 A-B). Secretory elements.
Large, conspicuous, secretory cavities noted in the phloem of Schisandra;
no secretory cavities clearly visible, though their presence suspected in the
phloem of Austrobaileya and Kadsura in the material available for examina-
tion. 'Resin' cells noted in the pith of Austrobaileya scandens, but none seen
in the corresponding position in Schisandra chinensis.

WOOD 1

Vessels large and solitary; 1-15 per sq. mm. in Kadsura and about 60 per
sq. mm. in Schisandra; with spiral thickening. Perforation plates scalariform,
with up to 15 bars in Schisandra and up to 7 bars in Kadsura (Garratt 746),
and with some simple perforations and occasional reticulate plates (Lemesle
1351) in Kadsura. (See also under 'Young Stem'.) Intervascular pitting
scalariform in Schisandra and typically opposite in Kadsura though Lemesle
y

(1351) refers also to scalariform pitting; pits to ray cells often unilaterally
compound. Tyloses present. Mean member length in Kadsura i -i mm. (100).
Parenchyma in terminal bands 1-3 cells wide. Rays 1-3 cells wide, up to
i mm. high; heterogeneous., with uniseriate margins of 1-7 cells.
Enlarged
oil cells present in the marginal rows. Fibres with conspicuous bordered

pits. Mean length 1-0-1-7 mm -

B. ILLICIUM
SUMMARY
Since it is a matter of opinion whether Illicium should be included in the
Schisandraceae or treated as a separate family, the following brief description,
which refers mainly to Illicium verum Hook, given independently of that
f., is

of the other genera included in the Schisandraceae. The wood exhibits the
following features. Vessels solitary, perforation plates scalariform, with
numerous fine bars, intervascular pitting scalariform to opposite, members
very long. Parenchyma sparse, paratracheal, and sometimes scattered along
the ring-boundary. Rays up to 3 cells wide, with numerous uniseriates,
markedly heterogeneous. Fibres with conspicuous bordered pits, of medium
length to moderately long.
LEAF
Glabrous. Cuticle on both surfaces very thick; locally striated on the upper
and more universally on the lower surface. Upper epidermis composed of
cells with sinuous anticlinal walls. Stomata confined to the lower surface;
surrounding cells not very clearly differentiated from those of the remainder
of the epidermis, but some of them containing bodies resembling plastids or
secreted material. Mesophyll differentiated into palisade and spongy por-
tions,but palisade consisting of 1-3 layers of almost cubical cells.
Petiole (Fig. 10 D) in transverse sections through the distal end of material
1
Based mainly on the description given by McLaughlin (1467).
z4 SCHISANDRACEAE
examined at Kew, exhibiting a solitary, shallow, crescentic, vascular strand.
Secretory cells with translucent contents, readily stained with haematoxylin,
abundant in the petiolar ground tissue.

Axis
YOUNG STEM
Cuticle thick. Primary cortex including infrequent, unbranched, stone
cells, the latter being mostly solitary but a few in small clusters. Pericycle
with little or no sclerenchyma. Xylem in the form of a continuous cylinder
traversed by uniseriate rays, the constituent cells of the latter filled with
granular contents readily stained by haematoxylin. Vessels small, angular,
mostly solitary but many in tangential or oblique pairs. Groundwork of
xylem composed of radial rows of angular fibres having bordered pits with
crossed apertures. Pith. Peripheral part consisting of cells with lignified
pitted walls and wide lumina. Central pith cells thin-walled. Secretory
elements. Abundant cells with translucent contents, readily stained with
haematoxylin, present in the primary cortex, phloem, and pith.

WOOD (Fig. 8 c and H)

Vessels very to moderately small (mean tangential diameter 25-100 /x);
almost exclusively solitary apart from apparent tangential pairs due to over-
lapping ends; 60-160 per sq. mm. Perforation plates scalariform, with very
numerous (up to 150) fine bars, which often anastomose. Intervascular
pitting scalariform to opposite; pits to ray cells similar. Tyloses present.
Mean length about 1*3 mm. Parenchyma very scanty; usually paratracheal
only, apart from occasional cells along the growth rings where these are
distinct, and limited to a few isolated cells in contact with the vessels;
McLaughlin (1467) describes the parenchyma as diffuse. Strands usually of
4-6 very tall cells. Rays up to 2 or 3 cells wide and up to 2 mm. high in some
species; uniseriates numerous and composed of high upright cells; 10-14
per mm. heterogeneous (Kribs's Type I), with marginal rows of very tall
;

upright cells, usually in 2-4 rows, but 10 or more rows not uncommon.
McLaughlin notes the occurrence of some rounded cells suggestive of oil
cells. Fibres with conspicuous bordered pits on both radial and tangential

walls, the apertures of the pits exserted. Kanehira (1206) refers to septate
fibres in /. anisatwn Linn. Walls moderately thick. Mean length about
1*6 mm.
TAXONOMIC NOTES (for Groups A and B)
Smith (2137) and Bailey and Nast (91) state definitely that Illicium should
not be included in the Winteraceae, but are undecided whether it should be
placed in the Schisandraceae or made the basis of a distinct family, the
Illiciaceae. According to Bailey and Nast, Illicium^ besides differing in wood
structure as noted above, is also unlike the Winteraceae in the following
respects, (i) The pollen is of a fundamentally different type, (ii) The primary
vascular system of the stem is a continuous pseudo-siphonostele, whereas in
the Winteraceae the vascular bundles form a less continuous ring, (iii) The
unilacunar node structure of Illicium contrasts with the uniformly trilacunar
nodes of the Winteraceae. (iv) The stomatal depressions of Illicium are not
 SCHISANDRACEAE 25
filled with the same granular material which occurs in those of the Win-
teraceae. (v) There are differences in floral anatomy and morphology,
(vi) There are differences in chromosome numbers. McLaughlin (1467)
considers that Kadsura and Schisandra together with the Magnoliaceae sensu
stricto form a natural group both morphologically and anatomically and that

they should be retained in the Magnoliales.

The wood structure indicates that Illicium is distinctly the most primitive
of the genera with vessels and may be regarded as intermediate in this respect
between the Magnoliaceae on the one hand and Kadsura, Schisandra, and the
Winteraceae on the other.
Illicium can, however,
hardly be regarded as a link between these families
as it differs in certain respects, e.g. in the absence of oil cells and the presence
of paratracheal parenchyma, from all the other groups.
McLaughlin (1467), who
refers to earlier investigations by van Tieghem
and Diels respectively, confirms their suggestion that Illicium should be
segregated from Drimys and its allies and made the type of a monotypic
family. McLaughlin also suggests transferring this family from the Mag-
noliales to the Hamamelidales.

ECONOMIC USES
A fragrant oil distilled from the fruits of Star Anise (Illicium verum Hook, f.)
is used medicinally. A chemical investigation of /. religiosum Siebold. has
has been made by Sze (2226).

GENERA DESCRIBED
GROUP A
FOR GENERAL ANATOMY. Austrobaileya,* Kadsura,* Schisandra.*
FOR WOOD STRUCTURE. Kadsura, Schisandra.
GROUP B
FOR GENERAL ANATOMY AND WOOD STRUCTURE. Illicium.*
* Kew
Represented in the slide collection.

LITERATURE
(t) On General Anatomy
Bailey and Nast 91, Ozenda 1646, Rehder 1902, Smith 2137, Sze 2226.
(ii) On Wood Structure
Garratt 746, 747, 747A, Kanehira 1206, Lemesle 1351, McLaughlin 1467, Record 1851.

7. WINTERACEAE
(FiG. 8 on p. 20; FIG. 10 on p. 32)

SUMMARY
(i) GENERAL
A family of trees and shrubs occurring chiefly in certain parts of South-east
Asia and South America. Drimys is the only genus which occurs in both the
Old and New Worlds. Smith (2136, .2137), who has recently revised the
taxonomy of the family, points out that the American species are all herma-
phrodite, whereas those from the Old World are dioecious. The lower surface
26 WINTERACEAE
of the leaf often appears punctate, owing to the stomata being situated in
depressions filled with a white, very finely granular material, believed to be
of a different chemical nature from the wax which occurs on the leaves of the
Magnoliaceae. The stomata are accompanied by 2-6 cells, parallel with the
guard cells. Our knowledge of the anatomy of the family has been con-
siderably extended by the recent work of Bailey and Nast (86-91). Features
in the leaf to which they draw attention include the trilacunar nodes; the
taxonomic value of the amount of sclerenchyma around the vascular bundles
of the veins and sheathing the terminal veinlets, increasing sclerification being
interpreted as evidence of a phylogenetic advance the occurrence of solitary
;

and clustered, often branched, sclerenchymatous idioblasts in the mesophyll ;

and the presence of mesophyll cells with reticulate thickenings in certain

species. The taxonomic value of the idioblasts is limited by variations in their
frequency, and cannot be fully assessed without more extended examination.
In young stems the vascular bundles are capped externally by fibres and
accompanied internally by elongated, thick- walled cells, the pericyclic fibres
becoming united by sclerosed parenchymatous cells to form a more or less
continuous ring in older stems. The late secondary phloem includes irregular
patches of sclerenchymatous tissue. Various types of scattered sclerenchyma-
tous idioblasts and clusters of sclerosed cells are common in the cortex and
pith, especially in members of the family from the Old World. Crystals
are infrequent, but have been observed in a few species. Spherical secretory
cells also occur.

(ii)
WOOD
Vessels absent. Parenchyma diffuse or in fine lines and sometimes
terminal. Rays up to 10 cells wide and very high, with numerous uniseriates,
markedly heterogeneous, occasionally with oil cells. Tracheids with 2-3
rows of circular pits, or occasionally with scalariform pits, on the radial walls ;

extremely long.

LEAF
Dorsiventral glabrous. Lower epidermis papillose, e.g. in most specimens
;

of Drimys brasiliensis Miers, but this character is not reliable for diagnostic
purposes; papillae also variable in length. Cuticle ranging from finely alveolar
to coarsely granular or warty. Epidermal cells also vary in size, form, arrange-
ment, wall thickness, and in the nature of the pitting, but these characters are
also unreliable for diagnostic purposes owing to variation within a species.
Stomata confined to the lower surface, and restricted to depressions filled
with a white granular material, the latter differing in chemical composition
from the wax in the Magnoliaceae and Schisandraceae. Granular material in
the stomatal depressions varying in appearance according to the drying con-
ditions during the preparation of herbarium specimens. Stomata rubiaceous,
accompanied by 2-6 cells parallel to the guard cells.
Mesophyll, in the Wintera section of Drimys, as well as in species of
Bubbia and Zygogynum, including branched sclerenchymatous idioblasts,
more numerous in some species than others, but frequency also varying in
different collections of a single species. Mesophyll cells with reticulate

thickenings noted by Bailey and Nast in Belliolum pancheri (Baill.) v. Tiegh.

 WINTERACEAE 27
and B. vieillardi v. Tiegh. but not in other species of Belliolum examined by the
same authors. Similarly thickened cells also noted in certain species of Bubbia.
Vascular bundles of the coarser veins and terminal veinlets more or less
sheathed with sclerenchymatous elements in Belliolum, certain species of
Bubbia, and in the Tasmannia section of Drimys. Vascular bundles of the
large veins also embedded in sclerenchyma in the Wintera section of Drimys,
but terminal veins in this group of species not accompanied by sclerenchyma.
Considerable variation in the amount of sclerenchyma associated with the
vein endings noted in Zygogynum. Different members of the Winteraceae
form a series showing increasingly complex vascular systems in the petiole
and lamina, the more specialized species being characterized by more numer-
ous vascular bundles and by a tendency for the separate bundles to become
fused to form an arc. Vascular system relatively simple (with an arc of vas-
cular bundles) in Drimys (Fig. 10 F) and Pseudowintera, moderately specialized
in Belliolum and Bubbia and still more advanced in Exospermum and Zygogy-
num. Shape and arrangement of the vascular bundles varying not only within
a species but in different leaves from an individual plant. Crystals relatively
infrequent, recorded only in Bubbia clemensiae A. C. Sm. and Exospermum
stipitatum (Baill.) v, Tiegh. Spherical secretory cells generally present.

Axis
YOUNG STEM
Cork arising in the epidermis in Drimys. Late-formed phloem stated by
Bailey and Nast to contain irregular patches of sclerenchymatous tissue.
&
Xylem of Drimys winteri J. R. G. Forst. examined at Kew appearing, in
transverse sections, as a more or less continuous cylinder, devoid of vessels
and traversed by narrow medullary rays. Ray cells become conspicuous in
sections stained with haematoxylin owing to the presence of deeply coloured
contents. Bailey and Nast (88) describe the vascular bundles in the stem as
being each provided with an external cap of slender, thick-walled fibres and
subtended internally by elongated, lignified, thick-walled cells, the external
caps of fibres becoming converted to a more or less continuous ring in older
material through sclerification of the intervening parenchymatous elements.
Scattered sclerenchymatous cells of varied form and clusters of sclereids
observed by Bailey and Nast in the cortex and pith of some members of the
family, but none seen in others. Cortex and pith of Old World representatives
of the family generally more sclerified than those from the New World,
especially in coriaceous species of Belliolum, Bubbia, Exospermum, and
Zygogynum. Crystals infrequent, but crystalliferous cells noted by Bailey
and Nast in association with the medullary and cortical sclerenchyma in
Belliolum, Exospermum, Pseudowintera, and Zygogynum. Scattered secretory
cells noted at Kew in the phloem and in the peripheral part of the pith of
Drimys winteri. Nodes trilacunar (with 3 foliar bundles related to 3 lacunae
in the vascular system of the stem) in all members of the family examined by
Bailey and Nast.
WOOD (Fig. 8 D)
Vessels absent. Parenchyma rather scanty and diffuse in Drimys winteri
]. R. & G. Forst. and Zygogynum (1467), more abundant and in uniseriate
28 WINTERACEAE
tangential lines in Bubbia haplopus (Burtt) A. C. Smith, Drimys colorataRaoul,
and D. axillaris Forst. (2256), and terminal in Zygogynum (1467) and the
lattertwo species of Drimys (2256). Strands usually of 8-10 tall cells. Rays
of two distinct sizes; the larger typically up to about 10 cells wide and
commonly over 5 mm. high, but much wider (20 cells?) in Pseudowintera
(88); uniseriates numerous, often more than i mm. high and composed of
tall cells; 9-13 rays per mm.; markedly heterogeneous (Kribs's
upright
Type with up to 10 or more marginal rows of tall upright cells; the inner
I)
cells sometimes square rather than procumbent in Drimys, but procumbent
in Bubbia haplopus\ often with sheath cells. Oil cells reported in the rays of
Zygogynum (1467). Tracheids usually with large, circular, bordered pits in
the radial walls, the pits most commonly in two or three rows and with
included apertures; the pits tend to be elongated or even scalariform in the
early wood tracheids in some species of Drimys and Bubbia, particularly in
the overlapping end- walls, the ratio of round to scalariform pits varying even
from specimen to specimen (88). Pits to ray cells similar to the pits to other
tracheids. Mean length (Bubbia and Drimys) 4*0-4-3 mm. and, according to
McLaughlin (1467) up to 5-1 and 6-3 mm. respectively in Drimys and
Zygogynum.
TAXONOMIC NOTES
(i)
BASED ON WOOD STRUCTURE
Bailey and Nast (88, 91), in their study of the morphology of this group,
have shown that, with the exclusion of Illicium, 'the Winteraceae become a
homogeneous, natural aggregation of obviously closely related plants'. Within
the group these authors distinguish the following trends of structural
specialization 'toward reduction or elimination of wood parenchyma in Sect.
:

Wintera of Drimys, toward excessively widened multiseriate rays in Pseudo-

wintera, and toward reduction of cell size, particularly in dwarfed or micro-
phyllous species, e.g. in Sect. Tasmannia of Drimys'.
The occasional scalariform pitting that occurs in the tracheids of Drimys
figured largely in the controversy of about 1918, associated with the names of
Bailey and Jeffrey, as to whether Drimys, Tetracentron, and Trochodendron
represent degenerate 'evascularized* Dicotyledons, whose ancestors possessed
true vessels in their secondary xylem, or whether they have descended directly
from ancestors that possessed scalariform tracheids in their secondary xylem
(99, 1167, 2256). Modern work on the significance of the length of the
cambial initial and vessel members (see Introduction) has provided a means of
testing such hypotheses and Bailey and Nast, writing in 1945, state that 'the
primitive character of the cambium and xylem in the Winteraceae, Trocho-
dendron and Tetracentron rules out any possibility of these plants having
developed vessels and subsequently having lost them'. Elsewhere they claim
that 'when the summation of evidence from all organs and parts of the plants
is taken into consideration, there are no convincing arguments for
deriving
the Trochodendraceae from the Winteraceae or vice versa, or even for
inferring that these families are closely related genetically. Nor can one
assume that the other ranalian families were derived from these vesselless
families.'
The same authors also point out that comparisons between the Winteraceae
 WINTER ACE'AE 29
and the Coniferae overlook important anatomical differences, such as occur
in the rays, and are thus misleading. They conclude that 'if the vesselless
wood of the Winteraceae is to be compared with that of the Gymnosperms,
itshould be with the secondary xylem of Pteridospermae and Bennettitales
rather than with that of the Coniferae, Ginkgoales or Cordaitales*.

(ii)
BASED ON GENERAL ANATOMY
Besides the evidence of wood structure, Bailey and Nast (91) consider that
other anatomical facts indicate that lllidum should not be included in the
Winteraceae, a view which is also supported by Smith (2137) on morpho-
logical grounds. This subject is discussed more fully under Schisandraceae,
Bailey and Nast (1944) also say that:
'The available evidence indicates that internal foliar characters are unstable and
variable in the Winteraceae, particularly in Belliolum, Bubbia and Zygogynum. . . .

In the Wintera section of Drimys, increasing coriaceousness is attained largely by

the formation of large, armed sclereids interspersed throughout the mesophyll.
On the contrary, in the Tasmannia section of Drimys, sclerification progresses along
the veins and veinlets, the bulk of the mesophyll remaining thin-walled. In Bellio-
lum, Bubbia and Zygogynum increasing coriaceousness commonly involves intensi-
y

fied sclerification along the veins and veinlets, and not infrequently throughout the

mesophyll. In the more coriaceous species of Bubbia and Zygogynum, all three
trends of sclerification may occur simultaneously.'

ECONOMIC USES
Winter's Bark (Drimys winteri J. R. & G. Forst.) has been valued for its
antiscorbutic properties, but is not now of much importance. True Winter's
Bark has frequently been confused with that of other plants. The following
microscopical characters were noted in specimens in the Kew Museums.
Exterior of the young bark covered by about 4 or 5 layers of very thick- walled
cork cells, the thickening being uniform on all of the walls, and the lumen
filled with gum-like deposits. Cork plentiful in old material, but very uneven
and giving the exterior of the bark a very rough appearance. Cork bounded
internally by a pareiichymatous cortex, the inner boundary of this region
being marked by groups of very large, thick-walled stone cells, the latter
sometimes associated with groups of fibres very much smaller in diameter but
also thick-walled. These parenchymatous elements collectively form a very
much interrupted ring. Smaller, isolated stone cells also present immediately
within the cork layer. Solitary, vertically elongated, secretory cells with pale-
yellow contents abundant in the phloem on the inside of the sclerenchyma
ring. Phloem in old bark very much broader and containing radially arranged
groups of stone cells. More numerous secretory cells also occur throughout
the phloem. Unlignified phloem elements of the older bark tend to be in
radial rows.
GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Belliolum, Bubbia, Drimys,* Exospermum, Pseudowintera, Zygogynum.
* Kew slide collection.
Represented in the

(ii) FOR WOOD STRUCTURE

Bubbia, Drimys, Zygogynum.
30 WINTERACEAE
LITERATURE
(i) On General Anatomy
Bailey 75, Bailey and Nast 86, 87, 88, 89, go, 91, 92, Jeffrey and Cole 1167, Knight
1250, Nast 1580, Smith, A. C. 2x36, 2137, 2139.

(ii) On Wood Structure

Bailey and Nast 88, 91, 94, Cozzo 494, Dadswell and Record 533, Hess 950, McLaughlin
1467, Thompson 2256.

8. CERCIDIPHYLLACEAE
(FiG. 7 on p. 14; FIG. 10 on p. 32)

SUMMARY
(i) GENERAL
Trees from China and Japan, with leaves somewhat similar to those of the
Hamamelidaceae. The sole genus is Cercidiphyllum of which C. japonicum
Sieb. et Zucc. has been examined.

(ii) WOOD
Vessels very small and numerous, with spirals in the tips of members, per-
foration plates scalariform with numerous bars, intervascular pitting scalari-
form to opposite, members very long. Parenchyma very sparse, terminal
and sometimes diffuse. Rays 1-2 cells wide, heterogeneous, and often fused
vertically. Fibres with conspicuous bordered pits, moderately long.

LEAF
Dorsiventral. Lower epidermis
sub-papillose. Leaf margins provided
with glandular emergences containing a gummy substance. Stomata
confined to the lower surface; ranunculaceous. Petiole (Fig. 10 N) in trans-
verse sections through the distal end, exhibiting a solitary, U-shaped vascular
strand with very much incurved ends. Petiolar vascular strand accompanied
on the outside by an interrupted ring of fibres. Large cluster crystals present
in the mesophyll as well as in the cortical region of the petiole.

Axis
YOUNG STEM (Fig. 10 M)
Cork originating in the outer part of the cortex. Pericycle including
isolated bundles of fibres. A
slightly interrupted ring of similar fibres also
arising in the phloem. in the form of a continuous cylinder traversed
Xylem
by narrow rays; including numerous vessels, very angular in transverse sec-
tion and tending to exhibit a radial arrangement, but tangential or oblique
pairs as well as small clusters also common. Individual vessel-elements,
where adjacent, separated from one another by very thin walls. Perforation
plates scalariform with numerous, very fine bars. Pith composed of thick-
walled mostly filled with starch. Perimedullary cells smaller than those
cells,

constituting the greater part of the pith, Large cluster crystals present in
the young primary cortex but tending to disappear from older material. Many
of the medullary ray cells, where traversing the phloem, contain large, con-
spicuous, solitary crystals.
 CERCIDIPHYLLACEAE 31
WOOD (Fig. 7 i-j)
Vessels very small (mean tangential diameter 25-50 p) and angular;
solitary and in short radial multiples and tangential pairs due to overlapping
ends (Fig. 71); more than 100 per sq. mm. and crowded; coarse spirals
present in the vessel tips. Perforation plates scalariform, with 20-50 bars
(1467). Intervascular pitting very scarce; when present, opposite to scalari-
form; pits to ray cells similar to the intervascular pitting, often unilaterally
compound. Tyloses sometimes present. Mean member length i -6 mm. (100).
Parenchyma sparse and limited to the boundaries of the growth rings in the
material examined, but McLaughlin (1467) refers to diffuse parenchyma, the
terminal parenchyma varying from occasional cells to discontinuous uniseriate
bands. Rays up to 2 cells wide and less than i mm. high; often with 2 or
more multiseriate parts separated by uniseriate upright cells and with the
latter nearly as wide tangentially as the multiseriate parts (Fig. 7 j) uniseriates ;

moderately numerous and composed of both procumbent and upright

cells; about 13 rays per mm.; heterogeneous (Kribs's Type II B), with 1-4
marginal rows of square to upright cells. Cells often containing brown
deposits and sometimes crystals of calcium oxalate (1467). Fibres with
conspicuous bordered pits, which are equally numerous on both radial and
tangential walls and of about the same size as the opposite intervascular
pitting. Walls of medium thickness and usually markedly flattened in the
last 2-8 rows, forming a distinct band of latewood from which vessels tend
to be absent. Mean length 1-9 mm. (100).

TAXONOMIC NOTES
McLaughlin (1467) considers that this family is more closely allied to the
Hamamelidales than to the Magnoliales.

GENUS DESCRIBED
Cercidiphyllum.*
* Kew
Represented in the slide collection,

LITERATURE
On Wood Structure
Bailey and Tapper 100, Kanehira 1206, 1209, McLaughlin 1466, 1467, Record 1843,
1851.

9. EUCOMMIACEAE
(Fie. 7 on p. 14; FIG. 10 on p. 32)

SUMMARY
(i) GENERAL
The family is represented by a single Chinese tree Eucomrnia ulmoides Oliv.
The most noteworthy anatomical feature is the occurrence of laticiferous
cells containing a substance similar to rubber. This material is liberated
from the bark, young twigs, and leaves when broken or removed from the
plant. The laticiferous elements occur particularly in the phloem and adjacent
tissue of the pericycle, although they extend into the mesophyll of the leaf.
 MAGNOLIACEAE, A-C, G, and I; SCHISANDRACEAE, D-E, H, J, and L;
FIG. 10.
WINTERACEAE, F; EUCOMMIACEAE, K and O; CERCIDIPHYLLACEAE, M-N
A, Magnolia virginiana Linn. Petiole x 19. B, M. grandiflora Linn. Stem X 10 (secretory cavities
in cortex). C, Liriodendron tulipifera Linn. Petiole X 10. D, Illicium verum Hook. f. Petiole X 13
(secretory cavities in ground tissue). E, Austrobaileya scandens Bail. Petiole X 13. F, Drimys winteri
Forst. Petiole X 10. G, Magnolia grandiflora Linn. Petiole x 10. H, Schisandra propinqua Hook. f.
et Thorns. Stem x 20. I, Michelia excelsa Blume. Petiole X 13. J t Schisandra chinensis (Turcz.) Baill,
Petiole x 13. K, Eucommia
~"
Oliver. Petiole X 1 9. L, Schisandra propinqua Hook, f et Thorns.
imia ulmoides Oliv .

X 20. M, Cercidiphyllum japonicum Sieb. et Zucc. Stem X 19. N, C.japonicum Sieb. et Zucc.
Petiole X20.
Petiole X29.9. O, Eucommia ulmoides Oliver. Stem x 19.
 EUCOMMIACEAE 33

According to Varossieau (2325) the laticiferous tubes are unicellular (in-

articulate), and differ in their mode of development from those of Cannabis
and Euphorbia.
(ii) WOOD
Vessels very small and nearly all solitary, semi-ring-porous, with spiral
thickening, perforations simple, intervascular pitting opposite to alternate,
members of medium length. Parenchyma diffuse and terminal. Rays up
to3-4 cells wide and low, with rather few uniseriates, almost homogeneous.
Fibres with distinctly bordered pits, moderately to very short.

LEAF
Dorsiventral. Hairs simple, unicellular. Mesophyll including groups of
silicified cells. Stomata confined
to the lower surface; ranunculaceous.
Petiole (Fig. 10 K), in transverse sections through the distal end, exhibiting
single, thick, crescent-shaped vascular strand, including xylem vessels
arranged in radial rows. Laticiferous cells mostly situated in the veins, but
sometimes in the mesophyll as well. Crystals absent.

Axis
YOUNG STEM (Fig. 10 o)
Cork originating in the epidermis. Primary cortex collenchymatous.
Pericycle containing a loose ring of fibres, together with elongated stone
cells with silicious contents and thin-walled, frequently paired, parenchyma-
tous cells each containing an areolate silica body. Xylem and phloem in
the form of continuous cylinders traversed by narrow rays. Pith hetero-
geneous, the walls of the peripheral cells being thicker than those of the cells
towards the centre. Laticiferous cells occur in the primary cortex, phloem,
and pith.
WOOD (Fig. 7 K-L)
Vessels very to extremely small (mean tangential diameter usually about
25-30 ju); nearly all solitary, but with occasional radial pairs; 250-350 per
sq. mm. (2262); semi-ring-porous; with spiral thickening. Perforations
simple, except in the immediate neighbourhood of the primary xylem (2158).
Intervascular pitting uncommon owing to the solitariness of the vessels; small,
opposite (2261) to alternate; pits to ray and wood parenchyma cells similar in
size and shape to the intervascular pits. Tyloses reported (2262) in a single

specimen. length 0-36-0-5 mm. (2262). Parenchyma apotracheal, as

Mean
among the fibres (Fig. 7 L) and along the boundaries
isolated cells scattered
of the rings. Strands usually of 4 cells. Rays up to 3, occasionally 4, cells
wide and seldom more than 0-5 mm. high; uniseriates rather few and com-
posed mostly of procumbent cells about 5 rays per mm. almost homogeneous
; ;

(intermediate between Kribs's Types Heterogeneous II B and Homo-

geneous I), with an occasional single marginal row of square to slightly
upright cells; the procumbent cells small (Fig. 7 K). Fibres with distinctly
bordered pits on both radial and tangential walls, the borders slightly smaller
than those of the intervascular pits. Walls moderately to very thick. A
few cells with spiral thickening present in some specimens. Mean length
o-67-O'84 mm. (2262).
4594 D
34 EUCOMMIACEAE
ROOT
Laticiferous cells present in the pericycle and phloem.

TAXONOMIC NOTES
Tippo (2261, 2262) places the Eucommiaceae in the Urticales near the
Ulmaceae and at about the same level of specialization. A similar position is
suggested by Varossieau (2324) on general morphological grounds. Among
the characters of the wood that Tippo considers as evidence of similarity with
Ulmaceae rather than Hamamelidaceae are simple perforation plates, ring-
porous vessels, ray type, and alternate intervascular pitting.
ECONOMIC USES
The bark of E. ulmoides has been used for medicinal purposes by the
Chinese. The rubber has never been extracted commercially owing to the
small quantity present in the latex.
GENUS DESCRIBED
Eucommia.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Harms 898, Parkin 1654, Varossieau 2324, 2325.
(ii) On Wood Structure
Record 1843, 1851, Tippo 2261, 2262, Varossieau 2324.

10. EUPTELEACEAE
(FiG. ii on p. 40)

SUMMARY
Following the taxonomic revision by Smith (2140), the Eupteleaceae are
here treated as a unigeneric family comprising 2 species of trees. These are
E. polyandra Sieb. et Zucc. from Japan and E. pleiosperma Hook. f. et Thorns,
from China and parts of India. Smith was unable to recognize E. francheti v.
Tiegh. as a separate species, since it cannot reliably be separated from
J?. pleiosperma by having a less papillose lower epidermis to the leaf than the

last The wood exhibits the following features. Vessels small,

species.
solitary,numerous, and with scalariform perforation plates; intervascular
pitting mostly opposite, members of medium length to moderately long.
Parenchyma in narrow terminal bands. Rays up to 10 cells wide and high,
heterogeneous. Fibres with small bordered pits, of medium length.

LEAF
DorsiventraL Hairs on young leaves simple, uniseriate, with a single or
several short basal cells, sometimes arranged bi-serially, together with one or
more elongated terminal cells with wide lumina. Lower epidermis fre-
quently papillose in E. pleiosperma Hook. f. et Thorns. Stomata on the
lower surface ranunculaceous and less specialized than in Tetracentron and
Trochodendron. Arm-palisade cells recorded in the mesophyll. Petiole of
E. pleiosperma (Fig. ii G) in transverse sections through the distal end of
 EUPTELEACEAE 35
material cultivated and examined at Kew, exhibiting a cylindrical vascular
strand, tending to become double with two groups of xylem on the adaxial
side and supported along the whole of the outer periphery by a well-developed
ring of fibres. One or two small accessory strands also present on the adaxial
side. Vessels in the xylem arranged in very definite radial rows. The follow-
ing additional particulars concerning the petiole structure have been recorded
by Nast and Bailey (93). Five-i i vascular strands (more numerous in E. pleio-
sperma than in E. polyandrd) enter the base of the petiole and, at certain levels
of the attachment of the leaf, commingle with those of the axillary bud to
form an arc of vascular strands confronting a single broad parenchymatous
region of the eustele. Vascular bundles first uniting nearer the distal end of
the petiole to form a more or less continuous arc, and, still closer to the lamina,
a vascular cylinder which extends into the midrib. No branched idioblasts,
either sclerotic or secretory, noted by Nast and Bailey, but secretory cells
with unidentified contents observed at Kew in the 'cortical* and 'medullary'
ground tissue of the petiole of E. pleiosperma^ and smaller, apparently tannini-
ferous elements in the phloem and medullary ray cells in the petiole of the
same species. A few cluster crystals noted in the parenchymatous ground
tissue of the petiole of E. pleiosperma.

Axis
YOUNG STEM (Fig. n i)
The following description applies mainly to E. pleiosperma. Cork arising
at a fairly deep level in the cortex. Pericycle including a continuous or
slightly interrupted sclerenchymatous ring, the latter consisting mostly of
fibres but with a small number of stone cells present as well. Phloem con-

taining no parenchymatous elements in the material examined. Xylem

containing very numerous, somewhat angular vessels, seldom more than
30 fjL in diameter. Vessels tending to be in radial rows in the first-formed
wood, but this character less marked in the later wood where they are solitary
or in variously arranged groups perforation plates scalariform with numerous
;

fine bars. Primary rays 1-3 wide, conspicuous; distal ends enlarged.
cells
Pith consisting of with starch. Large, con-
relatively thick-walled cells filled
spicuous, cluster crystals present in the cortex, especially in the inner part.
Secretory cells with unidentified contents also noted in the cortex.
WOOD
Vessels small (mean tangential diameter about 50 /x) and solitary; about
75 per sq.mm. in the material examined, but McLaughlin (1467) gives the
number as 120-250 per sq. mm. Perforation plates scalariform, with 20-90
bars. Intervascular pitting opposite to transitional, rarely alternate or scalari-
form; pits to ray cells unilaterally compound. With thin-walled tyloses.
Mean member length o-8-O'9 mm. Parenchyma in terminal bands 1-2 cells
wide. Rays up to 10 cells wide and 3 mm. high; uniseriates numerous, com-
posed mainly of upright cells, but occasionally with some square to slightly
procumbent cells; about 9 rays per mm.; heterogeneous, with 1-6 marginal
rows of square or upright cells (Kribs's Type II A) on the outside of mature
stems, more markedly heterogeneous Clype I) in small twigs (93). Fibres
with small, round, bordered pits, the apertures exserted; mean length
i '2-i -5 mm.
36 EUPTELEACEAE
TAXONOMIC NOTES
Although Euptelea has commonly been treated as a representative of the
Trochodendraceae, the recent researches of Smith (2140) and of Nast and
Bailey failed to reveal any important similarities between Euptelea and the
other two genera. The wood structure in particular is completely unlike that
of the Trochodendraceae. All three authors agree that Euptelea occupies a
rather isolated taxonomic position, although its affinities are regarded as
rather remotely ranalian.

GENUS DESCRIBED
Euptelea.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Nast and Bailey 93, Smith 2140.

(ii) On Wood Structure

Bailey and Nast 93, Kanehira 1206, McLaughlin 1466, 1467, Record 1851.

11. HIMANTANDRACEAE
(Fic. 7 on n
p. 14; FIG. on p. 40)

SUMMARY
According to Bailey, Nast, and Smith (94) the family consists of 2 principal
species of trees, i.e. H. belgraveana F. v. Muell. from New Guinea and
probably the North Moluccas and H. baccata'(F. M. Bailey) Diels from
Queensland. There is a third inadequately known species H. parviflora
(Bak. Norm, in New Guinea. The lower surface of the leaves bears an
f.)
indumentum of peltate scales. The wood exhibits the following features.
Vessels with faint spiral thickening, simple perforations, intervascular pitting
alternate, members moderately long. Parenchyma in concentric bands.
Rays mostly 2-3 cells wide, heterogeneous, and with intercellular spaces.
Fibres with bordered pits,of medium length to moderately long.

LEAF
Abaxial surface covered with a dense indumentum of peltate scales, those
of H, baccata smaller and less crowded than those of H. belgraveana. Upper
surface of mature leaves glabrous, but scales or stellate hairs noted by Bailey,
Nast, and Smith on the adaxial surface of immature leaves of certain specimens.
Stomata characteristically arranged in approximately circular clusters around
each of the peltate scales on the lower surface. Mesophyll said by the same
authors to include numerous, scattered clusters of sclereids, in all of the
specimens of H. belgraveana collected by Clemens, but sclereids infrequent in
or absent from other material which they examined. Petiole of H. bel-
graveana (Fig. 1 1 H), in transverse sections through the distal end of specimens
examined at Kew, exhibiting an outer cortical region composed of thick-
walled cells; inner cortical and medullary regions composed of cells with
much thinner walls apart from scattered groups of large, pitted, lignified
 HIMANTANDRACEAE 37
stone cells; a vascular system consisting of a circle of separate, collateral
bundles, each supported externally by groups of fibres. Petiole described by
Bailey, Nast, and Smith as having 3 vascular strands entering the base, the
traces dividing towards the distal end to form 6-8 bundles arranged so as to
appear as an interrupted circle when viewed in transverse sections. Pairs or
small clusters of crystals present in the cells of the lower epidermis, and in
strands accompanying the sheaths of sclerenchyma enclosing the vascular
bundles of the veins and veinlets. Spherical secretory cells abundant.

Axis
YOUNG STEM (Fig. n j)
The following description is based on an examination of H. belgraveana
grown at Kew, and on the notes recorded by Diels (583), and the account
published by Bailey, Nast, and Smith (94).
Cork arising in the sub-epidermis when still very young component cells ;

very strongly thickened on the inner tangential, and to a smaller extent on the
Primary cortex containing groups of very thick-walled cells
lateral walls.
which become attached locally to the outer part of the pericyclic scleren-
chyma. Pericycle with a composite, somewhat interrupted ring of scleren-
chyma. Phloem narrow, devoid of lignified elements in the young material
examined at Kew, but phloem of older stems described by Bailey, Nast, and
Smith (94) as stratified into fibrous and non-fibrous portions. Sieve tubes
similar to those of the Magnoliaceae. Primary vascular system appearing in
transverse sections as a circle of individually distinct vascular bundles, but
this character becomes less obvious with the onset of secondary thickening,
many of the rays being only i cell wide. Xylem including numerous vessels
up to about 45 p in diameter, tending to be in radial rows, with bordered pits
on the lateral walls and mixed scalariform perforation plates and simple
perforations; fibres with bordered pits; rays mostly 1-2 cells wide. Bailey >
Nast, and Smith record the occurrence of a higher proportion of scalariform
perforation plates and more frequent opposite pitting in the lateral walls of
vessels from the metaxylem and first-formed secondary xylem than in the
latersecondary xylem. (See 'Wood' below.) Nodes trilacunar according to
Bailey, Nast, and Smith. Pith broad, consisting of pitted cells, but also
including groups of stone cells which tend to form transverse diaphragms.
Isolated secretory cells with unidentified amorphous contents noted in the
primary cortex. Solitary crystals present in many of the cortical stone cells,
and, according to Bailey, Nast, and Smith, in parenchymatous cells adjacent
to the phloem fibres.

WooD 1
(Fig. 7 G-H)
Vessels medium-sized (100-200 fi mean tangential diameter); solitary, in
multiples of up to 4. cells and in irregular groups; McLaughlin notes very
indistinct spiral thickening. Perforations typically simple, but Record (1851)
lists this family as having both scalariform and simple perforation plates.

(See also under Young Stem'.) Intervascular pitting alternate, of medium

*

size; pits to ray and wood parenchyma similar. Members up to i mm. long.
Parenchyma in continuous bands 2-20, mostly 3-8, cells wide, and about
1
Based mainly on the description given by R, P, McLaughlin (1467).
38 HIMANTANDRACEAE
half a millimetre apart (Fig. 7 H). Strands usually of 8 cells. Rays up to 4,
mostly 2-3, cells wide, slightly more than i mm. high and heterogeneous,
with 2 or 3 (occasionally 4) marginal rows of upright cells. Intercellular
spaces moderately conspicuous. Fibres with bordered pits, which occur
mostly on the radial walls, with exserted apertures. Walls of medium thick-
ness. Up to 2 mm. long.

TAXONOMIC NOTES
Himantandra is treated in the Engler system as well as in that of Hutchinson
(1113) as a separate family related to the Magnoliaceae. This treatment is
fully supported by the more recent work of Bailey, Nast, and Smith. The
presence in Himantandra of secretory cells, of sclerenchymatous diaphragms
in the pith, the occurrence of vessels with simple perforations and scalariform
perforation plates and the ring of separate bundles in the petiole are all
characters which suggest affinities with the Magnoliaceae. Peltate scales of
the kind which occur in Himantandra have not been recorded in the Mag*
noliaceae and other families which are usually regarded as being closely
related. It is interesting to note that stellate hairs and scales occur in certain
species of Hibbertia (Dilleniaceae) which Hutchinson (1113) regards as
derived from the Magnoliales. From a study of the wood, McLaughlin (1467)
concludes that Himantandra would be more appropriately placed with the
Annonales than the Magnoliales.

GENUS DESCRIBED
Himantandra.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Bailey, Nast, and Smith 94, Diels 583, Hutchinson 1113.

(ii) On Wood Structure

McLaughlin 1467, Record 1843, 1851.

12. LACTORIDACEAE
SUMMARY
A
small family consisting of the single shrub Lactoris fernandeziana Phil,
from Juan Fernandez. The wood exhibits the following features. Vessels
small, perforations simple, intervascular pitting alternate, members very short.
Parenchyma diffuse. Rays very broad and high, composed of upright cells.
Fibres with small bordered pits, extremely short.

LEAF
Cells of the lower
epidermis described by Solereder as papillose. Secre-
tory cells present in the spongy mesophyll.
 LACTORIDACEAE 39
Axis
STEM
Pericycle including crescent-shaped strands of fibres. Vascular bundles
separated from one another by medullary rays 2-3 cells wide. Tannin sacs,
frequently arranged in longitudinal rows, present in the pith.
WOOD 1

Vessels small than 100 ft mean tangential diameter); solitary and in

(less
radial multiples; up to 18 per sq. mm. Perforations simple. Intervascular
pitting alternate and small. Members less than 0-3 mm. Parenchyma
apotracheal, diffuse. Rays extremely high and as broad as the intervening
areas of fibres; composed entirely of upright cells. Fibres with numerous,
small, irregularly distributed, bordered pits; up to 0-5 mm. long.

TAXONOMIC NOTES
The genus Lactoris in the Piperaceae in the Bentham and
was included
Hooker system, but it is now generally regarded as having affinities with the
Magnoliaceae. McLaughlin, however, in a study of the wood anatomy of the
Magnoliales in 1933, came to the conclusion that the Lactoridaceae should be
transferred to the Piperales.

GENUS DESCRIBED
Lactoris.
LITERATURE
On Wood Structure
McLaughlin 1467, Record and Hess 1886.

13. TROCHODENDRACEAE
(Fie. 7 on p. 14; FIG. u on p. 40)

SUMMARY
The family is here treated as unigeneric, and comprising the single genus
Trochodendron as proposed by Smith (2139), the sole representative being
T. aralioides Sieb. et Zucc., a tree which occurs in Japan and Formosa, and is
cultivated elsewhere. One of the most striking anatomical features is the
diverse forms of sclerenchymatous idioblasts which occur in the leaf and in the
inner part of the primary cortex of the stem. The wood exhibits the following
characters. Vessels absent.The greater part of the tissue formed of extremely
long tracheids with circular to scalariform pits on the radial walls. Paren-
chyma diffuse. Rays of 2 sizes, the larger up to 12 cells wide; markedly
heterogeneous.
LEAF
Dorsiventral. Upper epidermis composed of cells with very thick outer
walls. Stomata (Fig. 1 1 lower surface, subtended by cuti-
F) confined to the
cular vestibules, and each accompanied by 2 horseshoe-shaped subsidiary
cells, the latter forming a circle around each of the stomata. Subsidiary cells,
at least in living specimens, also clearly demarcated from the remainder of
the epidermis by the presence of deposits of a gum-like substance secreted in
1
Based on the descriptions by McLaughlin (1467) and Record and Hess (1886).
 FIG. ii. TROCHODENDRACEAE, A, C-D, and F; TETRACEN TRACEAE, B and E;
EUPTELEACEAE, G and I; HIMANTANDRACEAE, H and J
A, Trochpdendron aralioides S.etZ. Cortical tissue showing lacunae and sclereids Xioo. H>Tetracentron
sinensis Oliv.Stomata from lower epidermis of leaf x 167. C, Trochodendron aralioides S. et Z.
Petiole X 13. D, T. aralioides S. et Z. Stem X 8. E, Tetracentron sinensis Oliv. Petiole x 13. F,Trocho-
dendron aralioides S. et Z. Stomata from lower surface of leaf X 167. G, Euptelea pleiosperma Hook,
f. et Thorns. Petiole X 28. H, Himantandra belgraveana Diels. Petiole x 19. I, Euptelea pleiosperma
Hook. f. et Thorns. Stem X 19. J, Himantandra belgraveana Diels. Stem x 12.
In Figs. C and D well-developed intercellular spaces and numerous sclerenchymatous idioblasts
occur in the inner but not in the outer part of the cortex.
 TROCHODENDRACEAE 41
the cells. Mesophyll consisting of 2, or locally of 3, layers of palisade tissue
and very lacimar spongy tissue. Intercellular spaces of the spongy tissue
occupied by very large, branched, sclerenchymatous idioblasts (Fig. n A).
Vascular bundles of the veins embedded in the mesophyll, the larger ones
accompanied by fibres. Inner cortical region of the petiole (Fig. 1 1 c) very
lacunar, with idioblasts, somewhat resembling those of the lamina, in the
intercellular spaces.
Idioblasts in the inner lacunar tissue of the petiole, described by Foster
(695, 696) as occasionally in longitudinal series 01*3 connected cells, but more
usually isolated by parenchymatous elements. The laminar idioblasts, accord-
ing to the same author, vary from 'radiately branched elements with dichoto-
mous arms to bizarre asymmetrical, cruciform and fibre-like forms'. Idioblasts
vary in frequency in different specimens; small clusters of irregular thick-
walled types noted by Foster in certain material from Formosa 'above the
juncture of the veins supplying each marginal tooth'. Vascular system of the
petiole appearing, in transverse sections through the distal end of material
grown at Kew, as a solitary, slightly interrupted, arc-shaped strand, mostly
composed of xylem consisting of radial rows of tracheids.
According to Bailey and Nast (92), in large leaves 5-7 vascular strands enter
the base of the petiole, coalescing at a higher level to form an arc, sometimes
accompanied by 2 adaxial bundles, this structure extending upwards through
the petiole and midrib of the lamina. In small leaves 1-3 bundles enter the
base of the petiole. Secretory cells with unidentified contents noted at Kew
in the petiolar ground tissue.

Axis
YOUNG STEM (Fig. n
D)
Cork said to arise in the sub-epidermis. Inner part of the primary cortex
very lacunar; containing branched sclerenchymatous idioblasts in the inter-
cellular spaces. Pericycle including an almost continuous ring of scleren-

chyma, consisting mainly of fibres. Phloem devoid of mechanical elements

in the young material examined at Kew, but secondary phloem from old bark
said by Bailey and Nast (92) to include irregular masses of dense, non-fibrous,
crystalliferous sclerenchyma. Vascular bundles more or less individually
distinct in transverse sections; separated by conspicuous primary rays 1-3
cells wide, the component cells of the rays having relatively thick walls.
Xylem consisting mostly of radially arranged tracheids with horizontal
bordered pits. Pith including a fairly high proportion of thick-walled pitted
secretory cells. Scattered secretory cells with gum-like contents also noted
in the primary cortex. Solitary crystals recorded in the primary cortex, but
none observed in material examined at Kew. Occasional cells containing

solitary crystalsnoted in the region of the pericyclic fibres. Nodal anatomy.

According to Bailey and Nast (92), in large leaves 5-7 strands enter the base
of the petiole from a restricted portion of the vascular cylinder of the stem
and, in smaller leaves, 1-3 bundles occur in the corresponding position.

WOOD (Fig. 7 E-F)

Vessels absent. Parenchyma occurring in the late wood as scattered cells
and short tangential lines (Fig. 7 E). Rays of 2 distinct sizes, the larger up to
42 TROCHODENDRACEAE
12 cells wide and 1-2 mm. high; uniseriates numerous, composed of upright
cells;about 9 rays per mm. heterogeneous (Kribs's Type I-II A), with up to
;

10 marginal rows of square or upright cells; the cells of the multiseriate parts
definitely procumbent. Tracheids forming the greater part of the wood;
with bordered pits, mostly on the radial walls, the pits in the early wood
elongated (scalariform), those of the late wood circular, with slit-like aper-
tures that sometimes exceed the borders; pits to ray cells typically circular
(opposite), but sometimes transitional to scalariform in the early wood. With
a sharp distinction between the thin-walled early wood tracheids and the
thick-walled and radially flattened cells of the late wood.

TAXONOMIC NOTES
The between Trochodendron and the Magnoliaceae, Winteraceae,
affinities

Schisandraceae, &c., have been much discussed. The recent work of Smith,
together with that of Bailey and Nast (92), indicates that whilst there is a
close relationship between Trochodendron and Tetracentron^ these genera are
not very closely related to the Magnoliaceae, Degeneriaceae, Himantandra-
ceae, Winteraceae, Schisandraceae, Cercidiphyllaceae, and Eucommiaceae.
Whether Trochodendron and Tetracentron should be placed in the same or in
2 closely related families is a matter of opinion. The recent work of Croizat
(501) indicates that there are considerable differences between the 2 genera.
In Croizat's opinion the 2 genera have no 'Ranalian' affinities, but both of
them are 'ultimately allied to the hamamelidoidsaxifragoid plexus*.

GENUS DESCRIBED
Trochodendron.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Bailey and Nast 92, Bailey and Thompson 99, Croizat 501, Foster 695, 696, Nast and
Bailey 1581, Smith 2139, Varossieau 2324.

(ii) On Wood Structure

and Nast 93, Bailey and Thompson 99, Dadswell and Record 533, Kanehira 1206,
Bailey
1209, McLaughlin 1466, 1467, Record 1783, 1843, 1851, Thompson 2256, Varossieau
2324, Wierletak 2422.

14. TETRACENTRACEAE
(FiG. ii on p. 40)

SUMMARY
The
family here treated as unigeneric, and comprising the single genus
is
Tetracentron as proposed by Smith (2139), the sole representative being
T. sinense Oliv., a small tree which occurs in China and Upper Burma. As
demonstrated by Bailey and Nast (92) Tetracentron differs from the closely
related Trochodendron in the absence from the former of sclerenchymatous
idioblasts, their place being taken by large, branched, secretory idioblasts
with 'resinous* contents readily stained with Sudan IV. The wood exhibits
the following features. Vessels absent. The greater part of the tissue formed
 TETRACENTRACEAE 43
of extremely long tracheids with circular to scalariform pits on the radial
walls. Some shorter and wider vascular tracheids present. Parenchyma
diffuse. Rays up to 4 cells wide; markedly heterogeneous.
LEAF
Dorsiventral. Stomata(Fig. B) confined to the lower surface in some
1 1 ;

instances accompanied on either side by a subsidiary cell parallel to the pore,

but in others surrounded by 4 or 5 well-defined subsidiary cells. (This last
arrangement may be a development from the first caused by subdivision of the
2 initial subsidiary cells. Some indication that this may be so is afforded by
the occasional occurrence of a single subsidiary cell on one side of a stoma
and a subdivided one on the other.) Cuticle sometimes striated in the region
of the stomata. M esophyll consisting of i layer of palisade cells and a very
lacunar spongy tissue below. Larger veins projecting considerably below the
general level of the abaxial surface. Vascular bundles of the smaller veins
embedded in the mesophyll, each of them partly or wholly surrounded by
sclerenchyma. Petiole (Fig. u
E), in transverse sections through the distal
end of material examined at Kew, exhibiting a single crescent-shaped vas-
cular strand with the ends somewhat incurved or almost meeting so as to
form a cylinder. Xylem in the petiolar vascular strand consisting chiefly of
radial rows of tracheids. According to Bailey and Nast (92), 3 separate bundles
enter the base of the petiole. The same authors noted, in transverse sections
of the petiole taken at the level of the stipular flanges, 3 conspicuous vascular
strands which coalesce towards the distal end to form a single, almost closed
vascular cylinder. Vascular cylinder again dividing at the base of the lamina
to form the 5-7 bundles in the principal veins. Secretory elements. Large,
branched, secretory idioblasts with 'resinous' contents readily stained in
Sudan IV, noted in the leaf. These secretory elements said to be unlike the
spherical secretory cells of the Winteraceae, Magnoliaceae, &c. A
few solitary
and clustered crystals noted at Kew in the 'cortical' and medullary' ground
*

tissue of the petiole.

Axis
YOUNG STEM
Cork arising in the sub-epidermis. Pericycle including a composite and
somewhat interrupted ring of fibres and stone cells. Phloem devoid of
sclerenchyma in the young stems examined at Kew, but Bailey and Nast (92)
refer to tangentially orientated clusters of sclerenchymatous elements in the
bark of old stems. Xylem in the form of a continuous cylinder traversed by
rays 1-3 cells wide; chiefly composed of radial rows of tracheids, somewhat
angular as seen in transverse section, each element up to about 30 /a in
diameter, with large, horizontal bordered pits. Pith composed of relatively
thick-walled pitted cells, containing starch and occasional clustered crystals.
Solitary crystals also noted in some of the pericyclic stone cells. Secretory
elements. Numerous with gum-like but unidentified contents noted in
cells
the phloem of very young stems. Occasional large cells with translucent con-
tents observed in the cortex of material cultivated at Kew. Bailey and Nast
refer to large, branched, secretory idioblasts with 'resinous' contents, readily
stained in Sudan IV, in the outer part of the cortex. Nodal anatomy.
Trilactmar according to Bailey and Nast.
 44 TETRACENTRACEAE
WOOD
Vessels absent. Parenchyma occurring in the late wood as scattered cells
and small tangential groups of 2 or 3 cells. Rays up to 4 cells wide and i -5 mm.
high; uniseriates numerous and composed of square to upright cells; about
16
rays per mm.; heterogeneous, with up to 20 marginal rows of square or
upright the cells of the multiseriate parts square to procumbent.
cells;
Tracheids forming the greater part of the wood; with bordered pits in the
radial walls, those of the early wood elongated (scalariform), those of the late
wood circular, usually with slit-like apertures that exceed the borders; pits to
ray cells typically circular (opposite), but sometimes transitional to scalariform
in the early wood. With a gradual change from thin walls in the early wood
to moderately thick walls in the late wood; the outer rows radially flattened.
Up to 4-5 mm. long (1466). Vascular tracheids with numerous, alternate,
circular pits on both radial and tangential walls, occurring in longitudinal
series; wider and shorter (o*i8-i'28, mean 0-47 mm., 1466) than the other
tracheids and resembling short vessel members, except for the absence of
perforations; Thompson and Bailey (2256) note these cells as occurring
particularly just below the internodes.
TAXONOMIC NOTES, see under TROCHODENDRACEAE
GENUS DESCRIBED
Tetracentron.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Bailey and Nast 92, Croizat 501, Smith 2139.

(ii) On Wood Structure

and Thompson 99, Kanehira 1209, McLaughlin 1466, 1467, Record 1843, Tang
Bailey
2231, Thompson and Bailey 2256,

15. ANNONACEAE
(Fic, 12 on p. 46; FIG, 13 on p. 48; FIG. 14 on p. 54)

SUMMARY
(i) GENERAL
Trees, shrubs, or climbers which occur in tropical and sub-tropical regions.
The hairs are mostly simple but stellate types and small peltate scales also
occur. The cells of the epidermis, especially of the leaf, contain solitary or
clustered crystals (Fig. 12 A) in several genera, their distribution being of
considerable diagnostic value. Secretory cells are constantly present in the
leafparenchyma and sometimes in the parenchymatous tissues of the young
stem as well Stone cells, frequently in the form of diaphragms, are
generally present in the pith, while sclerosed elements of various kinds also
occur in other parenchymatous tissues of the leaf and axis.

(ii)
WOOD
Vessels usually few, perforations simple, intervascular pitting alternate
and usually minute to small, pits to parenchyma similar, members of medium
 ANNONACEAE 45
length. Parenchyma apotracheal, in numerous fine lines, sometimes with
a little vasicentric in addition, often storied, Rays typically wide and high,
with few uniseriates, 3-16 (mostly 4-8) cells wide, slightly heterogeneous to

homogeneous, commonly containing oil or mucilage cells. Fibres with small

bordered pits, of medium length.

LEAF
Generally dorsi ventral, but see also below under 'Mesophyll'. Hairs
(Fig. 12 B) mostly simple, but stellate and peltate types also occur. Cells of
the lower epidermis provided with papillae interconnected by cuticular
ridges in Annona glauca Schum. et Thonn., Cleistopholis glauca Pierre, C.
staudtii Engl. et Diels, Enantia kummeriae Engl. et Diels. Epidermis multi-
seriate in species of Annona Cleistopholis, Ellipeia, Miliusa, Mitrephora,
>

Pachypodanthium, Xylopia; sometimes including solitary or cluster crystals.

Stomata rubiaceous; confined to the lower surface. Mesophyll in Arta-
botrys suaveolens Blume. including 2 rows of palisade cells towards the upper
and i towards the lower surface, with spongy tissue between them. Mesophyll
including fibres parallel to the surface of the leaf in certain species of Annona^
Anaxagorea, Asteranthe, Guatteria, Heteropetalum, Popowia, Sagerea, Unona y

Uvaria, and similar cells arranged vertically in Heteropetalum brasiliense

Benth. Branched sclerenchymatous cells also recorded as frequent in Annona,
Duguetia, Guatteria, Habzelia, Unona. Midrib provided with medullary
bundles within a ring of collateral strands in Cananga odorata Hook. f. et
Thorns, including a closed vascular cylinder in Uvaria zeylanica Linn,
according to Worsdell (2469), and a V-shaped strand in Artabotrys suaveolens.
Vascular bundles of the small veins vertically transcurrent. Petiole (Fig.
14 A~B), in transverse sections through the distal end, exhibiting an arc of
widely spaced collateral bundles, the number of strands varying in different
genera and species, e.g. with an arc of 3 large and several small bundles in
Monodora myristica Dun. Additional medullary strands present in the petiole
of Cananga odorata. According to Ozenda (1647) a single vascular arc enters
the base of the petiole in most Annonaceae, the solitary arc dividing into
3 strands near the base of the petiole. Two additional lateral strands are also
given off, but reunite more towards the distal end of the petiole. Transverse
sections through the middle part of the petiole thus show an arc of 5 bundles,
and sections through the distal end an arc of 3 bundles. Uvaria differs from
Annona and related genera in having more numerous petiolar vascular strands
arranged in a circle, the structure thus recalling that of the petiole of the
Magnoliaceae. Large stone cells scattered in the parenchymatous tissues of
the petiole in Monodora myristica and Uvaria virens N. E. Br. Mucilage
cells recorded in certain species of Annona> Rollinia, and Habzelia. Crystals
(Fig. 12 A) solitary and clustered; present especially in the epidermis and to
a lesser extent in the'mesophyll and around the vascular bundles of the veins.
Considerable diagnostic value was attached by Solereder to the distribution
of crystals in the epidermal cells, the following more or less distinct but over-
lapping types being recognized, (i) A single solitary or clustered crystal
present in each epidermal cell. This type recorded in 182 species belonging
to 25 genera, including species of Annona, Artabotrys^ Asimina, Bocagea,
Duguetia, Goniothalamus, Melodorum, Rollinia, and Stormia. (ii) A single,
46 ANNONACEAE
oblique, solitary crystal of variable form and size present in each epidermal
cell. This type recorded in 56 species belonging to 10 genera including species
of Asimina (i sp.), Duguetia (i sp.), Melodorum, Miliusa, and Orophea.
(iii) A
single clustered or solitary crystal confined to some of the epidermal
cells,the intervening cells being filled with brown contents, (iv) single, A
oblique, solitary crystal present in some but not all of the epidermal cells, the
intervening cells being filled with brown contents, e.g. in Sageraea. (v) Epi-
dermal crystals confined to small cells above the veins, e.g. in species of
Alphonsea, Ephedranthus, and Mitrephora. (vi) Vascular bundles sheathed by
crystalliferous cells in Cyathocalyx (i sp.) and Stelechocarpus (i sp.).

FIG. 12. ANNONACEAE

A, Epidermal Mitrephora obtusa Hook. f. et Thorns. B, Peltate hair, with a piece
crystal-cells of
of the epidermis from the lower side of the leaf of Duguetia bracteosa Mart. By Solereder.

Axis
YOUNG STEM (Fig. 14 D)
Cork arising superficially in the few examined species of Artabotrys (Santos,
1990), Asimina^ Cananga, Monodora, Stormia, Uvaria. Cortex containing
variously shaped stone cells in many species, e.g. in Asimina triloba Dun. and
Monodora myristica Dun. Pericycle usually including strands of fibres
situated externally to the phloem. Phloem, in transverse sections, appearing
as triangular groups with the broad portions directed towards the interior;
becoming stratified owing to the development of tangential groups of fibres.
Portions of the rays between the phloem groups also triangular, but with
inwardly directed apices. Xylem generally in the form of a closed ring
traversed by narrow rays, but sometimes interrupted by the conspicuous and
relatively broad primary rays; including vessels with simple perforations.
The fine tangential lines of apotracheal parenchyma, present in the mature
secondary xylem (see Wood* below), are not always well defined in the young
stem. Pith segmented by diaphragms of stone cells in all examined members
of the family except species of Asimina and Monodora. Small plates of stone
cells sometimes occur sporadically in the pith even in the absence of
complete
septa. Secretory cells, with tanniniferous contents, common in the cortex.
 ANNONACEAE 47
WOOD (Fig. 13 A-F)
Vessels very small than 50 ^ mean tangential diameter), e.g. in some
(less
species of Afphonseopsis, Malmea, Orophea, Oxandra, and Popowia, to large
(more than 200 ji), e.g. in some species of Cananga, Cleistopholis, Guattaria,
Leptopoda, Rottinia, and Unona\ solitary and in short multiples, often with
some radial pattern owing to the arrangement of the solitary and paired vessels
between the large rays radial multiples of 4 or more cells moderately common
;

in Goniothalamus p.p. and Hornschuchia and with numerous clusters and a

tendency to an ulmiform pattern in the late wood of Asimina', 1-45 per sq,
mm.; fewest (2 or less per sq. mm.) in the species with large vessels and in
some species of Platymitra, Unonopsis, and Xylopia; most numerous (30 or
more per sq. mm.) in some species of Malmea and Oxandra; ring-porous in
Asimina and semi-ring-porous in Pachypodanthium spiral thickening reported
;

(1886) in Asimina. Perforations simple, horizontal to slightly oblique. Inter-

vascular pitting typically alternate, but, according to Record and Hess (1886),
sometimes opposite, e.g. in Cymbopetalum', minute to moderate-sized; vessels
seldom touching the rays, and pits to ray cells very rare; pits to parenchyma
cells similar to the intervascular pitting, sometimes unilaterally compound

(533) an d occasionally with coalescent apertures, e.g. in Anaxagorea acuminata

St. Hil.; without pits to fibres (1154). Solid deposits often present; calcium
carbonate observed by Molisch in Annona laevigatea Mart. Mean member
length 0-3-0-6 mm. Parenchyma apotracheal, in numerous, closely spaced,
fine lines, usually uniseriate, sometimes 2 or 3 cells wide (Fig. 13 A and D) and
often reticulate the cells often disjunctive with very distinctly grouped pits in
;

the radial walls; vasicentric parenchyma in addition sometimes present in the

genera with larger vessels, but rather sparse. Storied in Annona p.p., Asimina,
Cyathocalyx, Goniothalamus, Heteropetalum, Hexalobus, Miliusa, Monodora,
Popowia, and Rollinia, and rather vaguely storied in Cleistopholis, Guatteria,
Mitrephora, Polyalthia, and Stenanthera. Large, thin- walled mucilage cells
present in Cananga and Cymbopetalum (1886). Strands usually of 4 to
8 cells, but strands of 2 cells sometimes numerous, e.g. in Asimina and
Hornschuchia. Rays typically wide and high most commonly up to 4-8 cells
;

wide, but rays up to 13-15 cells wide recorded (1886) in Anaxagorea, Annona,
Guatteria, and Stenanona and only 3-4 cells wide in some species of Alphonsea,
Cymbopetalum, Disepalum, Diclinanona, Fissistigma, Hornschuchia, and Mal-
mea', up to 6 mm. high in some species, the larger rays often showing evidence
of dissection into smaller units uniseriates typically very few and sometimes
;

extremely few, e.g. in some species of Cleistopholis, Hexalobus, Monodora,

Pachypodanthium, Popowia, and Unona, though moderately numerous in
some species of Cyathocalyx, Heteropetalum, Oxandra, and Uvaria, and
usually composed of mixed upright and procumbent cells, with the latter
predominating, but sometimes wholly of either procumbent or square cells.
Rays 2-9, mostly 3-5, per mm. Cells often very irregular in size as seen in
tangential sections (Fig. 13 c), with occasional rows of large, almost square
cells interspersed among the smaller procumbent cells, e.g. in some species
of Bocagea, Griffithia, Griffithianthus, and Polyalthia. Typically almost or
quite homogeneous (Kribs's Types I and II), though sometimes distinctly
heterogeneous, e.g. in Miliusa velutina Hook. f. et Thorns. with marked dif-
;

ferences in the size and shape of the procumbent cells even in species of the
 48 ANNONACEAE
same genus, possibly associated with differences in habit; sometimes with
very narrow procumbent cells, from which occasional marginal rows may
stand out as distinctly higher; sometimes composed almost entirely of large,

FIG. 13. ANNONACEAE, A-F; EUPOMATIACEAE, G-H

A, Oxandra lanceolata Baill. B, Aberemoa asterotricha Diels, with large oil cell. C, Grtffithianthus
fuscus Merrill. D, Anaxagorea acuminata St. Hil. E, Xylopia ferruginea Baill. F, Alphonsea maingayi
Hook. f. ct Thorns. G, Eupomatia laurina R. Br. H, E. laurina R. Br. O.C., Oil or mucilage cell.

nearly square, cells, from which the marginal rows are not very clearly dis-

tinguished in tangential section. The rays with small procumbent cells some-
times homogeneous, e.g. in Duguetia, Hexalobus, Rollinia and Stenanthera. y

Typically without crystals, but with numerous, small crystals in some species
of AlphonseopsiSy Goniothalamus, and Hornschuchia\ Gonggrijp (794) reports
silica in Cyathocalyx. Oil or mucilage cells scattered irregularly among the
 ANNONACEAE 49
ray cells m Aberemoa
(Fig. 13 B), Cananga (1154), Cleistopholis Cyathocalyx,

p.p., Cymbopetalum (1886), Diclinanona, Duguetia, Mezzettia, Monodora^

Pachypodanthium, and Unona. Fibres with small, but often distinctly bordered
pits, which are equally numerous on both radial and tangential walls; the
borders about the same size or slightly larger than those of the intervascular
pitting. Walls thin to moderately thick. Mean length 0*9-1*5 mm.

TAXONOMIC NOTES
It is interesting to note that the secretory cells in the leaf of the Annonaceae
are of the same type as those found in the Magnoliaceae. Worsdell (2469) has
stated his opinion that the vascular structure of Annonaceae recalls that of the
Magnoliaceae and Paeonia spp.
The wood anatomy
of the family is very uniform, and the genera, with the
exception of Asimina, are not easily distinguishable. The simple perforations
and mean member length of the vessels, the frequently storied parenchyma
and the scarcity of uniseriate rays all indicate a moderately high level of
specialization.
ECONOMIC USES
The Custard Apple is Annona reticulata Linn. Artabotrys suaveolens Blume
from the Philippine Islands contains an alkaloid which may be of medicinal
value. Artabotrys odoratissimus R. Br., from India and Ceylon, is cultivated
for its fragrant flowers. Cananga odorata Hook. f. et Thorns, is used for per-
fumery purposes. The aromatic and pungent fruits of Xylopia aethiopica A.
Rich are used by West African natives as a condiment and in medicine.
Lancewood, Oxandra lanceolata (Sw.) Baill. is the only timber of commer-
cial importance. Schneider (2044) notes that in the Philippines some species
are used locally for house- building, but are not distinguished commercially.
Pearson and Brown (1679) include 3 genera among the commercial timbers
of India, Miliusa, used for tool handles, oars, &c., Polyalthia, used for matches
and packing-cases, and Sageraea, used for bentwood and tool handles.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aberemoa, Anaxagorea, Annona, Artabotrys, Asimina,* Asteranthe,
Cananga,* Cleistopholis, Duguetia, Ellipeia, Guatteria, Habzelia, Hetero-
petalum, Miliusa, Mitrephora, Monodora,* Pachypodanthium, Rollinia,
Stormia, Unona, Uvaria,* Xylopia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Aberemoa, Alphonsea, Alphonseopsis, Anaxagorea, Annona, Asimina,
Bocagea, Brieya, Cananga, Cleistopholis, Cyathocalyx, (Cymbopetalum),
Dennettia, (Diclinanona), Disepalum, Drepananthus, Duguetia, Enantia,
Fissistigma, Goniothalamus, Griffithia, Griffithianthus, Guamia, Guatteria,
Heteropetalum, Hexalobus, Hornschuchia, Malmea, Meiogyne, Mezzettia,
Miliusa, Mitrephora, Monodora, Orophea, Oxandra, Oxymitra, Pachypo-
danthium, Platymitra, Polyalthia, Popowia, Rollinia, Saccopetalum, Sageraea,
Sapranthus, Stelechocarpus, Stenanona, Stenanthera, (Trivalvaria), Unona,
Unonopsis, Uvariastrum, Uvariodendron, Xylopia.
4504 r
So ANNONACEAE
LITERATURE
(i) On General Anatomy
Ozenda 1647, Santos 1990, Worsdell 2469.
(ii) On Wood Structure
Bastos 147, Benoist 169, den Berger 179, 182, 183, Besson 186, Burgerstein 310, Cooper
and Record 461, Coster 481, Dadswell and Record 533, Desch 574, Garratt 747, 747A,
Gonggrijp 794, Hess 963, Hopkinson 1083, Howard 1088, Janssonius 1154, Jentsch 1175,
1177, Jones 1191, Kanehira 1206, 1209, 1214, Knbs 1283, Lecomte 1334, Normand 1607,
Pearson and Brown 1679, Pereira 1687, Pfeiffer, H. 1712, Pfeiffer, J. Ph. 1713, Record
1783, 1790, 1818, 1843, 1851, Record and Hess 1886, Riera 1937, Stone 2202, 2207,
Tupper 2295, Williams 2430.

16. EUPOMATIACEAE
(Fic. 13 on p. 48; FIG. 14 on p. 54)

SUMMARY
(i) GENERAL
A
family of Australian shrubs included in the single genus Eupomatia. The
rubiaceous stomata in the leaf and secretory sacs in the pith of the young
axis are two of the most significant anatomical features.

(ii) WOOD
Vessels small, with a
radial pattern, perforation plates scalariform, inter-
vascular pitting scalariform to opposite members very long. Parenchyma
;

rather sparse, diffuse, and paratracheal. Rays very broad and high, with
sheath cells. Fibres septate, of medium length to moderately long.

LEAF
Weakly dorsiventral almost glabrous. Cells of the epidermis with straight
;

or somewhat curved anticlinal walls those of the lower epidermis with rather
;

more sinuous anticlinal walls. Stomata confined to the lower surface always ;

rubiaceous, but sometimes with more than one pair of subsidiary cells parallel
to the pore. Mesophyll including a single layer of very short palisade cells,
the remainder consisting of homogeneous, somewhat spongy parenchyma.
Midrib including a deep, crescent-shaped vascular strand strongly supported
by sclerenchyma on the adaxial side and between the arms of the crescent.
Vascular bundles of the smaller veins embedded in the mesophyll. Petiole
(Fig. 14 c), in transverse sections through the distal end, exhibiting a deep
crescent of widely spaced vascular bundles, with a group of stone cells in the
centre of the petiole between the arms of the vascular crescent. Ozenda (1647)
observed 7 vascular strands in the base of the petiole, and concluded that the
nodal structure resembles that of the Magnoliaceae. Secretory cells with
amorphous contents scattered in the ground tissue of the petiole, and a few
secretory idioblasts, sometimes with yellowish granular contents, scattered
throughout the mesophyll. Ground tissue of the petiole also including small
cluster crystals.
 EUPOMATIACEAE 51

Axis
YOUNG STEM (Fig. 14 E)

Epidermis with very thick cuticle. Cork formation not seen in material
examined by the author. Outer part of the primary cortex collenchymatous.
Pericycle including well-developed but widely spaced groups of fibres with
wide lumina. Phloem and xylem traversed by relatively broad rays. Pith
wide, composed of large, thin-walled parenchymatous cells, but supported
by a few scattered stone cells. Secretory elements. Somewhat elongated
secretory cells with amorphous contents present in the cortex, and smaller
more numerous ones in the phloem. Elongated, slightly sinuous, secretory
sacs with contents, each sac surrounded by small parenchymatous
amorphous
cells, present in the pith. Small cluster crystals abundant in the ground
tissue of the very young stem.

WOOD (Fig. 13 G~H)

Vessels small (mean tangential diameter about 70 /x), solitary and in short
to long radial multiples, which often give a distinct radial pattern (Fig. 13 H).
Perforation plates scalariform, with many fine bars (20 to more than 100).
Intervascular pitting small to moderate-sized, opposite or scalariform; pits to
ray cells similar. Mean member length about 1-2 mm. Parenchyma rather
sparse, diffuse, and scanty paratracheal (Fig. 13 H). Rays of 2 distinct sizes,
the larger up to 12, usually 6 to 9, cells wide and up to 5 mm. high; uniseriates
rather few and low, composed of high upright cells; about 10 rays per mm.;
multiseriate rays composed almost entirely of square and upright cells; sheath
cells usually present, sometimes very high. Fibres with simple or indistinctly
bordered (747) pits, septate and with moderately thick walls. Mean length
about i -6 mm.

TAXONOMIC NOTES
Although Eupomatia has features in common with the Annonaceae it is
generally treated as the only genus in a distinct family. Both Garratt (747)
and Lemesle (1353) consider that differences in the wood structure support
this separation. Lemesle concludes from the anatomy of the wood and the
floral morphology that the Eupomatiaceae are among the most primitive of

angiosperms, more primitive even than the Magnoliaceae.

GENUS DESCRIBED
Eupomatia.*
* Kew
Represented in the slide collection.

The above description of the general anatomy is based on material of

E. bennettii F. Muell, cultivated at Kew.

LITERATURE
(i) On General Anatomy
Lemesle 1354, Ozenda 1647.

(ii) On Wood Structure

Garratt 747, Lemesle 1353, Record 1843,
 (52)

17. MENISPERMACEAE
(Fic, 14 on p. 54; FIG. 16 on p. 60; FIG. 91 on p. 402.

SUMMARY
(i) GENERAL
Thefamily includes shrubs and small trees but most members are lianes.
It occurs chiefly in tropical and sub-tropical regions. The vascular bundles
in young stems are individually distinct even with the naked eye, being
separated by the broad primary medullary rays which may be lignified or
unlignified. In the xylem, the vessels have wide lumina, and simple perfora-
tions, whilst the ground tissue consists of prosenchymatous elements with
bordered pits. The pericycle includes an undulating, generally continuous
ring of sclerenchyma formed from arcs of fibres, belonging to the separate
bundles, with groups of stone cells between them. Anomalous secondary
thickening is very common, and is due to the formation from extrafascicular
cambia, of one or more concentric or eccentric rings of bundles. (For further
details see under 'Wood'.) Crystals of calcium oxalate are common, and

may be solitary, clustered, or secreted in the form of raphides, styloids,

and crystal-sand. Secretory sacs, with various kinds of contents, are
characteristic of certain genera, whilst idioblasts and other types of isolated
sclerenchymatous elements also occur. Hairs include the following kinds,
(i) Uniseriate, consisting
of 2 or more cells, (ii) Glandular types, which may
be shaggy, long and uniseriate or small, (iii) Unicellular, ellipsoidal, or club-
shaped. The stomata are very variable, especially in the nature of the
surrounding epidermal cells, and no single type can be regarded as typical
of the family as a whole.

(ii) WOOD
Woodstypically with included phloem of the 'concentric* type. Vessels
very small to medium-sized, nearly all solitary and seldom touching the rays,
perforations simple, intervascular pitting alternate, pits to parenchyma similar
or elongated and grouped, members of medium length to moderately short.
Parenchyma (a) conjunctive, (b) apotracheal, diffuse or in short bands.
Rays interfascicular only. Fibres with numerous bordered pits, of medium
length.
LEAF
Generally dorsiventral; centric in Coccylus leaeba DC. according to Beau-
quesne (162).
Hairs, (i) Uniseriate, bicellular, in species of Abuta, Adeliopsis, Albertisia,
Antizoma, Bania, Carronia, Chondodendron, Cissampelos, Cocculus, Cyclea,
Detandra, Husemannia, Hyperbaena, Limacia, Macrococculus, Menispermum^
Pachygone, Paraphora, Pericampylus, Pleogyne, Pycnarrhena, Sciadotenia,
Syrrhonema, Tiliacora. (ii) Uniseriate, of more than 2 cells, in species of
Anamtrta, Arcangelisia, Calycocarpum, Chasmanthera, Coscinium Disciphania, y

Heptacyclum, Menispermum, Parabaena, Stephania, Tinospora. (iii) Small,

unicellular, ellipsoidal or club-shaped glandular hairs, in species of Kolobo-
petalum, Mieniophyton, Tinospora. (iv) Long, uniseriate glandular hairs, in
 MENISPERMA CEA E 53

species of Calycarpum, Parabaena. (v) Shaggy glandular hairs recorded in

Jateorhiza.
Epidermis, (i) Cells specially large in species of Cocculus. (ii) Cells partly
sclerotic in Anomospermum reticulatum Eichl., Hyperbaena laurifolia Urb.,
Triclisia loucoubensis BailL (iii) Papillose: (a) on both surfaces in Cissampelos

capensis Thunb., C. pareira Linn, (sometimes), Cocculus leaeba DC. and

Stephania rotunda Lour; (b) on the lower surface only, in some instances
confined to isolated cells above the veins, in Cissampelos fluminensis Eichl.,
C. glaberrima St. Hil., Hypsipodes subcordatus Miq., Jateorhiza columba (Lam.)
Miers., Menispermum canadense Linn., M. dauricum DC., Stephania abyssinica
Walp. Stomata exhibiting no characteristic structure; generally confined to
the lower surface, but recorded on both surfaces in species of Antizoma and
in Cocculus cebatha DC. Rubiaceous in Cocculus carolinus DC. and C. lauri-
folius DC. Rosettes of subsidiary cells recorded in certain species of Albertisia,
Anamirta, Bania, Burasaia, Carronia, Cocculus, Cyclea, Fawcettia, Hepta-
cyclum, Husemannia, Hyperlaena, Leichardtia, Limacia^ Macrococculus^ and
thin-walled subsidiary cells in Chlaenandra. Subsidiary cells narrow in
Parabaena, Pleogyne, Pycnarrhena, Sarcopetalum. Stomata in groups in
Macrococculus pomiferus Becc., Antizoma spp. Hydathodes recorded by
Engler (632) in Anamirta and related genera. Hypoderm
mucilaginous in
certain species of Adetiopsis,Anomospermum, Cissampelos, Cyclea, Limacia^
Stephania. MesophylL Thickenings or swellings present on the cell walls
in Adeliopsis, Canonia, Cissampelos, Cocculus, Cyclea, Limacia, Pachygone,
Pericampylus, Stephania, Syrrhonema. Spongy mesophyll partly or wholly
composed of thick- walled, generally lignified cells, e.g. in Albertisia papuana
Becc., Anamirta cocculus Wight et Arn,, Husemannia protensa F. v. Mull.,
Parabaena sagittata Miers. Idioblasts or similar isolated sclerosed cells
recorded in Abuta concolor Poepp. et Endl., Chlaenandra ovata Miq., Chon-
dodendron platyphyllum Miers., Coscinium blumeanum Miers., Detandra
paraensis Eich. 'Spicular cells' of various types present in Adeliopsis decumbens
Benth., Anamirta cocculus Anomospermum japurense Eich., Arcangelisia
',

lemniscata Becc., Burasaia congesta Decne., B. gracilis Decne., B. madagas-

cariensis DC., Heptacyclum zenkeri Engl., Limacia cuspidata Hook. f. et Th.,
and L. microphylla Miers.
Vascular bundles of the veins generally accompanied by sclerenchyma
except in certain species of Antizoma, Cissampelos, Desmonema, Diploclisia,
Disciphania, Miersiophyton, Stephania, Tinospora, Lateral and smaller veins
vertically transcurrent in species of Abuta, Anomospermum, Coscinium, Hyper~
baena, Limacia, Macrococculus, Menispermum, Pachygone, Paraphora, Peri-
campylus, Pycnarrhena, Sarcopetalum^ Triclisia.
Transverse sections through the central portion of the petiole (Fig. 14 F)
exhibiting a ring of isolated bundles in certain species of Anamirta, Arc-
angelisia, Cissampelos, Cocculus, Coscinium, Fibraurea, Limacia, Menispermum^
Pachygone, Pycnarrhena, Stephania, Tiliacora, Tinomisdum, Tinospora. Swell-
ings sometimes present at the distal and basal ends of the petiole or at the distal
end only. Anatomical structure of the swollen regions differing from that of
the remainder of the petiole. No correlation was found by Rudolph (1966) to
exist between species resembling one another in respect of the structure of
these swellings and those resembling one another in fruit and seed structure.
54 MENISPER MA CEA E
Crystals secreted in the form of needles, prisms, rhombohedra, solitary
types or clusters, raphides and crystal-sand, (i) Large solitary crystals
recorded in Aspidocarya, Burasaia, Chasmanthera, Chlaenandra, Fawcettia,
Hypsipodes, Kolobopetalum Leichardtia, Syntriandrium, Tinospora. (ii) Small
y

H
FIG. 14. ANNONACEAE, A-B and D; EUPOMATIACEAE, C and E;
MENISPERMACEAE, F-H
A, Cananga odor at a Hook. f. et Thorns. Petiole xg. B, Asimina triloba Dun. Petiole X 15. C,
Eupomatia bcnnettii F. Muell. Petiole X 8. D, Monodora myristica Dun. Stem X 7. E, Eupomatia
bennettii F. Muell. Stem x 8. F, Stephania rotunda Lour. Petiole x 33. G, Menispermum canadense
Linn. Stem x$. H, M. canadense Linn. Young stem x 15.

rhombohedral crystals in Abuta> Anomospermum, Burasaia, Heptacyclum,

Husemannia, Hyperbaena, Limacia,
Macrococculus, Pachygone, fleogyne,
Pycnarrhena, Tiliacora\ said to be particularly abundant in epidermis of
Anamirta cocculus Wight et Arn., Desmonema and Tiliacora warneckei EngL
(iii) Styloid-shaped or prismatic crystals recorded in Cissampelos, Cycled,

Diplodisia macrocarpa M.izrB.,Jateorhiza columba (Lam.) Miers., Pericampylus

 MENISPERMACEAE 55
incanus Miers., Stephania. (iv) Large styloid-like, geniculate, hemitropic
crystals present in Leichardtia clamboides F. v. Mull, (v) Large cluster crystals
recorded in Chondodendron platyphyllum Miers. (vi) Small cluster crystals in
Macrococculus pomiferus Becc. and Pericampylus incanus Miers. (vii) Raphides
recorded by Santos (1992) in Anamirta cocculus Wight et Arn. and Arcan-
gelisia flava (Linn.) Merrill, (viii) Paired crystal cells occur in Leichardtia
clamboides. (ix) Sphaero-crystalline masses of an unknown chemical sub-
stance recorded in the lower epidermis of Cocculus laurifolius DC, and Sarco-
petalum harveyanum F. v. Muel.
Secretory sacs in the lamina generally occur in association with the
sclerenchyma of the veins; comparatively rare in the mesophyll. Secretory
sacs also present in the petioles as well as in the axis. (They are believed,
according to Solereder and to Santos (1992), to arise from vertical rows of
cells of which the transverse walls break down, but this view has been disputed
by Beauquesne (162), who considers them to be truly unicellular. Nature of
the contents very variable.)
(i) Elongated sacs with yellowish or brown contents recorded in Albertisia,
Anamirta, Burasaia, Chasmanthera, Cissampelos, Cocculus, Desmonema, Disci-
phania y Fawcettia, Hypsipodes, Jateorhiza, Kolobopetalum y Limacia, Para-
baena, Syntriandrium^ Tiliacora, Tinospora. (ii) Short sacs with reddish-brown
contents said to occur in "Antizoma calcarifera Miers.', 'A, lycioides Miers.',
Cissampelos fasciculata Benth., C. pareira Linn., DiploclisiamacrocarpaNliers.,
Tiliacora racemosa Colebr., Triclisia loucoubensis Baill. Secretory receptacles
of intercellular origin and often enclosed by the arc-shaped cells of the
spongy tissue, recorded in Anomospermum japurense Eichl. and Tinospora
bakis Miers.
Rosettes of silicified cells present in Coscinium blumeanum Miers. Silicious
contents recorded in tubular cells at the ends of the smaller veins of Arc-
angelisia lemniscata Becc. Berberin recorded in Coscinium fenestratum Colebr. ,

Jateorhiza palmata (Lam.) Miers., Menispermum canadense Linn.

Axis
YOUNG STEM (Fig. 14 G-H)
Cork formation observed in only a few species; developing very late in
Menispermum canadense Linn. According to Moller, as cited by Solereder,
cork formation in this species is at first superficial and confined to certain
areas of the stem, *and then spreads peripherally as it penetrates deep into the
1
cortex Primary cortex sometimes including groups of stone cells, e.g. in
.

Anamirta cocculus Wight et Arn. Pericycle including crescents of fibres

opposite and often in contact with the phloem strands, but fibres sometimes
becoming united by stone cells to form a composite, continuous, sinuous ring
of sclerenchyma. Composite ring said by Solereder to persist in thick stems of
Cissampelos pareira Linn, and Hyperbaena domingensis Benth., but the arcs
of fibres become split into groups separated from one another by stone cells
in the last of these species. Vascular bundles separated by broad, lignified
or unlignified primary medullary rays. Vessels large and conspicuous, with
simple perforations. Pith generally broad. Inner part, e.g. according to
Solereder in Anamirta cocculus, Coscinium blumeanum Miers. and Limacia
 56 MENISPERMACEAE
velutina Miers., consisting ofempty cells, but peripheral part composed of
vertically elongated, pitted cells, sometimes with thin transverse septa.
Fibrous cells resembling phloem fibres sometimes present on the inner side
of the groups of primary xylem, especially in species, e.g. Aspidocarya uvifera
Hook. Thorns, and Cocculus laurifolius DC., which exhibit no clear
f. et
between the inner and outer part of the pith. Crystals and
differentiation
secretory sacs of the same types as described under 'Leaf '.

WooD 1
1 6 D and F and 91 A)
(Figs.
Vessels small (mean tangential diameter less than 100 p) to medium-sized
(100-200 ft), and occasionally very small (25-50 7*), e.g. in Cocculus\ mostly
solitary, but occasionally in tangential groups of 2-4, which may give a tan-
gential pattern, e.g. in Tiliacora glycosmantha Diels.; 5-40 per sq. mm.;
seldom in contact with the rays. Perforation plates simple, slightly oblique.
Intervascular pitting rather scarce, alternate and moderate-sized; pits to
parenchyma usually round with slit-like apertures, but, particularly at the
ends of the vessel members, sometimes large, irregular in shape and elongated,
simple or with narrow borders, a cluster of such pits often resembling an
irregular scalariform perforation plate. Myers (1577) has described thick-
walled and pitted tyloses in Menispermum, Mean member length O'3-o*45 mm.
Parenchyma (a) conjunctive, between the successive layers of xylem and
phloem bundles, and including layers of isodiametric or radially elongated
stone the layers usually 2-3 cells wide, but often wider between the
cells,

phloem strands; sometimes containing numerous crystals; (b) apotracheal,

diffuse, and in short tangential lines; strands usually of 2-4 cells. Rays
interfascicular only, and usually not continuous radially from one layer of
bundles to the next. Up to 10-23 ce M s wide and usually extremely high; cells
all slightlyprocumbent to all square, with little distinction between central
and marginal cells, and sometimes almost homogeneous. Fibres with numer-
ous, distinctly bordered pits on both radial and tangential walls pits to vessels ;

similar, but more numerous; such cells often have wider lumina and are
irregular in shape (especially in Abuta and Tiliacora) and could possibly be
moderately thick. Mean length i-i-
classified as vasicentric tracheids; walls
i -2 mm. Included (interxylary) phloem (Figs. 91 A and 16 D and F) of the
'concentric* type (c.L drcumvallatum) observed or reported (85, 1712, 1851,
2158) in Abuta, Anomospermum, Cebatha, Chasmanthera, Chondodendron,
Cissampelos, (Root), Clyphea, Cocculus, Disciphania, Jateorhiza, Menispermum^
Pachygona, Pericampylus, Telitoxicum, and Tiliacora. The anomaly consists
of successive bundles of xylem and phloem repeating the structure of the
young stem; the bundles separated by tangential bands of parenchyma and
large interfascicular rays. Solereder lists the following lianes as having normal
structure: species of Anamirta, Cissampelos, Cocculus, Coscinium, Jateorhiza,
Menispermum^ and Tinospora.

RHIZOME
With a stele consisting of a single ring of bundles surrounded by a ring of
sclerenchyma in Menispermum canadense Linn.
1
Based entirely on material with anomalous structure.
 MENISPERMACEAE 57
ANOMALOUS STRUCTURE, see under *WOOD'
ROOT
In Cocculus leaeba DC. and
'Tinospora tuberculata Beaume.' described by
Beauquesne (162) as somewhat resembling the
aerial stem, but with the

xylem groups of the bundles united at the centre, thereby practically or

completely obliterating the pith, and giving the xylem a stellate appearance,
with phloem at the ends of the radii. For root structure of Jateorhiza palmata
(Lam.) Miers. see 'Economic Uses'.

TAXONOMIC NOTES
The
general stem structure, with the separate vascular bundles, suggests
that the members of this family have affinities with the woody genera of the
Berberidaceae. This is also confirmed by the occurrence of berberin in certain
genera of Menispermaceae. In the respective systems of Bentham and Hooker,
Engler, and Hutchinson, the Menispermaceae are placed close to the Ber-
beridaceae. This shows that the vegetative anatomy and the floral structure
both point to the same conclusion concerning the affinities of this family.
Garratt (747) states that the wood anatomy of this family indicates that it has
no affinity with the Myristicaceae.

ECONOMIC USES
The (Lam.) Miers., which is known com-
sliced root of Jateorhiza palmata

mercially as CalumbaColombo Root, is used medicinally for treating

or
indigestion. Commercial samples of Calumba Root consist of thick slices
about 2-6 cm. in diameter, somewhat depressed at the centre and covered
externally by a thin, brownish, wrinkled layer of cork. The greater part of the
ground tissue of the root is composed of parenchyma with abundant starch.
Other diagnostic characters include groups or a loose ring of yellow, thick-
walled, pitted stone cells in the outer cortical tissues; prismatic crystals in
some of the stone cells phloem in the form of narrow, radiating groups of
;

somewhat disorganized, unlignified tissue; radial groups of wide xylem vessels

with yellowish walls, embedded in parenchymatous ground tissue; more
irregularly distributed vessels towards the centre of the root. The walls of
the stone cells, and to a lesser extent of the xylem elements, are coloured
green on treatment with 60 per cent, sulphuric acid.
The root of Chondodendron tomentosum Ruiz, et Pav. known as Tareira
brava', has tonic and diuretic properties. The fruits of Anamirta paniculata
Colebr. are used in the preparation of ointments.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Abuta, Adeliopsis, Albertisia, Anamirta, Anomospermum, Antizoma,
Arcangelisia, Aspidocarya, Bania, Burasaia, Calycarpum, Carronia, Chas-
manthera, Chlaenandra, Chondodendron, Cissampelos, Cocculus,* Cos-
cinium, Cyclea, Desmonema, Detandra, Diploclisia, Disciphania, Fawcettia,
Heptacyclum, Husemannia, Hyperbaena, Hypsipodes, Jateorhiza, Kolobo-
petalum, Leichardtia, Limacia, Macrococculus, Menispermum,* Miersio-
phyton, Pachygone, Parabaena, Paraphora, Pericampylus, Pleogyne,
58 MEN1SPERMACEAE
Pycnarrhena, Sarcopetalum, Sciadotenia, Sinomenium,* Stephania,* Syn-
triandrium, Syrrhonema, Tiliacora, Tinomiscium, Tinospora, Triclisia.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Abuta, Chondodendron, Cocculus, Hyperbaena, (Menispermum), Teli-
toxicum, Tiliacora.
LITERATURE
(i) On General Anatomy
Beauquesne i6, Engler 632, Fiebng 684, Holm 1066, 1073, Myers 1577, Rudolph
1966, Sabnis 1977, Santos 1989, 1992, Shirley and Lambert 2089, Short 2093.
(ii) On Wood Structure
Chalk and Chattaway 362, Dadswell and Record 533, Garratt 747, Janssonius 1154,
Myers 1577, Pfeiffer, H. 1712, Record 1843, 1851, Record and Hess 1886, Williams 2430.

18. BERBERIDACEAE
(Fic. 15 on p. 58; FIG. 1 6 on p. 60; FIG. 17 on p. 65)

SUMMARY
(i) GENERAL
A
family occurring chiefly in North Temperate regions and consisting
partly of herbs with rhizomes and partly of shrubs which are usually spiny.

FIG. 15. BERBERIDACEAE

A-B, Crystals in the upper epidermis of the leaf of Mahonia japonica
DC. (after Vesque).

The spines of Bzrberis are mainly composed of sclerenchymatous fibres. In

transverse sections of young internodes of the woody genera the vascular
bundles are separated by the broad primary medullary rays, whilst a ring of
fibres, believed to be pericyclic in origin, forms a tube at the periphery of the
stem. In the aerial stems of the herbaceous or semi-herbaceous genera,
Achlys, Diphylleia, Epimedium, Hydrastis, Jeffersonia, Leontice, Podophyllum,
and Vancouveria, on the other hand, the bundles are widely spaced and
generally scattered. A
somewhat intermediate type is represented by Nandina
Thunb. because it possesses a ring of widely spaced bundles sur-
domestica
rounding large pith which occupies most of the cross-sectional area of an
a
old stem. The structure of the individual vascular bundles recalls that of the
bundles in the monocotyledons. The stomata are generally ranunculaceous.
The tissues are frequently coloured yellow by the presence of berberin.
Solitary or cluster crystals of calcium oxalate often occur.
 BERBERIDACEAE 59

(ii) WOOD
Vessels very small and usually with some ulmiform or radial pattern,
occasionally ring-porous, with spiral thickening, perforations simple, inter-
vascular pitting alternate, members moderately to extremely short. Paren-
chyma absent. Rays typically wide (6-12 cells), high and with few or no
homogeneous or almost so. Fibres with small simple pits, some-
uniseriates,
times with occasional septa and sometimes storied, moderately to extremely
short.

LEAF
Usually dorsiventral. Hairs infrequent, mostly unicellular or uniseriate.
Glandular hairs with a uniseriate stalk and ellipsoidal head occur in Epimedium.
Brown dots, resembling lenticels, recorded on the lower surface in Berberis
feddeana Schn., but similar bodies said not to be consistently present in other
species of Berberis. Lower epidermis papillose in certain species of Berberis,
Epimedium, Jeffersonia, and Mahonia. Stomata generally confined to the
lower surface ranunculaceous. Sclerenchyma sometimes present below the
;

upper epidermis in Berberis and Mahonia spp. Grooves present in the lower
surface of Berberis empetrifolia Lam. Mesophyll not clearly differentiated
into palisade and spongy portions in Epimedium according to Solereder, and
in Hydrastis according to Maue (1460). Midrib with a varying number of
vascular bundles, often collectively surrounded by a sclerenchymatous sheath,
especially in the Berberidoideae. According to Solereder the number of
bundles in the principal veins has taxonomic value as follows. One bundle in
Caulophyllum, Jeffersonia, and Leontice several in Achlys, Berberis, Diphylleia,
;

Epimedium, Hydrastis, Mahonia, Nandina, and Podophyllum. This is of

doubtful value, however, because it is also stated by Holm (1017) that in
Jeffersonia there are 2 bundles in the main veins. Petiole. Widely spaced
vascular bundles recorded in Epimedium, Podophyllum, and Vancouveria. An
arc of a few petiolar vascular strands noted at Kew in transverse sections
through the distal end in a few species of Berberis', a circle of bundles in the
corresponding region in Epimedium and Podophyllum, but with additional
medullary bundles in the last of these genera; an adaxially flattened ring of
bundles noted in Mahonia aquifolium Nutt. Wide secretory canals recorded
in the phloem of Caulophyllum thalictroides (L.) Michx. by Holm (1016).
Yellowish or greenish prismatic crystals (Fig. 15 A-B) of an organic substance
of unknown composition recorded in the upper epidermal cells of certain
species of Mahonia.
It has been claimed by Maue (1460) that the leaves of the genera Berberis ,

Epimedium, Hydrastis, and Mahonia can be distinguished from one another

by the following characters. Berberis by the polygonal cells of the upper
epidermis, oval unlignified stomata, and the absence of hairs; Epimedium by
the possession of multicellular hairs Hydrastis by the fact that the spongy and
;

palisade regions of the mesophyll are not distinct, and by the presence of
unicellular, lignified hairs on both surfaces Mahonia by sinuous anticlinal
;

walls to the cells of the upper epidermis, rounded, lignified stomata and the
absence of trichomes; Podophyllum by the presence of calcium oxalate crystals
in the mesophyll, and a single layer of palisade tissue.
 60 BERBERIDACEAE
Axis
YOUNG STEM (Fig, 17 A, B, and E)
Cork arising in the outer part of the pericycle in some species of Berberis,
but more superficial in others; sub-epidermal in
Podophyllum. Cork cells
usually comparatively thin-walled and with wide lumina. Pericycle generally

FIG. 16. BERBERIDACEAE, A-Band E; LARDIZABALACEAE, C;

MENISPERMACEAE, D and F
A, Berberi$ darmnii Hk. T.S. B, B. aristata DC. Ray. C, Akebia quinata Decne. T.S. D, Abuta
concolor Poepp. et Endl. T.S. E, Berberis tenuifolium Lindl. Ray. F, Tiliacora
glycosmantha Diels.
1 .o.

including a continuous ring of fibres in Berberis according to Harvey-Gibson

and Horseman (916), but observations at Kew suggest arcs of fibres external
to the phloem strands to be more common. The stem of Berberis aristata DC.
is stated by Short
(2095) to be distinguishable from that of allied species of
Berberis by the presence of large crescents of pericyclic fibres
adjoining each
phloem strand. Vascular bundles separated by broad primary rays in
Berberis, and Mahonia. Transverse sections of aerial stems exhibiting widely
spaced and generally scattered bundles in herbaceous and semi-herbaceous
 BERBERIDACEAE 61

genera such as Achlys, Diphylleia, Epimedium, Hydrastis, Jeffersonia, Leontice,

Podophyllum, and Vancouveria. Vessels mostly with simple perforations;
scalariform plates also occur in Herberts. Phloem with sieve plates on the
bevelled ends of the sieve tubes, and sometimes on the lateral walls as well.
Peripheral part of the pith of the woody genera consisting of cells with thicker
walls than those at the centre.

WOOD (Fig. 1 6 A-B and E)

Vessels very small (25-50 p, mean tangential diameter), the late-wood
vessels of Berberis sometimes little wider than the fibres and angular in cross-
section; commonly in irregular clusters (Fig. 16 A), which tend to be grouped
in ulmiform bands in Berberis, e.g. in B. aquifolium Linn., and in radial lines,
in at least the early rings, in Mahonia tenuifolium (Lindl.) Britton, and in
radial lines of almost solitary vessels in Nandina domestica Thunb. seldom
;

touching the rays; 25-50 per sq. mm. ring-porous in some species of Berberis,
;

e.g. B. aristata DC. and B. vulgaris Linn., and semi-ring-porous in some other
species of Berberis and in Nandina; with spiral thickening. Perforations
typically simple, slightly oblique scalariform plates with very few bars reported
;

by Solereder to be occasionally present in Berberis Tupper (2295) refers to

;

exclusively scalariform plates in Berberis. Imperfect vessel members frequent ;

these may be the spirally thickened 'tracheids* of Nandina, to which Solereder

refers. Intervascular pitting alternate, moderate-sized. Pits to ray cells very
rare, similar to the intervascular pitting. Mean
member length O'i5-o-27 mm.
Parenchyma absent. Rays typically high and wide; usually up to 6-12 cells
wide, but up to 25 cells wide in B. kavakana Hay (1206) commonly more than
;

2 mm. and sometimes more than 5 mm. high, the tall rays often showing
evidence of subdivision into smaller units; uniseriates very few or lacking;
about 3 rays per mm. homogeneous (Kribs's Type II) or with only a slight
;

difference between marginal and central cells in Berberis, though with

moderately distinct sheath cells in B. darwinii Hook. (Fig. 16 B); composed
entirely of small square cells in Mahonia tenuifolium with numerous crystals
;

in Mahonia. Fibres with moderately numerous, small, simple pits on the

radial and tangential walls. Occasionally septate in Berberis and Nandina.
Walls thin to moderately thick. Sometimes storied in Berberis and Mahonia.
Mean length O'3~o-75 mm.
RHIZOME
A single ring of vascular bundles commonly present in Achlys, Caulo-
phyllum, Diphylleia, Epimedium, Hydrastis, Jeffersonia, and Podophyllum, but
in all of these genera the bundles are sometimes in two rings or scattered.
Additional cortical bundles, consisting mainly of sclerenchyma, present
in species of Achlys, Diphylleia, and Podophyllum. Xylem of Vancouveria
hexandra C. Morr. et Dec. embedded in a ring of sclerenchyma except in
parts of old specimens, according to Harvey-Gibson and Horseman (916).
Pericycle generally devoid of fibres in herbaceous forms (except Epimedium),
and Leontice altaica Pall. Two zones of pericyclic fibres recorded in
Epimedium, Nandina, and a few species, of Mahonia.
The microscopical features of commercial Podophyllum rhizome are
described under 'Economic Uses*.
62 BERBERIDACEAE
ROOT
Very secondary thickening in the roots of Podophyllum. Endodermis
little

well defined in this genus. Mycorrhiza recorded in the thin roots of Caulo-
phyllum thalictroides (L.) Michx, and 'Jeffenonia diphylla (L.) Pers.'.

TAXONOMIC NOTES
The stemsof the woody genera Berberis and Mahonia are, at first sight,
very dissimilar in structure from the aerial stems of herbaceous genera such
as Podophyllum. It is quite possible that these differences are partly associated
with the diverse mechanical requirements of herbaceous and woody plants
from the taxonomic standpoint, it seems probable that,
respectively, although,
if woody genera have had a common ancestor, it must
the herbaceous and
have been somewhat remote from the present-day representatives of the
family. Himmelbaur (976), on the other hand, has expressed the view that
the stem anatomy of the Berberidaceae is such as to suggest that all the
members of the family have been derived from one type. The similarity of
the herbaceous genera with scattered bundles to certain of the Ranunculaceae
and hence to the Monocotyledons has already been described under Ranun-
culaceae. Harvey-Gibson and Horseman (916), however, take exception to
Solereder's statement that the stem structure of Jeffersonia is Monocoty-
ledonous. The stem structure of Berberis and Mahonia is rather suggestive
of that of the Menispermaceae. The presence of berberin in the Meni-
spermaceae and Berberidaceae is also noteworthy in this connexion.
The anatomy of the wood of the shrubby genera indicates a moderately
high level of specialization, particularly the very short vessel members, the
homogeneous rays and the libriform fibres, which are sometimes storied.

ECONOMIC USES
The fruits of numerous
species of Berberis and Mahonia are edible, whilst
the roots of Berberis spp. are used for medicinal purposes. A
dye is extracted
from the yellow bark and roots of Berberis spp. The dried rhizomes of
Podophyllum peltatum Linn, and P. hexandrum Royle from North America
and India, respectively, are of medicinal value because they contain podo-
phyllin. Microscopical features of the rhizome of P. peltatum include the
following. Epidermis of axially elongated cells filled with reddish-brown,
tanniniferous contents. Cork arising in the hypoderm, consisting of 2 or 3
layers of cells. One or two layers of large, somewhat collenchymatous, empty
cells present beneath the cork. Cortex and pith parenchymatous, most of the
cells containing simple or compound starch grains. Isolated cluster crystals
scattered throughout the parenchymatous tissues. A
few of the parenchyma-
tous cells filled with a yellow secretion. Endodermis not distinct in the
rhizome, although well defined in the root. Pericycle, in some instances,
marked by small strands of fibres situated externally to the vascular bundles.
(In at least one sample in the Kew Museums these fibres were very infrequent.)
Vascular system consisting of a circle of 20 to 30 collateral vascular bundles,
separated from one another by broad rays. Many of the xylem vessels filled
with yellow contents. Groups of sclerenchymatous cells sometimes present
on the inside of the vascular bundles.
 BERBERIDACEAE 63
The rhizome of the Indian drug Podophyllum hexandrum Royle (syn. P.
emodi Wall), which also occurs in commerce, is generally distinguishable
from P. peltatum, according to Wallis and Goldberg (2350, 2351), by the
following characters, (i) Absence of epidermal cells with brown contents,
(ii) The smaller
size of the cluster crystals, (iii) The almost complete absence
of pericyclic fibres. (This last character appears to be somewhat unreliable,
as samples occur in the Kew Museums in which pericyclic fibres were more
numerous in the Indian than in the American species. A more reliable method
of distinguishing the rhizome of the 2 species in question is by grinding up
the rhizome with 90 per cent, alcohol, filtering, and adding a few drops of a
copper acetate solution. This treatment gives a green coloration with the
American drug and a brownish-yellow one with the Indian. This mode of
distinction seems to be effective even with material stored for 50 years in a
museum.)
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Achlys, Berberis,* Caulophyllum,* Diphylleia, Epimedium,* Hydrastis,
Jeffersonia, Leontice, Mahonia,* Nandina, Podophyllum,* Vancouveria.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Berberis, Mahonia, (Nandina).

LITERATURE
(i) On General Anatomy
Blaque and Maheu 205, Ewert 667, Harvey-Gibson and Horseman 916, Himmel 975,
Himmelbaur 976, Holm 1016, 1017, 1021, 1063, Kumazawa 1298, Maue 1460, Short
2095, Stearn 2189, Wallis and Goldberg 2350, 2351, Worsdell 2469.
(ii) On Wood Structure
Dadswell and Record 533, Howard 1088, Kanehira 1206, 1209, Record 1843, 1851,
Record and Hess 1886, Tupper 2295, Yamabayashi 2478.

19. CIRCAEASTERACEAE
This family is represented by the single species Circaeaster agrestis Maxim,
from the Himalayas. The plant is a herb with a basal rosette of leaves. It has
not been examined microscopically in recent years, and the following features
are taken from Solereder's account.
Cells of the upper epidermis of the leaf elongated parallel to the midrib.
Stomata small, ranunculaceous; confined to the lower surface. Vascular
bundles of the veins not accompanied by sclerenchyma. Vascular system of
the stem, like that of the main root, consisting of a diarch xylem plate with
only a small amount of secondary wood and phloem produced on both sides.
The genus was included in the Chloranthaceae in the Bentham and Hooker
system, but was placed amongst the Berberidales by Hutchinson. The
taxonomic position of the family is very uncertain.

GENUS DESCRIBED
Circaeaster.
 (64)

20. LARDIZABALACEAE
(Fic. 16 on p. 60; FIG. 17 on p. 65)

SUMMARY
A small family consisting mostly of lianes, but including some erect shrubs.
It is confined to Eastern Asia and temperate South America. The most
constant anatomical characters are the widely spaced vascular bundles of
the axis, usually separated by broad, generally lignified primary medullary
rays; the circle of separate bundles, visible in transverse sections through the
distal end of the petiole; the frequent occurrence of solitary crystals in the

pericyclic fibres of the axis or in the parenchymatous cortical cells immediately

adjacent. The outer part, or, in some instances, the whole of the pith, con-
sists of thick-walled, pitted parenchyma.
The wood exhibits the following features. Vessels large, sometimes with
spiral thickening, perforations wholly simple or with a few scalariform
plates, intervascular pitting alternate, members of medium length to very
short. Parenchyma very sparse or absent. With broad primary rays only.
Fibres with simple pits,sometimes septate and storied, extremely short in
Akebia.

LEAF
Dorsiventral. Epidermis on both surfaces consisting of cells with sinuous
anticlinal walls, especially on the lower side strongly papillose on the lower
;

surface in all examined species except Holboellia coriacea Diels. A well-

developed hypoderm present in Holboellia but not observed in the other
genera. Stomata confined to the lower surface; ranunculaceous. Mesophyll
including several layers of palisade cells. Veins embedded in the mesophyll,
often tending to be vertically transcurrent by sclerenchymatous elements.
Petiole (Fig. 17 c and D), in transverse sections through the distal end,
exhibiting, e.g. in Decaisnea fargesii Franch., a ring of individually distinct
vascular bundles, each accompanied on the outside by a well-developed
strand of fibres, the latter usually with wide lumina; similar, but with
fibres forming a continuous ring, e.g. in Akebia lobata Decne. Solitary

crystals present in the cells immediately external to the pericyclic fibres in

Akebia, but much more numerous in A. quinata Decne. than in A. lobata
Decne. observed also in the sub-epidermis in the first of these species. No
;

crystals around the vascular bundles of the veins observed in Holboellia

coriacea.

Axis
YOUNG STEM (Fig. 17 F)
Cork arising superficially. Peiicycle including well-defined strands of
fibres outside the phloem groups. Vascular system invariably consisting of

widely spaced bundles, separated by conspicuous, generally lignified rays.

Phloem masses well developed but containing no sclerenchyma. Xylem
including vessels of wide diameter; perforations generally simple, but only
scalariform plates observed at Kew in Decaisnea fargesii Franch. (See also
 LARDIZABALACEAE 65
Wood'.) Outer part of the pith composed of strongly lignified, pitted cells
in Akebia and Holboellia, the centre being occupied by thin- walled cells; the
whole tissue composed of lignified cells in Stauntonia hexaphylla Decne.

FIG. 17, BERBERIDACEAE~A-to and E; LARDIZABALACEAE, C-D>nd F

A, Podophyllum peltatum Linn. S.em X8. B, Epimedium alpinum Linn. Aerial stem X 15. C,
Decaisnea fargesii Franch. Petiole x 13. D, Akebia lobata Decne. Petiole x6o. E, Berberis thunbergii
DC. Stem x 15. F, Akebia lobata Decne. Stem X3O.

Solitary crystals abundant in the pericyclic fibres and pith of Holboellia

coriacea Diels; similar crystals, mostly situated in cells immediately external
to the pericyclic fibres, noted in Akebia and Lardizabala. Clustered crystals
observed only in the central part of the pith of Holboellia coriacea.

WOOD (Fig. 16 c)
Vessels of two distinct sizes in Akebia quinata Decne. very large and
solitary or very small and in tangential multiples and clusters (Fig. i6c);
Solereder refers to spiral thickening in the smaller vessels of Holboellia and
66 LARDIZABALACEAE
Lardizabala biternata Ruiz, et Pav. Perforations simple, except for a few
scalariform plates with few bars in Holboellia (2158). (See, however, under
*

Young Stem*.) Intervascular pitting alternate, large; pits to ray cells some-
times large and simple. Storied in ^Akebia quinata. Mean length about
o*23-O'55 mm., shortest in Akebia. Parenchyma very sparse or absent.
Rays limited to broad and high primary rays, which are unlignified and,
according to Solereder, do not become closed by interfascicular wood, but
form longitudinal plates that split up the wood into segments. Fibres with
large bordered pits in Akebia and Lardizabala (2158), but with simple pits in
Holboellia] with fine septa and storied in Akebia quinata. Mean length about
0*4 mm. in Akebia.

i TAXONOMIC NOTES
This small family is closely related to the Berberidaceae in which it was
included by Bentham and Hooker. It has also been suggested by Harvey-
Gibson and Horseman (916) that it is in some respects intermediate between
the Berberidaceae and Menispermaceae. The widely spaced vascular bundles
and general appearance of the stems are very alike in all 3 families. The
resemblance to the woody Ranunculaceae, such as Clematis, and some of the
Aristolochiaceae are also noteworthy, although it should be remembered that
these similarities may be partly bound up with the climbing habit of the plants.
Sargentodoxa, which differs somewhat from the Lardizabalaceae, is here
treated as the sole genus in a distinct family.

ECONOMIC USES
The Akebia lobata Decne. are eaten in Japan, and the sliced stems
fruits of
of A. quinata Decne. used medicinally in China.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Apart from a few scattered statements in the literature, the above descrip-
tion was based mainly on fresh material of the following species grown at
Kew: Akebia lobata Decne.,* A. quinata Decne.,* Decaisnea fargesii Franch.,*
Holboellia coriacea Diels,* Stauntonia hexaphylla Decne.*

* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Akebia, (Holboellia), Lardizabala.

LITERATURE
(i) On General Anatomy
Bowman 250, Harvey-Gibson and Horseman 916.

(ii) On Wood Structure

Garratt 747, Record 1851.
 (6?)

21. SARGENTODOXACEAE
SUMMARY
Sargentodoxa cuneata Rehd. et Wils., originally named Holboellia cuneata
Oliv. and included in the Lardizabalaceae, was given the status of a distinct
family by Stapf (2186). The species occurs in Central China. According to
Lemesle (1355) the anatomy of the stem was described by R^aubourg, and
the following description is based mainly on the last author's account, as
summarized and amplified by Lemesle.

Axis
STEM
Epidermis composed of quadrangular cells, but rounded on the outer
surface; outer wall thick and cutinized. Primary cortex consisting of about
12 layers of flattened cells. Pericycle completely sclerosed, forming a broad,
sinuous ring. Cork arising in the innermost layer of the pericycle. Vascular
system consisting of 4 large, distinct bundles alternating with smaller ones.
(According to Solereder the large bundles constitute an inner ring.) Xylem.
Vessels up to 200 ft in diameter. Ground tissue of the xylem consisting of
radial rows of fibres having circular bordered pits with obliquely crossed

apertures in both the radial and tangential walls. (Lemesle calls these fibres
'tracheides i ponctuations areotees'.) Pith parenchymatous, but including
a group of sclerosed cells at the centre. Large secretory cells with tannini-
ferous contents reported by Solereder to occur in association with the vascular
bundles.
TAXONOMIC AND PHYLOGENETIC NOTES
In Lemesle's opinion, the floral characters and the nature of the fibres
which constitute the ground tissue of the xylem indicate that Sargentodoxa is
a very primitive angiosperm. Comparisons, on rather slender evidence, are
even made with the Cycadaceae and Bennettitales. Sargentodoxa is clearly

differentiated from the Lardizabalaceae by its deep-seated cork.

GENUS EXAMINED
Sargentodoxa.
LITERATURE
On General Anatomy
Henderson 952, Lemesle 1355, Stapf 2186.

22. NYMPHAEACEAE
(Fig. 18 on p. 68; Fig. 19 on p. 72)

SUMMARY
Aquatic herbs which include the familiar water-lilies. The family occurs in
Tropical and North Temperate regions. Its members possess closed, scattered
vascular bundles resembling those of the Monocotyledons. As in most aquatic
herbs there are numerous intercellular spaces in the parenchymatous
tissues. Air passages arise in the position of the primary xylem owing to the
breakdown of the latter. True vessels are absent. Branched sclerenchymatous
68 NYMPHAEACEAE
idioblasts (Fig. 18 A) are common. Hairs uniseriate, of several cells,
some Calcium oxalate frequently deposited on the cell"
secreting mucilage.
walls. Clustered crystals occur in the stem and leaf of Nelumbium and
rhombic crystals of brasenin recorded in Brasenia. Articulated laticiferous
tubes or sacs, with thin suberized walls, are present in the parenchymatous
tissue of all organs, as well as in the vascular bundles; those of Nuphar are

nearly isodiametric, generally solitary or 2-3 in a linear row, but those occur-

FIG. 18. NYMPHAEACEAE

A, Transverse section through the leaf of Nymphaea marliadi Hort. the crystals in the walls of the
;

idioblasts and of the spongy parenchyma are drawn larger than they are in reality. B, Transverse
section through the wall of an air passage from the petiole of Nelumbium speciosum Willd., with a cell
bearing a clustered crystal. C, Intercellular space of Brasenia peltata Pursh., filled with mucilage
hairs. A-B, by Solereder, C, after Schrenk.

ring in the bundles are more elongated. The laticiferous sacs are arranged
together in long rows in Brasenia and Cabomba those of Euryale> Nymphaea^
;

and Victoria tubular and of considerable length, although consisting of only

a few cells. Rhizomes and stolons are often polystelic. Root hairs arise

from specialized cells. This feature is common amongst the Monocotyledons

but rare amongst the Dicotyledons. For a detailed description of the anatomy
of Nymphaea Conard's (457) monograph should be consulted.

LEAF (Fig. 18 A-C and Fig. 19 C-E)

Dorsiventral, with abundant intercellular spaces in the spongy tissue.
Hairs uniseriate; generally confined to young organs, becoming detached
from older ones. Hairs consisting of clothing and mucilaginous kinds.
Clothing hairs frequently uniseriate with 2-3 basal cells, the latter being
much shorter than the remainder and provided with suberized lateral walls.
Mucilage hairs shorter than the clothing hairs, but provided with an enlarged
terminal cell. Stomata ranunculaceous; generally confined to the upper
surface; closed by the interlocking of the cuticular ridges of the guard cells
 NYMPHAEACEAE 69
in Euryale, Nuphar, Nymphaea, and Victoria. Floating leaves of Nymphaea
alba Linn, with stomata evenly distributed over the greater part of the upper
surface of the leaf lamina, but absent from the region of attachment of the
petiole to the lamina. Stomata, in the region immediately surrounding that
from which they are absent, provided with guard cells twice as large as those
elsewhere. Large stomata less numerous than small ones. Average number
mm. under normal light intensity 460, maximum recorded 524,
per sq.
minimum 150. Submerged leaves, although apparently similar in external
appearance, are of two kinds, with and without stomata respectively. Stomatal
distribution in terrestrial forms similar to that of submerged plants. Stomatal
frequency is reduced by intense illumination according to Roshardt (1956,
1957). In Nuphar luteum Sibth., Griiss (830, 831) found stomata to be absent
from or infrequent in aerial leaves, those on the lower surface being confined
to the regions above the marginal bundles. Hydathodes present in the centre
of the upper surface in Nelumbium speciosum Willd., and at the points of
junction of the anastomosing veins of 'Brasenia pcltata Pursh'. Haustoria,
believed to serve for the uptake of water and dissolved mineral substances,
present in Nuphar luteum, Nymphaea lotus Linn, and Victoria regia Lindl. ;

those of Nuphar luteum consisting of 2 small basal cells with lignified walls,
bounded externally by a swollen one extending to the exterior of the leaf; a
large swollen cell present internally to the 2 lignified basal ones. Haustoria
of Nymphaea lotus and Victoria regia similar but differing in details. About
200 haustoria per sq. mm. present in floating and 9-112 per sq. mm. in
aerial leaves of Nuphar luteum according to Griiss. Branched sclerenchyma-
tous idioblasts (Fig. 18 A) varying in shape within a single species, and which
may be stellate, girder- or H-shaped, recorded by Solereder in Euryale,
Nuphar, Nymphaea, and Victoria', sometimes covered with minute crystals
of calcium oxalate. Idioblasts stated to be absent from Brasenia, Cabomba,
Nelumbium. Walls of the idioblasts of Nuphar and Nymphaea pitted where in
contact with adjacent cells. Spines, containing vascular tissue, present in
Euryale. Transparent dots of unknown physiological significance, but
believed to consist of pits formed by the death of the tissues in the regions
where present, occur in the leaves of Victoria. The pits in old leaves become
filled with algae or deposits of lime. Fat frequent in guard and mesophyll
cells of Nuphar luteum, becoming converted to starch under inadequate
illumination. Petiole, in Wansverse sections, exhibiting normally orientated,
double bundles in Brasenia and Cabomba', with mixed simple and double,
but normally orientated bundles in Euryale, Nymphaea, and Victoria] with
mixed normal and inversely orientated bundles in Nelumbium. For dia-
phragms in the petiole see under 'Axis'.
Enlarged mucilaginous cells (Fig. 18 c) sometimes project into the inter-
cellular air spaces, e.g. in Brasenia. Clustered crystals (Fig. 18 B) recorded

by Solereder in certain small cells in the partitions between the air lacunae in
the petiole (and stern) of Nelumbium, the crystalliferous cells projecting into
the intercellular cavities.
The structure and distribution of the vascular bundles and air canals in the
petioles and peduncles of different species of Nymphaea are of considerable
interest, and should be useful in identifying species. For details see the plate
on p. 59 of Conard's (457) monograph.
70 NYMPHAEACEAE
Axis
With abundant intercellular spaces of schizogenous origin, arranged in
circles in Euryale, Nymphaea, and Victoria, but distribution reticulate in

Nuphar. Diaphragms traverse the intercellular spaces in Nuphar luteum

Sibth., Nymphaea alba Linn., N. lotus Linn., and Victoria regia LindL, but
are not universally present in all the organs of these species absent from the
;

petiole and peduncle of Nymphaea, although present in the corresponding

regions of Victoria regia. Diaphragms of Nymphaea alba and Victoria regia
found by de Bruyne (300) to be very delicate, consisting of a single layer of
elongated cells united by lateral prolongations; those in the petiole and
peduncle of Nuphar luteum consisting of a kind of spongy tissue occupying
the whole lumen of the canal. Vascular bundles in transverse sections,
like those of the Monocotyledons, appearing irregularly distributed, without
cambium or true vessels, but provided with long tracheids possessing spiral
or annular thickenings. Primary xylem becoming disorganized, and thus
giving rise to schizogenous cavities, each bundle then consisting of a canal
and a phloem strand. Sclerenchyma absent from the bundles. Peduncle of
Brasenia and Cabomba with 3 simple, normally orientated bundles arranged
in a triangle; simple and double, but normally orientated bundles occur in the
peduncle ofEuryale, Nymphaea, and Victoria', normal and inversely orientated
bundles in the corresponding position in Nelumbium. Barclaya and Nuphar
possess simple bundles throughout.

RHIZOME
Frequently containing a confused mass of bundles. Polystelic in certain
species including Victoria regia LindL, where each stele consists of a ring of
about 20 bundles. Bundles in the stolons of Nymphaea mexicana Zucc. (syn.
Castalia flava Leitnr.) arranged in 4 or 5 widely separated groups of steles,
each enclosed in an endodermis and surrounding a protoxylem group. Two
steles, each consisting of a pair of bundles, observed by Arber (29) in the
stolons of Cabomba. Rhizomes almost universally astelic according to Van
Tieghem and Schoute as cited by Solereder, Nelumbium alone possessing a
general endodermis around the central cylinder. Rhizomes stated in the
literature to have double bundles in Brasenia and Cabomba simple, normally
;

orientated bundles in Euryale, Nymphaea, and Victoria-, mixed bundles

exhibiting normal and inverse orientation respectively in Nelumbium.

ROOTS
With numerous diaphragms in Nuphar luteum Sibth., Nymphaea alba
Linn., N. lotus Linn., and Victoria regia Lindl.; those of Nuphar luteum said
by de Bruyne (300) to be composed of a number of cells loosely attached to
the walls of the canals, thus giving the appearance of a membrane with holes
at the margins as seen in transverse section3. Root hairs arising from

specialized cells, as in many Monocotyledons.

TAXONOMIC NOTES
The most interesting features are the closed scattered bundles of the type
characteristic of the Monocotyledons, the absence of vessels from the xylem.
 NYMPHAEACEAE 71
and the occurrence of polystely. The absence of sclerenchyma and the
presence of abundant intercellular spaces are characters which are probably
correlated with the aquatic habitat of the family, and possess but little
taxonomic significance.

ECONOMIC USES
The Water Lilies are cultivated as decorative plants. Arber (29) has
recorded that the long peduncles of Water Lily flowers are sold in bazaars at
Cairo as tobacco pipes.
GENERA DESCRIBED
Barclaya, Brasenia, Cabomba, Castalia, Euryale, Nelumbium, Nuphar,*
Nymphaea,* Victoria.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Arber 29, de Bruyne 300, Conard 457, Green and Buck 808, Grass 830, 831, Gwynne-
Vaughan 853, Roshardt 1956, 1957.

23. SARRACENIACEAE
(FiG. 19 on p. 72)

SUMMARY
An American family of herbs with rhizomes bearing a rosette of radical,
tubular leaves, the latter provided with specialized glands and hairs which
serve for entrapping insects and other small organisms which are subsequently
digested by the plant as food.

LEAF (Fig. 19 A-B)

Distal portion tubular, with a petiolar region below. Upper part of petiole
expanded on one side as a laminar 'wing*. External epidermis provided with
secretory glands, accompanied in some species by upwardly directed hairs.
Glands especially numerous on the laminar wings in Heliamphora. Passing
from the apex downwards several distinct zones may be recognized on the
inside of the leaf, (i) Glandular zone occupying the inner surface of the lid
of Heliamphora and Sarracenia, and of the hood of Darlingtonia. Glands,
stomata (not accompanied by subsidiary cells) and downwardly directed hairs
present in this region, the glands of Heliamphora being of 2 distinct sizes.
(2) Slippery zone, consisting of epidermal cells with downwardly directed
projections, overlapping like fish scales. Glands sometimes present, but sto-
mata and hairs absent. (3) Eel-trap zone occupying the greater part of the
tube, with long, downwardly directed, needle-shaped hairs, and numerous
glands, but no stomata. The extreme basal portion of the leaf is provided
with a smooth surface. (Glands absent from the inside of the leaf of Darling-
tonia except for the hood according to Solereder.) [Solereder also states that
in Heliamphora the slippery zone is lacking, and the stomata in the eel-trap
zone are elevated. This is not in agreement with Macfarlane's (1409) descrip-
tion in which it is stated that in Heliamphora there is a smooth region below
 a.c

FIG. ig. SARRACENIACEAE, A-B; NYMPHAEACEAE, C-E

A, Sarracenia purpurea Linn. Petiole, through wing, x 9. B, S. purpurea Linn. Base of petiole
X 15. C, Victoria regia LindL Lamina x 15. D, V. regia Lindl. Lamina XSQ. E, Nuphar luteum
(L.) Sm, Outer part of petiole x 50.
a.c. Air canals. a.s. Air spaces, p.t. Palisade tissue. v,b. Vascular bundle.

Fig. C shows the lamina with a rib projecting from the lower surface. In Fig. D only the upper
part of one of the ribs is shown.
 SARRA CENIA CEAE 73
the glandular, hairy inner surface of the lid, whilst at the base of the tube
there are a few thickened hairs. There appear to be considerable variations
in different species of all the genera.]
Modified circular areas, in which the epidermis of the outer is separated
from that of the inner surface only by specially thin portions of the mesophyll,
occur in the upper part of the pitchers of Sarracenia drwnmondii Croom,
S* minor Walt., and S. psittacina Michx. Each of these areas is surrounded by
a fence of hairs. Scale or autumnal leaves of Sarracenia spp. bear numerous
glands on both upper and lower surfaces. Stomata absent from the upper
epidermis in all species, but infrequently on the lower (outer) epidermis in
S. purpurea Linn. The anatomical facts suggest that the scale leaves are
reduced pitchers. Petiole, in transverse sections, exhibiting an irregular
crescent of collateral vascular bundles, each bounded internally and externally
by sclerenchyma but embedded in loose, starch-containing tissue, the starch
grains being far more numerous in some of the cells than in others. According
to Macfarlane the bundles in the cylindrical portion of the petiole are arranged
in a discontinuous circle, bounded externally by a cortex of cells containing
chlorophyll. Other irregularly distributed vascular strands also occur within
the circle just mentioned. In the section of the petiole of S. purpurea used in
the preparation of Fig. 19 B the demarcation between the outer ring of bundles
and the scattered ones towards the centre
is not very clear. Two sets of
bundles present in the laminar wing with their xylem groups inwardly
directed (Fig. 19 A).

RHIZOME
Epidermis with brown pigment in the cell cavities; bounded internally by
2-3 layers of hypoderm, the outer cells being lignified. Cortex broad.
Vascular bundles collateral, of varying size and shape, arranged in a dis-
continuous ring and separated by rays of unequal width. Ring of bundles
more nearly closed in Darlingtonia than in Sarracenia. Bundles bounded
internally and externally by sclerenchyma. Xylem containing a few vessels
with spiral thickening, and more numerous vessels with scalariform perforation
plates.
ROOT
Primary structure pentarch or hexarch. Schweiger (2063) has recorded the
following information concerning a root of Sarracenia chelsoni Hort. i mm.
thick. Surface bounded by a 3 -layered hypodermis with thickened walls and
brown contents.Cortex spongy, with abundant intercellular spaces, bounded
internally by an endodermis of small cells with scarcely thickened walls.
Xylem arranged in 10 large groups. Metaxylem of S. purpurea Linn, said to
consist of tracheids, food-storing cells, and large vessels with oblique, scalari-
*
form septa*. Roots of Darlingtonia differ only in minor respects.

TAXONOMIC NOTES
The plants are so specialized in structure in correlation with their peculiar
mode of life that it is rather difficult, on anatomical grounds, to decide which

of the positions to which the family has been assigned in different systems of
classification is most likely to be correct. Schweiger (2063), who made careful
anatomical comparisons between Sarracenia and Cephalotus (Cephalotaceae),
74 SARRACENIACEAE
decided that such resemblances as exist between these 2 genera are due to
similarities in their mode of life and have no taxonomic significance.

GENERA DESCRIBED
Darlingtonia, Heliamphora, Sarracenia.*
* Kew
Represented in the slide collection,

LITERATURE
On General Anatomy
Arber 32, Lloyd 1383, Macfarlane 1409, Russell 1970, Schweiger 2063, Uphof 2315.

24. PAPAVERACEAE
(FiG. 20 on p. 76; FIG. 25 on p. 92)

SUMMARY
(i) GENERAL
Mainly herbs, but certain genera tending to be shrubby (Dendromecori),
or small trees (Bocconia). The family occurs mainly in temperate and sub-
tropical regions of the Northern Hemisphere. The stem of most Papaveraceae
exhibits, in transverse sections, a single ring of widely spaced vascular bundles,
which are nearly always collateral. The xylem groups frequently tend to
be V-shaped. Several rings of bundles sometimes present in Papaver.
Hairs scanty, uniseriate, biseriate, or multiseriate. Shaggy hairs occasional.
Glandular hairs absent. Stomata ranunculaceous. Petiole, in transverse
sections, exhibiting an arc of vascular bundles not accompanied
commonly
by sclerenchyma. Variously coloured latex present, sometimes in articulated
laticiferous tubes, but elsewhere in cells or sacs, the latter sometimes
arranged in longitudinal rows. Crystals of calcium oxalate recorded only in
Bocconia frutescens Linn, but said to be present in the floral leaves of a few
other species. Cluster crystals also noted in Romneya. Alkaloids are com-
mon in the family. These include such well-known substances as morphine
and codeine. Berberin, which is characteristic of the Berberidaceae, is known
to occur in Argemone mexicana Linn.

(ii) WOOD
Vessels small and tending to be ulmiform, or medium-sized and without
pattern, sometimes ring-porous and with spiral thickening, perforations
simple, intervascular pitting alternate, members of medium length. Paren-
chyma vasicentric. Rays all multiseriate,up to 5-12 cells wide, high,
heterogeneous. Fibres with simple pits, very short.

LEAF
Usually dorsiventral. Simple, uniseriate hairs recorded in Chelidonium,
Glaucium, Roemeria; shaggy hairs in Bocconia, Macleaya> Meconella, Meconop-
sis y Papaver,
Platystemon. Spines present in Argemone mexicana Linn.
Thickenings recorded on the anticlinal walls of the cells of the epidermis of
Glaucium corniculatum (L.) Curtis. Lower surface slightly papillose in San-
guinaria canadensis Linn., and with conical papillae on both surfaces in
 P AP AVER ACEAE 75
Dendromecon rigida Benth. Epidermis frequently covered with wax, especially
in Bocconia and Macleaya. Stomata present on both surfaces in Papaver
pilosum Sibth. et Smith., P. spicatum Bois., and Roemeria dodecandra (Forsk.)

Stapf, but generally confined to the lower surface in other genera and species;
ranunculaceous. Hydathodes occur in groups on the lower surface of the
teeth at the margin of the leaf in certain species of Papaver. Mesophyll
generally including i to several layers of palisade cells, but not distinctly
differentiated into palisade and spongy regions in Hesperomecon platystemon
Greene (syn. Platystigma lineare A, Gray), Meconella californica Torr., M.
oregana Nutt., and Papaver somniferum Linn. Palisade tissue recorded on
both sides of the leaf in Eschscholtzia californica Cham. Petiole (Fig. 20 D-E),
in transverse sections through the distal end, exhibiting an arc of vascular
bundles in Chelidonium majus Linn. (Fig. 20 E), Eschscholtzia californica,
Macleaya cordata (Willd.) R. Br. Romneya trichocalyx Eastw. (Fig. 20 D), and
Papaver dubium Linn. Bundles sometimes very close together or almost fused
in Macleaya cordata. Sclerenchyma generally scanty in or absent from the
petiole, but each bundle is surrounded by a massive layer of fibres in Boc-
conia frutescens Linn. Crystals of calcium oxalate recorded only in Bocconia
frutescens, but clustered crystals observed at Kew in the stem of Romneya.

LATEX CANALS AND CELLS

Latex generally present throughout the plant in either (i) articulated
laticiferous tubes , frequently having sieve plates on the transverse or lateral
walls, or (ii) laticiferous sacs consisting of more or less elongated cells,
either solitary or arranged in longitudinal rows. (According to Solereder, the
nature of the latex-containing elements is of diagnostic value in separating
certain genera. While this may be true in certain instances, it has also been
shown by Holm (1027) t 'lat the nature of the laticiferous elements varies in
different parts of individual plants of Sanguinaria canadensis Linn. ). Articulated
laticiferous tubes recorded in Argemone, Chelidonium, Papaver, Roemeria, and
the stem of Sanguinaria canadensis, and probably occurring also in Meconopsis
and Platystemon. Laticiferous sacs recorded in Cathcartia, in the root, rhizome,
and leaf of Sanguinaria canadensis and in Stylophorum. Colour and consis-
tency of the latex somewhat variable; white in Meconopsis, Papaver, Roemeria',
lemon yellow in Argemone', orange in Chelidonium', at first watery and
somewhat reddish, but later becoming more turbid in Bocconia, Eschscholtzia,
Glaucium, Hypecoum\ watery at first but becoming yellow in Macleaya and
blood-red in Sanguinaria. Latex tending to disappear from the older parts of
the plant in Chelidonium and Glaucium. Experiments with latex tubes of
Papaveraceae undertaken by Simon (2103) suggest that these channels do not
serve for conduction, although it has been stated by Fedde (676) that the
mineral substances in the latex increase its absorptive capacity.

Axis
STEM (Fig. 20 A, G~H)
A collenchymatous exodermis is differentiated in Argemone mexicana Linn.,
Glaucium flavum Crantz. and Macleaya cordata (Willd.) R. Br. Peripheral
part of cortex including sclerosed cells in Argemone mexicana and Bocconia sp.
according to Harvey-Gibson and Bradley (915). Endodermis not usually
 76 PAPAVERACEAE
distinct. Pericycle generally sclerenchymatous, or at least including a ring
of thin-walled, lignified, pitted elements, often with arcs of fibres external
to the individual bundles; described by Harvey-Gibson and Bradley as
not sclerosed in Argemone mexicana and Romneya trichocalyx Eastw., and

FIG. 20. PAPAVERACEAE, A, D-E, G-H; FUMARIACEAE, B~C and F

A, Chelidonium majus Linn. Stem X 15. B, Fumaria muralis Koch. Stem x 9. C, Dicentra specta-
bilis Linn. Stem x 8. D, Romneya trichocalyx Eastw. Petiole x 9. E, Chelidonium majus Linn.
Petiole X4O. F, Dicentra spectabilis Linn. Petiole x 15. G, Papaver orientate Linn. Stem X 9. (The
circle in the pericyclic region represents thin-walled, lignified, pitted elements.) H, Romneya tricho-
calyx Eastw. Stem x 3.
h.p. Hollow pith. p.e. Thin-walled, lignified elements with pits.

weakly sclerosed in Eschscholtzia californica Cham., Papaver somniferum

Linn., P. orientate Linn. Most Papaveraceae provided with a single ring of
collateral vascular bundles (Fig. 20 A), the interfascicular tissues between
them sometimes tending to become lignified. Numerous bundles arranged
in concentric zones, in a manner recalling that of some of the Ranunculaceae,
frequently occur in Papaver spp., e.g. according to Harvey-Gibson and
Bradley (915) in P. orientate Linn. (Fig. 20 G) and P. somniferum Linn. In the
 PAP AVER ACE AE 77
more woody species, e.g. Romneya trichocalyx Eastw. (Fig. 20 H) the vascular
bundles are separated by broad primary medullary rays, but are less widely
spaced than in the herbaceous species. Vessels with simple perforations.

PEDUNCLE
Usually with a circle of 4-5 vascular bundles in Papaver dubium Linn, and
P. somniferum Linn., but several circles of bundles recorded by Friedel (713)
in P. orientate Linn. Anatomical structure of the peduncle very constant in

Meconopsis\ cortex narrow, component cells generally with cellulose walls,

but sclerenchymatous in M. chelidonifolia Bur. et Franch. pericycle generally
;

including sclerenchyma; pith composed of cells with cellulose walls, becoming

hollow; vascular bundles widely separated, arranged in a single circle, their
number varying according to the species, as well as in different individuals of
the same species, or at different levels in a single peduncle xylem groups not
;

generally U-shaped (compare 'Stem') except in M. chelidonifolia where they

are more definitely U-shaped than in the Ranunculaceae.

WOOD (Fig. 25 F-G and i)

Vessels very small (25-50 ^ mean tangential diameter) in Dendromecon

rigida Benth. and Romneya coulteri Harv., and with many tracheid-like vessels
in the latter, medium-sized in Bocconia frutescens Linn. in short radial;

multiples and irregular clusters, which form vague to distinct oblique lines
in Dendromecon and Romneya few, solitary and in radial pairs, and without
;

pattern in Bocconia; semi-ring-porous in Dendromecon; with spiral thickening

in Dendromecon and Romneya. Perforations simple. Intervascular pitting
alternate, rather large in Bocconia] pits to ray or wood parenchyma similar to
the intervascular pitting in Dendromecon and Romneya, larger, with wide
apertures and often elongated in Bocconia. Mean member length about
0-4 mm. Parenchyma paratracheal, vasicentric (Fig. 25 F) to sparse; strands
of 1-2 cells in Bocconia and Dendromecon, fusiform cells sometimes pre-
dominating. Rays all multiseriate, up to 5 and 7 cells wide in Dendromecon
and Romneya, and up to 12 cells in Bocconia] commonly more than 2 mm.
high; cells all square or upright in Dendromecon and Romneya, markedly
heterogeneous in Bocconia, tangential sections showing separate groups of
small procumbent cells in each ray surrounded by square to upright cells
(Fig. 25 G); with sheath cells; about 3 rays per mm. Dadswell and Record
(533) re fer to a few l ar ge open, radial canals in Bocconia. Fibres with
numerous, small, simple pits on both radial and tangential walls. Walls
moderately thin to rather thick. Mean length 0*6-0-75 mm.

RHIZOME
Described by Holm (1027) as showing a circular band of vascular bundles
and interfascicular cambium in Sanguinaria canadensis Linn. Sclerenchyma
recorded in the pericycle of this species.

ROOT
Primary structure generally diarch. Growth in thickness very limited and
confined to the stele in Sanguinaria canadensis Linn, according to Holm
(1027).
78 PAPAVERACEAE
TAXONOMIC NOTES
One
of the most interesting anatomical features is the tendency for the
vascular bundles in a few species to be scattered. The xylem groups are also
occasionally U-shaped. Both these characters suggest affinities with the
Ranunculaceae, and therefore with some of the Monocotyledons. The view
expressed by Friedel (713, 714) that the presence of a small number of
vascular bundles in the peduncle indicates a primitive position within the
family seems to have very little to support it, and Friedel himself appears
subsequently to have become very uncertain about this suggestion, judging
from the remarks in the second of his two papers just cited.
The laticiferous elements are generally believed to be homologous with
the secretory cells of the closely related Fumariaceae.
The wood anatomy of Dendromecon and Romneya is closely similar, but
that of Bocconia differs strikingly in several vessel characters.

ECONOMIC USES
Opium is the dried latex from the seed capsules of Papaver somniferum Linn.
The petals of Papaver rhoeas Linn, and the dried rhizome of Sanguinaria
canadensis Linn, are used medicinally.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Argemone, Bocconia, Cathcartia,Chelidonium,* Dendromecon, Eschscholt-
zia, Glaucium, Macleaya,* Meconella, Meconopsis, Papaver,* Platystemon,
Platystigina, Pteridophyllum, Roemeria, Romneya,* Sanguinaria, Stylo-
phorum.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Bocconia, Dendromecon, Romneya.

LITERATURE
(i) On General Anatomy
Fedde 675, 676, Friedel 713, 714, 715, 716, 717, Fritsche* 725, Harvey-Gibson and
Bradley 915, Holm 1027, Simon 2103, Werdemann 2413*

(ii) On Wood Structure

Dads well and Record 533, Record 1843, 1851, Record and Hess 1886.

25. FUMARIACEAE
(Fic. 20 on p. 76)

SUMMARY
A
family of herbs, mainly from North Temperate regions, closely related
to, and by someauthorities included in, the Papaveraceae. Some of its
members are scandent. Features of morphological interest include the basal
tubers which occur in various species of Corydalis such as C. cava (L.)
Scweigg. et Korte and C. solida (L.) Swartz. The morphological nature of the
tubers of C. solida is somewhat uncertain as they cannot be strictly regarded
 FUMARIACEAE 79
as stems or roots. According to Fritsche (725) young tubers arise internally
from pre-existing ones. True bulbs occur in Dicentra cucullaria (L.) Bernh.
It is also noteworthy that the seedlings of certain species of Corydalis and
Dicentra possess only one cotyledon. The primary structure of the root is
diarch.
In a general way the anatomical characters of the Fumariaceae are so
similar to those of the herbaceous members of the Papaveraceae that there is
no need to recapitulate them in detail. The type of petiole structure is shown
in Fig. 20 F for Dicentra, whilst transverse sections of young stems of Fumaria
and Dicentra are illustrated in Fig. 20 B-c. Beyond this it will suffice to say
that the most characteristic feature of the family is the presence of secretory
cells or idioblasts up to 10 mm. long, which have been recorded in different
parts of certain species of Adlumia, Corydalis, Dicentra, and Fumaria. The
nature of the contents of these cells is uncertain. In view of the close affinities
of the Fumariaceae and Papaveraceae it seems reasonable to suppose that they
are of the same nature as the laticiferous cells of the Papaveraceae.

GENERA DESCRIBED
Adlumia, Corydalis,* Dicentra,* Fumaria.*
* Kew slide
Represented in the collection.

LITERATURE
On General Anatomy
Fritsche* 725, Kloimweder 1246.

26. CRUCIFERAE
(Fio. 21 on p. 80; FIG. 22 on p. 84; FIG. 25 on p. 92)

SUMMARY
(i) GENERAL
A cosmopolitan family consisting predominantly of herbs, but a few species
tend to be shrubby. Well-defined, characteristic anatomical features exist
throughout the family. Secretory cells containing myrosin, which is
coagulated and stained red by Millon's reagent, or coloured violet with orcin
solution and concentrated hydrochloric acid, are especially noteworthy. They
are widely distributed throughout the Cruciferae, but their frequency is
partially controlled by nutrition and environment. They may occur in any
part of the plant and in practically any of the tissues. The nature and
distribution of idioblasts containing myrosin may be of taxonomic value (see
4

Leaf*). The stomata are surrounded by 3 subsidiary cells of which i is

usually much smaller than the other 2, the so-called cruciferous type. Cork
arises in the pericycle or inner part of the cortex of the stem. The inter-
fascicular tissue between the xylem strands of adjacent vascular bundles in
the stem becomes lignified at a very early stage, so that normal, parenchyma-
tous, primary medullary rays are generally absent. Variations of this basic
*

type of stem structure are described under Axis'. Hairs are always uni-
cellular, but may be simple, unbranched, Y-shaped, 2-armed, peltate, or
 8o CRUCIFERAE
dendroid; and the walls are sometimes encrusted with calcium carbonate.
Glandular hairs rare. Water-storage cells are common in the epidermis,

FIG. 21. CRUCIFERAE

A, Simple hair from the upper side of the leaf of Capsella bursa-pastoris (L.) Medic. B, Forked hair
and
id glandular hair of Hesperis glutinosa Vis. from the floral region. C, Two-armed calcined hair from
the leaf of Cheiranthw cheiri L. D, and D 2 Stellate hairs from the leaf of Matthiola livtda DC. in
,

surface view and from the side. E, Stellate hair from the lower side of the leaf of Capsella bursa-pastoris
(L.) Medic. F, Peltate hair of Alyssum lepidotum Boiss. G, Dendroid hair from the lower side of the
leaf of Alyssum saxatile Linn. H and J, External glands of Matthiola lividai H, Entire glandular hair
seen from the side; J, Longitudinal section through the glandular head. A, E, and G after Vesque;
F after O. Bachmann the remainder by Solereder.
;

especially of the leaf. Crystals of calcium oxalate very infrequent, but

crystals, believed to be related to hesperidin, recorded in Capsella bursa-
pastoris (L.) Medik. Anomalous structure occurs in a few species.
 CRVCIFERAE 81

(ii)
WOOD
Vessels small, perforations simple, intervascular pitting alternate with
horizontal apertures, members very to extremely short. Parenchyma para-
tracheal, extremely sparse. Rays up to 2-4 cells wide, sometimes high,
heterogeneous. Fibres with small bordered pits, extremely short.

LEAF
Mesophyll structure variable; centric in Anastatica hierochuntica Linn.,
Crambe maritima Linn., Moricandia arvensis (L.) DC. Dorsiventral in leaves
on the lower part of the stem of Capsetta bursa-pastoris (L.) Medik. and
Lepidium sativum Linn. Upper cauline leaves of the same species with palisade
tissue towards both surfaces. Palisade tissue towards both surfaces in Cakile
edentula (Big.) Hook. Palisade and spongy mesophyll not differentiated in
Cakile maritima Scop. Wax commonly excreted on both surfaces. Palisade
tissue generally consisting of 1-3 layers.
Hairs (Fig. 21 A~J) very variable in form, but always unicellular apart from
the relatively infrequent glandular types; frequently calcified. Non-glandular
hairs often simple, but branched, T-shaped, and stellate kinds also occur, the
T-shaped variety having been recorded in species of Draba> Erysimum,
Farsetia, Lobularia, and 4-armed ones in Lesquerella spp. Glandular hairs,
consisting of a multicellular stalk and a head of i or more cells, recorded in
species of ^nchonium, Bunias, Chorispora, Descurainea, Hesperis, Matthiola,
Parrya, Sterigma. Simple and branched hairs occur either separately or
together. Glandular hairs generally to be found amongst other kinds. Treitel
(2276) found the structure and dimensions of the hairs to be the most impor-
tant anatomical features for the identification of leaves of different genera and
species.
Stomata (Fig. 22 E) typically cruciferous except sometimes, e.g. in
Sabularia\ generally occurring on both surfaces, but, according to Treitel
(2276), confined to the lower surface in Lunaria annua Linn. (syn. L. biennis
Moench.) and L. rediviva Linn. Stober (2201) found stomata to be more
numerous on the cauline than on the rosette leaves of Capsella bursa-pastoris
and Lepidium sativum, but those of rosette leaves larger and more elongated
than the ones on the cauline leaves in these 2 specie^. According to the same
author the size and shape of stomata vary considerably not only in these
2 species, but also in different individuals of either of them, or even in different
parts of a single leaf. Epidermis. Solitary water-storage cells interspersed
amongst normal epidermal ones in Heliophila spp., Isatis tinctoria Linn.,
Senebiera coronopus Poir. Small groups of these cells sometimes present as
well in the above species. Water-storage cells form a network on the surface
of the leaf in some species, e.g. Diplotaxis acris (Forsk.) Boiss., D. tenuifolia
(L.) DC., Eremobium aegyptiacum (Spreng.) Hochreut. (syn. Malcomia
aegyptiaca Spreng.),- Moricandia arvensis (L.) DC., Raphanus sativus Linn.,
Savignya parviflora (Del.) Webb. The cells in some species form elongated
sacs, e.g. Heliophila spp. Uniseriate, unbranched groups of cells, smaller
than those of the surrounding tissues, recorded by Schweidler (2060, 2061)
in the mesophyll of certain species, including Iberis pinnata Linn., /. umbellata
Linn., and Moricandia arvensis (L.) DC. Biseriate and branched groups less
4594 n
82 CRUCIFERAE
frequent. When in the palisade region each series has its long axis at right
angles to the epidermis, but parallel to the epidermis when in the spongy
mesophyll. No special contents noted in these cells.
Vascular bundles of the larger veins nearly always surrounded by collen-
chyma. Transverse sections through the distal ends of the petioles from a
random selection of different members of the family revealed 3 main but
intergrading types of vascular structure, (i) Asingle principal bundle accom-
panied by subsidiary strands in the wings in Alyssum saxatile Linn., Aubrietia
deltoidea (L.) DC. (Fig. 22 B) (main bundle supported by sclerenchymatous
masses with wide lumina to the cells), Cheiranthm cheiri Linn. (Fig. 22 c),
Draba sp. (ii) A crescent or U-shaped group of separate bundles in Armoracia
lapathifolia Gilib. (syn. Cochlearia armoracia Linn.) (In this species the
xylem vessels in each of the larger bundles form a U-shaped group bundles
;

strongly supported by sclerenchyma; many scattered subsidiary bundles also

present), Eunias orientate Linn. (Fig. 22 K), Lepidiwnsativum Linn. (Fig. 22 A),
Lunaria rediviva Linn., Matthicla incana (L.) R. Br. (a very open arc),
(iii) Larger
individual bundles of the main U-shaped group, each consisting
of a ring or cluster of vascular strands, scattered accessory strands often being
present as well, in Brassicajuncea (L.) Czernjaew, B. oleracea Linn. (Fig. 22 F),
Crambe maritima Linn. (Fig. 22 D), C. orientalis Linn. Bundles frequently
more numerous at the base than at the distal end of the petioles, e.g. 50-100
present at the base of Brassica oleracea. Considerable variation in petiolar
vascular structure sometimes occurs in different species of a large genus such
as Brassica. Crystals of calcium oxalate rare, but large ones recorded in a
species of Crambe, and in Sisymbrium altissimum Linn., although seldom
present in other members of the Sisymbrieae. Secretory cells* Various
attempts have been made to classify the Cruciferae according to the frequency
and distribution of the myrosin cells or idioblasts. Thus Heinricher (940)
recognized 5 types, (i) Myrosin cells distributed throughout the leaf paren-

chyma, (ii) Most of the myrosin cells situated in the parenchymatous sheaths
around the vascular bundles of the veins; only a small proportion distributed
in the mesophyll. (iii) Myrosin cells restricted to certain cells
of the sheaths
around the vascular bundles of the veins, (iv) Myrosin cells confined to the
veins, but occurring only in the mechanical tissue situated on the inside of the
parenchymatous sheaths, (v) Myrosin cells occurring in the mesophyll, but
predominantly in the sub-epidermis. Heinricher also divided the family into
7 sections according to the distribution of the myrosin cells in the stem.
Schweidler (2060, 2062) adopted a somewhat different arrangement which
takes into account the nature of the contents of the idioblasts as well as their
distribution. He recognized 3 groups, (i) Exo-idioblastae with chlorophyll-
containing idioblasts exclusively in the mesophyll. (ii) Endo-idioblastae with
idioblasts containing albumen but no chlorophyll, present exclusively in
association with the bundles, (iii) Heteroidioblastae, with idioblasts both in
the mesophyll and conducting strands. Using the nature and distribution of
the idioblasts as a taxonomic character, Schweidler made suggestions con-
cerning the possible close affinity of certain genera, and indicated that in other
instances species now included in a single genus should be separated. Details
can best be obtained from his papers, but since the frequency of these bodies
can be modified to some extent by varying the nutrition and environmental
 CRUCIFERAE 83
conditions, seems to be desirable that their taxonomic significance should
it

be investigated experimentally before their value is assessed.

Axis
STEM (Fig. 22 G-J, L)
Epidermis, usually composed of cells with strongly thickened inner walls;
sclerotic in Arabis procurrem Waldst. 2-Iayered in Diplotaxis harra
et Kit. ;

(Forsk.) Boiss, Zilla spinosa (L.) Prantl (syn. Zilla myagroides ForsL). Outer
part of primary cortex frequently collenchymatous. Palisade chlorenchyma
recorded in the outer part of the cortex of desert plants such as Eremobium
aegyptiacum (Spreng.) Hochreut. (syn. Malcomia aegyptiaca Spreng.) and
Zilla spinosa (L.) Prantl. Vertically elongated, unlignified but spirally to
reticulately thickened cells present in the outer cortex of the xerophytic species
Oudneya africana R. Br. Stem furrowed in Notothlaspi australe Hook, f.,
according to Betts (187), the cells below the furrows being smaller than those
elsewhere in the cortex and containing abundant chlorophyll. Cork of woody
species originating in the inner part of the cortex or in the phloem (Arabis
procurrens and Aubrietia deltoidea (L.) DC.). Phellogen in the perennating
stems of 'Alyssum arduinf arising, according to Hollendonner (988), amongst
mechanical cells at the bases of the petioles when the leaves become detached;
and also between the annual flowering and perennial sterns. A
second
phellogen subsequently arises in the bark around groups of sclereids which
then form islands. Endodermis well defined in species of Brassica, Cap-
sella Kernera, Lepidium, Nasturtium, Sinapis, and other genera. Pericycle
y

including well- developed strands or, in many genera including Brasstca,

Draba, Lesquerella, and Lunaria, a continuous ring of fibres.
Vascular structure showing a considerable range of variation (see below).
Vessels of variable diameter; perforations simple. Wood fibres with simple
pits. Pith generally parenchymatous, frequently occupying a large proportion
of the total diameter of the stem, becoming hollow in many species septate ;

in old branches of Diplotaxis harra (Forsk.) Boiss., outer portion prosenchy-

matous in Zilla spinosa (L.) Prantl. Adventitious shoots, stated by Wilson
(2438) to be exogenous in origin, arise from roots of Nasturtium austriacum
Cranz. (syn. Rorippa austriaca Spach.).
The xylem and associated tissues of the Cruciferae exhibit a considerable
range of variation, which is largely associated with differences in the maximum

transverse area attained by the stems of different species (Fig. 22 G-J, L).
Attention was drawn to this range of structure by Dennert, who recognized 7
distinct types which were cited by Schulz (2055). The following summary
was, however, drawn up after examining the slides in the collection at Kew.
In species with thin stems the xylem constitutes a closed cylinder as seen in
transverse section, the vessels being evenly distributed. This type of structure
is
accompanied by a considerable reduction or absence of recognizable
medullary rays. It occurs, for instance, in Alyssum spinosum Linn., Arabis
albida Stev., Draba sp., Iberis sempervirens Linn. (Fig. 22 G), Isatis tinctoria
Linn., Lepidium sativum Linn, (old stems), Matthiola incana (L.) R. Br. In
the above instances the cambium has the form of a continuous ring. This
applies also to species of Brassica, Crambe, Lunaria, and probably to many
other genera as well, but in these plants the xylem does not form a continuous
 FIG. 22. CRUCIFERAE
A, Lepidium sativum Linn. Petiole X 15. B, Aubrietia deltoidea (L,) DC. Petiole x 19. C, Cheir-
anthus cheiri Linn. Petiole x 15. D, Crambe maritima Linn. Petiole x 4. E, Iberis sempervirens Linn.
Stomata on lower surface X 167. F, Brassica oleracea Linn. Petiole X 9. G, Iberis sempervirens Linn.
Stem X 10. H, Lunaria rediviva Linn. Stem X 8. I, Alyssum spinosum Linn, Stem x 10. J, Bunias
orientalis Linn. Stem X 3. K, B. orientalis Linn. Petiole x 10. L, Armoracia lapathifolia Gilib.
Stem xg.
l.x. Lignified xylem. u.x, Unlignified xylem.
 CRUCIFERAE 85

ring since the primary bundles remain separated from one another by
lignified sclerenchymatous tissues, containing no vessels but nevertheless
produced by the cambium. This sclerenchymatous tissue could be interpreted
either as secondary xylem devoid of vessels or as lignified medullary ray tissue
produced by the cambium. In young stems of Brassica and certain other
genera the cambium is at first confined to the primary bundles, but subse-
quently extends into the interfascicular zones and so gives rise to the
structure just described. In other instances the primary bundles remain
individually distinct, but are separated from one another by interfascicular
sclerenchyma produced by lignification of the primary tissues, the cambium
taking no part in its formation. This structure is to be seen in young stems
of Alyssum saxatile Linn, and Brassica juncea (L.) Czernjaew as well as in
older material of Bunias orientalis Linn. (Fig. 22 j). Amongst the species
which have been examined its maximum development is attained in
Armoracia lapathifolia Gilib. (syn. Cochlearia armoracia) (Fig. 22 L), where
the cambium strands are still confined to the separate bundles even in fully
matured specimens.
Another interesting feature of the xylem is provided by species in which
the vessels of the first-formed wood are very minute compared with those in
the wood of older material. This is to be seen especially in species with thin
or wiry stems such as Arabis albida Stev., Aubrietia deltoidea (L.) DC., and
Iberis sempervirens Linn. Species with more fleshy stems, such as Bunias
orientalis Linn.,sometimes exhibit this feature as well, but to a less marked
extent. In plants such as Alyssum spinosum Linn. (Fig. 22 i), there are alter-
nating concentric zones of xylem with large and small vessels respectively,
the smaller ones being embedded in zones of relatively thin-walled ground
tissue. In these instances it looks as if the cambium periodically produces a

'juvenile' form of xylem. In Alyssum spinosum the smaller vessels are spirally
thickened, whilst the larger ones are provided with large, horizontal, bordered
pits. The structure of the wood in old stems of Alyssum saxatile Linn, is also
somewhat similar, since the xylem includes irregular islands of parenchyma.
The vessels in the first-formed wood in this species are minute. (See also
'Anomalous Structure'.)
It must be emphasized that, although the variation in the vascular structure
of the stems of the Cruciferae may serve as an aid to the identification of
genera and species, similarity of structure must not necessarily be regarded
as evidence of affinity. The structure appears to be related to the mechanical
and physiological requirements of the different species rather than to their
taxonomic relationships.

WOOD (Fig. 25 H and L)

Vessels small, mostly solitary, but with some radial multiples and clusters
(Fig. 25 L), sometimes with a slight tangential pattern in Brassica; seldom
touching the rays; about 40-50 per sq. mm. Perforations simple. Inter-
vascular pitting alternate, sometimes elongated horizontally, with horizontal
apertures; sometimes with striations due to coalescent apertures; pits to
parenchyma similar. Mean member length 1-8-2-4 mm -
Parenchyma
paratracheal, limited to rare cells about the vessels (Fig. 25 L). Rays up to
2-4 cells wide; less than i mm. high in Schouwia (1493), more than 2 mm. in
86 CRUCIFERAE
Brassica\ uniseriates few; 2-6 rays per mm.; markedly heterogenous and
composed mainly of square to upright cells. Fibres with small bordered pits,
equally numerous on both radial and tangential walls. Mean length about
0-4 mm. (1493)-

ROOT
Primary structure usually diarch, but tetrarch structure recorded in
Cochlearia. The layer of cells immediately external to the endodermis has
characteristic thickenings in a considerable number of unrelated genera, but
none have been recorded in others. Adventitious roots, stated to be exogenous
in origin, arise from subterranean stems or stems in contact with the soil in
Nasturtium austriacum Cranz (syn. Rorippa austriaca Spach.) according to
Wilson (2438).

ANOMALOUS STRUCTURE
Concentric zones of unlignified xylem present in the stems of Alyssum
spinosum Linn., Brassicafructiculosa Cyrillo, and Vellaspinosa Boiss. Ground-
work of wood composed of irregular masses of lignified and unlignified tissue
in Alyssum saxatile Linn. Scattered phloem islands recorded by Pfeiffer
(1712) in the xylem of certain species of Cochlearia Linn., Brassica Linn., and
Raphanus Linn. Medullary bundles, sometimes concentric in structure,
occur in the rhizome of Armoracia lapathifolia Gilib. (syn. Cochlearia
armoracia Linn.) and Raphanus sativus Linn. Secondary interxylary bundles
said to occur in the unlignified xylem of the rhizome of Armoracia lapathifolia
and in the root of Brassica napus Linn., B. campestris Linn. var. rapa Harm,
(syn. B. rapa Linn.), and Raphanus sativus Linn. Cortical bundles recorded
in Eruca saliva Mill, and Lepidium latifolium Linn. An inner cambium
recorded in the pith of 'Alyssum arduinf.

ECONOMIC USES
Useful vegetables such as cabbages, cauliflowers, turnips, swedes, &c., are
obtained from the genus Brassica. The seeds of other species of Brassica
yield commercial oils and other products such as Rape, Colza, and Mustard.
The family also includes the horseradish (Armoracia lapathifolia Gilib.) and
Woad (Isatis tinctoria Linn.).
The fleshy root of the horseradish (Armoracia lapathifolia) is bounded
externally by a few layers of thin-walled cork cells. A
broad layer of spongy
tissue is present on the inside of the phellogen and extends inwards to a
rather indistinct layer of cambium. The spongy tissue is composed of axially
elongated thin-walled parenchymatous cells, appearing circular in transverse
sections except where compressed against adjacent cells. Occasional, usually
isolated, stone cells occur in the spongy tissue immediately within the phello-
gen. The spongy tissue probably represents the cortex and phloem, but there
is no distinct line of demarcation between the two. Xylem consisting of a
broad region of isolated or small groups of vessels up to 60 or 70 \L in diameter,
embedded in spongy parenchymatous ground tissue. Vessels provided with
horizontal bordered pits and simple perforations. Groups of meristematic
cells occur in association with and sometimes partly or wholly surround
individual vessels or vessel clusters. Strands of similar, actively dividing cells
 CRUCIFERAE 87
also occur in the spongy ground tissue independently of the vessels. These
meristematic cells probably represent centres of proliferation of the parenchy-
matous tissue rather than true carnbial regions. Medullary rays absent.
Centre of the root occupied by a layer of very lacunar, parenchymatous pith.
The enlarged, tuberous portion of the stem of Brassica oleracea Linn. var.
gongy lodes Linn., known as Kohlrabi, is somewhat anomalous in structure.
The normal circle of bundles is very poorly developed, whilst the primary
rays are exceptionally broad and consist of thin-walled parenchyma. The
greater part of the tissue is made up of pith with a system of concentric
bundles embedded in it. The medullary bundles coalesce with one another
and are continuous with the normal bundle system of the stem at the base and
apex of the swollen portion. The developmental anatomy of Kohlrabi has
been described by Orsos (1640).

TAXONOMIC NOTES
A
well-defined family of herbs and small shrubs with several distinctive
anatomical characters. One of the most interesting is the universal occurrence
of myrosin cells, which are sometimes stated to be homologous with the
laticiferous elements of the Papaveraceae and other related families.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Alyssum,* Anastatica, Arabis, Aubrietia,* Brassica,* Bunias,* Cakile, Car-
damine, Cheiranthus,* Cochlearia,* Crambe,* Diplotaxis, Draba,* Eremo-
bium, Eruca, Goldbachia, Heliophila, Iberis,* Isatis,* Kernera, Lepidium,*
Lesquerella, Lunaria,* Matthiola,* Moricandia, Nasturtium, Notothlaspi,
Oudneya, Peltaria, Raphanus, Savignya, Senebiera, Sinapis, Sisymbrium,
Subularia, Vella, Zilla.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

1 2
Brassica, (Schouwia).
LITERATURE
(i) On
General Anatomy
Betts 187-90, Boodle 232, Bunton 309, Heinricher 940, Hollendonner 988, Orsos 1640,
Pfeiffer, H. 1712, Sabnis 1977, Schulz 2051-3, 2055, Schweidler 2060-2, Simmonds
2101, Soeding 2157, Starr 2188, Stober 2201, Treitel 2276, Wilson 2438, Wright 2475,
Zohary and Fahn 2511.
(ii) On Wood Structure
Messeri 1493.

27. CAPPARIDACEAE
(Fic. 23 on p. 88 ;
FIG. 24 on p. 90; FIG. 25 on p. 92; FIG. 26 on p. 96)

SUMMARY
(i) GENERAL
This mainly tropical family consists of herbs, shrubs, and small trees, while
a few lianes are included in Cadaba, Capparis, and other genera. Very few
1
Brassica oleracea var. ramosa (DC.) Alefeld.
*
Schouwia schimperi Jaubert et Spach. From the description given by Messeri (1493).
88 CAPP ARID ACE AE
characters are constant throughout the family. The stomata are ranuncu-
laceous. Crystals of calcium oxalate are usually small and clustered or pris-
matic; large solitary crystals rare, but recorded in Forchhammeria spp.

FIG. 23. CAPPARJDACEAE

Hairs of: A, Capparis spinosa L. B, C. quintflora DC. C, C. xeylanica L. ; D, Cadaba capparoides
; ;

DC.; E, Capparis verrucosa Jacq.; F-G, Steripkoma paradoxum (Jacq.) Endl.; H, Capparis ferntginea
L. J, C. domingensis Spreng.; K, Cadaba farinosa Forsk. L, Atanusquea emarginata Miers; M, Cap'
; ;

pans breyma Jacq. J, By Solereder; the rest after Vesque.

%

Myrosin cells have been observed in Capparis^ Cleome, Gynandropsis, and

Polanisia. Anomalous structure occurs in certain genera.

(ii) WOOD
Vessels very small to medium-sized, often in clusters or long radial
multiples, perforations simple, intervascular pitting alternate, small and
vestured, pits to parenchyma similar, members moderately to extremely short;
 C APP ARID ACE AE 89
small tracheid-like members sometimes numerous. Parenchyma para-
tracheal, usually sparsely vasicentric; fusiform cells often numerous and
occasionally forming bands; sometimes storied. Rays up to 2-5 cells wide,
apart from interfascicular rays when present, typically homogeneous or nearly
so,but sometimes composed entirely of square to upright cells. Fibres with
small simple pits, moderately to extremely short, sometimes storied. Included
phloem of the 'concentric* type present in some genera.
LEAF
Dorsiventral, isobilateral, or centric.
Hairs (Fig. 23 A-M).

(a) Simple unicellular, (i) Short, usually with thick walls, in Boscia
octrandra Hochst, B. salicifolia Oliv., Capparis cynophallophora Linn., and
Maerua crassifolia Forsk. (ii) Long in Capparis mollis H.B. et K. and Stixis
suaveolens (Roxb.) Pierre, (iii) Ribbon-shaped
in Capparis rothii Oliv. (iv)
Thin-walled, ascus-shaped hairs, with wide lumina and pointed at both ends
in Capparis galatea Fres., C. rupestris Sibth. et Sm., C. sarmentosa Cunningh.,
C. spinosa Linn, (v) Malpighian hairs (Fig. 23 B) in Capparis quiniflora DC.
and C. zeyheri Turcz. (vi) Many-rayed hairs (Fig. 23 c) in Capparis
avicenniaefolia H.B. et K., C. dealbata DC., C. diversiflora Wight et Arn., C.
erythrocarpa Isert., C.foetida Bl., C. heyneana Wall., C. horrida Linn., C. moonii
Wight., C. tenera Dalz., and C. zeylanica Linn.
(4) Uniseriate hairs in Capparis dtrifolia Lam., C. lanceolaris DC., and
C. tomentosa Lam.

(c) Emergences resembling shaggy hairs in Cadaba capparoides DC.

(Fig. 23 D), C. indicaLamk., and C. scaposa DC. (syn. C. paptllosa Steud.).
(d) Short, shaggy hairs in Morisonia americana Linn.
(i) With ray cells all lying in the same plane
(<?) Stellate hairs, in Steriphoma

paradoxum (Jacq.) Endl. (Fig. 23 F-G) and S.peruvianum Spruce, (ii) Brush-
shaped with ray cells radiating in all directions in Capparis angustifolia
H.B. et K. C. indica Fawcett et Rendle (syn. C. breynia Jacq.), C. detonsa Triana
,

et Planch., C. ferruginea Linn. (Fig. 23 H), C. incana H.B. et K., and C. yco
Mart, (iii) Candelabra hairs (Fig. 23 j) in Capparis domingensis Spreng.
(/) Peltate hairs of various types in certain species ofAtamisquea (Fig. 23 L),
Cadaba, Capparis (Fig. 23 M), Morisonia.
(g) Glandular hairs common in the Cleomeae, but rare in the Cappareae.

Variations in the striation of the cuticle stated to be of taxonomic value.

Lower epidermis papillose in species of Cleome, Crataeva, Isomeris, and
Wislizenia refracta Engelm.; locally papillose in Courbonia glauca DC.
Epidermal cells sclerosed in Capparis yco Mart. outer walls perforated in
;

Capparis baducca Linn. Hypoderm recorded in Forchhammeria and some

species of Capparis. Stomata ranunculaceous in the mature leaf; generally
present on both surfaces of isobilateral or centric leaves, but usually confined
to the lower surface in dorsiventral leaves. Mesophyll centric or dorsiventral.
Centric structure recorded in certain species of Boscia, Cadaba, Capparis,
Cleome, Courbonia, Maerua. Dorsiventral in certain species of Boscia, Cadaba,
Capparis, Crataeva, Maerua, Morisonia, Ritchiea, Roydsia, Steriphoma,
90 CAPPARIDACEAE
Thylachium, Tovaria. Mesophyll consisting wholly of short palisade tissue
in Capparis galatea Fresen. and Cadaba glandulosa Forsk. Fine transparent
striae, formed by a peculiar fissuring of the leaves on drying, occur in certain
species of Capparis and Forchhammeria. Variously shaped sclerenchymatous
cells (Fig.246-0) frequently present in the mesophyll of certain species of
Boscia, Cadaba^ Capparis (lowest layer of cells in the spongy parenchyma
sclerosed in C. zeylanica Linn. (Fig. 24 A)), Courbonia, Niebuhria, Thylachium.

FIG. 24. CAPPARIDACEAE

A, Transverse section through the leaf of Capparis zeylanica L. B, Transverse section through the
Jacq., with elongated idioblasts converging towards
leaf of Capparis adoratissima the base of a hair.
C, Idioblast of the leaf of Capparis domingensis Spreng. D, Transverse section of the leaf of Capparis
galeata Fres. with reservoir-tracheids. E, Upper epidermis of the leaf of Capparis cynophallophora L.
with crystals of gypsum. F, Upper epidermis of the leaf of Capparis polyantha Tr. et PI., likewise
containing crystals of gypsum. C, By Solereder; the remainder after Vesque.

Sclerenchymatous cells sometimes extending between the epidermal cells

and thus coming into contact with the leaf surface in Capparis and Boscia spp.
Smaller veins nearly always embedded in the mesophyll and sometimes
accompanied by sclerenchyma or collenchyma; vertically transcurrent in
Morisonia; the smaller veins of Capparis yco Mart, and C. zeylanica Linn,
also become vertically transcurrent owing to the development of sclerenchyma
on both sides of the vascular bundles. Veins in certain species of Cadaba and
Capparis end in pitted storage tracheids (Fig. 24 D). Midrib usually with
bundles arranged in a straight band or in an arc with the convex surface
towards the lower side; provided with a ring of bundles in Capparis pachaca
H.B. et K., Morisonia, and Steriphoma, the ring being formed by the presence
of a second arc of bundles having the xylem and phloem inversely orientated.
Petiole, in transverse sections through the distal end of the limited material
examined at Kew, exhibiting the following types of vascular structure. A
small crescent of very closely placed vascular bundles with strongly incurved
ends in 'Capparis linearis Jacq.' (Fig. 26 B) similar, but more clearly dissected
;
 CAPP ARID ACE AE 91
into separate bundles which form an almost closed cylinder in Steriphoma
ellipticum (DC.) Spreng (Fig. 26 A); a wide but shallow arc of about 14
individually distinct bundles in Cleome spinosa Jacq. (Fig. 260); a solitary
arc but with the ends detached so as to appear as distinct strands, and with
the conspicuous xylem vessels closely packed in very definite radial rows in
Crataeva religiosa Forst. (Fig. 26 c).
Crystals mostly secreted in the form of solitary prisms or clusters in the
mesophyll of certain species of Capparis. Solitary crystals occur throughout
the epidermis in species of Cadaba, Capparis (Fig. 24 E-F), Cleome, Forch-
hammeria, Maerua, Morisonia, Niebuhria. The crystals not always composed
of the same chemical substance. Sphaero-crystalline masses of gypsum occur
as wart-like elevations of the leaf in Cladostemon kirkii (Oliv.) Pax et Gilg and
Thylachium panduraeforme (L.) Juss. and large clustered crystals of gypsum
in the mesophyll of certain species of Capparis, Small rhomboid, octahedral,
or rounded crystalloids present in the epidermis and ground tissue of the
petiole of Boscia\ colourless, refractive, calcareous deposits recorded in
Capparis micrantha A. Rich and other members of the same genus. Myrosin
cells (see also 'Axis'), in Capparis, Cleome, Gynandropsts, Polanisia> occur
either free in the mesophyll or accompany the vascular bundles in the veins;
shape not very distinct from that of adjacent cells except for a tendency to be
vertically elongated and sometimes arranged in vertical groups in the cortex.
Myrosin also reported by Exbrayat-Durivaux (669) to occur in seedlings of
Capparis spinosa Linn., Cleome spp., and Dactylaena micrantha Schrad.

Axis
YOUNG STEM (Fig. 26 E)
Cork originating in the epidermis in Crataeva, in the sub-epidermis in
Cadaba glandulosa Forsk., and at various points in the cortex in Capparis.
Cork cells often remaining thin-walled, but becoming unilaterally thickened
in Crataeva. Aerenchyma arising in the innermost part of the primary
cortex of the lower part of the stem of specimens of Cleome spinosa Jacq. from
swampy habitats. Primary cortex with groups of stone cells in Crataeva and
Forchhammeria, those in the latter genus being especially numerous and filled
with large solitary crystals of calcium oxalate. Stone cells in young stems of
Capparis baducca Linn, arranged in a closed ring. Pericycle including
strands of fibres alternating with stone cells to form a closed ring in Cadaba
glandulosa and Roydsia or incomplete rings in Capparis and Crataeva.
Isolated groups of stone cells reported by Sabnis (1977) to occur in species of
Capparis and Cleome. Solitary crystals of calcium oxalate present in the peri-
cyclic stone cells of Cadaba and Forchhammeria. Secondary phloem devoid
of fibres in nearlyall investigated genera, but well-developed fibres recorded

in Boscia rehmanniana Pest by Adamson (6), especially in old material.

Myrosin cells (see also 'Leaf) recorded in the primary and secondary cortex
of the stem and root in Capparis, Cleome, Gynandropsis, Polanisia. Crystals,
see under 'Leaf, 'Cortex', and Tericycle'.

WOOD (Fig. 25 A-E, J-K, and M)

Vessels very small (25-50 fi mean tangential diameter), e.g. in some species
of Atamisquea, Isomeris, and Maerua, to medium-sized (100-200 /a), e.g. in
 M
FIG. 25. CAPPAWDACEAE, A-E, J~K, and M; PAPAVERACEAE, F-G and I;
CRUCIFERAE, H and L
A, Atanusquea emarginata Miers. B, Crataeva religiosa Forst. C, Boscia senegalensis Lam. D,
Cadaba trifokata Wight et Am. E, homeris arborea Nutt, F, Bocconia frutescew Linn. G, B. frutescens
Linn. H> Brassica oleracea Linn. var. ramosa (DC.) Alefeld. I, Dendromecon rigida Benth. J,
Crataeva reUgiosa Forst. K, Cadaba trifoliate Wight et Arn. L, Brassica oleracea Linn. var. ramosa
(DC.) Alefeld. T.S. M, ForchhammerialongifoUa Standl. T.S.
 CAPP ARID ACEAE 93
some species of Crataeva and in Maerua linearis (DC.) Pax (syn. Niebuhria
linearis DC.); solitary and in short radial multiples without any pattern in

Apophyllum, Atamisquea, Bucholzia, Capparis (most species), Crataeva, Maerua

rosmarinoides (Sond.) Gilg et Benedict and Maerua linearis (but not in other
species of M.); with a marked radial pattern in Boscia, Cadaba (Fig. 25 K),
Maerua (most species), and Steriphoma', with both radial multiples and clusters
arranged in tangential lines in Isomeris', commonly in clusters in Apophyllum,
Cadaba, Capparis, Cleome, and Steriphoma, the clusters sometimes consisting
of i or 2 moderate-sized vessels surrounded by many minute, tracheid-like
vessels, e.g. in Cadaba trifoliata W. et A. and Capparis aphylla Roth. 2-35 per ;

sq.mm.; spiral thickening reported (Solereder 2158) in the tips of the

members in Cleome. Perforations simple, horizontal, or oblique and often
vertical on the side- walls of the smallest members; imperfect vessel members
very common in the genera with clustered vessels. Intervascular pitting
alternate, very small to minute, obscurely vestured (Bailey 73, 78); pits to
ray cells similar to the intervascular pitting. Often filled with solid deposits.
Mean length 0-15-0-3 mm. Parenchyma paratracheal, except in Cadaba
and Forchhammeria most commonly vasicentric and sparse (Fig. 25 j) some-
; ;

times more abundant and aliform or confluent, e.g. in Capparis p.p. (Jans-
sonius 1154), Crataeva p.p., Maerua p.p., and Steriphoma; apotracheal, in
4-seriate bands composed mainly of fusiform cells in Cadaba, and diffuse in
Forchhammeria in Bucholzia coriacea Engl. the parenchyma is of 2 types,
;

relatively thick-walled strands of 2 cells about the vessels (vasicentric) and

broad (4), continuous bands of thin-walled fusiform cells in numerous con-
;

centric bands in Steriphoma (Record and Hess 1886), terminal bands present
in some species of Capparis and Crataeva. Strands typically of 1-2 cells;
fusiform cells common in Bucholzia, Cadaba, Crataeva, and Isomeris. Tend-
ing to be storied in Atamisquea, Capparis, e.g. C. salicifolia Gris. (Cozzo 493),
Crataeva, Isomeris, and Maerua p.p. Rays usually up to 2-5 cells wide and
seldom more than i mm. high; occasionally up to 6 cells in Crataeva and
Isomeris very wide and high interfascicular rays present in Forchhammeria',
;

uniseriates typically few, except where the maximum width is only 2 or 3 cells,
e.g. in species of Atamisquea, Boscia, Bucholzia, Morisonia, Steriphoma, and
Stuebelia (Record and Hess 1886), and composed of square cells in Atamisquea,
but mostly of procumbent cells in Boscia and Bucholzia', 3-9 rays per mm.;
typically homogeneous, though with occasional single rows of square cells in
Boscia and some species of Capparis and Steriphoma', composed almost
entirely of square cells in Atamisquea, Cadaba, and Maerua rosmarinoides,
distinctly heterogeneous in Cleome (Chattaway 376). Crystals present in
unspecialized cells in Atamisquea, Capparis (Janssonius 1154), Morisonia, and
Steriphoma and occurring in almost every ray cell in Atamisquea emarginata
Miers. Fibres typically with very small simple or faintly bordered pits, but
with small, distinctly bordered pits in Forchhammeria', pits mostly on the
radial walls except in Forchhammeria and Isomeris and usually absent from the
walls touching vessels. Occasionally septate in Morisonia (Chattaway 376).
Walls thin to thick; sometimes with alternating bands of thinner and thicker-
walled fibres in Capparis. Occasionally storied. Mean length 0-2-0*9 mm -

Included (interxylary) phloem of the 'concentric* type (c.L circumvallatum)

occurs in certain genera (see p. 94).
94 CAPPARIDACEAE
ANOMALOUS STRUCTURE
Anomalous stem structure, consisting of successive rings of growth,
recorded in Boscia (Adamson 6), Cadaba, Forchhammeria^ Maerua^ Stixi$\
typically with successive layers of secondary xylem containing numerous rays
and often including more than one growth ring, but in Forchhammeria with
successive bundles of xylem and phloem repeating the structure of the young
stem, separated by large interfascicular rays. Secondary bundles arise in the
phloem in Maerua crassifolia Forsk. (syn. M. uniflora Vahl.) and in the
pericycle internally to the sclerenchyma in Forchhammeria trifoliata Radlk.,
but externally to the pericyclic fibres in Forchhammeria pallida Liebm.
Phloem of secondary growth zones forming continuous rings in Forchhammeria
and Maerua.
In Boscia rehmanniana Pest, according to Adamson (6), a succession of
irregular zones of phloem and xylem is produced from secondary cambia
arising externally to and at various distances from the phloem. The secondary
cambia nearest to the phloem behave normally by developing in a centrifugal
manner. Those farther out are in the form of 2 active zones which respectively
develop in centrifugal and centripetal directions, the new tissues formed in
the last of these ways exhibiting inverse orientation. Centripetal develop-
ment ceases sooner than the centrifugal. By repetition of this mode of growth
the wood assumes a complex structure. When the cambia cease to be active
they often become decomposed with the formation of gum, and subsequently
split along these lines of weakness. The branches are also easily detached
from the trunk along the planes of cleavage. Capparis lianes are normal in
structure.

TAXONOMIC NOTES
The
presence of myrosin cells in certain genera suggests that the Cappari-
daceae and Cruciferae may have affinities with one another, and also with the
Resedaceae where similar cells also occur. The considerable diversity of
structure in the family is partly correlated with the various habits of the
plants. Pax and Hoffmann (1678) include Koeberlinia in this family, but this
does not appear to be supported by its anatomical structure. It is described
in this book under Koeberliniaceae (p. 331). The wood anatomy of the only
species available of Forchhammeria (F. longifolia) differs significantly from the
rest of the family.

ECONOMIC USES
Capers are the pickled flower- buds of Capparis spinosa Linn.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Boscia, Cadaba, Capparis,* Cladostemon, Cleome,* Courbonia, Crataeva,*
Forchhammeria, Gynandropsis, Isomeris, Maerua, Morisonia, Polanisia,
Ritchiea, Steriphoma,* Thylachium, Tovaria, Wislizenia.
* Kew
Represented in the slide collection,

(ii) FOR WOOD STRUCTURE

Apophyllum, Atamisquea, Boscia, Bucholzia, Cadaba, Capparis, (Cleome),
 CAPP'ARID ACE AE 95
Crataeva, Forchhammeria, Isomeris, Maerua, (Morisonia), (Niebuhria),
(Roydsia), (Steriphoma), (Stuebelia).

LITERATURE
(i) On General Anatomy
Adamson 6, Exbrayat-Durivaux 669, Pax and Hoffmann 1678, Sabnis 1977*

(ii) On Wood Structure

Bailey 73, 78, Benoist 169, Burgerstein 310, Chalk and Chattaway 362, Chattaway 376,
Cozzo 493, Desch 574, Howard 1088, Janssonius 1154, Kanehira 1206, 1209, Lecomte
1334, Pfeiffer, H. 1712, Record 1783, 1843, 1851, Record and Hess 1886, Record and Mell
1894, Williams 2430*

28. RESEDACEAE
(Fie. 26 on p. 96)

SUMMARY
An herbaceous family which occurs in the North Temperate zone, par-
ticularly in the Mediterranean region. The hairs are simple, unicellular
trichomes, frequently rounded at the apex. Structures intermediate between
hairs and papillae also occur in some species and are believed to serve for the
storage of water. Myrosin cells are especially characteristic of the family,
and sometimes occur in association with the stomata. The epidermis of the
leaf consists of cells of 2 distinct sizes. The stomata are ranunculaceous.
The mesophyll is not clearly differentiated into spongy and palisade regions.
Crystals of calcium oxalate are generally absent.

LEAF
Hairs simple, unicellular, sometimes seated on short emergences; glan-
dular types absent. Cells of the epidermis often of 2 distinct sizes, the larger
ones in some instances projecting as papillae above the leaf surface, e.g. in
certain species of Caylusea, Oligomeris, and Reseda. Inner walls of the
epidermal frequently mucilaginous in species of Caylusea, Ochradenus^
cells

Oligomeris^ Reseda.Mucilaginous cells also recorded in the mesophyll of

Randonia africana Cosson, and near the leaf veins in Caylusea. Stomata
ranunculaceous, but sometimes accompanied by myrosin cells; recorded as
being present on both surfaces in certain species of Oligomeris and Reseda;
those of the xerophyte Reseda muricata Presl. not specially protected against
excessive transpiration according to Evenari (665). Mesophyll not clearly
differentiated into palisade and spongy regions, sometimes centric in Ochra-
denus and Oligomeris. Vascular bundles of the veins not accompanied by
mechanical tissue. Petiole, in transverse sections, exhibiting a single vascular
strand in Astrocarpus, but with an arc of bundles in certain species of Reseda.

Axis
YOUNG STEM (Fig. 26 F)
Cells of the epidermis becoming sclerosed in old stems of Ochradenus.
Palisade chlorenchyma and mucilaginous cells present in the cortex of
Ochradenus baccatus Del. in which the leaves are reduced. Myrosin cells
 FIG. 26. CAPPARIDACEAE, A-E; RESEDACEAE, F.
A, Steriphoma eUipticum (DC.) Spreng. Petiole X 25. B, 'Capparis linearis Jacq.' Petiole X 25.
C, Crataeva religiosa Forst. Petiole X 2^. D, Cleome spinosa Jacq. Petiole X 25. E, Capparis linearis
l

Jacq/ Stem X 25. F, Reseda luteola Linn. Stem x 8.

s.c. Secretory canal, u.c. Layer of stone cells with U-shaped thickenings.
 RESEDACEAE 97
exhibiting a red colour when treated with Millon's reagent, and often distin-
guishable from their neighbours by their greater length, recorded in the stem
of Reseda spp. Pericycle including strands of fibres in Ochradenus and
Reseda. Xylem and phloem forming closed cylinders traversed by narrow,
primary medullary rays. Vessel perforations simple.

WOOD
The secondary xylem is very similar in Ochradenus and Reseda. Vessels
with small lumina and simple perforations. Fibres with simple pits. Rays
1-2 cells wide.

ROOT
Woody or fleshy in different species. Containing myrosin cells similar
to those in the stem. Primary structure diarch.

ECONOMIC USES
The well-known Mignonette of gardens is Reseda odorata Linn.

TAXONOMIC NOTES
The general structure of the stem, as well as the occurrence of myrosin
cells, suggests that the Resedaceae have affinities with the Cruciferae. Myrosin
cells also occur in some of the Capparidaceae. It is interesting to note that
the stomata are not cruciferous.

GENERA DESCRIBED
Astrocarpus,* Caylusea, Ochradenus, Oligomeris, Randonia, Reseda.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Bolle aan, Evenari (Schwarz) 665.

29. CISTACEAE
(FiG. 27 on p. 98; FIG. 28 on p. 100; FIG. 29 on p. 106)

SUMMARY
(i) GENERAL
A family of woody or somewhat wiry herbs, but also including some shrubs.
Many species occur in the Mediterranean region. Most members of the
family are especially characterized by the nature of the hairs, which are of
the following kinds, (i) Simple, unicellular, rigid, appearing, when mature,
to have a double structure in their basal portions, (ii) Tufted, each component
often similar in structure to the simple hairs of type (i). (iii) Peltate, (iv) Various
forms of uniseriate and frequently capitate glands. The xylem and phloem
are in the form of a continuous cylinder around the pith. Calcium oxalate is
usually present in the form of abundant cluster crystals.
98 CISTACEAE
(ii) WOOD
Vessels small and numerous, occasionally with a tendency to arrangement
in radial or tangential rows or to be semi-ring-porous; numerous; perforations
simple, intervascular pitting alternate and small, pits to ray cells similar,
members moderately to very short. Parenchyma absent or very rare. Rays
most species, but sometimes up to 3 cells wide, low and hetero-
uniseriate in
geneous. Fibres with bordered pits, except in part of i genus, extremely
short.

FIG. 27. CISTACEAE

A, Basal part of a simple hair of Cistus creticus L. B, Tufted hair of Cistus creticus L. C-D, Tufted
hairsfrom the bracts of Cistus ladaniferus L, E, Peltate hair of Helianthemum squamatum (L.) Pers.
F-G, Glandular hairs: F, Cistus ladaniferus L.; G, Cistus creticus L. By Solereder.

LEAF
Dorsiventral or centric. Hairs of several kinds, (i) Simple, unicellular, rigid
hairs which appear, when mature, to have a second hair included in their basal
portions (Fig. 27 A-B). The basal portion of these hairs in Fig. 27 A~B, as
well as the apparent secondary ones included within them, are stained yellow
when treated with eau de Javelle, whilst Schultze's solution shows that the
distal part of the outer hair is of cellulose, but that the basal portion of the
outer and the whole of the inner hair are lignified. (According to Solereder,
whose views are supported by Card (742), this peculiar structure originates
after the young hair has become thickened, when the protoplasm retreats to
the base and secretes a cellulose cap which projects acutely towards the apex
 CIST ACE AE 99
of the hair, although it is enclosed within it. Contrary to Solereder's statement,
1
microchemical tests show that the 'cap of the inner hair is lignified. It seems
probable that the mode of development of these hairs would be worth
reinvestigating.) (ii) Tufted hairs (Fig. 27 C-D), consisting of several members,
each of which is often of the same type as the solitary ones described above,
but with their bases sunk in groups in the epidermis. Sometimes individual
hairs of a group are relatively short, and their basal portions concrescent.
(iii)
Peltate hairs (Fig. 27 E) also occur, (iv) Glandular hairs (Fig. 27 F-G),
which are uniseriate and sometimes capitate, but of very variable form,
several types sometimes occurring in a single species.
According to Gard (742) the nature of the hairs may be of
*
some value in
separating species of Cistus. Solitary hairs of the double* type recorded
especially on Lechea, to some extent in Hudsonia, and sporadically in other
genera. Tufted hairs widespread, probably occurring in all genera, but most
prevalent in the Cisteae (sensu Janchen) although known in Hudsonia as well ;

exhibiting a considerable range of forms in different genera and species.

Stellate hairs, peltate scales, and intermediate kinds of structures occur, the
scales sometimes giving a silvery appearance to those parts of the plants
covered by them, e.g. in Halimium atripliciifolium (Lam.) Spach and Helian-
themum squamatum (L.) Pers. Scales particularly thick- walled in Cistus
ladaniferus Linn. The downy covering of hairs, present especially in various
species of Fumana, said by Janchen (1142) to be formed from glands which
have lost their secretory function. Epidermis composed of cells with curved
or straight anticlinal walls; upper epidermal cells large in Hudsonia tomentosa
Nutt. according to Starr (2188). Epidermis mucilaginous in certain species
of Helianthemum. Stomata present on both surfaces in most species, but
apparently absent from the upper side in Cistus ranunculaceous. Mesophyll
;

always including well-developed palisade tissue. Vascular bundles of the

veins not accompanied by sclerenchyma, but in the more xerophyllous
species of Cistus said by Gard (742) to be embedded in columns of cells with
thick cellulose walls, the columns extending from the upper to the lower
epidermis and appearing as a white network surrounding green meshes of
chlorenchyma in whole leaves when examined by transmitted light. Veins
accompanied by mechanical tissue in Hudsonia ericoides Linn. Petiole of
Cistus, in transverse sections through the distal end, exhibiting an arc of 3 or
more, often about 5, collateral bundles, the one at the centre being larger
than the lateral ones; all of them embedded in ground tissue which is fre-
quently lacunar. Petiole of Helianthemum supplied by a single arc-shaped
bundle embedded in ground tissue with fewer intercellular spaces. A single
petiolar strand stated to occur also in Lechea. This generic distinction is not
absolute. (The leaves of Cistus are persistent, and their structure is stated to
vary with increase in age. When 2 years old they are, according to Gard (742),
always thicker than first-year leaves, owing to an increase in the number of
palisade layers and enlargement of the intercellular spaces. The hairy cover-
ing is more dense in young leaves, the glandular hairs in particular being
more numerous and their secretory function more active.) Silica bodies,
resembling cystoliths, recorded in the lamina of Cistus. Cluster crystals of
calcium oxalate abundant.
 ioo CISTACEAE
Axis
YOUNG STEM (Fig. 28 i)
First-formed cork arising in the epidermis in Cistus, but later-formed
phellogens more deeply seated. Cork described by Card (742) as specially

FIG. 28. V1OLACEAE, A-F and H; CISTACEAE, G and I

A, Viola lutea Huds. Stem x n. B, Hymenanthera obovata T. Kirk. Petiole x 17. C, H. obovata
T. Kirk. Stem x 17, D, Viola lutea Huds. Petiole X 17. E, V. odorata Linn. Petiole X 17. F, Melt-
cytus ramiflorus Forst. Petiole x 17. G, Cistus laurifolius Linn. Petiole X 17. H, Melicytus ramiflorus
Forst. Stem x 9. I, Cistus laurifolius Linn, Stem x 17.

well developed in desert species of Helianthemum belonging to the section

Eriocarpum. Cortex partly sclerenchymatous in Hudsonia tomentosa Nutt.
according to Starr (2188). Water-storage tissue frequently occurs in the
inner part of the cortex, e.g. in Helianthemum squamatum (L.) Pers. or in the
 CISTACEAE ici

endodermal region in H. apenninum (L.) Mill, and H. salidfolium (L.) Mill.

Endodermis clearly defined in young stems of certain species of Helian-
themum, but sometimes less obvious in Cistus. Pericycle including small
strands of thick-walled fibres, apparently more numerous in Cistus than in
Helianthemum in the species examined. Xylem and phloem present in the
form of a closed cylinder surrounding the pith. Vessels small, with simple
perforations. Wood fibres with abundant bordered pits. Rays narrow; stated
by Solereder to be absent from Lechea and according to Janchen (1142)
generally from Hudsonia. Pith fairly broad, consisting of moderately thick-
walled pitted cells, many filled with contents which stain deeply with haema-
J

toxylin* Contrary to Solereder s statement occasional canals with gum-like

contents were observed at Kew in the pericyclic region in old stems of a
cultivated variety of Helianthemum. Cluster crystals abundant. Solitary
types less frequent.

WOOD 1
(Fig. 29 A and F)
Vessels extremely to very small exclusively solitary or in occasional pairs
; ;

tending to be in radial rows in Hudsonia and in tangential rows in Helianthe-

mum, section Eriocarpum very numerous, e.g. with more than 100 per sq. mm.
;

in Cistus semi-ring-porous and with spiral thickening in some species of

;

Cistus, e.g. in C. laurifolius Linn. Perforations simple and oblique. Inter-

vascular pitting alternate and very small; pits to ray cells similar. Solid
deposits sometimes abundant in Cistus tyloses rare, except in Hudsonia.
;

Mean member length 0*2-0-3 mm. (Cistus). Parenchyma absent or rare.

According to Janchen, absent from Grocanthemum, most species of Fumana,
several species of Helianthemum and Lechea, but metatracheal parenchyma
present in Cistus. Vestal, however, states that parenchyma is absent from all
these genera and suggests that the low uniseriate rays in some species might
easilybe mistaken for wood parenchyma. Rays, according to Vestal, always
present, exclusively uniseriate in Helianthemum, Hudsonia, and Lechea p.p.
and 1-3 cells wide in Cistus and Lechea p.p. Vestal queries Piccioli's state-
;

ment, as quoted by Solereder, that rays are absent from Lechea; Janchen
(1142) states that rays are completely absent from Hudsonia, except from
H. montana Nutt. very low, usually less than 0-5 mm. high; uniseriates often
;

of only 1-3 cells and composed of mixed procumbent and upright cells;
rays very numerous, e.g. 15-25 per mm. in Cistus', heterogeneous (Kribs's
Types II B and III). Fibres with small bordered pits on both radial and
tangential walls in Cistus, but, according to Janchen, with simple pits in
Helianthemum, section Eriocarpum walls thin to thick. Mean length about
;

0-4 mm. in Cistus.

TAXONOMIC NOTES
The peculiar nature of the hairs is an interesting and characteristic feature
of the family. According to Solereder, hairs with the curious double type of
structure are known only in the Combretaceae besides the Cistaceae. Card
(742) has made a special study of the value of anatomical characters in
1
Based mainly on the descriptions by Janchen (1142) and Vestal (2329).
102 CISTACEAE
classifying the species of Cistus. His views are well summarized in his own
words as follows :

'Nous concluerons que la subdivision des Cistes par les considerations anato-
miques ne coincide pas en tous points avec celle qui a pour fondements les caract&res
exterieurs. L'anatomie fait apercevoir certaines affinites, certaines liaisons, que
n'indique pas la classification des Floristes. Quelle que soit la mdthode employee
Je resultat obtenu n'est pas satisfaisant; et cela parce que certaines especes ont

probablement disparu alors que d'autres sont encores restees inconnues/

ECONOMIC USES
Labdanum, Ladanum, a resinous substance used in perfumery, exudes
or
from the leaves and branches of certain species and hybrids of Cistus,
notably C. ladaniferus Linn., C. villosus Linn. var. creticus (L.) Boiss., and
C. cyprius Lam. (= C. ladaniferus L,mn.xlaurifolius Linn.). The dried,
crushed leaves and young stems of Cistus sp. are sometimes used as a substitute
for or adulterant of culinary herbs such as Marjoram (Origanum marjorana
Linn.). Cistus leaves have been detected in herb samples submitted to Kew
for identification. The characteristic stellate hairs provide an important

diagnostic feature.
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Cistus,* Fumana, Halimium,* Helianthemum,* Hudsonia, Lechea.
*
Represented in the Kew slide collection.

(ii) FOR WOOD STRUCTURE

Cistus, (Crocanthemum), (Fumana), (Helianthemum), (Hudsonia),
(Lechea).
LITERATURE
(i) On General Anatomy
Card 742, Janchen 1142, Ponzo 1733, Starr 2188.

(ii) On Wood Structure

Janchen 1x42, Record 1843, 1851, Vestal 2329.

30. VIOLACEAE
(Fic. 28 on p. 100 ;
FIG. 29 on p. 106)

SUMMARY
(i) GENERAL
Mostly herbs or shrubs, but including a few lianes. The family occurs in
temperate and tropical regions. Certain species of Viola, especially amongst
those from Chile, exhibit unusual habit forms and other ecological specializa-
tions such as reduced leaves. The stomata are either cruciferous or rubia-
ceous. The hairs are simple and may be unicellular or uniseriate. Glandular
leaf teeth occur in Viola. The epidermis is frequently mucilaginous, and
some of the cells are frequently filled with brown contents. Calcium oxalate
is usually present as solitary or clustered crystals. Inulin rare. The stem,

in most of the very limited number of species which have been examined, is
 VIOL ACE AE 103

generally characterized by closed cylinders of xylem and phloem, the

former containing a few vessels of small diameter, whilst the phloem is
generally devoid of sclerenchyma. In Viola some species are provided with
closed rings of xylem and phloem, but other members of the genus possess
individually distinct bundles, separated from one another by narrow bands
of prosenchymatous elements with wide lumina. This difference in organiza-
tion does not appear to be entirely determined by whether a particular species
is woody or herbaceous. In very young stems of Hymenanthera the bundles

are sometimes just sufficiently separated to be individually distinct, but with

increase in age the xylem and phloem form almost completely closed rings.
Sclerenchyma is frequently present in groups or almost continuous bands in
the pericycle, although certain species of Viola are devoid of mechanical
elements in this region. The pith is generally solid in woody species, but
frequently becomes hollow in herbs.

(ii) WOOD
Vessels small and numerous, occasionally with a radial pattern, sometimes
semi-ring-porous and with spiral thickening, perforation plates simple,
scalariform, or both, intervascular pitting usually opposite or scalariform, but
sometimes alternate, pits to ray cells similar and often simple; members
usually of medium length, sometimes very long. Parenchyma absent or as
rare cells about the vessels. Rays absent from Viola in the other genera of
;

2 distinct the larger up to 4-10 cells wide; high; often composed

sizes,
entirely of square and upright cells. Fibres with simple or bordered pits,
septate; usually short to medium, sometimes very long.

LEAF
Generally dorsiventral, but centric in certain species of Hybanthus. Hairs
consisting exclusively of simple unicellular or uniseriate trichomes, apart from
the glandular shaggy hairs of the leaf teeth in Viola. The latter stated some-
times to secrete lime. Cuticle usually thin, sometimes striated. The surface
is provided with a thin coating of wax in certain species of Viola. Epidermis

with the inner walls of the cells frequently mucilaginous in certain species of
Leonia, Rinorea, and Viola (but not in Hawaiian species of this genus examined
by Skottsberg, 2123); mucilaginous cells sometimes appearing as transparent
dots in the Cells containing a reddish, or, more rarely, yellow gum-resin
leaf.

present in the epidermis of certain species of Agatea, Anchietea, Corynostylis,

Isodendrion, Paypayrola, and Viola; practically the whole epidermis consisting
of cells of this kind in some species, but only occasional cells of this kind
present in others. (See also under 'Secretory Elements'.) Some of the
epidermal cells of V. mauiensis Mann, considerably larger than their neigh-
bours and serving for water-storage according to Skottsberg (2123). Similar
cells not observed by the same author in V. robusta Hillebr. but tending to be
formed in V. luciae Skottsberg, the last species being a hybrid between the
first two.
A hypoderm of several layers present in Hymenanthera latifolia Endl.
Stomata distinctly cruciferous in Hybanthus, Leonia, Paypayrola^ Rinorea
(pro parte); indistinctly cruciferous in Amphirrhox, Hymenanthera, Rinorea
(pro parte) rubiaceous in certain species of Anchietea, Corynostylis, Noisettia,
;
 104 VIOL ACE AE
Isodendrion, Melicytus\ transitional between cruciferous and rubiaceous in
certain species of Hybanthus and Viola, Stomata present on both surfaces or
confined to the lower side ;
confined to the upper surface in Viola tridentata
Menz. and V. muscoides Philippi. ; either deeply sunk or in the same plane as
the epidermis itself; situated at the bases of small pits or canals in some species
of Viola. Mesophyll including no clearly defined palisade tissue in certain
species of Viola> but clearly differentiated into palisade and spongy portions
in the Hawaiian species of Viola examined by Skottsberg (2123). A
projecting
network, consisting of strands of tissue containing 2-3 layers of palisade cells,
recorded by Solereder on the upper surface of an undetermined species of
Viola from Peru. Arm-palisade cells stated by Sabnis (1977) to occur on the
abaxial side in Viola stocksii Boiss. Small vascular bundles generally not
accompanied by sclerenchyma in most herbaceous and certain woody species,
but accompanied by sclerenchyma strands or sheaths in certain species of
Melicytus, Paypayrola, and Rinorea. Midrib of smaller leaves with only i
vascular bundle, but 2 present in the larger leaves, the upper bundle having
the xylem and phloem inversely orientated more complex vascular arrange-
;

ments also occur in large leaves. Petiole, in transverse sections through the
distal end, exhibiting a single large, crescentic, collateral vascular strand in

Hymenanthera obovata Kirk (Fig. 28 B) with a single crescent-shaped strand

;

either alone or accompanied by smaller lateral ones in the limited number of

Viola spp. (Fig. 28 D and E) so far investigated. Transverse sections through
the distal end of a petiole of Melicytus ramiflorus Forst. (Fig. 28 F) recently
examined at Kew showed an almost closed vascular cylinder, flattened and
slightly open on the adaxial side, the abaxial part of the cylinder tending to be
dissected into separate bundles, the whole vascular strand being surrounded
by an interrupted ring of fibres with wide lumina.
Calcium oxalate present as solitary and clustered crystals in the mesophyll
and petiolar ground tissue. (The existence of 8 types of crystal structure and
distribution is recorded by Solereder (2158) and Melchior (1483), but this
appears to need reinvestigation. The taxonomic significance of these types
also needs to be reassessed because some of the genera in which they are
stated to occur are probably members of the Ochnaceae rather than of the
Violaceae.) Sphaerocrystalline masses of an organic substance of unknown
composition recorded in certain species of Agatea and Rinorea. Secretory
elements. Skottsberg (2123) records the occurrence of secretory cells with
brown contents in the mesophyll but not in the epidermis of the Hawaiian
plants Viola luciae Skottsb. and V. mauiensis Mann., but none seen' in V.
robusta Hillebr. The old stipules in the same species wholly composed of
these cells. (See also under 'Epidermis' above.) Various kinds of ecological
specializations, which are believed to reduce transpiration, occur in Viola,
notably in the sections Andinium and Tridens.

Axis
YOUNG STEM (Fig. 28 A, c, and H)
Epidermis composed of tabular cells with thick outer walls, but locally
elongated in a palisade-like manner in Viola stocksii Boiss. according to
Sabnis (1977). Epidermal cells, oval with thin walls in V. cunninghamii
Hook. f. large, rectangular with thin walls in Hymenanthera dentata R.Br. var.
;
 VIOL ACE AE 105

alpina Kirk. ; inner walls gelatinized in the Hawaiian species of Viola examined
by Skottsberg (2123). A
hypoderm of large thin-walled cells noted at Kew
in Melicytus ramiflorus Forst. Cork described by Betts (188) as having the
form of a wide band in Hymenanthera dentata. Seen to arise superficially
in Hymenanthera obovata Kirk, and Melicytus ramiflorus^ component cells
with greatly thickened inner tangential walls in the last species. Cortex
collenchymatous in Viola stocksii according to Sabnis (1977); parenchymatous
in V. cunninghamii (Betts 188); consisting of thick- walled cells containing
oil drops in Hymenanthera dentata. Endodermis described by Betts (188)
as well defined, and consisting of large cells with thin suberized walls in
Viola cunninghamii, Pericycle including strands of fibres, in Amphirrhox,
Hybanthus, Hymenanthera, Melicytus, Paypayrola, and Rinorea, the strands
often becoming connected by stone cells to appear as a complete ring in
transverse sections. Sclerenchyma said by Sabnis (1977) to be absent from
the pericycle in Viola stocksii and by Betts (188) from V. cunninghamii.
Phloem according to the last 2 authors forming a closed ring in Hymenanthera
dentata, Viola cunninghamii, and V. stocksii. Xylem constituting a closed
ring with very few vessels in Hymenanthera dentata according to Betts (188).
The existence of this structure also observed by the same author in other
species of Hymenanthera as well as in Viola cunninghamii. Primary vascular
bundles of Melicytus ramiflorus Forst. (Fig. 28 H) examined at Kew appeared
individually distinct in transverse section, but united by lignified medullary
ray tissue. Vessels in the secondary xylem of slightly older stems of the same
species mostly in radial multiples, but a few solitary and in irregular clusters.
Vascular bundles also individually distinct and separated from one another
by narrow bands of prosenchymatous elements with wide lumina in certain
species of Viola (Fig. 28 A). Vessels with simple perforations and some
scalariform plates (see 'Wood'). Pith generally solid in woody, but some-
times becoming hollow in herbaceous species; the cells sometimes con-
taining brown deposits in woody species. Pith composed of moderately
thick-walled pitted cells in Melicytus ramiflorus. Solitary and clustered
crystals often abundant.
Skottsberg (2123) has recently recorded the following additional informa-
tion concerning the stem structure of the Hawaiian species Viola robusta
Hillebr., V. mauiensis Mann., and V. luciae Skottsb., the last being a hybrid
between the first 2 species. Amongst the characters noted in old stems of
V. robusta were the following. Primary cortex well developed. Endodermis
not clearly differentiated. Pericycle containing starch. Secondary phloem in
the form of a cylinder. Xylem dense, all constituent elements more or less
rectangular in transverse sections. No medullary rays visible in longitudinal
sections, but radiating rows of cells devoid of vessels observed in transverse
sections. Vessels narrow, infrequent, some of those near the primary xylem
provided with more or less well-defined scalariform perforation plates.
Occasional solitary vessels with brown contents noted. Short rectangular
cells with thick, lignified, pitted walls present between the protoxylem strands
and also forming a cylinder around the pith. Pith solid, even in the thickest
stems, but sometimes ruptured in dried material; composed of large, thin-
walled, amyliferous, parenchymatous cells. Cluster crystals present in the
cortex and pith. Stem of V, mauiensis similar in structure, but phloem
 I

FIG. 29. CISTACEAE, A and F; CANELLACEAE, B-C and G-H; BIXACEAE, D-E;
VIOLACEAE, I-J; COCHLOSPERMACEAE, K-L
A, Cistus laurifolius Linn. B, Warburgia ugandensis Sprague. C, Cinnamosma fragrant Baill. Show-
ing oil cell. D, Bixa orellana Linn. E, B. orellana Linn. F, Cistus laurifolius Linn. G, Warburgia
ugandensis Sprague. H, Cinnamosma fragrans Baill, I, Amphirrhox longifolia Spreng. J, A. longifolia
Spreng. K, Cochlospermum orinocoense Steud. L, C. williamsii Macbride.
i.e. = intercellular canal.
 VIOL ACE AE 107

containing numerous brown cells, and a circle of similar cells in the wood
parenchyma. Characters of the hybrid V. luciae intermediate between those
of the parents.

WOOD (Fig. 29 i and j)

Vessels usually very small (mean tangential diameter 25-50 /A), but occa-
sionally medium-sized (100-200 ^), e.g. inAgateaviolarisA. Gray exclusively
;

solitary in Agatea violaris and Anchietia (1886) and almost so in Viola

chamissoniana Gilg. solitary and in short radial multiples in the other genera
;

and occasionally in radial rows, e.g. in Hybanthus (1886) and Paypayrola

longifolia Tul.; usually about 25-70 per sq. mm.; very few, according to
Betts (188), in Hymenanthera dentata R.Br. var. alpina Kirk.; with spiral
thickening in Hybanthus and Hymenanthera; semi-ring-porous in Hymenan-
thera. Perforation plates usually exclusively simple in Agatea, Anchietia
(1886), Hybanthus, Hymenanthera, Melicytus, and Viola, and exclusively
scalariform in Ahodeia p.p. (2158), Amphirrhox, Gloeospermum, Leonia, Pay-
payrola, and Rinorea, but Solereder lists the following as having both simple
and scalariform plates: Anchietea, Ahodeia p.p., Corynostylis, Hymenanthera,
lonidium and Noisettia\ Janssonius (1154) notes a few simple perforations in
Ahodeia cymulosa Miq. the scalariform plates with few to numerous bars and
;

sometimes accompanied by irregular, coarsely reticulate plates. Intervascular

pitting typically alternate to opposite, but scalariform in Paypayrola and
sometimes in Leonia (1886); the pits usually moderate-sized to rather large,
but very small in Hybanthus and Melicytus\ pits to ray cells similar to the
intervascular pitting or larger, often simple or with very narrow borders.
Pitted tyloses sometimes present, e.g. in Agatea and Amphirrhox. Mean
member length 0-4-0*9 mm., but as much as 1*3 mm. in Paypayrola (1283).
Parenchyma absent or extremely sparse; when present, paratracheal, as a
few cellsabout the vessels; Dadswell and Record (533), however, describe
the parenchyma as 'sparingly paratracheal or diffuse' and Williams (2430)
refers to fine lines between the rays in Leonia glycycarpa Ruiz, et Pav. Rays
absent from Viola; in the other genera of 2 distinct sizes, the larger up to
4-10 cells wide and often more than 3 mm. high; reported to be wholly
uniseriate or only partially biseriate, in Gloeospermum sprucei Eichl. (2430)
and in Hymenanthera dentata var. alpina (188); uniseriates numerous and
composed of upright cells; usually about 12-15 rays per mm.; the large rays
usually composed almost entirely of square to upright cells, with uniseriate
marginal rows of 4-10, or sometimes more, cells; often with a tendency to
sheath cells (Fig. 29 j) commonly containing numerous single crystals.
;

Perforated ray cells sometimes present (358). Fibres septate and with simple
or indistinctly bordered pits in most genera, but with small bordered pits in
Agatea and Viola and, according to Solereder, with both simple and bordered
pits in Anchietea and Corynostylis. Walls thick, often with a gelatinous layer.
Mean length usually 0-5-1-2 mm. (100), but up to 2-5 mm. in Paypayrola
(Martin-Lavigne, 1450).

PEDUNCLE
The following features have been recorded by Skottsberg (2123) concerning
the Hawaiian species of Viola which he examined. Transverse sections of
 io8 VIOL ACE AE
Viola robusta Hillebr. show 4 wide vascular bundles separated by medullary
rays. Secondary tissues probably not formed in this species. Structure in
V. mauiensis Mann, differing somewhat from that of V. robusta as follows.
Epidermis and sub-epidermis composed of cells with thickened, gelatinized
walls. Secondary xylem in the form of a closed cylinder with numerous
strands of primary xylem on the inside. Secretory cells with brown contents
in the cortex, and with yellowish contents in the phloem. Transverse sections
of thick specimens of the scape of the herbaceous V. kauaiensis Asa Gray
exhibit considerable development of secondary xylem and medullary rays
with lignified cell walls. Numerous cluster crystals in the cortex and pith
of this species.

ROOT
Inulin stated by Solereder to be present in Hybanthus (lonidium)
ipecacuanha (L.) BailL

TAXONOMIC NOTES
Further investigation, especially of the woody members of this family,
seems to be desirable in order that its affinities may be more definitely estab-
lished. The genera Lavradia, Schuurmawia, and Sauvagesia, which were
included by Bentham and Hooker in the Violaceae-Sauvagesieae, are described
in this book under the Ochnaceae, where they have been included by Gilg (770)
and Hutchinson (1113). They differ anatomically in certain important respects
from the Violaceae.
Taylor (2237) has drawn attention to the similarity in wood anatomy
between this family and the Flacourtiaceae, and has suggested that the
Turneraceae also belong to this complex. Taylor also points out that,
may
although placed high in the Archichlamydeae, these families exhibit many
primitive features in their wood anatomy.

ECONOMIC USES
The root of Hybanthus (lonidium) ipecacuanha (L.) BailL is used as a sub-
stitute for true Ipecacuanha root (Cephaelis ipecacuanha (Brot.) A. Rich)
family Rubiaceae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Agatea, Amphirrhox, Anchietea, Corynostylis, Hybanthus, Hymenan-
thera,* Isodendrion, Leonia, Melicytus,* Noisettia, Paypayrola, Rinorea,
Viola*
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Agatea, (Alsodeia), Amphirrhox, (Anchietea), (Corynostylis), Gloeosper-
mum, Hybanthus, Hymenanthera, (lonidium), Leonia, Melicytus, (Noisettia),
Paypayrola, Rinorea, Viola.
 VIOL ACE AE 109
LITERATURE
(i) On General Anatomy
Betts 1 88, Gilg 770, Hutchinson 1113, Melchior 1483, Meyer 1504, Sabnis 1977,
Sayeedud-Din 2012, Skottsberg 2123, Thompson, W. P. 2254.
(ii) On Wood Structure
den Berger 182, Betts 188, Chalk and Chattaway 358, Dadswell and Record
Bailey 100,
533, Janssonms 1154, Kribs 1283, Martin-Lavigne 1450, Pfeiffer, J. Ph. 1713, Record
1843, 1851, Record and Hess 1886, Taylor 2237, Thompson 2254, Williams, LI. 2426,
2430-

31. CANELLACEAE
(Fie. 29 on p. 1 06; FIG. 30 on p. 118)

SUMMARY
(i)
GENERAL
Glabrous, aromatic trees, from tropical America, E. Africa, and Madagascar,
which are characterized especially by the presence of secretory cells in the
parenchyma of the axis and leaf as well as in the wood. The stomata are
generally rubiaceous, although no definite subsidiary cells are present in
Cinnamosma fragrans Baill. Bundles of fibres usually occur in the pericycle,
although they are sometimes rather late in developing. well-developed A
phelloderm generally formed. Calcium oxalate is most often secreted in
is

the form of clustered crystals, although solitary ones occur as well. The
medullary rays are narrow in the wood but broaden considerably in the
phloem.

(ii) WOOD
Vessels small, solitary, and few, perforation plates scalariform, intervascular
pitting opposite or occasionally elongated, members very to extremely long.

Parenchyma diffuse, or about the vessels, or both; containing numerous oil

or mucilage cells. Rays wholly uniseriate or up to 3 or 4 cells wide, homo-
geneous or nearly so, or of square and upright cells only, oil or mucilage cells
absent from most species. Fibres with conspicuous bordered pits, moderately
to very long.

LEAF
Epidermis with straight anticlinal walls. Stomata confined to the lower
surface; rubiaceous in Canella and Cinnamodendron, but ranunculaceous in
Cinnamosma fragrans Baill. A single layer of hypoderm also occurs in C.
fragrans. Mesophyll said by Solereder to include no palisade tissue in species
of Canella and Cinnamodendron hitherto examined. Vascular bundles of the
veins accompanied above and below by strands of fibres. Midrib including
a single vascular bundle. Petiole stated to exhibit 3 vascular bundles accom-
panied by sclerenchyma in Cinnamosma; a single, very open, arc-shaped
strand containing radial groups of small vessels in the xylem observed at Kew
in transverse sections through the distal end in Canella alba Murray (Fig. 30 H).
Vascular strand in the last species accompanied by rather scanty sclerenchyma.
Secretory cells with yellow contents always present, appearing as transparent
dots in cut leaves. Calcium oxalate usually secreted in the form of clustered
no CANELLACEAE
crystals, -but small solitary crystals sometimes present as well. Small
crystals situated in the epidermis of Canella.

Axis
YOUNG STEM (Fig. 30 K)
Cork originating in the sub-epidermis in Canella and Cinnamodendron.
A layer of phelloderm present in older stems, the inner cell wails being
provided with U-shaped thickenings in Canella. Pericycle containing
bundles or a continuous ring of fibres in Canella and Cinnamosma', pericyclic
sclerenchyma stated to be absent from Cinnamodendron. The amount of
sclerenchyma appears to vary with age in Canella and probably in the other
genera as well. Phlc ^,m characterized by sieve-tubes with scalariform sieve-
plates. Xylem constituting a closed cylinder traversed by narrow rays.
Vessels with scalarifcrm perforation plates. Secretory cells, similar to those
in the leaf, present in the cortex and xylem.

WOOD (Fig. 29 B-C and G-H)

Vessels moderately small (mean tangential diameter 50-100 /x) or slightly
larger; usually exclusively solitary, apart from apparent tangential or oblique
pairs due to overlapping ends; sometimes with a few radial pairs in Warburgia
and occasionally in short radial rows of vessels, which, however, are not in
actual contact (1886); usually about 4-10 per sq. mm., but more numerous
in Capsicodendron. Perforation plates exclusively scalariform, with 10-60 bars
(2329). Intervascular pitting rare, opposite, moderately large to large, some-
times horizontally elongated; pits to ray cells similar to the intervascular
pitting. Mean member length 1-2-2-3 mm Parenchyma apotracheal in
*

Capsicodendron and Warburgia, as isolated cells scattered among the fibres

and in short tangential lines (Fig. 29 G) paratracheal in Canella and Pleoden-
;

dron, rather scanty and limited to the abaxial sides of the pores; both types
present in Cinnamosma, the paratracheal parenchyma often vasicentric
(Fig. 29 H). Strands commonly of 12-16 cells. Oil or mucilage cells present
in Canella, Capsicodendron, Cinnamosma, and Warburgia. Rays i (occa-
sionally 2) cell wide in Canella, Capsicodendron, Cinnamosma, and Pleo-
dendron, commonly 2-3 (up to 4) cells wide in Warburgia', usually low, but
occasionally i mm. high in Canella and Warburgia', uniseriates moderately
numerous in Warburgia and composed of cells similar to those of the multi-
seriate rays;about 10-15 rays per mm.; homogeneous (Kribs's Type I), with
only occasional marginal rows of square cells, e.g. in Warburgia, or composed
almost entirely of square to upright cells, e.g. in Cinnamosma', commonly
containing crystals. Oil or mucilage cells in the rays observed only in Cinna
mosma (Fig. 29 c). Fibres with numerous, conspicuously bordered pits,
equally numerous on both radial and tangential walls and of about the same
size as the intervascular pitting. Walls moderately thin to thick. Mean

length 1-5-27 mm.

TAXONOMIC NOTES
Vestal (2329) in a study of the wood anatomy of the Guttiferae and their
allies, concludes that the Canellaceae do not belong to this group of families
and suggests that they could well be placed near the Myristicaceae and the
 CANELLACEAE in
arboreal Ranales. Garratt, on the other hand, in a study of the woods of the
Myristicacae (747), concludes that, despite certain points of similarity, the
Canellaceae seem rather far removed from this family. In another study
(747 A) Garratt concludes that, on the basis of wood anatomy, no relationship
with the Monimiaceae can be inferred.
The most outstanding feature of the wood, the oil or mucilage cells of the
rays and wood parenchyma, suggests affinity with the Magnoliaceae and
the Lauraceae. Though other characters lend little direct support, most of the
points of difference between the Canellaceae and the Lauraceae, except for
the common occurrence of septate fibres in the latter, might be explained as
due to different levels of specialization. Vessel member length and some other
characters suggest that the Canellaceae are much less highly specialized than
the Lauraceae and are even less specialized than the Magnoliaceae. Affinity
with the Lauraceae would imply relationship to the Monimiaceae and
Hernandiaceae, all of which have anatomical features in common.

ECONOMIC USES
The bark of Canella Murray is used for medicinal purposes. The bark
alba
of this species characterized by phelloderm consisting of stone cells strongly
is

thickened on the radial and inner walls; numerous oil cells; starch grains
simple or 2-3 compound, individual grains being up to 20 (usually 5-10)^
in diameter; cluster crystals of calcium oxalate; fibres occasionally present in
the pericycle but absent from the phloem. The scented wood of Cinnamosma
fragrans Baill. is exported from Zanzibar to Bombay, and probably Siam,
where it is used in religious ceremonies. Further details concerning this wood
have been recorded by Metcalfe (1496).

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Canella,* Cinnamodendron, Cinnamosma.
*
Represented in the Kew slide collection.

(ii)
FOR WOOD STRUCTURE
Canella, (Capsicodendron), Cinnamosma, (Pleodendron), Warburgia.

LITERATURE
(i) On General Anatomy
Gilg 772, Greenish and Walli? 812.

(ii) On Wood Structure

Howard 1088, Kanehira 1209, Record 1783, 1790, 1843, 1851,
Garratt 747, 747 A,
Record and Hess 1886, Record and Mell 1894, Metcalfe 1496, 1497, Vestal 2329.
 (112)

32. BIXACEAE
(FiG. 29 on p. 106; FIG. 30 on p. 118)

SUMMARY
A small tropical family consisting of the single genus Bixa. Owing to the
somewhat conflicting statements in the literature, the following description
of the leaf and young stem of the small tree Bixa orellana Linn, has been based
mainly on original observations on material grown at Kew. The wood exhibits
the following features. Vessels moderately small, with numerous small
multiples, perforations simple, intervascular pitting alternate and minute, pits
to parenchyma similar, members moderately short. Parenchyma diffuse,
storied. Rays mostly 1-3 cells wide, heterogeneous, the uniseriates storied.
Fibres with bordered pits, storied, moderately short.

LEAF
Dorsi ventral. Hairs described by de Boer (214) as either tufted or peltate,
the former consisting of small groups of unicellular trichomes and the latter
of a circular disk supported by a short quadricellular base. Stomata confined
to the lower surface; ranunculaceous about 115 per sq. mm. Petiole
;

(Fig. 30 G) almost circular in transverse section. Blocks of sclerenchyma,

appearing as an almost continuous ring because so closely packed together,
present immediately on the outside of the petiolar vascular strand, the latter,
in transverse sections through the distal end of the petiole, appearing to con-
sist of an almost continuous ring of phloem situated externally to a more

interrupted ring of xylem. Enclosed within the vascular ring is a central mass
of parenchyma with a subsidiary, collateral vascular strand, unaccompanied
by sclerenchyma, embedded in it. Xylem of the central strand appearing as
2 separate groups at least in some sections. Secretory canals, surrounded
by an epithelium of small cells and filled with dark refractive contents,
situated in the parenchymatous portion of the petiole both within and external
to the main vascular ring. Secretory cells present in the mesophyll and in
the peripheral tissues of the petiole. Cluster crystals of calcium oxalate
abundant.

Axis
YOUNG STEM (Fig. 30 j)
Cork superficial in origin.^ Pericycle including an almost continuous or
somewhat interrupted, composite ring of sclerenchyma. Phloem appearing
in transverse sections as strands separated from one another by the triangular
distal ends of the medullary rays, the widest portion of each ray being in
contact with the pericyclic sclerenchyma. Phloem stratified into alternating
hard and soft zones by approximately concentric rings of strands of fibres.
Xylem in the form of a closed cylinder traversed by narrow rays. Secretory
canals present in the pith; and secretory cells in the outer part of the
cortex. Cluster crystals of calcium oxalate occur in the cortex and pith.

WOOD (Fig. 29 D-E)

Vessels moderately small to medium-sized (mean tangential diameter 90 /x
in material examined) solitary and in numerous radial multiples of 2 or 3 cells
; ;
 BIXACEAE 113
about 20 per sq. mm. in material examined. Perforations simple, transverse
to oblique. Intervascular pitting alternate, minute and crowded. Pits to ray
cells similar to the intervascular pitting. Mean member length about 0-3 mm.

Parenchyma apotracheal, as scattered cells (Fig. 29 E) and short lines from

ray to ray. Strands usually of 4 cells. Storied. Rays up to 3, occasionally 4,
cells wide; mostly low, but the multiseriate rays sometimes up to i mm., and

commonly 2-3 stories, high; uniseriates moderately numerous and composed

of upright, square, and procumbent cells; about n
rays per mm.; hetero-
geneous (Kribs's Type II B), the multiseriate rays usually with not more than
i or 2
marginal rows of upright cells. Uniseriate rays storied. Fibres with
rather few, small, bordered pits on both radial and tangential walls, the
borders slightly larger than those of the intervascular pits. Storied. Mean
length about 0-8 mm.

TAXONOMIC NOTES
The stratification of the phloem into hard and
soft portions, the broadening
of the rays where they traverse the phloem, and the complex vascular struc-
ture of the petiole all suggest that Bixa has affinities with the Cochlosper-
maceae and Tiliaceae. The anatomy of the wood is in agreement with 'this
view, and the absence of septate fibres and the presence of diffuse parenchyma
distinguish the wood of Bixa very clearly from the woods of the Flacourtiaceae.
Secretory canals are also to be found at least in certain members of all these
families. The classification of Bixa and certain genera, now included in the
Flacourtiaceae, together in one family as in the system of Bentham and
Hooker presents difficulties on other anatomical grounds, besides the differ-
ences in wood structure just mentioned above. Solereder (2158) evidently
appreciated this difficulty also, because he begins his account of the Bixaceae
as understood by Bentham and Hooker by remarking that 'Anatomical
characters common to all members are almost entirely wanting in the Order*.

ECONOMIC USES
The
seeds of Bixa orellana Linn, yield Annatto dye which is not very
durable. It is used for colouring butter and cheese, &c.

GENUS DESCRIBED
Bixa.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Beauvisage 163, de Boer 214, Pilger 1722.

(ii) On Wood Structure

Record 1809, 1843, 1851, Record and Hess 1886, Record and Mell 1894, Stone 2206,
Vestal 2329.
 ("4)

33. COCHLOSPERMACEAE
(Fic. 29 on p. 1 06)

SUMMARY
(i)
GENERAL
The anatomy of the small tropical trees and shrubs included in this family
needs further investigation. Some of the following facts concerning the leaf
and young stem have been taken from Pilger's (1723) account.

(ii)
WOOD
Vessels medium-sized to large and rather few, perforations simple, inter-
vascular pitting alternate and rather large, pits to parenchyma often large and
simple, members of medium length. Parenchyma mostly as broad apotra-
cheal bands, but also vasicentric, storied. Rays up to 5 or 6 cells wide,
uniseriates common and storied, inultiseriates 2 or more stories high, hetero-
geneous. Fibres with small bordered pits, often storied, of medium length.
Radial intercellular canals usually present.

LEAF
Hairs mostly long, simple, unicellular, but peltate forms also occur in
Cochlospermum. Epidermis often composed of mucilaginous cells. Stomata
confined to the lower surface in Amoreuxia and Cochlospermum but recorded
on both sides in Sphaerosepalum alternifolium Bak. Vascular bundles of the
smaller veins not accompanied by sclerenchyma. Petiole with 3 bundles
entering the base in Amoreuxia, Cochlospermum, and Sphaerosepalum, but
exhibiting a closed cylindrical strand in transverse sections through the distal
end. Subsidiary, medullary bundles such as those in Bixa (Family Bixaceae)
absent from the petiole. Secretory cells with yellow or reddish resinous
contents present in the mesophyll of Amoreuxia and Cochlospermum*, large
mucilage cells recorded in the corresponding position in Sphaerosepalum.
Large prismatic crystals also present in the mesophyll of Sphaerosepalum.

Axis
YOUNG STEM
Pericycle including isolated strands of fibres. Phloem traversed by the
triangular distal ends of the medullary rays as in some of the Bixaceae and
Tiliaceae; stratified into sclerosed and soft portions in Cochlospermum.
Secretory cells with yellow or reddish resinous contents recorded in the
primary cortex of Amoreuxia and Cochlospermum, and large mucilage cells in
the corresponding position in Sphaerosepalum. Mucilage canals present in
the inner part of the cortex and the periphery of the pith in Amoreuxia and
Cochlospermum. Large prismatic crystals present in the primary cortex of
Sphaerosepalum.

WOOD (Fig. 29 K-L)

Vessels medium-sized (100-200 /x mean tangential diameter) to large
(more than 200 ju,); solitary and in small multiples; about 5-7 per sq. mm.
Perforations simple. Intervascular pitting alternate, rather large and often
 COCHLOSPERMACEAE 115
with horizontal apertures pits to ray and wood parenchyma usually larger than
;

the intervascular pits, elongated horizontally and simple. Tyloses sometimes

abundant. Mean member length about 0*6 mm. Parenchyma predominantly
apotracheal; in regular to irregular bands 2-8 cells wide and in narrow sheaths
about the vessels (vasicentric) (Fig. 29 L). Strands usually of 4 cells. Storied.
Cell walls very thin and apparently unlignified the cells of the bands some-
;

times much dilated, e.g. in Cochlospermum vitifolium (Willd.) Spreng. Strands

usually of 4 cells. Rays typically up to 5 or 6 cells wide, but with occasional
very wide and high rays in some species; uniseriates moderately numerous,
composed of both procumbent and upright cells and often storied; multi-
seriate rays usually 2 or more stories high; about 6 rays per mm.; hetero-

geneous (Kribs's Type II B) with 1-2 marginal rows of square to upright

cells. Fibres. Pits with very small borders, mostly on the radial walls. Walls

moderately to very thin. Often storied and sometimes markedly 'bauchig*.

Mean length about 1*1 mm. Intercellular canals usually present in some of
the rays, but not common (Fig. 29 K).

TAXONOMIC NOTES
Amoreuxia and Cochlospermum have characters in common with the
Bixaceae as understood in the systems of Engler and Prantl and Hutchinson.
In Dr. Hutchinson's opinion, however, Sphaerosepalum should not be included
in the Cochlospermaceae, its affinities appearing to lie rather with the
Guttiferae. It will be seen above that there are anatomical differences between
Spliaerosepalum and the two other genera, but Sphaerosepalum needs further
anatomical investigation before an opinion on its taxonomic position can be
expressed by an anatomist.

ECONOMIC USES
Cochlospermum gossypium DC. from India yields Katira Gum. A yellow
dye is obtained from the roots of Cochlospermum tinctorium Rich.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Amoreuxia, Cochlospermum, Sphaerosepalum.

(ii) FOR WOOD STRUCTURE

Cochlospermum.
LITERATURE
(i) On General Anatomy
Pilger 1723.

(ii) On Wood Structure

Record 1851, Record and Hess 1886, Vestal 2329, Williams 2430.
 (n6)

34. FLACOURTIACEAE
(Fic. 30 on p. 118; FIG. 31 on p. 120)

SUMMARY
(i) GENERAL
A
family of woody plants which are mostly shrubs or small trees; occa-
sionally they attain a considerable size. The family is mostly tropical, although
a few genera are known from South Africa, New South Wales, Chile, whilst
Idesia grows in Japan. The anatomy does not exhibit any particularly striking
or noteworthy features, but a more complete examination might well be
undertaken, as many of the *genera do not appear to have been investigated.
Hairs. Simple, unicellular, sclerenchymatous trichomes; tufted; 2-armed;
with thin partitions between the component cells.
stellate; peltate; uniseriate,
The stomata are rubiaceous or cruciferous. The petiole, in the few species
investigated, is provided at the distal end with either i vascular bundle or
with 3 which may become united to form an arc which is open on the adaxial
side. Schiaogenous secretory cavities occur in many of the genera which
were included by Bentham and Hooker in the Samydaceae. Calcium oxalate
is present as clustered or solitary crystals, sometimes situated in specialized
the epidermis. The nature and distribution of the
cells (crystal-idioblasts) in
calcium oxalate crystals is often helpful in recognizing genera and species.
The presence of large solitary prisms in some of the stone cells of the pericycle
of the young stem is characteristic of certain genera, whilst in others there is an
abundance of cluster crystals in the phloem. Cluster crystals were also found
to be very numerous in the parenchymatous tissues of the petiole in all of the
species available for examination. Further details are recorded under 'Axis'.
The pericycle in the young stem includes strands of fibres which are
sometimes united by stone cells to form a composite ring, which in some
genera is rather broad. The phloem is not stratified Into hard and soft
portions, and the medullary rays are not known to be broadened where they
traverse the phloem except in Erythrospermum. The phloem and xylem in
young stems are in the form of closed cylinders traversed by narrow rays.

(ii) WOOD
Vessels usually moderately small; solitary and in twos and threes; occa-
sionally with spiral thickening, perforations usually all simple, but some gen-
era with a few, and others with wholly, scalariform plates; intervascular
pitting (a) scalariform, opposite or large and alternate, with pitting to ray
cells large and often scalariform, or (b) very small and alternate, with pits to

ray cells similar; members moderately to very long. Parenchyma absent or

very sparse; when present paratracheal. Rays usually 3-5 cells wide, but
sometimes only i or 2, and occasionally up to 15, cells wide; high to very
high; markedly heterogeneous, sometimes composed entirely of square and
upright cells. Fibres typically septate; pits simple to distinctly bordered,
limited to the radial walls; of medium length to very long.

LEAF
Generally dorsiventral but sometimes centric; consisting entirely of palisade
tissue in Xylosma ellipticum Tul. Hairs of several types, (i) Simple, uni-
 FLACOURTIACEAE 117
cellular in Carriereaand Itoa; unicellular with blunt protuberances in Hopel-
stigma. (ii)Simple, unicellular, sclerenchymatous trichomes in Banara,
Byrsanthus, Calantica, Casearia, Homalium, Samyda, Tetrathylacium, Zue-
lania. (iii) Tufted hairs in certain species of Banara, Casearia, Ryania.

(iv) Two-armed hairs in Banara sp. and Homalium sp. (v) Stellate hairs
recorded in species of Banara, Homalium, Kiggelaria, Patrisia, Pineda.
(vi) Peltate hairs in Mayna and probably other members of the Oncobeae.
(vii) Uniseriate hairs with
thin walls between the component cells in Casearia,
Samyda, Zuelania. (viii) Uniseriate, glandular hairs, sometimes very long
and provided with a rounded terminal cell, in Hopelstigma. Wart-like emer-
gences, stated to function as hydathodes, recorded on the petiole at the ends
of vascular bundles in Scolopia. Epidermis characterized by the presence
of crystal-idioblasts in species of Bembida, Camptostylus, Casearia (many spp.
idioblasts sometimes very large), Dasylepis, Erythrospermum, Itoa, Laetia,
Ludia, Lunania (many spp.), Oncoba, Ophiobotrys, Osmelia, Poggea, Pyramido-
carpus, Rawsonia, Samyda, Scolopia, Xylosma, Zuelania (in some members of
the Apocynaceae similar crystals occur in the epidermis). Pits in the walls of
the epidermal cells not uncommon, and also secondary tangential divisions
of the cells in Scolopia and probably other genera. Epidermis composed
of mucilaginous cells in species of Gerrardina, Paropsia, and Neumannia',
papillose in Idesia sp. and on the lower surface in Berberidopsis. Inner and
lateral walls of the epidermal cells thickened in Laetia coriacea Spruce,
Cuticle exhibiting markings comparable with those on etched glass recorded
in Aphaerema. Banara, Bembicia, Byrsanthus, Calantica, Casearia, Dissomeria,
Euceraea, Gerrardina, Homalium, Lunania, Ophiobotrys, Osmelia, Pyramido-
carpits, Samyda, Tetrathylacium, Zuelania. Wedge-shaped ridges of cuticle,
stated to be very striking in surface view, recorded in Banara brasiliensis
(Gray) Benth. Wax present on the surface in certain species of Laetia and
Oncoba. Hypoderm of one to several layers present beneath the upper
epidermis in species of Banara, Byrsanthus, Casearia (many spp.), Dasylepis,
Erythrospermum, Euceraea, Homalium, Itoa, Kiggelaria, Laetia, Ludia, Xylosma,
and Zuelania. Stomata generally confined to the lower surface, but recorded
on the upper surface as well in Casearia and Lunania rubiaceous in species
;

of Banara, Bembicia, Berberidopsis, Byrsanthus, Calantica, Carrierea, Casearia,

Dissomeria, Euceraea, Flacourtia, Homalium, Itoa, Laetia, Ludia, Mayna,
Oncoba, Pangium, Samyda, Xylosma, Zuelania', cruciferous in species of
Centroplacus, Neumannia, Osmelia, and Rawsonia\ with a tendency towards
the cruciferous type in Erythrospermum, Hydnocarpus, Kiggelaria, Mayna.
Stomata appearing as islands owing to the contrast between the thin-walled
subsidiary cells and the surrounding thick-walled pitted cells of the remainder
of the epidermis on the lower surface of Banara portoricensis Urb. Mesophyll
containing sclerenchymatous fibres in certain species of Calantica, Casearia,
Homalium, Zuelania. Simple and branched sclerenchymatous idioblasts
(spicular cells) recorded in species of Centroplacus, Erythwspermum, Patrisia,
and Ryania. Arm-palisade cells recorded in Casearia. Upper layers of the
spongy mesophyll strongly thickened and filled with brown contents in many
species of Lunania. Vascular bundles of the smaller veins generally accom-
panied on both the upper and lower sides, or wholly surrounded by a thick
ring of sclerenchyma. Vascular bundles in the larger veins of Erythrospermum
 o
FIG. 30. FLACOURTIACEAE, A-C, E-F, I, L-O; BIXACEAE, G and J;
CANELLACEAE, H and K; THYMELAEACEAE, D
A, Dovyalis caffra (Hook. f. et Harv.) Warb. Petiole x 15; sclerosed cells on the adaxial side of the
vascular arc not shown. B, Oncoba spinosa Forfik. Petiole X 15. C, Azara gilliesii Hook, et Am. Petiole
X 15. D, Solmsia calophytta Baill. Petiole X 15. E, Gerrardinafoliosa Oliver. Petiole X 15. F Homalium
t

africanum Benth, Petiole X p. G, Bixa orellana Linn. Petiole X 10. H, Camlla alba Murr. Petiole
X 15. I, Azara dentata Ruiz, et Pav. Petiole x 15. J, Bixa orellana Linn. Young stem X 10. K,
*
Canella alba Murr. Stem X 15. L, Azara dentata Ruiz, et Pav. Stem X 15. M, Homatium africanum
Benth. Stern X 9. N, Berberidopsis corallina Hook. f. Stem x 15. O, Asteropeia sphaerocarpa Baker.
Stem X 7.
b.f. Phloem fibres. c.e. Cutinized epidermis, h.p. Hollow pith. /. Lenticel. m.c. Mucilage canal.
p.r. Primary rays broadening in the phloem, p.s. Pericyclic sclerenchyma. s.c. Secretory cells, s.r.
Secretory canal.
 FLACOURTIACEAE 119

sp.forming an almost closed ring. Midrib of Dipentodon sintcus Dunn, with

crescent-shaped groups of xylem and phloem, with additional phloem groups
on the adaxial side between the ends of the crescent, the whole strand being
surrounded by a broad cylinder of fibres. Petiole, in transverse sections
through the distal end of material examined at Kew, exhibiting the following
types of structure, (i) Median petiolar vascular strand solitary and crescentic.
(a) Vascular strand shallow, with sclerosed cells between the ends of the
vascular crescent in Dovyalis caffra (Hook. f. et Harv.) Warb. (Fig. 30 A).
(b) Vascular strands thick, slightly crescentic, sometimes with sclerosed cells
between the ends of the crescent in Azara dentata Ruiz, et Pav. (Fig. 301), and
Flacourtia prunifolia H.B. et K. Structure somewhat similar in Homalium
africanum Benth. (Fig. 30 F). (c) Vascular strand of Gerrardina foliosa Oliv.
deeply crescentic and strongly supported by thick- walled fibres in the pericyclic
region, inwardly directed strands of fibres extending towards each other from
the ends of the vascular crescent without quite meeting, (d) Vascular strand
crescentic with strongly incurved ends in Berberidopsis corallina Hook. f. An
arc-shaped vascular strand, accompanied by smaller ones towards the wings,
also noted in Carrierea calycina Franch. (2) Vascular arc dissected, (a) Median
vascular strands of Kiggelaria africana Linn, slightly interrupted and provided
with inwardly directed ends, the xylem including well-marked radial multiples
of thick-walled vessels, (b) Vascular system of Azara gilliesii Hook, et Arn.
(Fig. 30 c) dissected into several distinct strands (3) Median vascular strand
cylindrical, (a) Vascular strand circular in Hydnocarpus wightiana Blume and
Oncoba spinosa Forsk. (Fig. 30 B), the xylem including very well-marked, radial
multiples of thick- walled vessels, (b) Vascular strand cylindrical, but adaxially
flattened or slightly concave in Poliothyrsis sinensis Oliv., the thick- walled
xylem vessels being in well-marked radial multiples.
Cluster crystals very common and often abundant; observed in the cortical
region of the petiole in species of Azara, Berberidopsis, Dipentodon, Dovyalis,
Flacourtia, Gerrardina, Homalium, Hydnocarpus, Kiggelaria, Oncoba, Polio-
thyrsis, Bodies resembling cystoliths recorded in Homalium.
Rawsonia.
(See also under 'Epidermis'.) Secretory cavities (described as resin glands
by Gilg, 773) not always conspicuous in dried material, present in the
mesophyll of most species of Casearia, and commonly or occasionally also in
Laetia, Lunania, Osmelia, Samyda, Zuelania.

Axis
YOUNG STEM (Fig. 30 L-O)
Structure rather imperfectly known. Cork superficial in origin in most
of the species investigated; consisting of thin- walled cells in species of
Azara, Dovyalis, Hydnocarpus, Poliothyrsis. Pericycle including isolated
strands of fibres, sometimes united to form a composite ring. Isolated strands
of fibres recorded in species of Azara, Dasylepis, Dendrostylis, Grandidiera,
Gynocardia, Oncoba. A
composite and continuous ring of sclerenchyma
present in species of Banara, Casearia, Dipentodon, Homalium, Hydnocarpus,
Poliothyrsis, Ryana, Taraktogenos\ a sclerenchymatous ring composed of
slightly elongated parenchymatous cells recorded in a species of Physena.
Phloem of Erythrospermum, like that of the Bixaceae and Cochlospermaceae,
traversed by enlarged distal ends of the medullary rays, the rays appearing
 FIG. 31. PITTOSPORACEAE, A-C; FLACOURTIACEAE, D-L
A, Pittosporum ferrugineum (Dryand.) Ait. B, Bursaria sptnosa Cav. C, Pittosporum ferrugineum
(Dryand.) Ait. D, Idesia polycarpa Maxim. E, Homatium tomentosum Benth. F, Casearia gladnfornns
Mast. G, Ryania speciosa Vahl. H, Dovyalis xyssyphoides E. Mey. I, Homalium longifolium Benth,
J, H. smytkei Hutch, et Dalz. K, Ryania speciosa Vahl. L, Taraktogenos kunstleri King.
 FLACOURTIACEAE 121

triangular in transverse sections. Rays in other investigated members of the

Flacourtiaceae not enlarged in the phloem. Phloem and xylem, in most of
the genera examined, forming continuous cylinders traversed only by narrow
rays. Thick, longitudinal strands of stone cells present in the phloem of
Homalium africanum Benth. Vessels mostly with simple perforations but
occasionally with scalariform plates, the latter occurring especially in the
primary wood, e.g. in Homalium africanum (see also 'Wood'). Pith very
heterogeneous in Berberidopsis corallina Hook. f.
Crystals frequently very abundant, their nature and distribution often
characteristic of and helpful in identifying genera and species.
The following more detailed observations were made on material grown
at Kew.
Solitary crystals in the inner part of the primary cortex of Casearia sp.,
(i)

Dovyalis caffra (Hook. f. et Harv.) Warb., Flacourtia prunifolia H.B. et K.,

Homalium africanum Benth., Hydnocarpus wightiana Blume, Oncoba spinosa
Forsk.
(ii)Solitary crystals in some of the stone cells of the pericycle in Casearia
sp., Dovyalis caffra, Flacourtia prunifolia, Homalium africanum, Hydnocarpus
wightiana, Oncoba spinosa, Poliothyrsis sinensis Oliv.
(iii) Solitary crystals
abundant in the soft tissues of the phloem of Homalium
africanum and Oncoba spinosa,
(iv) Large solitary crystals in the pith of Azara dentata Ruiz, et Pav.,
Azara gilliesii Hook, et Arn., Dovyalis caffra, Flacourtia prunifolia, Hydno-
carpus wightiana, Oncoba spinosa, Poliothyrsis sinensis, Rawsonia lucida Harv.
et Sond.
(v) Cluster crystalsabundant in the cortex of Berberidopsis corallina Hook,
f. and Kiggelaria africana Linn.
Cluster crystals infrequent in the cortex of Dipentodon sinicus Dunn.,
(vi)
Dovyalis caffra, Homalium africanum Benth., Poliothyrsis sinensis, Rawsonia
lucida.

(vii) Cluster crystals abundant in the phloem in Azara dentata, Azara

gilliesii, Casearia sp., Dovyalis caffra, Homalium africanum, Hydnocarpus
wightiana (not very numerous), Kiggelaria africana.
(viii) Cluster crystals in the pith of Azara gilliesii, Hydnocarpus wightiana,

Kiggelaria africana, Poliothyrsis sinensis (only in very young stems), Rawsonia

lucida.

(ix) Crystals of all kinds relatively scarce in Gerrardina spp.

The above information concerning crystals could usefully be elaborated by

further investigation.

Secretory cells, whose contents are stained deeply with haematoxylin,

abundant in the parenchymatous tissues of Berberidopsis corallina and
Rawsonia lucida.

WOOD (Fig. 31 D-L)

Vessels usually moderately small (50-100 mean tangential diameter),
fj.

sometimes slightly larger, e.g. in some species of Ahernia, Mesaulospermum,

Pangium, Peridiscus, Tetrathylacium, and Trichadenia, or smaller, e.g. in some
species of Carpotroche, Dovyalis, Rawsonia, Trimeria, and Xylosma. Solitary
122 FLA CO UR TIA CEAE
and in radial multiples of 2 or 3 cells; multiples of 4 or more cells sometimes
moderately common locally or in particular species, e.g. of Flacourtia, Hydno-
carpus, Kiggelaria, Trimeria, and Xylosma-, individual vessels or groups often
separated radially by only a single flattened fibre sometimes in small clusters.
;

Usually 10-50 per sq. mm., but sometimes fewer (5-9) in species of Pangium,
Ryparosa, and Trichadenia and more numerous (50-110) in species of Azara,
Gossypiospermum, Ludia, Prockia, Rawsonia, Ryania, Samyda, Trimeria,
and Xylosma. Spiral thickening reported in Azara, Hisingera, Olmediella,
Poliothyrsis, and Xymalos (1851). Perforation plates simple and oblique in
most genera, but with some scalariform plates in addition (usually with few
bars and sometimes rare), in Aberia (2158), Ahernia, Azara, Caloncoba,
Casearia p.p., Dendrostylis (2158), Dovyalis, Gynocardia, Hasseltia, Homalium,
Lunania, Olmediella^ Oncoba, and Ryparosa*, with occasional scalariform
plates in the neighbourhood of the primary wood in Banara (2430), Carpo-
troche, Casearia (2430), Grandidiera (2158), Idesia, Lindackeria, Mayna, and
Trimeria (2158); all or predominantly scalariform in Asteriostigma, Azara p.p.
(2158), Bergsmia (1154), Carpotroche p.p. (2430), Dasykpis, Erythrospermum,
Hasseltiopsis (1886), Hydnocarpus, Peridiscus, Rawsonia, Scottellia, Tarak-
togenos, and Tetrathylacium. Janssonius (1154) refers to scalariform plates in
Pangium edule Reinw. Sometimes with a few reticulate or foraminate plates,
e.g. in Hydnocarpus and Rawsonia, and occasionally with scalariform or
reticulate plates to ray cells, even in woods in which the normal perforations
between members are simple, e.g. in Trichadenia. Intervascular pitting
scalariform in Hasseltia and opposite or transitional in Erythrospermum,
Hydnocarpus, Mayna, Peridiscus, and Ryparosa p.p., alternate in the others;
the alternate pitting very small to minute, and the pits to ray cells similar
[though sometimes with several pits subtended by one ray pit (unilaterally
compound), e.g. in Ryania] in Banara, Casearia, Flacourtia, Gossypiospermum,
Hecatospermum, Homalium, Laetia (1886), Ludia, Lunania, Qphiobotrys, Osme-
lia, Prockia, Ryania, Samyda, Scolopia, Trichadenia, Trimeria, Xylosma, and
Zuelania\ pits large in the other genera with alternate pitting and the pitting
to rays in these and the genera with scalariform and opposite intervascular
pitting commonly elongated and often scalariform, e.g. in Ancistrothyrsus
(1886), Arechavaletaia (1886), Asteriastigma, Azara, Dasylepis, Carpotroche,
Dovyalis, Erythrospermum, Hasseltia, Hasseltiopsis (1886), Kiggelaria, Lin-
dackeria, Olmediella, Pangium, Peridiscus, Rawsonia, Ryparosa, Scottellia,
Tetrathylacium, Trichadenia, and Taraktogenos. Tyloses occasionally present
and, according to Record and Hess (1886), fairly abundant in Ancistrothyrsus
(sometimes sclerotic), Lindackeria, and Peridiscus. Gonggrijp (794) refers to
silica in the tyloses of Hydnocarpus and Taraktogenos. Mean member length
most commonly 0-7-1 -3 mm. and up to 1-7 mm. in Taraktogenos. Paren-
chyma absent or very sparse. When present, paratracheal and usually
limited to isolated cells touching the vessels (Fig. 31 j), but Record and Hess
(1886) describe the parenchyma as irregularly paratracheal, short aliform and
diffuse* in Ancistrothyrsus and as 'reticulate' in Peridiscus', Williams (2430)
refers to fine lines in Lindackeria and Lunania, but these were not present in
the material available to the author. Moderately abundant about the vessels
and scattered among the fibres in Pangium edule Reinw. In Aphloia what
appear in transverse sections to be diffuse parenchyma cells are thin-walled
 FLACOURTIACEAE 123

septate fibres (see below). Record and Hess (1886) note chambered crystals
in Olmediella. Rays most commonly up to 3-5 cells wide, but up to 6-8 cells in
some species of Banara, Rawsonia, and Scottellia and 10 or more cells wide
in Aphloia, Patrisia (2430), and Ryania\ seldom more than 2 cells wide in
some species of Casearia, Idesia, and Xylosma and almost exclusively uni-
seriate in Asteriastigma macrocarpa Bedd. Typically high and often over

3 mm. high, but seldom more than i mm. high in Dovyalis, Idesia, and some
species of Homalium. Uniseriates numerous and composed of high upright
cells. Rays 8-24, mostly 9-15 permm. Markedly heterogeneous (Kribs's
Type I), typically with 10 or more marginal rows of square or upright cells,
but sometimes with only 4-10 rows, e.g. in Homalium tomentosum Benth.,
Idesia polycarpa Maxim., Pangium edule Reinw., and Scottellia coriacea A.
Chev. The multiseriate rays sometimes composed entirely of square and
upright cells, e.g. in Casearia nitida (L.) Jacq. and Lindackeria dentata Gilg,
as also are the wholly uniseriate rays of Asteriastigma macrocarpa. Occasionally
with sheath cells (1154). Vertically fused rays common (Fig. 31 i). Nearly
always containing crystals, which are often abundant sometimes apparently ;

chambered 1 or chambered when occurring in the upright cells, e.g. in Banara,

Flacourtia, Homalium, Prockia, Scolopia, Trimeria, and Xylosma. 'Perforated
ray cells' reported in several genera (358). Fibres septate, except in Ancisto-
thyrsus and Peridiscus (1886) and septa rare or absent from Ludia. Pits
almost entirely limited to the radial walls, simple or with small, and often
very indistinct, borders, but with large, distinct borders in Aphloia and in
Ancistrothyrsus and Peridiscus (1886). In Aphloia the ground tissue consists
of thick-walled fibres with conspicuous bordered pits, among which are
scattered thin-walled septate fibres, which are distributed as single cells or
short uniseriate lines resembling diffuse parenchyma; the thick- walled fibres
also are septate in A. myrtiflora Galpin, but not in A. mauritiana Baker; the

parenchyma-like septate fibres have bordered pits and in both these and the
thick- walled fibres the pits are equally numerous in both radial and tangential
walls. Walls moderately thin to very thick in the other genera and often with
a gelatinous layer. Mean length O'Q-z-y, most commonly about 1-6 mm., and
2.0 mm. or more in Homalium, Pangium, Taraktogenos, and Xymalos.

DIONCOPHYLLUM PELTATUM Hutch, et Dalz.

An anomalous liane with successive bundles of xylem and phloem, without

regular arrangement.
Vessels very large, mostly solitary, but with some pairs and multiples;
about 2 per mm. Perforations simple. Intervascular pitting alternate and
to parenchyma similar. Paren-
large, often with coalescent apertures; pits
chyma (a) conjunctive, between the bundles; containing scattered, large,
solitary, and often "spindle-shaped stone cells; (b) paratracheal, as narrow
sheaths round the vessels (vasicentric) and (c) apotracheal, as a few cells
;

1
Record and Hess (i 886) draw attention to the fact
that these cells are not, strictly speaking,
'chambered' in the sense that each crystal is enclosed in a separate cell and not merely
separated from the next by a septum. The genuinely chambered groups of crystals that
occur in Banara are described by these authors as occurring in the fibres; in the material
examined of Banara guianensis Aubl. such crystals were observed only in the very tall upright
cells of the rays.
1
24 FLA CO UR TIA CEAE
scattered among the fibres. Rays uniseriate; short to moderately high; con-
tinuing through the individual bundles, but varying in direction from bundle
to bundle. Fibres with large, conspicuously bordered pits that are equally
numerous on both radial and tangential walls, the borders smaller than those
of the intervascular pitting. Walls moderately thick.

BARK
According to Sprague and Boodle (2173) the bark of Casearia praecox
Griseb. exhibits the following characters. Cork cells thickened on the inner
side, their cavities becoming nearly or quite obliterated. Secondary phloem
containing secretory canals and stone cells, the former provided with a thin-
walled epithelium, the contents being soluble in alcohol. The stone cells in
the older part of the secondary phloem form continuous or nearly continuous
zones alternating with soft tissue, but in the younger phloem are arranged in
rounded groups or tangential bands. Solitary crystals of calcium oxalate
present in the stone cells.

ANOMALOUS GENERA
The following genera are treated separately since their taxonomic positions
are uncertain, whilst the facts recorded about their anatomical structure
do not appear to conform, at least in certain respects, with those of the
Flacourtiaceae.

(i) ASTEROPEIA
LEAF
Dorsiventral. Hairs not observed. Epidermis strongly cutinized. Stomata
confined to the lower surface, each surrounded by about 12 cells similar to
those of the remainder of the epidermis; guard cells very strongly cutinized.
Aqueous hypoderm, about 2 or 3 cells wide, situated below the upper
epidermis in A. sphaerocarpa Baker, but 4 or 5 layers recorded by Beauvisage
(163) in other species. Mesophyll consisting of 3 or 4 layers of palisade cells
(with .corrugated anticlinal walls in herbarium material) and an extensive
spongy region, the latter containing rounded sclerenchymatous idioblasts with
short arms. Similar idioblasts also present in the collenchyma above the
median vein. Vascular bundles of the veins embedded in the mesophyll and
supported by sclerenchymatous sheaths. Petiole, in transverse sections
through the distal end, exhibiting an almost closed but dorsally flattened
vascular strand in which the phloem is as broad as or locally even broader
than the xylem, the whole vascular system being surrounded by an almost
continuous ring of sclerenchyma, and enclosing a few sclerosed elements in
the medullary region.

Axis
YOUNG STEM
Cuticle very thick. Cork arising in the sub-epidermis. Cortex containing
rounded or angular sclerenchymatous idioblasts sometimes with short arms,
either isolated or arranged in groups, Pericycle demarcated externally by
a composite, continuous ring of sclerenchyma and including a few stone cells
in the inner part. Phloem and xylem in the form of continuous cylinders,
 FLA CO UR TIA CEAE 125
traversed by uniseriate rays. Phloem apparently containing secretory cavities,
but their true nature could not be determined in the herbarium material
examined. Vessels mostly solitary, rather unevenly distributed, usually
circular or slightly oval in transverse section, seldom exceeding 50 /* in radial
diameter; perforations simple. Pith somewhat heterogeneous, including
solitary and grouped sclerosed cells.
With regard to the secondary
xylem of Asteropeia, the only material avail-
able was a specimen of A. rhopaloides Bak., which had not been
single
correlated with herbarium material. It differed markedly from the rest of the
family, e.g. in having minute, alternate intervascular pitting and pits to ray
cells, solitary vessels and exclusively uniseriate rays.

SPECIES OF ASTEROPEIA DESCRIBED

The above description is based on an examination of Asteropeia sphaero-
carpa Baker in the Kew Herbarium. It differs only in minor respects from
other species such as A. multiflora Dup-Th. and A. ambylocarpa Tul. pre-
viously describee! by Beauvisage (163).

(ii)
PSILOXYLON
Secretory cavities recorded in the mesophyll as well as in the cortex of
young stem. Cork arising in the pericycle, and consisting of layers composed
respectively of flattened, thin-walled cells and thickened cells with relatively
wide lumina. Pericycle demarcated by sclerenchyma consisting wholly of
stone cells. Secondary phloem devoid of fibres, but including layers of cells
filled with cluster crystals. Intraxylary phloem recorded.
Solereder considered that this genus should be placed in the Myrtaceae.

(iii) FROPIERA
The anatomy of this genus needs further investigation. The leaves probably
contain secretory cavities.

(iv) HASSELTIA AND PROCKIA

Phloem containing secretory cells with yellow contents resembling gum-
resin,and which are more abundant in Hasseltia than Prockia. Sclerenchyma-
tous elements said to be absent from the pericyle and phloem.

(v) PLAGIOPTERON
Secretory elements resembling latex canals are present in the phloem in
the veins of the leaf as well as in the phloem and pith of the axis. These
elements contain a rubber-like substance which can be drawn out in the form
of fine threads when the leaf is broken.

TAXONOMIC NOTES
Most of the genera which were included by Bentham and Hooker in the
Bixaceae and Samydaceae are characterized by stomata of the rubiaceous or
cruciferous types and the presence of crystal idioblasts in the epidermis of the
leaf. These characters together suggest that these genera should be combined
into a single family, which in practice conforms approximately with the
Flacourtiaceae as understood by Gilg (768). This is also in agreement with
126 FLACOURTIACEAE
the views of modern taxonomists who base their conclusions on external
morphological characters. The group thus constituted is also very homo-
geneous as regards its wood anatomy, though the genera differ considerably
in the degree of specialization of the vessels. Record and Hess (1886), how-
ever, note that 2 genera, Ancistrothyrsus and Peridiscus, seem to be rather out
of place. Only one line of subdivision suggests itself, the distinction between
the genera with minute intervascular and vessel-ray pitting and the others.
In the present account, however, the Flacourtiaceae-Paropsieae and Abatieae
in the sense used by Gilg (768) have been described under Passifloraceae.
Bixa has been regarded as the sole representative of the Bixaceae, whilst
Amoreuxia, Cochhspermum, and Sphaerosepalum have been described under
Cochlospermaceae. It will be seen that these last 4 genera differ from the
Flacourtiaceae, but are, with the possble exception of Sphaerosepalum, very
differentfrom one another. It seems to be largely a matter of opinion whether
the Bixaceae and Cochlospermaceae should be recognized as distinct or
united into one family. On anatomical grounds they are not very clearly
demarcated, except in so far as the woods of Bixa and Cochhspermum are
clearly distinguishable by their parenchyma. (See also under 'Bixaceae* and
'Cochlospermaceae'.) The genus Erythrospermum which tends to conform
with the Tiliaceae in the possession of rays which broaden out in the phloem,
on external morphological characters and in its wood anatomy, agrees rather
with the Flacourtiaceae. Dipentodon has been included in the Flacourtiaceae
rather than in the Celastraceae on the advice of Dr. T. A. Sprague, especially
as the anatomical structure conforms with this suggestion. The family also
includes several genera whose true taxonomic position has not yet been
finally established. These have been described separately, partly to emphasize
the points in which they differ from most members of the family, and partly
to facilitate their transfer to other families if this should become necessary in
the future. The genera concerned are Asteropeia, Fropiera, Hasseltia,
Plagiopteron, Prockia, and Psiloxylon. The Flacourtiaceae are distinguished
from the Bixaceae and Tiliaceae by the lack of mucilage receptacles in the
cortex and pith of the young stem, and from the Violaceae by the possession
of crystal-idioblasts in the epidermis.
The following views, based on a study of wood structure, have also been
expressed by Vestal (2239). He regards the Cistaceae as derived from the
Bixaceae and these in turn 'stem' from the Flacourtiaceae, with the Cochlo-
spermaceae as a parallel development also arising in the Flacourtiaceae. Of
these families he says *the Flacourtiaceae are the only ones that show in their
anatomy any relationship to the Dilleniaceae'.
Vestal opposes Hutchinson's view that this group is basic for the Theales
and Guttiferales, on the grounds that it is 'rather difficult to conceive of the
obviously primitive vessel members in the Theaceae, Actinidiaceae and
Saurauiaceae as coming from the highly evolved vessel members of the Bixales
series'. It is, however, open to question whether the vessels of the Flacourtia-
ceae can be correctly described as 'highly evolved* in view of the occurrence
of scalariform intervascular pitting, entirely scalariform perforation plates
and mean member lengths of over 2 mm. in several genera. (Cf. Actiniaceae
0*7 mm., Saurauiaceae 1-5 mm., and Theaceae mostly i'O~i*6 and 2-4 in
Adinandra.)
 FLACOURTIACEAE 127

ECONOMIC USES
Kei Apples are the Dovyalis caffra (Hook. f. et Harv.) Warb. The
fruits of
and Oncoba spinosa Forsk. are edible. Chaul-
fruits of certain Flacourtia spp.

moogra Oil is obtained from the seeds of Tamktogenos kurzii King, whilst a
similar oil is obtained from Hydnocarpus spp. West Indian Boxwood is
derived from Casearia praecox Griseb.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aphaerema, Asteropeia,* Azara,* Banara, Bembicia, Berberidopsis,*
Byrsanthus, Calantica, Camptostylus, Carrierea,* Casearia,* Centroplacus,
Dasylepis, Dipentodon,* Dissomeria, Dovyalis,* Erythrospermum, Euceraea,
Flacourtia,* Fropiera, Gerrardina,* Grandidiera, Gynocardia, Hasseltia,
Homalium,* Hopelstigma, Hydnocarpus,* Idesia, Itoa, Kiggelaria,* Laetia,
Ludia, Lunania, Mayna, Neumannia, Oncoba,* Ophiobotrys, Osmelia,
Pangium, Patrisia, Physena, Pineda, Plagiopteron, Poggea, Poliothyrsis,*
Prockia, Psiloxylon, Pyramidocarpus, Rawsonia,* Ryania (or Patrisia),
Samyda, Scolopia, Taraktogenos, Tetrathylacium, Tisonia, Trimeria,
Xylosma, Zuelania.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
('Abatia'), (Aberia, or Dovyalis), Ahernia, (Ancistrothyrsus), Aphloia (or
Neumannia), (Arechavaletaia), Asteriastigma, Asteropeia, Azara, Banara,
(Bennettia), (Bergsmia, or Ryparosa), Caloncoba, Carpotroche, Casearia,
Dasylepis, (Dendrostylis, or May no), Dioncophyllum, Dovyalis, Eleu-
therandra, Erythrospermum, (Grandidiera), Gossypiospermum, Gynocardia,
Hasseltia, (Hasseltiopsis), Hecatostemon, (Hisingera, or Xylosma), Homa-
lium, Hydnocarpus, Idesia, Kiggelaria, Laetia, Lindackeria, Lunania, Mayna,
Olmediella, Oncoba, Ophiobotrys, Osmelia, Pangium, (Patrisia), Peridiscus,
(Poliothyrsis), Prockia, Rawsonia, Ryania (or Patrisia), Ryparosa, Samyda,
Scolopia, Scottellia, Taraktogenos, Tetrathylacium, Trichadenia, Trimeria,
Xylosma, Zuelania.
LITERATURE
(i) On General Anatomy
Beauvisage 163, Braendlein 253, Gilg 768, 773, Harvey-Gibson and Horseman 916,
Sprague and Boodle 2173.
(ii) On Wood Structure
Bannan 133, Benoist 170, den Berger 177, 179, 182, Besson 186, Burgerstein 310, Chalk
and Chattaway 358, 362, Cooper 461, Coster 481, Desch 574, Giordano 786, Gonggrijp
794, Howard 1088, Janssonius 1154, Kanehira 1206, 1209, Kribs 1283, Lecomte 1334,
Pearson and Brown 1679, Record 1818, 1843, 1851, 1884, Record and Hess 1886, Stone
2206, Tang 2231, Tupper 2298, Vestal 2239, Williams 2430, Yamabayashi 2478.
 (128)

35. PITTOSPORACEAE
(Fic. 31 on p. 120; FIG. 32 on p. 130)

SUMMARY
(i) GENERAL
A
family of trees, shrubs, and climbers, occurring in the warmer regions of
the Old World, and especially well developed in Australia. A constant and
characteristic feature of the family is the presence of secretory canals in the
pericycle of the stem, roots, and leaves. Similar canals occur also in the
secondary phloem of older stems, but according to Solereder they are never
present in the primary cortex or pith, although they have been found in the
secondary cortex. The stomata are rubiaceous with 2 or more pairs of sub-
sidiary cells parallel to the pore. There is usually little or no sclerenchyma
in the family, mechanical support being provided by collenchymatous
elements. The hairs may be: (i) Simple, uniseriate trichomes with 2 or 3
basal cells and a long terminal cell, (ii) Two-armed hairs with a short stalk,
(iii) Club-shaped glandular
hairs. Transitions between these types also occur.
Calcium oxalate is generally present in the form of clustered and solitary
crystals, whilst styloids occur in the phloem.

(ii) WOOD
Vessels small; in numerous small multiples and clusters, often with a
diagonal pattern; perforations simple; intervascular pitting alternate and
small, pits to ray cells similar; members of medium length. Parenchyma
sparse, vasicentric. Rays up to 3-7 cells wide, with few uniseriates and almost
homogeneous. Fibres septate, of medium length to moderately short,

LEAF
Centric in a few species, but mostly dorsi ventral. Hairs, (i) Uniseriate
with 2 or 3 short basal cells and a long terminal cell in Billardiera, Marianthus,
Pronaya, Sollya. (ii) Two-armed trichomes with a short stalk, the terminal
cell sometimes being deciduous, in Bursaria, Citriobatus, Pittosporum.
Club-shaped glandular hairs in Hymenosporum and certain species of
(iii)
Bursaria and Pittosporum. Epidermis with strongly thickened outer walls.
Upper epidermis papillose in Marianthus tennis Benth. Cells of the epidermis
horizontally divided in certain species of Pittosporum; the inner layer elongated
in a palisade-like manner in P. phillyraeoides DC. Stomata confined to the
lower surface rubiaceous, provided with 2 or several subsidiary cells parallel
;

to the pore, the subsidiary cells generally having thinner walls than the
remaining epidermal cells. Petiole, in transverse sections, e.g. of Pittosporum
tenuifolium Gaertn. (Fig. 32 D), generally exhibiting an arc of 3, 5, 7 or more
vascular strands, but a solitary vascular bundle recorded in certain species
with small leaves; bundles not usually accompanied by sclerenchyma.
Crystals of calcium oxalate clustered or solitary and sometimes present in
specially large sacs. Secretory canals (see also under 'Axis') present.

Axis
YOUNG STEM (Fig. 32 F)
Cork arising superficially in Pittosporum and Solly a; component cells with
 PITTOSPORACEAE 129
thin to thick walls and filled with gum-like contents in the same 2 genera.
Primary cortex generally collenchymatous especially in the outer portions.
Secondary phloem also tending to become collenchymatous; sieve tubes with
comparatively small lumina and scalariform sieve plates. Xylem in the form
of a cylinder traversed by rather broad, lignified rays in Pittosporum, and by
narrower rays in Sollya heterophylla Lindl. Vessels with simple perforations.
Pith lignified. Secretory canals (see also 'Leaf '), with colourless, yellow or
reddish contents, present especially in the inner part of the cortex and peri-
cycle. Secretory canals stated to occur, although less frequently, in the
phloem or pith of Billardiera, Bursaria, Cheiranthera, Citriobatus, Hymeno-
$porum> Marianthus, Pittosporum, Pronaya, Sollya. Crystals, similar to those
in the leaf, sometimes situated in large sacs, chiefly in the phloem. Styloids
also recorded in the phloem.

WOOD (Fig. 31 A-C)

Vessels very to moderately small (mean tangential diameter 40-100 ^);
solitary, in short radial multiples and in numerous clusters sometimes with a
;

distinct oblique or tangential pattern (Fig. 31 c); 25-50 per sq. mm. with
spiral thickening in at least the tips of the members in some species of
Bursaria, Hymenosporum, and Pittosporum\ Solereder lists several other
genera as having spiral thickening or spiral striations. Perforations simple,
oblique. Intervascular pitting alternate, small to very small, pits to ray cells
similar. Often containing a yellow substance in Pittosporum. Mean member
length 0-45-0-9 mm. Parenchyma scanty, paratracheal, occurring as a few
about the vessels. Very occasionally containing a few chambered crystals.
cells
Strands usually of 4 cells. Rays up to 3-7 cells wide; less than i mm. high;
uniseriates rare, composed mainly of procumbent cells; 5-12 rays per mm.;
almost homogeneous (Kribs's Types Het. II B to Horn. I), usually with a
single marginal row of square cells (Fig. 313). Crystals present in the marginal
cells, often in rounded, slightly larger cells in Citriobatus and Pittosporum.
Fibres septate, but the septa sometimes rather few. Pits small and almost
entirely limited to the radial walls, simple or with very small borders. Walls
moderately thin. Mean length o-8-i-i mm.
ROOT
Secretory canals similar to those in the stem, present in the pericycle,
situated externally to the xylem, as well as the phloem groups. Secretory
canals stated to inhibit the development of lateral roots on the outer side of
the xylem groups. Lateral roots in consequence arising between the secretory
passage opposite a xylem group and the neighbouring secretory passage
opposite a phloem group. Lateral roots stated to originate in a similar manner
in the Araliaceae and Umbelliferae.

TAXONOMIC NOTES
Solereder (2158) and Guenot (832) have suggested on anatomical grounds
that this family has affinities with the Umbelliferae and Araliaceae, an idea
which was propounded by Van Tieghem and followed up by the authors just
mentioned, but Pritzel (1758) thinks that the floral differences are too great to
support this suggested relationship Pritzel mentions that possible relationships
.
 FRANKENIACEAE, A; POL YGALACEAE, B and G-I;
FIG. 32.
TREMANDRACEAE, C and E; PtTTOSPORACEAE, D and F
A, Frankenia laevu Linn. Stem x 17. B, Barnhartia floribunda Gleason. Petiole X9. C, Platytheca
Steetz. Stem X 17. D Pitto$porumtenuifoliumGaertn. Petiole X 33, E, Tetratheca thymifolia
t

m, Stem X 17. F, Pittosporum temafolium Gaertn. Young stem X33. G, Moutabea guianenris Aubl.
filioides t

Petiole XQ, H, Carpolobia alba G. Don. Stem x 17, 1, Polygala oppositifolia Linn. Stem X 17.

Fig. E. Abundant secretory cells with tannimferous contents ih all unlignified tissues.
c. Thick outer wall of epidermis, e. Epidermis, s.c Secretory canals. u,f. Unlignified fibres.
 PITTOSPORACEAE 131
between the Pittosporaceae on the one hand and such various families as
Celastraceae, Escalloniaceae, Polygalaceae, Rhamnaceae, Tremandraceae,
and Vochysiaceae on the other have been proposed from time to time. The
septate fibres and paratracheal parenchyma of the Pittosporaceae suggest the
Araliaceae as the nearest of the above families and make any relationship with
the Escalloniaceae improbable. There are according to Pritzel difficulties
about all of the possible relationships which have just been mentioned, so he
prefers to regard the Pittosporaceae as a somewhat isolated family.

ECONOMIC USES
The woods of certain species of Pittosporum are of economic importance.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Billardiera, Bursaria, Cheiranthera, Citriobatus, Hymenosporum, Marian-
thus, Pittosporum,* Pronaya, Sollya.*
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
(Billardiera), Bursaria, (Cheiranthera), Citriobatus, Hymenosporum,
(Marianthus), Pittosporum, (Pronaya), (Sollya).
LITERATURE
(i) On General Anatomy
Guenot 832, Pritzel 1758.

(ii) OK Wood Structure

Baker 104, Brown, F. B. H. 280, Kanehira 1206, 1209, Record 1843, 1851, Yamabayashi
2478-

36. TREMANDRACEAE
(FiG. 32 on p. 130)

SUMMARY
This family of slender, heath-like shrublets, endemic in Australia, is fairly
uniform in respect of its anatomical characters. The wood exhibits the follow-
ing features. Vessels small, sometimes with spiral thickening, perforations
simple. Parenchyma scanty paratracheal or absent. Rays 1-2 cells wide,
composed of upright cells.Fibres with simple and bordered pits.

LEAF
Flat or linear, with margins frequently curved inwards towards the abaxial
side. The shape of the leaf as seen in transverse section is stated to be of
value for the identification of species. Hairs, (i) Simple, unicellular, stiff
and thick-walled in Tetratheca. (ii) Stellate in Tremandra. (iii) Glandular,
especially in Tetratheca and to some extent in Platytheca. Mesophyll includ-
ing i layer of palisade cells. Inner walls of the cells of the epidermis
frequently mucilaginous, except in Tremandra. Stomata confined to the
lower surface; ranunculaceous. Petiole, in transverse sections, exhibiting
a single vascular bundle, or i large and 2 small bundles. Calcium oxalate
present in the form of solitary, and less frequently of clustered crystals*
132 TREMANDRACEAE
Axis
YOUNG STEM (Fig. 32 c and E)

Primary cortex of species of Tetratheca with reduced leaves containing

palisade parenchyma. Bundles of sclerenchymatous fibres present in the
3 angles of species of Tetratheca with triangular branches. composite, A
continuous ring of sclerenchyma recorded in the pericycle of Tetratheca
ciliata Lindl. and T. thymifolia Sm. strands of sclerenchyma observed in
;

Platytheca galioides Steetz. Xylem in the form of a continuous cylinder

traversed by narrow medullary rays. Stem almost wholly composed of
secondary xylem in Platytheca galioides Steetz. (Fig, 32 c). Vessels in the
same species small and angular; small but less angular in Tetratheca thymifolia.
Perforations mostly simple, but occasional scalariform plates also recorded (see
*

Wood*}. Pith very narrow, consisting of cells with moderately thick, lignified
walls in Platytheca galoides\ much broader, somewhat heterogeneous and
composed of a mixture of large empty thin-walled cells mixed with smaller
ones filled with secreted material in Tetratheca thymifolia. Secretory cells
with unidentified contents abundant in the unlignified tissues of Tetratheca
thymifolia.

WOOD 1

Vessels very to extremely small; solitary and with a tendency to chains and
multiples; with spiral thickening in Platytheca galioides Steetz. and Tetratheca
spp. Perforations simple, but with a few scalariform plates near the primary
wood in Tremandra stelligera R.Br. Intervascular pitting circular to oblong,
usually in a single row owing to the small lumina of the vessels. Pits to ray
and wood parenchyma often simple and oblong. Parenchyma scanty para-
tracheal or absent. Sometimes containing crystals. Rays 1-2 cells wide and
composed almost entirely of upright cells; seldom more than 10 cells high.
Fibres with small bordered pits in Tetratheca glandulosa Labill. and with
both simple and bordered pits in the other genera. With fine spiral thickening
in Tetratheca efoliata F, v. Muell.

TAXONOMIC NOTES
The affinities of this family have never been well established. Its anatomical
features do not afford much assistance in arriving at a conclusion. In these
circumstances the family must be regarded as somewhat isolated.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Platytheca,* Tetratheca,* Tremandra.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
(Platytheca), (Tetratheca), (Tremandra).

LITERATURE
On Wood Structure
Heimsch 938, Record 1800, 1851*
1
Based almost entirely on the descriptions given by Solereder and by Heimsch (938).
 (133)

37. POLYGALACEAE
(FiG, 32 on p. 130; FIG. 33 on p. 136; FIG. 91 on p. 402)

SUMMARY
(i) GENERAL
A family of herbs, shrubs, or rarely small trees occurring in both temperate
and tropical regions. Hairs generally simple and unicellular, except in
Bredemeyera and Xanthophyllum where they are sometimes uniseriate. The
stomata are generally ranunculaceous, but sometimes tend to be rubiaceous.
The midrib and petiole generally possess one vascular strand. Calcium
oxalate occurs as solitary or clustered crystals. Lysigenous secretory
cavities and oil ducts occur in Polygala. The xylem in transverse sections
of young stems generally appears as a closed ring, even in herbaceous species,
although according to Holm (1001, 1018), in certain North American species
of Polygala there are about five collateral bundles in very young stems. For
Anomalous Structure see under 'Wood*.

(ii)
WOOD
Vessels small to large, usually exclusively solitary, perforations simple,
intervascular pitting alternate, pits to ray cells usually similar, rarely elongated
and simple, members of medium length. Parenchyma usually predominantly
paratracheal in the Polygaleae, scanty or, less commonly, aliform or confluent;
predominantly apotracheal, diffuse or banded, in the Moutabeae and Xantho-
phylleae. Rays typically exclusively uniseriate or up to 2 or 3 cells wide, but
occasionally wider; heterogeneous. Fibres with bordered pits, of medium
length to moderately short. Included phloem of the 'concentric' type
present in the climbers.

LEAF
Hairs almost exclusively unicellular, but sometimes becoming uniseriate
by septation in Bredemeyera and Xanthophyllum. Unicellular hairs with thick-
walled apices recorded by Holm (1001, 1018) in Polygala senega Linn.
Structure of the mesophyll very varied, (i) Consisting of one uniform tissue
in a few species of Mundtia and Carpolobia. (ii) Leaves generally dorsiventral
both in species with a relatively broad lamina, as well as in some of those
which are lanceolate or linear with incurved margins, (iii) Centric in a few
species of Comesperma (Bredemeyera), Muraltia, and Polygala possessing
lanceolate or linear leaves with the margins incurved. Palisade parenchyma
with folded lateral walls, and spongy parenchyma partly sclerosed in Monnina
speciosa Tr. et Planch. Bundles of diamond-shaped bodies stated by Sabnis
(1920) to be present in the palisade cells of PolygaL erioptera DC. Cells of
the epidermis generally with straight anticlinal walls except in a few species
mentioned by Solereder; those on the lower surface provided with knob-like
papillae in species of Securidaca, or sometimes with much thickened papillae
in Muraltia and Polygala. Outer wall of epidermal cells sometimes strongly
thickened. Cuticle smooth or with granular thickening; rarely striated.
Hypoderm recorded only in Moutabea guianensis Aubl. Long sclerosed cells
occur in the palisade tissue of the same species. Stomata present on both
134 POLYGALACEAE
surfaces or confined to the lower side, this difference being only of specific
diagnostic value ; generally ranunculaceous, but rubiaceous in certain species
of Securidaca and Xanthophyllum. Stomata absent from the scale leaves of
the saprophytic genus Epirrhizanthes (Salomonia). Vascular bundles of the
veins of species with small leaves not generally accompanied by sclerenchyma.
Veins vertically transcurrent in Phlebotaenia sp. Midrib usually with a single
band or horseshoe-shaped vascular bundle, except in Moutabea with two
strands, the upper one inversely orientated. Petiole, in transverse sections
through the distal end, nearly always exhibiting a single, usually crescent-
shaped bundle, e.g. in Barnhartia floribunda Gleason (Fig. 32 B), but with
an almost completely closed vascular ring in Bredemeyera, Polygala, and
Securidaca, or a completely closed ring in Carpolobia, Moutabea (Fig. 32 G),
and certain species of Xanthophyllum. Mostly solitary but sometimes clustered
crystals usually present in the leaf and axis, but none recorded in Bredemeyera
and Xanthophyllum. Numerous lysigenous cavities recorded by Sabnis
(1977) in the mesophyll towards the lower side of the leaf in Polygala erioptera
DC. Oil drops and oil ducts stated by Holm (1018) to occur in certain
North American species of Polygala, but not in other species of this genus
from the same part of the world.

Axis
YOUNG STEM (Fig. 32 H-I)
Epidermis strongly thickened in xerophilous species, but thin-walled in
species having leaves adpressed to the stem; cells sometimes divided hori-
zontally in Securidaca, and, according to Holm (1018), containing oil drops in
certain species of Polygala. Primary cortex frequently collenchymatous, but
containing stone cells in species of Atroxima, Barnhartia, Carpolobia,
Moutabea, Polygala strands of fibres also present in the cortex of herbaceous
;

shoots of Comesperma and Securidaca, as well as in the ribs on the stems in

certain species of Comesperma and Polygala. Palisade chlorenchyma recorded
by Sabnis (1977) in certain species of Polygala. Endodermis indistinctly
visible'in Polygala senega Linn, but not seen by Holm (1018) in other North
American species of this genus. Pericycle generally containing fibres either
in the form of strands or as a continuous ring; their distribution varying
considerably in different species of the single genus Polygala. The vascular
cylinder is also surrounded by fibres in the saprophytic genus Epirrhizanthes
(Salomonia). Xylem, in transverse sections, generally appearing as a closed
ring in herbaceous species. Five primary collateral bundles, which sub-
sequently become connected by cambium but without vessels arising in the
interfascicular region, present in certain North American species of Polygala.
Vessels usually small, sparse, and often scattered in the older wood of young
stems, perforations simple. More numerous, spirally thickened vessels present
in the protoxylem of species of Moutabea, Securidaca, Swertia. Solitary
crystals observed in the cortex and/or phloem of species of Atroxima, Barn-
hartia, Moutabea, Securidaca. Clustered crystals observed only in the primary
cortex of Polygala oppositifolia Linn, but possibly occurring elsewhere as well.
Lysigenous oil ducts recorded by Holm (1018) in the cortex of certain North
American species of Polygala.
 POLYGALACEAE 135
WOOD (Fig. 33 and 91 B)
Vessels usually small to medium-sized (60-150^ mean tangential diameter)
in most genera, large (more than 2OO/x) in some species of Securidaca and
Xanthophyllum, and sometimes very to extremely small in small stems and
twigs; typically exclusively solitary or nearly so, but in short multiples in
Bredemeyera p.p. and in 'pore chains' in Monnina (938) and with some
clusters in Monnina (938) and Phlebotaenia; usually about 7-12 per sq. mm.,
but fewer than 5 per sq. mm. in some species of Securidaca and Xantho-
phyllum', spiral thickening observed in Mundtia spinosa DC. and recorded by
Solereder for Polygala eckloniana Presl. and Muraltia ononidifolia Eckl. et
Zeyh. Perforations simple, transverse or slightly oblique; Heimsch (938)
notes a few net-like plates in one species of Polygala. Intervascular pitting
rare in genera with solitary vessels, alternate, moderate-sized to small, and
minute in some species of Xanthophyllum; pits to ray cells similar to the
intervascular pittting in most genera, but in Carpolobia p.p. and Moutabea
(938) small bordered pits are accompanied by relatively large, circular or
oblong pits with narrow borders or simple. With a few sclerotic tyloses in
Moutabea (938). Mean member length 0-4-0*5 mm. Parenchyma pre-
dominantly paratracheal in Badiera, Bredemeyera, Carpolobia, Monnina,
Mundtia, Phlebotaenia, Polygala, and Securidaca p.p.; usually scanty and
limited to a few cells round the vessels, but tending to be aliform in some
species of Bredemeyera and Carpolobia, e.g. in C. goetzei Giirke, aliform to
confluent in Phlebotaenia, and as long uniseriate wings attached to the
vessels on the abaxial sides in Carpolobia parvifolia Stapf f. (Fig. 33 E). Some
diffuse parenchyma may occur in addition in Phlebotaenia and Securinaga and
in the latter predominates in some species. According to Heimsch (938),
however, the parenchyma in Carpolobia and Polygala is diffuse, and, accord-
ing to Record and Hess (1886), that of Monnina is irregularly diffuse and
terminal. Apotracheal parenchyma appears to be characteristic of Moutabea
and Xanthophyllum, diffuse in the former and in broken to continuous uni-
seriate bands in the latter (Fig. 33 F). Chambered crystalliferous strands

present in Carpolobia and Phlebotaenia. Parenchyma strands usually of 4 cells,

but commonly of 8 cells in Xanthophyllum. Rays exclusively uniseriate, or
only occasionally biseriate (Fig. 32 A, c, and G), in some species of Bredemeyera
(twigs of B. lucida A. W. Benn.), Carpolobia, Mundtia, Phlebotaenia, Securi-
daca, and Xanthophyllum up to 2 or 3 cells wide in the other genera and in
;

Securidaca p.p., but with occasional larger rays in the latter and with rays up
to 5 or 6 cells wide in Monnina (1886) and Moutabea (938); usually low, but
sometimes more than i mm. high in Xanthophyllum uniseriates numerous in
9
,

species with multiseriate rays, and usually composed of procumbent and

upright cells; usually 11-19 raY s P er mm heterogeneous (Kribs's Types II A
-

and B and III), sometimes with 4 or more uniseriate rows of erect marginal
cells; the 'procumbent* cells sometimes almost square and occasionally, e.g. in
Securidaca, with square and upright cells only. Fibres with distinctly bordered
pits that are equally numerous in both radial and tangential walls, the borders
of about the same size as those of the intervascular pitting. Walls moderately
thin to thick. Mean length o-8-i-i mm. Vasicentric tracheids present in
Securidaca. Included (interxylary) phloem of the 'concentric' type (c.
I.

drcumvattatum) present in species of Bredemeyera (1886), Moutabea (1886),

 136 POLYGALACEAE
and Securidaca. Successive layers or arcs of xylem and phloem repeating the
structure of the young stem are separated by rather broad tangential bands of
conjunctive parenchyma (Fig. 91 B); the conjunctive tissue is unlignified and
often includes a band of cells similar to pericyclic fibres (Fig. 33 B); xylem
layers seldom interrupted by interfascicular rays.

FIG. 33. POLYGALACEAE

A, Carpolobia parvtfolia Stapf f. B, Securidaca diversifolia (L.) Blake. C, S. virgata Sw. D, S. longi-
pedunculata Fresn. E, Carpolobia parvifolia Stapf f. F, Xanthophyllum affine Korth. G, X. affine
Korth.
/. Cells resembling pericyclic fibres, u.p. Unlignified parenchyma.

ANOMALOUS STRUCTURE, see under 'WOOD'

ROOT
Thin Polygala lutea Linn, stated to be devoid of hairs, but
lateral roots of

provided with a papillose epidermis. The root of Polygala senega Linn, is

described under 'Economic Uses*.
 POLYGALACEAE 137
TAXONOMIC NOTES
Krameria. Heimsch (938) has shown that, in its wood anatomy, Krameria
differs markedly from the Leguminosae, but would not be out of place in the

Polygalaceae. See also under 'Krameriaceae', p. 538.

Diclidanthera. O'Donell (1630), in a study of the pollen grain morphology
and wood anatomy of Diclidanthera, concludes that this genus belongs to the
Polygalaceae. Unpublished work by Sprague and Metcalfe also supports this
view that Diclidanthera is related to the Polygalaceae. See also under
'Diclidantheraceae', p. 138.
Xanthophyllum. This genus differs in its wood parenchyma from all the
other members of the Polygalaceae this would tend to support the treatment
;

of this genus as a monotypic family. Jauch (1157), on the other hand, con-
siders that Xanthophyllum should be retained in the Polygalaceae, this opinion
being based on the evidence of floral anatomy and the structure of pollen
grains.
Heimsch (938) considers the wood anatomy to show that the Polygalaceae
form part of an interrelated group, the other members of which are the
Trigoniaceae, Tremandraceae, Zygophyllaceae, Malpighiaceae, and Vochy-
siaceae, and that, though they are more specialized in their wood structure,
these families are generally related to the Linaceae, Humiriaceae, and
Erythroxylaceae.
Jauch (1157) suggests that the occurrence of lysigenous canals in certain
American species indicates a possible affinity between the Polygalaceae and
Anacardiaceae.

ECONOMIC USES
Senega or Snake Root, imported from South Canada and the U.S.A. for
medicinal use, is derived from Polygala senega Linn. Certain Indian species
of Polygala also possess roots with medicinal properties (see 'Root* above).
A fibre is obtained in East Africa from Securidaca longipedunculata Fres. The

following anatomical features of Polygala senega have been recorded by Holm

(1018), Steiger (2191), and in the British Pharmaceutical Codex. Young roots
normal in structure, but becoming eccentric and sometimes winged when
mature. Epidermis sparingly hairy. Some of the cells of the cortex contain-
ing a yellowish substance. Endodermis thin- walled. Cork in older roots
formed from the cambium, the primary cortex and endodermis then becoming
disorganized. The secondary cortex also originating from the cambium,
frequently very unequally developed, instances being recorded of its con-
sisting of 20 layers of cells on one side of the root and only 10 on the other.
Secondary xylem and phloem present in the form of collateral bundles. Small
strands of phloem also occur in the cortex at some distance from the main
stele. The xylem consists of numerous pitted tracheids and less frequent
smaller vessels, the latter often containing a yellowish oily substance. Rays
long and narrow on one side of the root, but short and broad on the other.
Parenchyma of the root containing oil but no starch. Fibres and calcium
oxalate crystals absent.
Commercial samples of Indian Senega root submitted to Kew for identi-
from the American species in having abundant cells with
fication differed
brown contents in the inner part of the cortex. On another occasion a
138 POLYGALACEAE
substitute for Polygala senega was received from the Middle East. This sample
showed a succession of interrupted concentric rings of xylem and phloem
situated externally to the central xylem mass. Although this type of structure
is known in other members of the family it has never been definitely recorded
in Polygala itself, but Holm(1018) implies that it may sometimes occur in
this genus also. A specimen with the same structure as that of the Middle
East sample was found as an admixture in a sample of P. serpentaria from
South Africa.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Atroxima,* Barnhartia,* Bredemeyera, Carpolobia,* Epirrhizanthes (Salo-
monia), Monnina, Moutabea,* Mundtia, Muraltia, Phlebotaenia, Polygala,*
Securidaca,* Xanthophyllum.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Badiera, Bredemeyera, Carpolobia, (Monnina), (Moutabea), Mundtia,
(Muraltia), Phlebotaenia, Polygala, Securidaca, Xanthophyllum.

LITERATURE
(i) On General Anatomy
Holm xoox, 1018, Jauch 1157, O'Donell 1630, Sabnis 1977, Steiger 2191.

(ii) On Wood Structure

den Berger 179, Chalk and Chattaway 362, Cooper 461, Heimsch 938, Lecomte 1334,
O'Donell 1630, Pfeiffer, H. 1711, 1712, Record 1843, 1851, Record and Hess 1886.

38. DICLIDANTHERACEAE
SUMMARY
The anatomy of the Brazilian shrubs and trees belonging to the genus
Diclidanthera is known. The following facts concerning
rather imperfectly
D. laurifolia Mart, have been recorded by Solereder and those concerning
D. pendutiflora Mart, observed at Kew, The wood exhibits the following
features. Vessels mostly solitary, with simple perforations and alternate
intervascular pitting. Vasicentric parenchyma. Wide* heterogeneous rays.
Fibres with bordered pits.

LEAF
Dorsiventral. Hairs D.
laurifolia Mart, simple, with thin septa.
in
Hypoderm of 1-2 layers of large ceils well developed below the upper
epidermis, Mesophyll of D. penduliflora Mart, including about 3 layers of
palisade tissue. Midrib, in the same species, containing a single vascular
bundle, strongly supported on the abaxial and to a smaller extent on the
adaxial side by thick-walled fibres. Vascular bundles of the veins, again in
the same species, embedded in the mesophyll and entirely sheathed by thick-
walled fibres. Solitary crystals in both species, numerous in cells adjacent
to the vascular bundles.
 DICLIDANTHERACEAE 139
Axis
YOUNG STEM
Cork in D. laurifolia Mart, arising superficially; said to be composed of
stone cells. Pericycle in both D. laurifolia Mart, and D. penduliflora Mart,
including a broad, continuous, composite ring of sclerenchyma. Secondary
phloem said to include pitted, sclerenchymatous elements in D. laurifolia^
but no sclerosed elements observed in the phloem of young twigs of D.
penduliflora. Xylem in D. penduliflora appearing, in transverse sections, as
a continuous cylinder traversed by narrow, mostly uniseriate medullary rays.
Vessels, in the same species, somewhat unevenly distributed being entirely
absent from some segments of the xylem, mostly solitary, but some in tan-
gential or oblique pairs or clustered only simple perforations known to occur,
;

Pith fairly broad in D. penduliflora\ consisting of moderately thick- walled,

pitted cells. Stone cells recorded in the pith of D. laurifolia.

WOOD 1

Vessels mostly solitary, perforations simple, intervascular pitting alternate.

Parenchyma vasicentric. Rays tending to break up into smaller units;
heterogeneous (Kribs's Type II A-B). Fibres with large bordered pits.
TAXONOMIC NOTES
The taxonomic position of Diclidanthera is not well established. The
anatomical features, so far as they have been studied, tend to support the
unpublished view put forward by Dr. T. A. Sprague that Diclidanthera may
have affinities with the Polygalaceae. O'Donell (1630) concludes from a study
of the anatomy, and particularly of the anomalous secondary thickening, that
Diclidanthera should be in the Polygalaceae near Moutabea and Securidaca.

GENUS DESCRIBED
Diclidanthera.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy and Wood Structure
O'Donell 1630.

39. VOCHYSIACEAE
(FiG. 34 on p. 144)

SUMMARY
(i) GENERAL
A
small family, including large trees, shrubs, and climbers. It occurs
chiefly in tropical America but extends
to West Africa. Hairs exclusively

non-glandular; always unicellular but sometimes tufted, e.g. in Erisma. Large

nectaries have been recorded above the stipules in species of Qualea. The
stomata are said to be ranunculaceous in Erisma, Salvertia, Vochysia, and
rubiaceous in Qualea. The epidermis of the leaf is frequently composed of
1
Based entirely on the description by O'Donell (1630).
140 VOCHYSIACEAE
cells with mucilaginous inner walls. In the stem, cork arises in the outer

part of the cortex in Salvertia and Vochysia and in the pericycle in Erisma
and Qualea. Intraxylary phloem occurs in the form of a ring or as separate
strands in the perimedullary region, whilst in some species a ring and separate
strands occur together. Solitary crystals have been reported in Erisma and
Qualea and clustered ones in Callisthene, Salvertia, and Vochysia. Secretory
elements. Mucilage cells occur in the cortex and pith of Qualea, and muci-
lage canals in the pith of the stem and petiole as well as in the leaf veins in
Erisma, Salvertia, Vochysia.

(ii)
WOOD
Vessels medium-sized to large and few, perforation plates simple, inter-
vascular pitting alternate, pits to parenchyma similar or elongated, members
of medium-length to moderately short. Parenchyma paratracheal, aliform
and confluent, the latter often in broad continuous bands. Rays up to 2-8
(usually 3-5) cells wide, heterogeneous to homogeneous. Fibres with simple
pits, occasionally septate, of medium length. Traumatic vertical inter-
cellular canals moderately common. Included phloem of the 'foraminate*
type sometimes present.

LEAF
Dorsiventral except in Qualea glauca Warm, where
palisade tissue occurs
towards both surfaces. Hairs mostly simple, unicellular, but varying in
length and in the thickness of the walls, in Callisthene, Qualea, Salvertia, &c. ;

unicellular, 2-armed in certain species of Vochysia; tufted, and provided with

a variable number of rays in Erisma. Glandular hairs absent. Solitary extra-
floral nectaries recorded above the stipules in Erisma laurifolium Warm, and

Vochysia oppugnata Warm. Epidermis on both surfaces composed of poly-

gonal cells as seen in surface view; cells on the upper side in Erisma violaceum
Mart, remarkably small and in part divided by thin, vertical secondary walls ;

2-layered on the upper surface in certain species of Qualea and in Vochysia

rufa Mart.; frequently mucilaginous, for example, in species of Callisthene,
Qualea, Salvertia', cells of the lower epidermis papillose in Qualea glauca
Warm. Stomata confined to the lower surface; ranunculaceous in at least
certain species of Erisma, Salvertia, Vochysia rubiaceous stomata recorded
;

in Callisthene and Qualea. Mesophyll including spirally thickened storage

tracheids in Vochysia rufa Mart. Vascular bundles of the smaller veins
vertically transcurrent by thin-walled or, less frequently, by sclerotic cells.
Vascular bundles in large and small veins with or without an investment of
sclerenchyma according to the species. Only solitary crystals recorded in
Erisma and Qualea and clustered ones in species of Callisthene, Salvertia,
Vochysia. Secretory elements. Mucilage cells present in the parenchyma-
tous portions of the petiole in various species of Qualea, and mucilage canals
in the petiole and veins of species of Erisma, Salvertia, Vochysia.

Axis
YOUNG STEM
Cork arising in the outer part of the cortex in Salvertia and Vochysia and
in the pericycle in Erisma and Qualea. Cortex containing abundant scleren-
 VOCHYSIACEAE 141

chyma. Pericycle also including sclerenchyma, that of Qualea glaziovii

Warm, forming an almost closed ring. Pith including groups of sclerenchyma
in species of Erisma, Qualea, Salvertia, Vochysia. Intraxylary phloem
recorded in 29 species included in the genera Callisthene, Erisma, Qualea,
Salvertia, Vochysia', in the form of strands in species of Emma, Salvertia,
and Vochysia', constituting a continuous ring in the outer part of the pith in
Callisthene and Qualea] a continuous ring accompanied by separate strands
noted in species of Erisma and Qualea. For Interxylary phloem see under
'Wood'. Secretory elements. Mucilage cells common in the pith and
cortex of Qualea. Mucilage canals in the pith recorded in Erisma micranthum
Spruce, Salvertia convallariaeodora St. Hil. and numerous species of Vochysia.
Tannin cells said to be widely distributed in the pith and cortex.

WOOD (Fig. 34 C-G)

Vessels medium-sized (100-200 p mean tangential diameter) to large
(more than 200 fj) in some species of Qualea and Vochysia', solitary and in
radial multiples of 1-4 cells; usually about 2 per sq. mm. Perforations simple,
transverse or slightly oblique. Intervascular pitting alternate and vestured
(78), in Erisma medium-sized to rather large and pits to ray cells often oblong
and simple or with only very narrow borders; intervascular pitting rather
small and with pits to ray cells similar in Qualea and Vochysia. Tyloses and
solid white deposits sometimes present. Mean member length o*3-o-7 mm.

Parenchyma predominantly paratracheal, aliform and confluent in all three

genera; the confluent parenchyma forming regular, continuous bands 2-6 cells
wide in Erisma and Vochysia (Fig. 34 D and E) and rather less continuous
bands in Salvertia (Heimsch 938); usually with a few cells or groups of cells
scattered among the According to Heimsch (938) the parenchyma and
fibres.
vessels in Qualea grandiflora Martius together form an irregular, diagonal
reticulum. Occasionally containing chambered crystals in Erisma and Qualea.
Strands most commonly of 4 cells. Rays most commonly up to 3-5 cells wide,
but a few rays up to 6-8 cells wide in some species of Vochysia] up to I mm.
high in Vochysia hondurensis Sprague, but typically lower; uniseriates numer-
ous in Erisma and Vochysia and composed of upright cells, rather few and
composed of procumbent and upright cells in Qualea*, about 5-8 rays per mm. ;

heterogeneous (Kribs's Type II A and B) in Erisma and Vochysia, with up to

3 or 4 uniseriate marginal rows of square and upright cells; almost or quite
homogeneous (Kribs's Type Het. II B-Hom. I) in Qualea. Fibres with few,
small, simple pits that are almost confined to the radial walls; the pits some-
times with very small borders in Vochysia', a few cells sometimes septate in
Erisma and Vochysia (1894). Mean length 1-0-1-3 mm. Intercellular canals
of the vertical traumatic type observed or reported in Erisma (1712), Qualea
(1801) and Vochysia (Fig. 34 G). Included (interxylary) phloem of the
'foraminate' type common in Erisma (1712) and reported in Erismadelphus
(1851) and Qualea (1886), though rare in the latter; consisting of strands of
unlignifiedphloem scattered through the wood, surrounded and often linked
together by parenchyma (Fig. 34 E) strands of what appears to be soft bast
;

(Fig. 34 F) commonly present also in the rays of Erisma (362).

142 VOCHYSIACEAE
ANOMALOUS STRUCTURE, see under *WOOD*
TAXONOMIC NOTES
Heimsch (938) considers that in its wood anatomy this family forms part of
an interrelated group, the other members of which are the Polygalaceae,
Trigoniaceae, Tremandraceae, Zygophyllaceae, and Malpighiaceae, but notes
that the Malpighiaceae and Vochysiaceae differ from this complex in having
libriform fibres and vestured pits; the Vochysiaceae are further set apart by
the pronounced development of banded parenchyma and the occurrence of
intercellular canals.

ECONOMIC USES
The
timbers generally appear to be of little commercial importance, but
Yemeri, Vochysia hondurensis Sprague, from British Honduras shows some
promise as a plywood and for purposes similar to those for which Gaboon
Mahogany is used.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Callisthene, Erisma, Qualea, Salvertia, Vochysia.

(ii)
FOR WOOD STRUCTURE
Erisma, (Erismadelphus), Qualea, (Salvertia), Vochysia.

LITERATURE
On Wood Structure
Bailey 78, Benoist 170, Besson 186, Campbell 2514, Chalk and Chattaway 362, Heimsch
938, Howard 1088, Kanehira 1209, Knbs 1283, Memaud 1491, Pfeiffer, H. 1712, Record
1787, 1801, 1843, 1851, Record and Hess 1886, Record and Mell 1894, Stone 2202, 2207,
Williams, LL 2530.

40. TRIGONIACEAE
(FiG. 34 on p. 144)

SUMMARY
(i) GENERAL
A small family of treesand climbing shrubs which occur in tropical South
America. The hairs are exclusively simple and unicellular. The epidermis
of the leaf is frequently composed of cells with mucilaginous inner walls. The
stomata in Trigonia are commonly rubiaceous. In the young stem the cork
arises in the outer part of the cortex. The vascular bundles are simple,
xylem includes vessels with simple perforations embedded
collateral, whilst the
in prosenchymatous ground tissue with bordered pits. The family differs
from the Vochysiaceae in the absence of intraxylary phloem.

(ii)
WOOD
solitary, perforations simple, members moderately long. Paren-
Vessels
chyma apotracheal (diffuse and banded) or paratracheal (aliform). Rays up
to 4 cells wide, uniseriates moderately numerous, heterogeneous with many
marginal rows. Fibres with bordered pits, of medium length.
 TRIGONIACEAE 143
LEAF
Dorsiventral. Hairs; only simple, unicellular types known. Epidermis
often consisting of mucilaginous cells, the latter being polygonal in surface
view; those of Lightia licanioides Spruce specially small. Epidermis partly or
wholly 2-layered in Lightia licanioides and Trigoniastrum hypoleucum Miq.
A large-celled, mucilaginous hypoderm also recorded in the first of these
species. Stomata confined to the lower surface in all investigated species;
commonly rubiaceous in Trigonia. Mesophyll including branched scleren-
chymatous fibres in Lightia licanioides and Trigoniastrum hypoleucum. Vas-
cular bundles of the veins with or without an investment of sclerenchyma
according to the species. Crystals of calcium oxalate solitary or clustered.

Axis
YOUNG STEM
Cork arising in the sub-epidermis. Secondary phloem containing scleren-
chyma. Xylem consisting of a cylinder traversed by rays 3-5 cells wide, and
including vessels with simple perforations embedded in prosenchymatous
ground tissue with bordered pits. Intraxylary phloem absent. Solereder

reports Petersen as having recorded the occurrence of inversely orientated

medullary bundles in an undetermined species of Trigonia.
WOOD (Fig. 34 A-B)
Vessels usually exclusively solitary, but Heimsch (938) notes some pore
multiples in Trigonia, Perforations simple; Heimsch has observed some
vestigial scalariform plates in Trigonia. Intervascular pitting extremely rare;
pits to parenchyma and ray cells alternate and rather small or large and
elongated. Mean member length (Trigoniastrum) 0-9 mm. Parenchyma in
Trigoniastrum apotracheal,
1
diffuse and in short tangential lines; sometimes
with some paratracheal in addition (Fig. 34 A) similar or witlr distinct
;

apotracheal bands in Trigonia\ paratracheal in Lightia, vasicentric or 'abaxial,

with or without aliform extensions which may connect as many as four or
five vessels' (938). Sometimes containing chambered crystals. Strands com-

monly of 8-12 cells in Trigoniastrum. Rays up to 4 cells wide; uniseriates

moderately numerous and composed of upright cells or mixed procumbent,
square and upright cells; heterogeneous (Kribs's Type II A and B), some-
times with many uniseriate marginal rows, and varying in different species
from mostly procumbent to square and seldom upright, to all square or
upright. Sometimes containing chambered crystals. Heimsch (938) notes
spiral and reticulate thickening in the ray and wood parenchyma cells of
Trigonia sericea H. B. et K. Fibres with distinctly bordered pits in both radial
and tangential walls. Heimsch refers to septate elements as occurring in some
species. Walls usually thick. Mean length (Trigoniastrum) 1*3 mm.
TAXONOMIC NOTES
The Trigoniaceae were, in the Bentham and Hooker system, included in
the Vochysiaceae, from which they differ most conspicuously in the absence
1
A single specimen of Trigoniastrum hypoleucum Miq. from Java differed from the other
material of this genus examined or described in having paratracheal parenchyma similar to
that described below for Lightia.
 FIG. 34. TRIGONIACEAE, A-B; VOCHYSIACEAE, C-G; TAMARICACEAE H-I; t

FOUQUIERACEAE, J-K
A, Trigoniastrum hypoleucum Miq. B, T. hypoleucum Miq. C, Qualea rosea Aubl. D, O. roiaa
Aubl. E, Erisma tricolor Ducke. F, E. bicolor Ducke. G, Vochysia hondurensis Sprague, H, Tamarix
articulata Vahl. I, T. mannifera Ehr. J, Fouquiera splendens Engelm. K, F. iplendens Engelm.
i.e. Intercellular canals.
 TRIGONIACEAE 145
of intraxylary phloem and in having bordered
pits to the ground tissue
elements of the xylem.
Heimsch (938), on the basis of the wood anatomy, groups this family with
the Polygalaceae, Tremandraceae, Zygophyllaceae, Malpighiaceae, and
Vochy-
siaceae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Lightia, Trigonia, Trigoniastrum.

(ii) FOR WOOD STRUCTURE

(Lightia), (Trigonia), Trigoniastrum.

LITERATURE
On Wood Structure
Heimsch 938, Record 1843, 1851, Record and Hess 1886.

41. FRANKENIACEAE
(Fic. 32 on p. 130; FIG. 35 on p. 146)

SUMMARY
Herbs or shrublets, mostly from maritime regions or dry areas such as
steppes and deserts. They exhibit ecological specializations in correlation
with these habitats, such as leaves with revolute margins, or ericoid micro-
phylls which are sometimes adpressed to the stem. The structure of the
mesophyll is largely determined by the morphology and orientation of the
leaves. The most distinctive anatomical character is the universal occurrence
of epidermal salt glands of a special type (Fig. 35 A-C), generally situated
in pits. They consist of about 6 cells derived from the division of a single
epidermal cell, the 2 uppermost cells resembling a pair of guard cells as seen
in surface view. Hairs are simple trichomes, generally unicellular but some-
times tufted. Calcium oxalate is known to occur only in the form of clustered
crystals. Cork in the stem is sub-epidermal in origin in all of the genera
except Anthobryum. The phloem consists of very small elements. The
xylem contains vessels with very narrow lumina, and definite medullary rays
are lacking. The pith consists of large cells in young stems, but becomes
hollow when older.
LEAF
Generally dorsiventral in species with revolute margins, but tending to be
sub-centric in Frankenia portulacifolia Beatson (syn. Beatsonia portulacoides
Roxb.) and certain other species of Frankenia. The shape of transverse
sections of the leaf taken at corresponding levels is valuable for the identifica-
tion of species. Half the thickness of the mesophyll in species with dorsi-
ventral leaves consists of palisade tissue, but palisade tissue develops only
towards the lower surface in species with the upper surface adpressed to the
axis. Intermediate types of mesophyll with a certain amount of palisade
tissue towards both surfaces also recorded in some species of Frankenia, or
4594 L
146 FRANKENIACEAE
consisting wholly of palisade cells in other species such as F. corymbosa Desf.
Hairs mostly simple, unicellular, sometimes tufted; either thin- or thick-
walled; in some instances having the form of papillae, but in others con-
siderably longer. Some of the hairs bicellular and/or paired in Frankenia
portulacifolia Beatson. Epidermal glands (see 'Summary') sometimes paired
Cuticle often thick, especially in Niederleinia. Cells of the
in Niederleinia.
epidermis frequently enlarged. Stomata ranunculaceous; usually sunken;
in some species confined to grooves.

FIG. 35. FRANKENIACEAE

Gland of Frankenia pulverulenta L. A, In surface-view. B, In
: a
section cut parallel to the common vertical wall of the 2 gland-cells.
C, In a section cut perpendicularly to this vertical wall. By
Solereder.

Variations in the nature and distribution of the mechanical tissue stated

by Niedenzu (1596) to provide a reliable means for identifying species which
are not readily distinguished by floral characters. Sclerenchyma more or less
well developed in the phloem in F. thymifolia Desf., F. jamesii Torn, and
Niederleinia; vascular bundles accompanied by a sclerenchymatous sheath in
the Basigonia section of Frankenia mechanical elements consisting of 'spicular
;

cells* (i.e. bundles of branched sclerenchymatous cells around the veins) in

F. bracteata Turcz., &c., or of thick- walled xylem elements in F. campestris
Schau., &c. Certain of the palisade cells become thick-walled and serve a
mechanical function in F. portulacifolia Beatson. Principal veins embedded
in the mesophyll variations in their number and degree of development said
;

to be a feature of specific diagnostic value. The median vein alone well

developed in Hypericopsis persica (Jaub. et Spach) Boiss. and Niederleinia
juniperoides Hieron. although smaller subsidiary bundles also occur in the
upper part of the leaf sheath of Niederleinia^ and larger ones in Anthobryum.
Vascular bundles of the veins made up of xylem containing vessels with
narrow lumina and phloem composed of very small elements. Petiole in
Hypericopsis described by Surgis (2222) as exhibiting a similar vascular struc-
ture to that of the median vein of the lamina.

Axis
YOUNG STEM (Fig. 32 A)
The following features may be added to those given in the summary.
Epidermis narrow, without hairs, glands, or stomata in Niederleinia, but
 FRANKENIACEAE 147

according to Surgis (2222) possessing all of these structures in

Hypericopsis.
Cork pericyclic in origin in Anthobryum. Pericycle containing rings of
fibres in Hypericopsis seen to include strands of fibres in Frankenia laevis
;

Linn, when examined at Kew (Fig. 32 A), but a composite continuous ring
of sclerenchyma also recorded in the same species. Phloem and xylem in
the form of continuous cylinders in F. laevis. Vessels of the xylem in all
species small, scattered, with simple perforations; those of F. laevis up to
about 30 p, in radial diameter. Ground tissue of the xylem composed of
prosenchyma with simple pits.
(Surgis (2222) gives particulars of anatomical characters whereby individual
species in this family may be recognized. These can best be consulted in his
paper.)
ECONOMIC USES
Frankenia portulacifolia Beatson (syn. Beatsonia portulacoides Roxb.) is the
source of St. Helena tea.

TAXONOMIC NOTES
The family iswell defined by the possession of characteristic epidermal
glands, but it is interesting to note that similar glands also occur in the
Tamaricaceae and Plumbaginaceae. The glands are probably to be regarded
as an ecological specialization which may quite well have arisen independently
in each of these families. Other families to which it has been suggested that
the Frankeniaceae may be related include the Elatinaceae and Caryophyllaceae.
The anatomical structure of Anthobryum confirms that this genus belongs
to the Frankeniaceae and not to the Primulaceae in which it was at one
time included.
GENERA DESCRIBED
Anthobryum, Beatsonia, Frankenia,* Hypericopsis, Niederleinia.
*
Represented in the Kew slide collection.

LITERATURE
On General Anatomy
Niedenzu 1596, Surgis 2222.

42. CARYOPHYLLACEAE
(Including Illecebraceae of Bentham and Hooker)
(Fie. 37 on p. 154; FIG. 38 on p. 160)

SUMMARY
A family of annual or perennial herbs, most of which have a number of
anatomical characters in common. It occurs chiefly in north temperate
regions, but some species extend far to the north. The hairs are generally
simple, unicellular or uniseriate trichomes, sometimes provided with a uni-
cellular glandular head. Branched hairs also occur in certain genera (see
'Leaf'). Stomata are usually of the so-called caryophyllaceous type, being
accompanied by 2 subsidiary cells whose common wall is placed transversely
to the pore, but exceptions occur, especially amongst the genera included by
148 CAR YOPH YLLA CEAE
Bentham and Hooker in the Illecebraceae. Even amongst these the caryo-
phyllaceous type of stoma also occurs in Habrosia and Sderanthus. The leaf
may be dorsiventral or centric, whilst an interesting feature of certain arctic
species is that the palisade tissue is confined to the abaxial side. The amount
and distribution of the palisade tissue sometimes varies within a species
according to the nature of the illumination. Even arctic members of the
genus exhibit no anatomical specializations which serve for the reduction of
transpiration, a feature which Warming (2362) suggests is possibly correlated
with the obliquity of the sun's rays which fail to raise the temperature of the
leaves appreciably, whilst the atmospheric humidity is very great, fogs and
cloud being frequent. In this connexion it is important to realize, however,
that the cushion-like habit and common occurrence of narrow overlapping
leaves may also be important. Calcium oxalate is commonly secreted in the
form of large, conspicuous cluster-crystals, although solitary crystals and
crystal-sand occur rarely. A narrow or broad ring of sclerenchyma is nearly
always present outside the phloem. In some instances the prosenchymatous
elements of which these rings are composed have much thicker walls, and the
lumen is narrower towards the outside than in those towards the phloem.
When this is so the prosenchymatous nature of the inner elements is not
apparent in transverse sections, although their true nature is revealed in
longitudinal sections. Solereder described these sclerenchyma rings as
occurring in the cortex, but observations made at Kew suggest rather that
they are pericyclic, the true primary cortex being very narrow in some
instances. The xylem and phloem of the stem appear, in transverse sec-
tions, either in the form of a continuous cylinder, or as individually distinct
bundles separated by broad rays. Intermediate conditions between these two
types also occur. The arrangement of the vascular tissue appears to be
characteristic for species and therefore of diagnostic value. Secondary
medullary rays apparently do not occur. Vessels with simple perforations.
Anomalous secondary thickening, consisting of the formation of succes-
sive rings of bundles, occurs in old sterns, but more frquently in roots of
certain genera. Friedel and Yen (719) have described an interesting species
of Drypis which strongly resembles an Eryngium (Umbelliferae) in external
form. They show nevertheless that the internal structure of the 2 genera
differs considerably, that of the Drypis being strongly caryophyllaceous in
type.

LEAF
Dorsiventral or centric; central tissue serving for water-storage in Sphaero-
carpos. For further details see under 'Mesophyll* below. Hairs, (i) Uni-
cellular types recorded in certain species of Herniaria, Paronychia, Siphonychia.

(ii) Simple, long or short, uniseriate hairs frequent in most genera except
some of those included by Bentham and Hooker in the Illecebraceae. (iii) Uni-
seriate hairs with a glandular cell at the apex recorded in Dysphania, Habrosia,

Loeflingia, Silene (Fig.37 A), Spergula, Spergularia. Almost sessile glands

(Fig, 37 B) occur, for example, in Viscaria. (iv) Branched hairs occur in
Achyronichia, Cerastium, Pollichia sp., Polycarpaea> Polycarpon, and Stipu-
licida. Hair structure believed by Correns (476) to be of specific diagnostic
value in Cerastium. Epidermis composed of cells with straight or undulating
 CARYOPHYLLACEAE 149
anticlinal walls, cells elongated in species with long leaves. Epidermal cells

especially tall as seen in transverse section in some species;papillose in certain

species of Gypsophila, Polycarpon, and Saponaria, the nature of the papillae
being of specific diagnostic value in the last of these. Papillae in longitudinal
rows, chiefly near the midrib and margin in Achyronychia, Cometes, Corrigiola,
Habrosia, Loeflingia, Ortegia, Pollichia, Polycarpaea, Polycarpon, Scleranthus.
Cuticle smooth, punctate or variously striated in different species. Wax
thickly deposited on the surface in certain species of Cometes, Dicheranthus,
and Pteranthus. Stomata generally of the caryophyllaceous type (Fig. 37 c
and 38 B), but exceptions also recorded, notably in the Caryophyllaceae-
Polycarpeae except Loeflingia, and Illecebraceae of Bentham and Hooker.
The caryophyllaceous type of stoma is also rare in certain species of Arenaria,
Cerastium, and Stellaria with elongated epidermal cells. Stomata sometimes
tending to be of the cruciferous type. Stomata generally present on both
surfaces in the Illecebraceae of Bentham and Hooker, as well as in Honckenya
peploides (L.) Ehrh. (Syn. Arenaria peploides Linn.) (Warming 2362), certain
Cerastium spp., Colobanthus quitensis Bartl. (Betts 187) (most numerous on
the upper side), Polycarpaea corymbosa (L.) Lam. (Sabnis 1977), Saponaria
officinalis Linn. (Holm 1057). Groups of stomata, overlying patches of pale-
coloured parenchyma situated at the ends of veins, occur in the teeth of young
leaves of the genera included by Bentham and Hooker in the Caryophylleae.
Stomata in certain narrow-leaved arctic species have the pores parallel to the
longitudinal axis of the leaf. Hypoderm recorded towards the lower surface
in a species of Colobanthus, and a collenchymatous pseudo-hypoderm in
Saponaria (Simmler 2100). Mesophyll centric or dorsiventral, vascular
strands surrounded by water-storage cells in species with centric leaves.
Dorsiventral, with palisade tissue towards the upper (adaxial) surface in
Colobanthus quitensis Bartl. (Betts 187), an arctic species of Minuartia
(Warming 2362), Saponaria officinalis Linn. (Holm 1057), but, according to
Simmler (2100) palisade tissue sometimes occurs towards both surfaces in
this genus or alternatively the whole of the mesophyll is composed of rounded
cells, those towards the centre of the leaf being slightly larger. A normal
dorsiventral mesophyll also recorded in certain species of Siphonychia. Dorsi-
ventral mesophyll with palisade tissue towards the abaxial surface in a few
species of Cerastium, Melandrium, Minuartia, Silene, and Viscaria recorded
by Warming (2362). Distribution of palisade tissue variable in certain arctic
species, notably of Cerastium and Minuartia. Lacunae observed, also by
Warming, in the spongy mescphyll towards the adaxial surface in a few species
of Cerastium, Melandrium, Silene, and Viscaria. Stellate cells recorded in the
spongy mesophyll of Cerastium sp. Centric mesophyll mentioned by various
authors in Achyronychia and in certain species of Arenaria, Cardionema,
Cometes, Corrigiola, Dicker anthus, Dysphania, Gymnocarpos, Habrosia, Haya,
Hernieria, Illecebrum, Paronychia, Pollichia, Polycarpaea, Pteranthus, Scleran-
thus, Sclerocephalus, Siphonychia. Midrib, in transverse sections, exhibiting
one main vascular bundle accompanied by two smaller ones in Scleranthus
biflorus (Forst.) Hook, according to Foweraker (703). One bundle enters the
leaf in certain arctic species, but divides to form three in species with narrow
leaves, or a greater number in those with broader ones (Warming 2362).
Vascular bundles not usually accompanied by sclerenchyma except in certain
ISO CARYOPHYLLACEAE
species of Arenaria, Cardionema, Habrosia, Melandrium, Minuartia, Pollichia,
Scleranthus, and Silene. Calcium oxalate commonly present in the form of
large, conspicuous cluster crystals in many genera and species including
Arenaria, Corrigiola, Gymnocarpos, Minuartia sp., Pteranthus, Scleranthus,
Silene. The abundance of the crystals sometimes varies within a single species
in specimens from different localities. Crystal-sand also recorded in Dyspfiania,
Gymnocarpos, Habrosia, and other genera. Tanniniferous substances
present in Polycarpaea corymbosa Lam. according to Sabnis (1977).

Axis
STEM (Fig. 38 E and G)
Epidermis including longitudinal rows of papillose cells in Loeflingia,
Ortegia, Polycarpon, and Stipulicida. Cork usually arising in the pericycle,
e.g. in certain species of Dianthus, Honckenya, Lychnis, Polycarpaea, Sagina,
Saponaria, Silene, Stellaria; arising in the outermost part of the cortex in
Spergularia; sub-epidermal in Paronychia. Cortex not usually exhibiting
distinctive features, frequently narrow and sometimes containing assimilatory
tissue (Dianthus, &c.); the inner part consisting of small cells with very thick
mucilaginous walls, e.g. in a species of Colobanthus, but with an outer zone of
small suberized cells between this and the epidermis (Betts 187); consisting
of thick- walled cells, giving a dark-brown colour when treated with chlorzinc-
iodide, in Scleranthus biflorus (Forst.) Hook. (Foweraker 703); completely
sclerenchymatous in Drypis spinosa Linn. (Friedel and Yen 719). Endo-
dermis frequently well defined, e.g. in certain species of Arenaria, Corrigiola,
Dianthus, Lychnis, and Saponaria. Pericycle characterized by a sclerenchy-
matous ring of varying width in different genera and species. With a broad
ring up to about 10 cells wide in certain species of Dianthus, Lychnis Minuartia,
,

and Silene] with a ring 4-6 cells wide in certain species of Cometes, Corrigiola,
Gymnocarpos, Herniaria, Illecebrum, Lychnis, Paronychia, Pollichia, Pteran-
thus, Saponaria (not in very young stems), Scleranthus, and Silene ; with a ring
1-2 cells wide in Herniaria sp. Pericycle described by Friedel and Yen (719)
as wholly sclerenchymatous in Drypis. Outer part of the sclerenchymatous
ring more strongly lignified and composed of cells with narrower lumina than
the part towards the phloem, the prosenchymatous nature of the inner
elements being visible only in longitudinal sections, in certain species of
Gypsophila, Lychnis, Saponaria, and Silene. Ring of fibres accompanied by
groups of stone Polycarpaea corymbosa (L.) Lam. according to Sabnis
cells in

(1977). The pericyclic sclerenchyma becomes split up into separate strands

in species with a considerable amount of secondary thickening. Vascular
bundles individually distinct in transverse sections, but tending to become
united to form relatively large arcs in certain species of Arenaria, Corrigiola,
Drypis, Lychnis (Fig. 38 G), Paronychia, Polycarpaea, Saponaria (rarely in
this genus), Scleranthus, Silene, Spergula, Stellaria. Bundles not individually
distinct, but xylem and phloem forming continuous rings in certain species
of Dianthus, Gymnocarpos, Gypsophila (Fig. 38 E), Herniaria, Minuartia,
Paronychia, Pollichia, Sagina, Saponaria (most species), Silene, Spergularia.
Phloem forming a continuous ring, but accompanied by xylem in separate
strands in certain species of Arenaria, Buffonia, and Telephium. Vessels with
simple perforations. A large proportion of the phloem often consists of
 CARYOPHYLLACEAE 151

parenchyma. For anomalous thickening see below. Calcium oxalate, in

the form of large cluster crystals occurs in certain species of Dianthus,
Herniaria, Lychnis (chiefly in the endodermis), Saponaria (not in the endo-
dermis), Silene (in the endodermis). Solitary crystals rare, said to be confined
to Polycarpaea.

WOOD
Vessels with simple perforations. Parenchyma frequently constituting
a large proportion of the wood. Rays generally absent, except for the tissue
between the bundles in those species in which they are individually distinct.
Fibres with simple or bordered pits. The anatomy of the woody 'Dianthus
arboreus* (syn. D. aciphyllus Sieb.) has been fully described by Tellini (2240).

ROOT
Structure best known which is described
in Saponaria officinalis Linn,
under 'Economic Uses'. Roots of Silene vulgaris Linn,
grown in different
kinds of soil were found by Millner (1538) to differ in the amount of peri derm
formed, and to have more numerous vessels when grown in sand than in clay.
Minor differences between the root structure of closely related species of
Silene also recorded by the same author.

ANOMALOUS SECONDARY THICKENING

Concentric rings of xylem and phloem or of individually distinct vascular
bundles occur in the roots and/or sometimes the old stems of certain species
of Acanthonychia Achyronychia, Cardionema, Cerdia, Cometes, Corrigiola,
y

Dysphama, Haya, Leptogonium, Ortegia, Pollichia, Polycarpaea, Polycarpon

(Fig. 37 D), Pycnophyllum, Silene, Spergula, Spergularia, Stipulicida, Tele-
phium, and possibly other genera. For further details see Pfeiffer (1712).
Although secondary thickening is usually normal in Acanthophyllum, the
broad rings of secondary xylem are interspersed with narrow bands of un-
lignified parenchyma in certain species. In other species the initial ring of
normal bundles becomes split up into 4 isolated strands of xylem and phloem
by radial groups of parenchyma. Later on 4 meristematic areas arise in the
pith opposite the xylem strands and give rise to inversely orientated xylem
and phloem which eventually unite with the outer vascular tissues.

TAXONOMIC NOTES
The anatomical characters of the genera included by Bentham and Hooker
in the family Illecebraceae are so similar to those of the Caryophyllaceae that
both groups have been described together as in the system of Engler and
Prantl. It is not possible from the anatomical facts which are at present
available to decide to which of the tribes of the Caryophyllaceae the Illece-
braceae may belong, because the existing tribes of the first of these groups do
not themselves seeni to be very well defined on anatomical grounds. Further
research may enable this matter to be cleared up.
Another family whose floral characters resemble those of the Caryo-
phyllaceae is the Polygonaceae. The anatomical feature which seems to lend
most support to this relationship is the occasional occurrence of anomalous
secondary thickening in both families. It is also interesting to note that the
152 CARYOPHYLLACEAE
dominant form in which calcium oxalate is secreted in both families is the
cluster crystal. Other anatomical features of the two families are not suffi-
ciently distinctive to provide conclusive evidence of their close relationship,
although there are no known facts which make their close connexion seem
improbable.
The position in the Centrospermae in which the Caryophyllaceae (including
Illecebraceae) have been placed in the Engler system differs considerably
from the position to which the family is assigned in the Genera Plan-
tarum of Bentham and Hooker. One of the most outstanding features of
the Centrospermae is the widespread occurrence of anomalous secondary
thickening throughout the group. On these grounds there seems to be good
reason for including the Caryophyllaceae in the order since anomalies of the
same kind occur sporadically in the family. The widespread occurrence of
crystal-sand and cluster crystals in the Amaranthaceae brings this family
more into line with the Caryophyllaceae than some of the other families in
the Centrospermae such as the Phytolaccaceae where raphides predominate.
Evidence of affinities between the Caryophyllaceae and Portulacaceae is
afforded by the occurrence of saponin in certain members of both families.
On the other hand, the occurrence of interxylary phloem in the Portulacaceae
tends rather to demarcate this family from the remainder of the Centro-
spermae with the exception of the Basellaceae.

ECONOMIC USES
The family contains many well-known ornamental garden plants such as
Carnations, Pinks, Gypsophilas, &c. The Soap Root obtained from Saponaria
officinalis
Linn, contains saponin, and has in the past been used for medicinal
purposes. Its anatomical characters include the following. Root covered
externally by a layer of brown cork cells. Secondary phloem consisting of
sieve tubes and parenchyma. Xylem composed mainly of parenchyma, but
including scattered vessels, which are solitary or tending to be in radial rows.
Rays absent. Calcium oxalate, mostly in the form of conspicuous cluster
crystals but also occurring as crystal-sand, secreted in all parenchymatous
tissues. Starch absent. Saponin present. Structure somewhat similar in
Stlene, but vessels in the older roots becoming twisted.
The root of Corrigiola littoralis Linn, has sometimes been substituted for
that of Anacyclus pyrethrum (Family Compositae, see p. 802) which is used in
medicine. Transverse sections of the root of Corrigiola littoralis exhibit a
central xylem mass which tends to be radially dissected, surrounded by con-
centric rings of vascular bundles.

GENERA DESCRIBED
Achyronichia, Arenaria,* Buffonia, Cardionema, Cerastium, Cerdia, Colo-
banthus, Cometes, Corrigiola,* Dianthus,* Dicheranthus, Drypis, Dysphania,
Gymnocarpos, Gypsophila,* Habrosia, Haya, Herniaria,* Honckenya, Illece-
brum, Loeflingia, Lychnis,* Melandrium, Minuartia,* Ortegia, Paronychia,
Pollichia, Polycarpaea, Polycarpon, Pteranthus, Pycnophyllum, Sagina,
Saponaria,* Scleranthus,* Sclerocephalus, Silene,* Siphonychia, Spergula,
Spergularia, Stellaria,* Stipulicida,* Telephium, Viscaria.
* Kew
Represented in the slide collection.
 CARYOPHYLLACEAE 153
LITERATURE
On General Anatomy
Betts 187, Bonnet 230, Chkhubianishvili 397, Correns 476, Foweraker 703, Friedei and
Yen 719, Holm 1057, Joshi 1197, Milkier 1538, Pax and Hoffmann 1677, Pfeiffer 1712,
Privault 1759, Sabnis 1977, Simmler 2100, Starr 2188, Tellini 2240, Warming 2362*

43. PORTULACACEAE
(Fie. 36 on p. 154; FIG. 38 on p. 160)

SUMMARY
Herbs, or small shrubs, often succulent, and frequently containing abundant
mucilage in the parenchyma cells of the leaf and stem. The family is widely
distributed but occurs chiefly in the New World. The presence of mucilage
and the comparatively poor development of mechanical tissue make it very
difficult to cut freehand sections of members of this family. The hairs may
be (i) simple, unicellular in Calandrinia (ii) in the form of papillae, e.g. in
;

Spraguea, and Talinum; (iii) multicellular and shaggy, the superficial cells
sometimes being papillose (Fig. 360); (iv) branched in Calandriniopsis\
(v) uniseriate and glandular (Fig. 36 D). The stomata are somewhat variable,
but sometimes rubiaceous with 2 or 4 subsidiary cells parallel to the pore.
Crystals of calcium oxalate occur in the form of (i) clusters; (ii) solitary
prisms; (iii) crystal-sand consisting of acicular crystals. The structure of the
mesophyll is very variable and exhibits several interesting types of structure
J

probably to be regarded as ecological specializations (see 'Leaf ). The fleshy

leaves of Portulaca grandiflora Hook, are provided with a shallow groove in
the adaxial surface. The pericycle of the stem includes a sclerenchymatous
ring in some species, but in many others mechanical elements are lacking in
this region. The vascular bundles of the axis appear separate and indivi-

dually distinct in transverse sections of most species, although exceptions

occur. In the xylem the vessels are provided with simple perforations, and
the fibres of the ground tissue with simple pits.

LEAF
Cuticle thick in Ceraria namaquensis Pearson et Stephens but thin in
Portulacaria afra Jacq., according to Michell (1511). Hairs as described in
the 'Summary* above; srid to be absent from the Montioideae with the
exception of Wangerinia. Epidermis including solitary cells with the form
of bladder-like protrusions from the surface in Spraguea and Talinum.
Hypoderm of thick-walled, unlignified cells recorded beneath the lower
epidermis in Lenzia. Stomata (Fig. 363) present on both surfaces of the
leaf in most instances; confined to the lower surface in Montia australasica

(Hook, f.) Pax et Hoffmann (syn. Claytonia australasica Hook, f.) according
to Betts (187); most numerous on the lower surface in certain species of
Portulaca, Spraguea, and Talinum; rubiaceous, accompanied by 4 subsidiary
cells parallel to the pore in Calandrinia or by 2 in some of the other genera
and species. True subsidiary cells absent from Montia. Mesophyll of the
succulent leaves varying greatly in structure, (i) Dorsiventral, e.g. in Montia
australasica according to Betts (187). (ii) Consisting of assinulatory and
 D

FIG. 36. PORTULACACEAE

A, Transverse section of the leaf of Portulaca oleracea
Linn. B, Stomatal apparatus of Portulaca oleracea. C
Shaggy hair, and D, Glandular hair of Calandrinia urn-
btllata DC. After Vesque.

FIG. 37. CARYOPH YLLACEAE

A, Glandular hair of Silene viUosa, Forsk. B f Transverse section through the outer-
most part of the sticky region of the stem of Viscaria viscosa (Scop.) Aschers. C, Epi-
dermis of the leaf with stomata in Viscaria viscosa. D, Transverse section through the
root of Polycarpaeafragilis, Delile. By Solereder.
 PORTULACACEAE 155

large water-storage cells either intermingled or localized in separate zones.

Assimilatory cells forming a network between the water-storage cells in
Portulaca hirsutissima Cambess; confined to the sub-epidermal region on
either side of the leaf, the centre being occupied by water-storage cells in
Portulaca tuberosa Roxb. (iii) Consisting almost wholly of water-storage
cells, the vascular bundles alone being surrounded by green palisade cells in
certain species of Portulaca (Fig. 36 A).
Vascular bundles in the veins not accompanied by sclerenchyma but some-
;

times surrounded by a sheath of large, colourless, parenchymatous cells, e.g.

in Portulaca grandiflora Hook, and P. oleracea Linn. (Fig. 36 A). Petiole, in
transverse sections through the distal end, exhibiting a solitary vascular strand
in Portulaca grandiflora and P. oleracea^ the strand being surrounded by a
well-defined endodermis in the last named species. Crystals exclusively
clustered in Calyptridium, Lewisia, Monocosmia, Portulaca, Portulacaria^ and
Talinum. ; occurring as mixed solitary and clustered forms in Anacampseros,
Calandrinia, Spraguea\ crystal-sand, consisting of small needles and prisms
recorded in Calandrinia* No crystals observed in Montia. Mucilage cells
generally present, sometimes especially large in Anacampseros and Portulacaria.

Axis
STEM (Fig. 38 A and c)
Epidermis composed of dome-shaped cells with thick cuticle in Montia
australasica (Hook, f.) Pax et Hoffmann (syn. Claytonia australasica Hook, f.)
according to Betts (187), and, of polygonal cells in certain species of Portulaca
according to Sabnis ( 1 977). Bodies, believed to be composed of an organic sub-
stance, recorded by Kisser (1239) as being included within the walls between the
epidermal cells,and in those between the epidermis and adjacent collenchyma
in 'Portulaca gilliem\ Similar bodies apparently absent from herbarium
specimens of other members of the genus or in other genera of the family
examined at the same time. Cortex broad, consisting of large, thick-walled
cells with intercellular spaces between them in Montia australasica. Outer

part of the cortex consisting of thin-walled parenchyma filled with starch, but
the inner part containing chlorophyll in certain species of Portulaca. Scleren-
chymatous cells recorded by Michell (1511) in the cortex, opposite the
vascular bundles of species of Ceraria. Endodermis well defined in Montia
australasica not conspicuous in species of Ceraria. Pericycle frequently
;

without sclerenchyma, e.g. in certain species of Montia and Portulaca^ but

including mechanical elements in some of the Portulacoideae, e.g. in Calan-
drinia caulescent H. B. et K. A
continuous ring of sclerenchyma noted in the
pericycle of Lenzia. Vascular bundles collateral, appearing individually
distinct and arranged in a ring in transverse sections of the species of Calan-
drinia, Montia, and Portulaca available for examination; also recorded as
being separate in Ceraria and Lenzia. Bundles deeply seated and stated by
Sabnis (1977) to be separated by uniseriate rays in certain species of Portulaca.
Xylem and phloem recorded by Betts (187) as being in the form of a con-
tinuous cylinder in Montia australasica. Phloem accompanied towards the
outside by considerable masses of fibres in species of Ceraria. Xylem con-
taining tangential bands of parenchyma in Portulacaria afra Jacq. Vessels
small, with simple perforations sometimes tending to be in radial rows with
; ;
156 PORTULACACEAE
simple pits in Ceraria gariepina Pears, et Steph., according to Michell (1511);
with spiral thickening in Montia perfoliata (Don.) How, Portulaca grandiflora
Hook., P. oleracea Linn., and probably in other species as well, Intraxylary
phloem stated by Pax and Hoffmann (1676) to occur in Montia and its allies,

but observations at Kew show that it is not always well defined. Pith varying
in size in different species, generally consisting of thin-walled cells. Dark-
brown mucilage cells commonly present according to Michell (1511), those
in two species of Ceraria being in the form of a ring, or forming a network
amongst water-storage cells in the cortex. Drops of fixed oil present through-
out the stem in certain species of Ceraria, according to the same author.

BARK
Leathery, easily detached from the stem, and sometimes containing an
inflammable material in species of Ceraria thinner, but similar in structure
;

in species of Portulacaria according to Michell (1511).

TAXONOMIC NOTES
Theanatomical characters of the Portulacaceae as a whole are not suffi-
ciently distinctive to provide reliable evidence concerning the affinities of the

family. There seems no definite reason for not accepting the commonly held
view that the Portulacaceae and Caryophyllaceae are near relatives, both
included in the Centrospermae. The occurrence of saponin in certain of the
Portulacaceae as well as in some of the Caryophyllaceae provides additional
evidence of the close relationships between the two families.
An attempt has been made by Chorinsky (408), who compared the epidermal
outgrowths of Anacampseros with those of Pereskia and Rhipsalis, to establish
the existence of affinities between the Portulacaceae and Cactaceae. The
evidence afforded by this comparison is stated to indicate the existence of
affinities between these two families, but this fact, taken by itself, does not
appear to be very convincing.

ECONOMIC USES
Certain members of this family are commonly cultivated in English gardens,
but they are not of any economic importance. Species of Montia, Portulaca,
and Talinum have sometimes been used as pot-herbs, while Lewisia rediviva
Pursh is also edible.
GENERA DESCRIBED
Anacampseros, Calandrinia,* Calandriniopsis, Calyptridium, Ceraria,
Hectorella, Lenzia, Lewisia, Monocosmia, Montia,* Portulaca,* Portulacaria,
Spraguea, Talinum, Wangerinia.
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Betts 187, Chorinsky 408, Kisser 1239, Marroquin and Howard 1446, Michell 1511,
Pax and Hoffmann 1676, Sabnis 1977.
 44. TAMARICACEAE
(Fic. 34 on p. 144; FIG. 38 on p. 160)

GENERAL
SUMMARY
(i)
Shrubs or trees with slender branches and small needle-like or scale leaves,
which grow chiefly on desert or saline soils in the Mediterranean region and
Central Asia. The leaf is generally centric, the centre of the mesophyll being
composed of water-storage cells, whilst the mostly ranunculaceous stomata
are usually present on both surfaces or confined to the upper side. The pores
of the stomata are usually arranged transversely to the course of the veins.
External glands which secrete mineral salts have been recorded on the leaves
of all genera and on the stem of certain species of Tamarix. According to
Brunswick (296) the solitary or clustered crystals which are generally present
consist of gypsum and not of calcium oxalate.

(ii) WOOD
Vessels small to medium-sized;
solitary, in short multiples and in clusters;
semi-ring-porous, perforations simple, intervascular pitting alternate and
very small, pits to parenchyma similar, members extremely short. Paren-
chyma scanty paratracheal to vasicentric, storied, fusiform cells common.
Rays very wide (up to 25 cells wide) and very high, heterogeneous. Fibres
with simple pits, moderately to extremely short.

LEAF
Cuticle smooth, Hairs scanty, where
fairly thick, striate or punctate.
present unicellular with thick walls and narrow lumina. External glands
*

(see Summary') frequently embedded in the mesophyll. Epidermis com-

posed of cells with straight anticlinal walls; papillose in a few species of
Myricaria, Reaumuria, and Tamarix. Stomata present on both surfaces or
confined to the upper side, with the pore placed transversely to the course of
the veins in Hololachne, Myricaria, Reaumuria, and Tamarix, but exceptions
recorded in Reaumuria; generally ranunculaceous, but said to be rubiaceous
in a species of Myricaria. Mesophyll centric except in certain species of
Tamarix with an abortive lamina, palisade tissue in the latter being confined
to the lower surface of the leaf sheath which is directed towards the light.
Branched fibres, present in the mesophyll of Reaumuria oxiana Boiss., extend
between the elongated palisade cells to the epidermis. Storage-tracheids,
sometimes extending to the epidermis, also recorded in certain species of
Reaumuria and Tamarix. Vascular bundles of the veins not usually accom-
panied by sclerenchyma, except sometimes in Tamarix embedded in the ;

mesophyll. Solitary and cluster crystals very common; said by Brunswick

(296) to consist of gypsum and not of calcium oxalate; generally present
in
the mesophyll or beside the vascular bundles in Hololachm, Myricaria, and
Tamarix. Tannin often present in the palisade tissue.

Axis
YOUNG STEM (Fig. 38 D)
External glands, similar to those on the recorded by Sabnis (1977)
leaf,
in certain species of Tamarix. Cork superficial and durable in Tamarix;
158 TAMARICACEAE
formed from a succession of phellogens and easily detached in Reaumuria
according to Brunner (295). Cortex generally consisting of large or small,
thin-walled parenchymatous cells, the outer part composed of assimilatory
cells in certain species. Cortex also including water-storage tracheids with
pitted or scalariform thickenings, and isolated, enlarged, stone cells in certain
species of Tamanx. Pericycle containing strands of fibres, which form a
continuous ring in young stems but become more widely separated by paren-
chyma when older. Phloem present in the form of a continuous ring or in
strands. Secondary phloem usually containing massive strands of fibres,
which are convex towards the exterior and arranged in tangential series as
seen in transverse sections. Phloem fibres lacking in Reaumuria. Xylem
constituting a large proportion of the total diameter of the stem even when
young, being present in the form of a cylinder traversed by the fairly broad
primary rays in Tamanx (Fig. 38 D). Vessels with simple perforations. Pith
consisting of thick- walled cells in species of Tamanx. Crystals, stated by
Brunswick (296) to consist of gypsum, present in the pith and cortex of
Hololachne, Myricaria, and Tamanx. Tannin often abundant.

WOOD (Fig. 34 H-I)

Vessels medium-sized (mean tangential diameter usually 100-20 p) in
most species of Tamarix, small (less than 100 /x) in Myricaria and Tamanx
p.p. solitary, in short multiples and in clusters; 3-12 per sq. mm. semi-ring-
; ;

porous in some species of Tamarix and ring-porous in Myricaria (185).

Perforations simple, transverse. Intervascular pitting alternate, very small to
minute; pits to ray cells similar. Mean member length 0-1-0-2 mm. Paren-
chyma paratracheal, as a few cells to broad sheaths round the vessels and
vessel groups (Fig. 34 H). Strands of 1-2 cells; fusiform cells common.
Storied. Rays up to 25 cells wide in Tamarix articulata Vahl., but usually
up to 10-15 ceu s
*
w*de in other species; often up to 2 mm. high; uniseriates
rare or absent; rays very few, commonly not more than 2 or 3 per mm.;
heterogeneous, the rays often appearing as homogeneous in tangential sec-
tions (Fig. 34i), but with several rows of square cells
on the margins and
sheathing the rays. Numerous solitary crystals sometimes present, e.g. in
T. tetragyna Ehrb.; Brown (1679) re po rts occasional druses in T. articulata.
Fibres with few, simple pits, almost entirely limited to the radial walls.
Walls moderately thin. Mean length o-4-O'9 mm.
TAXONOMIC NOTES
The occurrence of similar external glands in the Frankeniaceae and Tamari-
caceae is taxonomically interesting, but it is
important to realize that the glands
are probably ecological specializations which may have arisen independently
*
in the two families. See also under Frankeniaceae'.

ECONOMIC USES
Tamanx galls are used in India in native medicines, as well as for dyeing or
tanning. They are occasionally used as a substitute for oak galls. Tamarix
manna, which obtained from twigs punctured by insects, is also used for
is
medicinal purposes in India. The wood is of no commercial importance.
It was commonly used for dowels by the Ancient Egyptians.
 TAMARICACEAE 159
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Hololachne, Myricaria, Reaumuria, Tamarix,* (Fouquiera is described
under Fouquieraceae).
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
(Myricaria), Tamarix.

LITERATURE
(i) On General Anatomy
Brunner 295, Brunswick 396, Niedenzu 1597, Sabnis 1977, Zemke 2505.
(ii) On Wood Structure
Chowdhury 411, Dadswell and Record 533, Greguss 2522, Howard 1088, Messeri 1493,
Pearson and Brown 1679, Record 1843, 1851, Trabut 2274.

45. FOUQUIERACEAE
(FiG. 34 on p. 144; FIG. 38 on p. 160)

SUMMARY
(i) GENERAL
A small family of peculiar dwarf shrubs or small trees confined to south-
western North America, consisting of the two genera Fouquiera and Idria.
Fouquiera includes several species of shrubs or small trees, 15-20 ft. high,
with numerous branches radiating from a short, often inconspicuous trunk.
Idria, of which only one species is known, is a stem succulent with a greatly
swollen trunk somewhat resembling an inverted parsnip, bearing fleshy
branches and spines, and sometimes attaining a height of 40 ft. when mature.
Young plants somewhat resemble kohl-rabi. The anatomical structure of the
two genera differs somewhat in correlation with their divergence in habit, so
they are here described separately.

(ii) WOOD
Vessels small, with a slight tangential or radial pattern, often ring-porous,
with simple perforations, members moderately short. Parenchyma diffuse.
Rays 4-6 cells wide, often composed almost entirely of square to upright
cells. Fibres with small bordered pits, of medium length.

FOUQUIERA
LEAF
Not definitely dorsiventral. Spines arising as described below for Idria.

Epidermis sometimes including cells with mucilaginous inner walls.

Stomata present on both surfaces; ranunculaceous.

Axis
STEM (Fig. 38 F and H)
Very striking in general appearance owing to the reticulate pattern of corky
layers on the surface. Young stems furrowed and provided with a narrow
 H
FIG. 38. PORTULACACEAE, A and C; CARYOPHYLLACEAE, B, E, and G;
TAMAWCACEAE, D FOUQUIERACEAE, F and H
;

A, Portulaca oleracea Linn. Stem X 23. B, Saponaria officinalis Linn. Stomata x 250. C, Montia
perfoliata (Don.)How. Stem X2j. D, Tamarixgallica Linn. Stem X 13. EtGypsophilapaniculatal^irm.
Stem x 8. F, Fouquiera splendens Engelm. Young stem X 10, G, Lychnis coronaria Des. Stem X 10.
H, Fouquiera splendens Engelm. Stem X 10.
Fig, B, stomata on lower surface of leaf. F, Showing furrows and ring of close vascular strands.
e.c. Slightly elongated cells with thickened walls. p.c. Layer of small somewhat prosenchymatous
cells, tu.t. Water-storage cells.
 FOUQUIERACEAE 161

layer of fibres with thick walls and narrow lumina immediately within the
epidermis except at the bases of the furrows. The fibrous layer, which is
described by Solereder as cork, becomes wider with increase in age owing to
secondary division of cells in the sub-epidermal region. Cork, consisting of
cells with thin walls and wide lumina, is subsequently formed
internally to the
fibres, the stem eventually becoming covered with bark consisting of paper-
like layers capable of burning with a smoky flame owing to the presence of

waxy material. Primary cortex, according to Scott (2069) consisting mainly

of parenchyma, but stated to contain assimilatory tissue of varying width;
also including collenchyma and groups of stone cells. A continuous network
of water-storage cells throughout the cortex is particularly evident in sections
mounted in chlor-zinc-iodide because the component cells remain unstained
in contrast to the adjacent starch-filled parenchyma. Vascular bundles
separated by wide primary rays in young stems. Phloem appearing in
transverse sections as irregular, triangular wedges including sieve tubes with
;

lateral sieve plates. Xylem exhibiting seasonal growth rings. Pith stated by
Scott (2069) to be horizontally septate. Abundant crystals, believed to con-
sist of silicates, recorded by Reiche (1908).

WOOD (Fig. 34 J-K)

Vessels very small (25-50 /z mean tangential diameter) or slightly larger;
solitary and in clusters (Fig. 34 K), and often with a tangential pattern or
irregular radial rows or groups (1886); about 60 per sq. mm.; semi-ring-
porous to distinctly ring-porous. Perforations typically simple, but Solereder
refers to occasional scalariforrn plates with few bars. Intervascular pitting
often elongated horizontally, varying from round and alternate to almost
scalariforrn in the smaller vessels some pits to ray cells similar to the inter-
;

vascular pitting, others larger and simple or with only narrow borders. Mean
member length 0*3 mm. (100). Parenchyma apotracheal, as numerous
scattered cells and short, uniseriate lines. Strands of 2-4 cells. Rays up to
6 (occasionally 8) cells wide often up to about i mm. high uniseriates rather
; ;

few; about 5 rays per mm.; uniseriate and multiseriate rays often composed
almost entirely of square to upright cells (Fig. 341). Fibres with small
bordered pits, mostly on the radial walls; walls thick and mostly gelatinous;
mean length 1*1 mm. (96).

ROOT
Bounded externally by laminated cork. Water-storage cells not seen by
Scott (2069) in the disintegrating cortex of old roots.
(The respective accounts of the anatomy of Fouquiera given by Reiche
(1908) and Scott (2069) differ in certain details.)

IDRIA
The following description is based mainly on Humphrey's (u 10) account.

LEAF
Leaves of wild specimens with palisade tissue on both sides and 2-4 layers
of spongy tissue at the centre. Palisade tissue sometimes confined to the
upper surface in greenhouse material. Stomata flush with the surface of the
4594 M
162 FOUQUIERACEAE
lamina. Veins embedded in the central spongy tissue. Petiole of a nearly
mature leaf containing a fairly thick, semicircular vascular strand, concave

upward as seen in transverse section. Spines, when first formed, attached to

the abaxial side of the lamina and extending down the branches for a short
distance; becoming separated from the lamina and petiole at the time of leaf
fall by an abscission layer, a small scar remaining to indicate where the
vascular strand passed from the cortex into the petiole.

Axis
MAIN TRUNK
Exterior bounded by cork consisting of many rows of yellowish, compressed
cells. Outer part of the cortex containing large groups of stone cells, the
latter forming an almost continuous ring round the stem inner part parenchy-
;

matous, with simple pits between the cells giving the appearance of a per-
forated Swiss cheese. Cortex also including water-storage cells. Phloem
appearing in transverse sections as slender or wedge-shaped groups, most of
the tissues becoming crushed, apparently owing to pressure against the
sclerenchymatous sheath in the cortex. Outer part of the xylem consisting
of a narrow, almost continuous cylinder, broken by narrow bands of radial
parenchyma extending into the phloem. Inner part of the xylem mainly
parenchymatous, containing numerous lacunae. Scattered vessels present in
the parenchyma in roughly concentric rows; probably pushed into these
positions by proliferation of the intervening parenchymatous cells. Vessels
of the secondary xylem with circular or slit-like pits in the radial and tangential
walls. Pith consisting of homogeneous parenchyma, the cells tending to be

vertically elongated.

BRANCH
The following regions recognizable in a typical branch 6-6 mm. in diameter,
(i) Cork
in the form of a thick layer, (ii) Cortex half as wide as the cork,
(iii)
A narrow phloem, (iv) Xylem, about as wide as the cork; not divided
into tracheal and parenchymatous portions as in the main stem, and consisting
of vessels, fibres, and parenchyma. Wood fibres with numerous small pits;
many of them septate. Vessels up to 38 p in diameter; minimum length of
the component members 280 \L pits in the lateral walls circular to slit-shaped.
;

Pith with radius about equal to that of the xylem, composed of rather thick-
walled parenchymatous cells with simple pits.

TAXONOMIC NOTES
There has been considerable disagreement concerning the taxonomic
position of the family. Fouquiera, in the Bentham and Hooker system, is
treated as a member of the Tamaricaceae, whilst in the Engler system
Fouquiera and Idria have been placed in a separate family close to the
Tamaricaceae. Reiche (1908), after reviewing the various taxonomic positions
which have been proposed for the family, could do no more than conclude
that it is a primitive member of the Sympetalae. Humphrey (mo) has also
reminded us that possible affinities with the Loasaceae and Polemoniaceae
have also been suggested.
The wood structure of Fouquiera indicates that there is no good reason for
 FO UQ UIERA CEAE 163

putting this genus in the same family asTamarix has highly

Tamarix.
specialized wood with short elements, stories, libriform fibres, and para-
tracheal parenchyma; Fouquiera is at a relatively low level of
specialization,
with fibre tracheids, &c. and apotracheal parenchyma. Hutchinson
(1113)
includes the Fouquieraceae and the Malesherbiaceae in the Tamaricales.

GENERA DESCRIBED
Fouquiera,* Idria.
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Humphrey mo, Hutchinson 1113, Reiche 1908, Scott, F. M. 2069.
(ii) On Wood Structure

Record 1843, 1851, Record and Hess 1886.

46. ELATINACEAE
SUMMARY
A
small, cosmopolitan family of herbs or shrublets, often aquatic, included
in the 2 genera Bergia and Elatine. A
secretion, brownish when dry, and
probably resinous in nature, is widely distributed in the tissues, or deposited
in a granular form on the surface of the stem.

LEAF
Hairs
entirely absent from Elatine and 10 species of Bergia examined by
Pettier (1735). Other species of Bergia are provided with uni- or bicellular,
and less frequently with uniseriate trichomes. Glandular hairs, consisting of
a multicellular short column bearing a rounded head, also occur in Bergia.
Six species examined by Pottier possessed only glandular hairs, but others
had mixtures in various proportions of glandular and non-glandular hairs.
Whole surface of the plant covered with conical hairs consisting of i to several
cells in Bergia arenarioides (Camb.) Fenzl. and Bergia suffruticosa (Delile)
Fenzl. according to Niedenzu (1595). Epidermis consisting of cells with
sinuous anticlinal walls, those on the upper generally less undulating than
those on the lower side, but varying considerably in this respect in different
species of Bergia. Epidermal cells often very variable in size, especially in
Bergia, the larger ones serving for water storage, and sometimes appearing
as glistening spots on the surface when viewed with a lens. Particularly large
cells, deeply stained by alum carmine, present in the epidermis of Elatine

californica Gray. Stomata present on both surfaces, variable in size, appear-

ing at first sight to be ranunculaceous, but careful examination shows them
nearly always to be accompanied by 2 cells parallel to the pore and with 2
more at right angles to them. Mesophyll of Bergia spp. composed mainly
of small isodiametric or slightly elongated (palisade) cells. Palisade cells
irregularly distributed, their frequency varying in different species; sometimes
arranged in definite layers. Base of the leaf in Elatine containing a lacunar
164 ELATINACEAE
tissue, but with the mesophyll so much reduced or even lacking towards the
apex that the upper and lower epidermis are contiguous. Vascular bundles
of the veins generally surrounded by loose or compact parenchymatous cells
in Bergia, but canals present in this region in a few species; accompanied by
sclerenchyma only in a few species of Bergia. Vessels arranged in more or
*
less radial rows. Secreted material see Summary'.

Axis
STEM
Stem of Elatine often irregular in outline as seen in transverse section, owing
to the presence of elongated hair-like cells. Suberized cells sometimes present
on the outside of old stems of Bergia. Cortex frequently containing lacunae,
especially in aquatic species of both genera, but their size and distribution are
said to be of considerable specific diagnostic value. Endodermis clearly
defined, consisting of thin- walled, or more rarely, of thick- walled cells.
Xylem exhibiting growth rings in old stems of Bergia. Vessels mostly some-
what quadrangular and arranged in radial rows as seen in transverse sections ;

perforations simple; thickening spiral or annular in the primary xylem, but

with horizontally elliptical pits in older material. Pith lacking in nearly all
species of Elajine. Secretory cells with tanniniferous contents recorded by
Sabnis (1977) in the cortex or pith of Bergia. For other secreted material see
Nummary'. Secreted material said by Niedenzu (1595) to be particularly
abundant phloem. Cluster crystals of calcium oxalate sometimes occur
in the
in the region of the endodermis and in the pith cells of Bergia.
Three types of stem structure have been recognized by Pottier (1735) in
different groups of Bergia spp. (i) Plant glabrous ; cortical intercellular spaces
very large; fibres absent from the pericycle; pith cells elongated and some-
what resembling phloem as seen in longitudinal section, (ii) Cortical lacunae
small; pericyclic fibres present; endodermis well developed and provided with
casparian thickenings, (iii) Cortex consisting entirely of assimilatory tissue.

ROOT
Central vascular cylinder (with very variable diameter in different species
of Bergia, greatly reduced in Elatine), attached to the epidermis only by
filaments of cells, hence becoming easily separated from the surrounding
tissues when sections are cut. Endodermis, pericycle, and phloem not well
defined.
TAXONOMIC NOTES
The taxonomic position of the family seems to be somewhat uncertain
judging from the different positions to which it has been assigned by various
authors. By some it is regarded as having affinities with the Caryophyllaceae
and by others as being more closely related to the Frankeniaceae and Tamari-
caceae. As with other mainly herbaceous plants which are inhabitants of
damp localities or even aquatic, the anatomical structure is not very helpful as
an index of taxonomic affinity, since the anatomy of the plants is largely
correlated with the nature of the environment. The anatomical structure is
not very thoroughly known, and it may be that a more complete investigation
would throw further light on the subject.
 ELATINACEAE 165
GENERA DESCRIBED
Bergia, Elatine.
LITERATURE
On General Anatomy
D'Almeida 536, Niedenzu 1595, Pettier 1735, Sabnis 1977.

47. HYPERICACEAE
(Fie. 39 on p. 166; FIG. 43 on p. 176)

SUMMARY
(i) GENERAL
Herbs, shrubs, or small trees occurring mostly in the tropics, but some
species of Hypericum extend to North Temperate regions. The genus Hyperi-
cum itself includes species which exhibit a great diversity of habit forms
ranging from slender herbs to large shrubs or small trees. H. elodes Linn, is
aquatic. Some of the herbaceous species possess an underground rhizome.
The most constant anatomical character is the schizogenous secretory
cavities, which are present in the leaf of all genera, where they appear as
opaque or translucent dots. These cavities are filled with an oil containing a pig-
ment which has been shown by Siersch (2096) to consist of an anthocyanidin,
probably in part a rhamnose glucoside of pelargonidin. Secretory canals
are present in the phloem, and sometimes also in the primary cortex, pericycle
and pith of the axis, as well as in the petiole and veins of the leaf, A key to
the identification of species of Hypericum based on the nature and distribution
of the secretory cavities has been drawn up by Clos (433). The leaf is
generally dorsiventral, with stomata, each of which is surrounded by 2 or
more cells (see 'Leaf '), confined to the lower surface except in rare instances.
The axis, in transverse sections, exhibits closed rings of xylem and
phloem, traversed by narrow but well-defined medullary rays. In the
herbaceous species the xylem and phloem are relatively narrow, whilst there
is a greater development of pith. The cortex in Hypericum is usually narrow

and includes secretory cells. Secretory cells also occur in the phloem and
pith. The pericycle includes a ring of sclerenchymatous fibres which is

usually closed. Solitary and clustered crystals of calcium oxalate occur.

(ii) WOOD
(a) Hypericum
Hypericum differs from the other genera in having very small, spirally
thickened vessels, no parenchyma, wholly uniseriate rays, and septate fibres
(in some species).

(b) Genera other than Hypericum

Vessels medium-sized, often with an oblique pattern, perforations simple,
intervascular pitting alternate, pits to ray cells usually similar to the inter-
vascular pitting and sometimes unilaterally compound, members of medium
length. Parenchyma in continuous apotracheal bands and sometimes vasi-
centric in addition. Rays up to 2-4 cells wide, heterogeneous to homogeneous.
1 66 HYPER1CACEAE
Fibres usually with simple or indistinctly bordered pits, of medium length.
Vasicentric tracheids often present.

FIG. 39. GUTTIFERAE, A-B and G-H; QUIINACEAE, C and E-F;

HYPERICACEAE> D and I
A, Calophyllutn calaba Jacq. Petiole x 10. B, C. oz&zfaz Jacq. Young stem X 10. C,Quiinarhytidopus
Tul. Petiole X 15. D, Hypericum androsaemum Linn. Stem X2. E, Quiina guianensis Aubl. Petiole
X 15. F, Q. rhyttdopus Tul. Stem x 9. G, Garcinia mangostana Linn. Petiole x 8. H, G. mangostana
Linn. Stem X8. I, Hypericum elodes Linn. Young stem x 15.
c.e. Cutinized epidermis. a.s. Air space, i.s. and l.s. Intercellular spaces, s.c. Secretory canals.

LEAF
Usually dorsiventral. Hairs seldom numerous mostly simple, unicellular ;

or uniseriate; stellate types with uniseriate stalks recorded in certain species

of Harungana, Psorospermum and Vismia. t
Glandular hairs unknown.
 HYPERICACEAE 167
Cuticle frequently thick, especially in species with leathery leaves. Epidermis
papillose in certain species of Cratoxylon, Harungana, and Hypericum.
Stomata, confined to the lower surface in most instances; rubiaceous in the
Cratoxyleae and Vismieae, but commonly surrounded by 3 or more cells in
Hypericum. All types, ranging from a stoma accompanied by 2 cells parallel
to the pore to one surrounded by about 9 radially orientated cells, have been
recorded in a single leaf of Hypericum. Hypoderm present in certain species
of Harungana, Hypericum, Psorospermum^ and Vismia. Mesophyll usually
including a single, but occasionally 2, layers of palisade tissue. Petiole of all
investigated species exhibiting, in transverse sections, a single arc-shaped
*
vascular strand. For secretory cavities and canals see Summary*.
Clustered crystals of calcium oxalate present in Harungana, Psorospermum,
and Vismia. According to Briquet (272) leaves of certain species of Hypericum
from arid regions in Brazil have thickened margins consisting of enlarged
epidermal cells or collenchyma.

Axis
YOUNG STEM (Fig. 39 D and i)

Cork generally arising in the pericycle ; consisting of alternating layers of

unsuberized and uniformly sclerosed cells with fairly wide lumina in Psoro-
spermum sp. Cortex in woody and semi- woody species of Hypericum narrow,
consisting of rather loose, small-celled parenchyma when young, but becoming
more compact when older. Cortex of the aquatic H. elodes Linn. (Fig. 39 i)
broad, with abundant intercellular spaces, the structure in transverse sections
resembling a network partly composed of secretory idioblasts, the latter being
especially numerous at the corners of the meshes. The endodermis is well
defined in Hypericum elodes and, when the stem is sufficiently young, in
terrestrial species of thesame genus. Pericycle containing a sclerenchyma-
tous ring, often closed in certain species of Cratoxylon, Eliaea, Harungana,
Hypericum, Psorospermum, and Vismia, but sclerenchyma absent from the
pericycle in certain herbaceous species of Ascyrum and Hypericum according
to Solereder. Aring of fibres, accompanied on the inside by stone cells with
thick inner tangential walls, recorded by Solereder immediately beneath the
epidermis of Cratoxylon polyanthum Korth. Xylem and phloem forming
closed rings in Hypericum, but there is a greater development of pith in the
herbaceous than in the woody species. Vessels with simple perforations.
Cells of the narrow medullary rays filled with secreted material in Hypericum.
Secretory canals (see 'Summary') always present in the secondary phloem
(not always very conspicuous), but sometimes occurring also in the pith,
pericycle, and outer part of the primary cortex. Their exact position may be
of specific diagnostic value according to Clos (433), but the subject needs
reinvestigation. Secretory canals are present in the phloem even in species
like Hypericum elodes (Fig. 39 i) whose structure is specialized in relation to
a marshy environment. Secretory cells, presumably tanniniferous (stained
very darkly with haematoxylin) scattered in the cortex, phloem, and pith of
Hypericum.

WOOD (Fig. 43 D-G and j)

Vessels typically medium-sized (100-200 p mean tangential diameter),
168 HYPERICACEAE
but extremely small (less than 25 ju) in Hypericum] solitary and in radial pairs,
with a distinct oblique, zigzag pattern in Psorospermum and Vismia (Fig. 43 j)
and with a tendency to an oblique arrangement in some species of Cratoxylon',
sometimes with a tendency to a tangential pattern in Hypericum\ typically
with fewer than 5 vessels per sq. mm., but slightly more numerous in Vismia
p.p. and sometimes up to 200 or more in Hypericum] occasionally with a slight
tendency to be ring-porous in Hypericum. Perforation plates simple, slightly
oblique; Vestal (2329), however, notes scalariform or clustered perforation
plates in some species of Hypericum. Intervascular pitting alternate, of
medium size to rather small; pits to ray cells usually similar to the inter-
vascular pitting, but often elongated and with narrow borders in Harungana,
and unilaterally compound in some species of Cratoxylon and in Harungana.
Tyloses present in some species, solid deposits sometimes moderately
abundant. Mean member length 0*4-0*5 mm. Parenchyma in continuous
to broken apotracheal bands, 2-6 cells wide in Cratoxylon (most species),
Psorospermum and Vismia (Fig. 43 j) similar bands occur in Harungana, but
;

tend to link the vasicentric sheaths of tracheids and parenchyma (Fig. 43 F) ;

the bands widely spaced and possibly terminal in Cratoxylon arborescens

Blume. Parenchyma absent from Hypericum. Strands usually of 4 cells, but
commonly of 8 cells in some species of Vismia strands with septate crystalli-
;

ferous cells reported (1886) in Vismia. Rays up to 4 cells wide in Psorosper-

mum and Vismia p.p. and tending to be of two distinct sizes in the former, but
more commonly only up to 2 or 3 cells wide; all uniseriate in Hypericum] less
than i mm. high and commonly less than high in Hypericum', uni-
5 cells
seriates typically numerous and composed mainly of upright cells, but rather
few in some species of Cratoxylon, e.g. C. arborescens. Usually about 10 rays
per mm., but up to 24 in Hypericum. Typically heterogeneous (Kribs's
Type II A) and commonly with 4-8 marginal rows of upright cells, but with
only i marginal row or homogeneous in Cratoxylon] composed entirely of
square and upright cells in the material of Hypericum examined, but Vestal
(2329) describes the rays of this genus as falling into Kribs's types Hetero-
geneous II B and III. Silica reported in Cratoxylon (794). Fibres typically
with simple or indistinctly bordered pits that occur mostly on the radial walls,
but with distinctly bordered pits in Psorospermum. Vestal (2329) notes the
occurrence of some septate fibres in Hypericum, particularly in the Asiatic
species, and has described the formation of the septa in H. androsaemum Linn.
(2331). Walls thin in Harungana, moderately thick to thick in the other
genera. Silica reported (794) in Cratoxylon. Mean length i-o-i*4 mm.
Vasicentric tracheids present in Harungana, some species of Hypericum
(2329) and Vismia, and less abundantly in Cratoxylon] together with very
small vessels, and sometimes parenchyma, they form a matrix linking together
and surrounding groups of larger vessels (Fig. 43 F and j).

ROOT
Secretory canals present in the pericycle and secondary phloem of
Hypericum.
ANOMALOUS STRUCTURE
Bausch's (154) reference to included phloem in Hypericum appears to be
an error.
 HYPERICACEAE 169

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
The family is very homogeneous, and has many
characters in common with
the Guttiferae. In fact the Hypericaceae are treated as a tribe of the Gutti-
ferae in Engler's system. The family also has affinities with the Theaceae,
although secretory canals are lacking in the Theaceae.

(ii)
FROM WOOD STRUCTURE
The genera other than Hypericum
have characters in their wood anatomy,
such as a diagonal pattern of the vessels and the occurrence of vasicentric
tracheids that strongly suggest a close relationship with the Guttiferae-
Calophylloideae.
itself, however, has little in common with this tribe. Its septate
Hypericum
fibres resemble those of the Clusieae, but apart from this there is not much
positive evidence of relationship the wood of Hypericum is much more highly
;

specialized than that of the Clusieae.

Vestal (2329) regards the Hypericaceae and Guttiferae as obviously closely
related and agrees with Engler, Wettstein, and others in considering them as
parts of the same family. On the basis of the wood anatomy and embryo
structure Vestal supports Hochreutiner's combination of the two groups
through Psorospermum and the tribes Calophylloideae and Clusioideae of the
Guttiferae, and regards Hypericum and its near relatives as derived from the
Clusioideae, and the arborescent Hypericaceae from the Calophylloideae.

ECONOMIC USES
Species of Hypericum are commonly cultivated for ornamental purposes.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Ascyrum, Cratoxylon, Eliaea, Harungana, Hypericum,* Psorospermum,
Vismia.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Harungana, Hypericum (Psorospermum), Vismia.

LITERATURE
(i) On General Anatomy
Briquet 272, Clos 433, Siersch 2096, Starr 2188, Vestal 2329, 2331.

(ii) On Wood Structure

Bailey 100, Bausch 154, Cooper 461, Desch 574, Foxworthy 705, Giordano 786,
Gonggrijp 794, Janssonius 1154, Kribs 1283, Lecomte 1334, Record 1843, 1851, Record
and Hess 1886, Vestal 2329, 2331, Williams 2430.
 (170)

48. GUTTIFERAE
(Fic, 39 on p. 166; FIG, 40 on p. 170; FIG. 41 on p. 172; FIG. 42 on p. 174; FIG. 43 on p, 176;
FIG. 44 on p. 182 ;
FIG. 49 on p. 208)

SUMMARY
(i) GENERAL
A family of tropical trees and shrubs with rather uniform anatomical struc-
ture, and which are characterized especially by the presence in all species of
schizogenous secretory receptacles, usually in the form of canals, but
less frequently of cavities. The canals in the axis are most commonly situated
in the pith and primary cortex, but more rarely in the primary or secondary
phloem as well. When present in the leaf they may accompany the vascular

FIG. 40. GUTTIFERAE

Hairs of: A, Calophyllum retuswn Wall. B, C, bracteatum Thw. C, C. pseudotacamahaca
; ;

Tr. et PI. D, C. calaba Jacq. (buds) E, C. tomentosum Wight. After Vesque.

; ;

bundles or run independently of them in the mesophyll. They also occur in

the petiole. The canals are replaced by secretory cavities in the leaf of certain
species. The distribution of the canals and cavities is of some value for the
identification of genera and species, but De Cordemoy (469) has shown that
allowance must be made for variations within a species in response to environ-
mental and especially to soil conditions. Secretory cells, which are readily
stained with haematoxylin, are common in the parenchymatous tissues and
ray cells of young stems. Hairs (Fig. 40) are rare in the family, and where
present consist of simple unicellular or uniseriate trichomes, which are more
rarely branched or dichotomously divided. Stellate hairs are also known from
a few genera. Stomata are rubiaceous. The xylem and phloem in young
stems appear, in transverse sections, in the form of continuous rings traversed
by narrow rays. Crystals of calcium oxalate are usually solitary or clustered,
or more rarely in the form of small prisms. They occur fairly widely in the
parenchymatous tissues. Anomalous structure recorded in Endodesmia.
 GUTTIFERAE 171
(ii)
WOOD
Group A (Clusieae)
Vessels often in small multiples, perforation plates simple or simple and
scalariform, intervascular pitting and pits to ray cells scalariform, members
moderately long to very long. Parenchyma paratracheal, sometimes accom-
panied by diffuse. Rays 4-9 cells wide, high, markedly heterogeneous.
Fibres commonly septate, with simple or slightly bordered pits, moderately
long. Without vasicentric tracheids or intercellular canals in the rays.
Group B (Calophylloideae)
Vessels exclusively solitary, often with a diagonal pattern, perforations
simple, intervascular pitting alternate, pits to ray cells commonly elongated,
members of medium length. Parenchyma apotracheal, in broad bands or
diffuse. Rays narrow (1-3 cells wide) and low, heterogeneous. Fibres with
bordered or simple pits, of medium length. Vasicentric tracheids nearly
always present. Intercellular canals occasionally present in the rays.
The genera of the Morononoideae and Garcinieae are intermediate, but
tend to be most like Group B, though lacking the characteristic vessel pattern
and vasicentric tracheids. In the Kielmeyeroideae some genera resemble
Group A, others Group B.
LEAF
Dorsiventral in most species; centric or sub-centric in certain species of
Calophyllum, Clusia, Garcinia, Kielmeyera, Moronobea, Pentadesma, Platonia,
Rheedia, Symphonia. Hairs infrequent or even absent; for types see 'Sum-
mary*. Stellate but unicellular trichomes recorded in Caraipa (Fig. 49 B) and
Marila. Cork warts recorded in Clusia sp. and Xanthochymus sp. The
height of the cells of the epidermis, the thickness of the cuticle and degree
of undulation of the anticlinal walls of the epidermal cells are somewhat
variable and valuable only for the identification of species. The thickness of the
cuticle is often considerable. Epidermal cells with marginal pits in the lateral
walls recorded in certain species of Garcinia, Kayea, Mammea, Mesua, Penta-
desma, Platonia, Poedloneuron, Rheedia, Tovomita. Lower epidermis papillose
in certain species of Garcinia. Some of the epidermal cells horizontally
divided in Calophyllum inophyllum Linn. Stomata (Fig. 41 c) confined to
the lower surface, except in certain species of Garcinia', rubiaceous. Sub-
sidiary cells filled with a brown homogeneous substance in Calophyllum
inophyllum; papillose in certain species of Chrysochlamys and Garcinia.
Hypoderm of one to several layers common on the upper side of the leaf,
especially in Calophyllum, Chrysochlamys, Clusia, Garcinia, Havetia, Kiel-
meyera (in K. petiolaris Mart., but not all other species of K. hypoderm cells
;

thick walled andpitted in this genus), Montrouziera, Oedematopus (spongy,

Fig. 41 B), Pilosperma, Platonia, Quapoia, Tovomita. Mesophyll tending
to be sclerotic in several genera. Palisade tissue with reticulately thickened
walls, almost prosenchymatous in shape in Clusia rosea Linn. (Fig. 41 A).
Storage tracheids recorded by Solereder in the leaf tissue of Endodesmia.
Vascular bundles of the smaller veins accompanied or surrounded by
sclerenchyma, except in a few species of Garcinia; embedded in the mesophyll
in a majority of the genera, but vertically transcurrent in Calophyllum,
Caraipa, Kayea, Kielmeyera, Mammea, Marila, Poedloneuron. Petiole
172 GUTTIFERAE
examined in only a few species, but in these exhibiting, in transverse sections,
six more
or less distinct types of vascular structure, (i) With a single, arc-
shaped vascular strand open on the adaxial side and not accompanied by
sclerenchyma in Calophyllum calaba Jacq. (Fig. 39 A), (ii) As (i) but ends
of the arc-shaped strand incurved and nearly meeting to form a closed tube
in species of Clusia, Garcinia, Hdploclathra kiantha Benth. (iii) Main vas-
cular strand in the form of a sinuous but completely closed cylinder in
Rheedia laterifolia Linn, (iv) Main vascular strand forming an almost closed
tube surrounding two collateral bundles in Mammea sp. (v) As (iv) but main
strand forming a completely closed tube surrounding a few collateral bundles

FIG. 41. GVTTIFERAE

A, Group of palisade cells from the mesophyll of Clusia rosea L. in a transverse section. B, Spongy,
hypoderm from the leaf of Oedematopus obovatus Tr. et PL C, Crystal-containing epidermis of the leaf
of Garcinia punctata Oliv, D, Transverse section through the leaf of Calophyllum buodfolium Vesque,
with a resin-canal accompanied by tracheids and sclerenchymatous fibres. A, by Solereder, the
remainder after Vesque.

in Garcinia mangostana Linn. (Fig. 39 G), Haploclathra paniculata Benth.,

and Mahurea palustris Aubl. or with a closed ring surrounding a solitary
;

strand in Caraipa laxiflora Benth. and Kielmeyera petiolaris Mart, (vi) Main
strand in the form of two superimposed U's in Marila racemosa Swartz.
Crystals of calcium oxalate usually present in the mesophyll in the form
of clusters. Solitary as well as clustered crystals present in Calophyllum,
Garcinia (Fig. 41 c), Kayea, Rheedia. Crystals exclusively solitary in Mammea,
Mesua, Pentadesma, Poeciloneuron. Small, usually prismatic crystals occur in
many species of Garcinia. Secretory canals or cavities universally present,
sometimes appearing as transparent lines or dots when examined with a lens, or
with the naked eye. Canals either following the vascular bundles (Fig. 44 E)
or running independently of them in the mesophyll; in the latter case some-
times (Calophyllum) accompanied by peculiar bundles of tracheids and by
sclerenchyma (Fig. 41 D). They also pass through the 'cortical' and 'medul-
lary' regions of the petiole. The nature, size, and distribution of the secretory
 GUTTIFERAE *73
canals or cavities are valuable for the identification of species, after
making
due allowance for variations in response to environmental conditions.

Axis
YOUNG STEM (Fig. 39 B and H)
Epidermis with
a thick cuticle or sclerosed in certain species of Clusia,
Garcinia, Mammea, and probably in other genera as well. Cork arising
immediately below the epidermis in all examined species of Catophyllum,
Caraipa, Clusia, Endodesmia, Haploclathra, Havetiopsis, Kielmeyera, Mahurea,
Mesua, Oedematopus, Pilosperma, Platonia, Tovomita. Outer tangential and
radial walls of component cells strongly thickened in Endodesmia. Cortex
commonly containing sclerosed cells; a ring of stone cells present in Mesua
ferrea Linn. Sclerosed cells not recorded in the cortex of Marila racemdsa
Swartz. An endodermis consisting of large, starch-containing cells, recorded
in Kielmeyera. Pericycle recorded as having or observed to include isolated
bundles of fibres in certain species of Calophyllum, Garcinia, Mammea,
Rheedia, Symphonia, Tsimatimia] but many genera and species, including
those of the Kielmeyeroideae, are provided with a composite, continuous
ring of sclerenchyma. Both types probably occur in stems of different ages
belonging to the same species. Xylem and phloem appearing, in transverse
sections, in the form of continuous rings, traversed by narrow rays in all
species which have been examined, notably in Calophyllum, Clusia, Garcinia,
Mammea as represented in the Kew slide collection. Phloem including fibres
in Caraipa and Marila. Vessels in young twigs seldom more than 50 p in
diameter, solitary or in short radial groups; perforations simple. Wood fibres
arranged in radial rows in Marila] very thick walled in Mahurea. Pith, apart
from canals, &c., composed of cellulose cells in some species but somewhat
lignified in others. Secretory canals universally present in the primary
cortex and pith, and often in the phloem as well, although the canals in the
phloem are usually smaller and less conspicuous than those elsewhere. Canals
stated to be absent from the phloem in Allanblackia, Clusia, Havetiopsis,
Oedematopus, Pentadesma, Pilosperma, Tovomita] present in the secondary
phloem in Mammea] those in the perimedullary region of Kielmeyera,
Mahurea, Marila, and in the pericycle of Mahure apalustris Aubl. specially
large. Secretory canals sometimes rendered obscure owing to
the protrusion
of the epithelial cells into the cavity in a manner resembling tyloses in vessels.
Secretory cells, readily stained with haematoxylin, common in the paren-
chymatous tissues, notably in species of Calophyllum, Clusia, Rheedia, and
probably in other genera as well.

WOOD
(a) Group A. Clusia Type (Fig. 42 A-D)
Clusieae: Chrysochlamys, Clusia, Havetiopsis, Tovomita, and Tovomitopsis.
Vessels usually medium-sized (100-200 ^ mean tangential diameter), but
moderately small (50-100 JJL) in Tovomita and Tovomitopsis] solitary and in
small multiples, without any pattern; usually 10-14 per sq. mm. Perforation
plates all simple or simple and scalariform; some scalariforrn plates, usually
with numerous, fine bars, present in at least some species of Chrysochlamys,
Clusia, Tovomita, and Tovomitopsis. Intervascular pitting scalariform. Pits
174 GUTTIFERAE
to ray cells almost all elongated horizontally. Solid deposits sometimes
abundant. Mean length 0*9-1*2 mm. Parenchyma typically paratracheal,
vasicentric; tending to be aliform in Tovomita and with some scattered cells;
sometimes scanty, e.g. in Chrysochlamys and Tovomitopsis (Fig. 42 A). Gum-
like deposits abundant; chambered crystals rather rare; druses, contained in

FIG. 42. GUTTIFERAE (Group A and miscellaneous)

A, Tovomitopsis multiflora Standl. B, Clusia cooperi Standley. C, Tovomitopsis multiflora Stand!.
D, Clusia odorata Seem, E, Symphonia globulifera Linn. F, Garcima mannii OHv. G, Rheedia edulis
Planch, et Triana. H, Garcinia rostrata Hassk.
i.e. Intercellular canal.

dilated cells, present in Clusia. Rays usually of two distinct widths, the larger
up to 4-9 (mostly 4-6) cells wide and typically more than i mm. high (often
considerably more than 2 mm.); uniseriates typically numerous and composed
of high upright cells, but very rare in Havetiopsis\ 5-15 rays per mm.;
markedly heterogeneous (Kribs's Type I), with many marginal rows of high
upright cells, the cells sometimes all square or upright, e.g. in Clusia and
 GUTTIFERAE 175
Tovomitopsis (Fig. 42 B and c), and the upright cells in the latter very high in
proportion to their width. Typically with abundant gum-like deposits.
Fibres septate in Chrysochlamys, Clusia, Havetiopsis, and Tovomitopsis. Pits
simple or with minute borders, and more numerous on the radial than on the
tangential walls. Walls thick to extremely thick. Mean length 1-4-1-9 mm.
Mahurea and Marila, of the Kielmeyeroideae, appear to have closer
affinities with this than with the Calophyllum group. Caraipa and
Haplo-
clathera, on the other hand, resemble the latter type and are described with it.
Mania is outstanding in having septate fibres distributed like diffuse and
banded parenchyma among fibre-tracheids with numerous bordered pits.
The main points of interest or of difference between Mahurea and Mania
and the genera of Group A are given below.
Vessels smaller and more numerous in Marila', rare scalariform plates
sometimes present in Marila intervascular pitting alternate pits to ray cells
; ;

small and circular; mean member length in Marila 0-8 mm. Parenchyma
absent from Mahurea and rare in Marila. Rays seldom much more than
i mm. high. Fibres septate in Mahurea and with some septate fibres in
Marila, the other fibres in Marila with numerous, distinctly bordered pits on
both radial and tangential walls mean length in Marila i 5 mm.
;

(b) Group B. Calophyllum Type (Fig. 43 H-I, K-O)

Calophylloideae Calophyllum, Kayea, Mammea, Mesua, and Poeciloneuron.
:

Vessels usually medium-sized (100-200 fi, mean tangential diameter) and

sometimes slightly larger rather smaller in Kayea almost exclusively solitary
; ; ;

with a marked oblique pattern in Calophyllum and Mesua, and sometimes in

Mammea africana Sabine (Fig. 43 o); usually about 5 per sq. mm., but more
numerous (10-50) in Kayea. Perforations typically simple and slightly oblique,
but Record and Hess (1886) note a few scalariform plates in Mammea.
Intervascular pitting rare, alternate; pits to tracheids moderately large; pits
to ray cellscommonly elongated and simple, with the long axes horizontal or
axial, and sometimes unilaterally compound; Record (1793) has reported
'cribriform membranes* in Calophyllum, but, according to Bailey (78), these
are not true vestured pits. Tyloses sometimes abundant, sclerosed in Mesua;
solid deposits sometimes present. Mean length 0-4-1-0 mm. Parenchyma
in broad, often widely spaced apotracheal bands in Calophyllum, Kayea, and
Mesua (Fig. 43 K) the bands sometimes locally discontinuous and showing
;

some tendency to association with the vessels; diffuse in Mammea (Fig. 43 o);
predominantly paratracheal in Poeciloneuron, sheathing the vessels on the
abaxial sides and extending as long, narrow wings (unilaterally paratracheal).
Chambered crystalliferous cells usually present and often abundant. Strands
typically of 8 cells. Rays usually up to 2-3 cells wide, but exclusively uni-
seriate in Mesua and most species of Calophyllum sometimes 4-5 cells wide
;

in Mammea', less than i mm. high; uniseriates numerous, usually composed

of both upright and procumbent cells; 8-17 rays per mm.; heterogeneous
(Kribs's Type II B), with 1-4 (occasionally more) marginal rows of square to
upright cells; commonly with more than 4 marginal rows in Mammea africana
and the uniseriates composed of upright cells only. Gum-like deposits
abundant. Fibres with numerous bordered pits on both radial and tangential
walls in Kayea, Mammea, and Poeciloneuron , the borders conspicuous in
 D

FIG. 43. QUIINACEAE, A-C; HYPERICACEAE, D-G and J;

GUTTIFERAE (Group B), H~I and K-O
A, Touroulia guianensis Aubl. B, Lacunaria jenmanii (Oliv.) Ducke. C, Qtdina panamensis Standl.
D, Visnria latifotta Chois. E, Hypericum acerosum H. B. et K. F, Harungana madagascariensis Lam.
G, Cratoxylon arborescent BK H, Mesua ferrea Linn. I, Calophyllum inophyllum Linn. J, Vismia
leonensis Hook. f. K, Calophyllum wallichianum Planch. L Kayea grandis King. M, Mammea africana
?
Sabine. N, Poeciloneuron iridicum Bedd. O, Mammea africana Sabine.
i.e. Intercellular canal.
 GUTTIFERAE 177
Mammea", with small bordered or simple pits on the radial walls in Colo-
phyllum and Mesua. Walls thick to extremely thick. Mean length i -0-1 -4 mm.
Vasicentric tracheids with conspicuous bordered pits observed in all the
genera; Record and Hess (1886), however, have not recorded tracheids in the
New World species of Mammea. Intercellular canals present in the rays of
Mammea (Fig. 43 M).
The genera of the Moronoboideae Montrouziera, Moronobea, Pentadesma,
Platonia, and Symphonia and of the Garcinieae Allanblackia, Garcinia,
Pentaphalangium, and Rheedia on the whole show greater affinities with this
than with the Clusia group (A). The main points of interest or of difference
from Group B are given below.
Vessels similar except that short radial multiples are moderately common
and there is no diagonal pattern, except for a slight tendency in Pentadesma.
Intervascular pitting medium-sized to large and often with coalescent aper-
tures in the Garcinieae; pits to ray cells not so markedly elongated and all
circular in the Garcinieae (except Garcinia). Mean member lengths, Gardnia
0-3-0-6 mm., Symphonia 0-6 mm. Parenchyma similar but the bands more
numerous and less regular (Fig. 42 H) often more closely associated with the
;

vessels and intermediate between confluent and metatracheal (Fig. 42 E).

Chambered crystals common in most of the genera; druses sometimes present
in Garcinia, e.g. G. globulosa Ridl., and Pentaphyllum. Rays wider (up to
4-6 cells) and distinctly higher; uniseriates few to very few, composed of
1-3 upright cells, procumbent cells only or mixed procumbent and upright
cells; rays varying from distinctly heterogeneous (Kribs's Type II B) with
1-4 marginal rows of upright cells, e.g. in Garcinia mannii Oliv., to almost
homogeneous (Kribs's Type I), e.g. in Allanblackia and Garcinia holtumi
Ridl. Silica reported (794) in Garcinia. Fibres with simple pits, more com-
mon on the radial than on the tangential walls. Mean lengths, Garcinia
1-3-2-0 mm., Symphonia 1-7 mm. Vasicentric tracheids absent. Inter-
cellular canals present in the rays of Rheedia (Fig. 42 G).
Caraipa and Haploclathra of the Kielmeyeroideae appear from descriptions
to be closely related to this group ;
both have a tendency to oblique lines of
vessels, almost exclusively uniseriate rays and parenchyma that is described
(1886) as unilaterally paratracheal with short to long extensions; Caraipa has
solitary vessels, fibres with distinctly bordered pits and Vasicentric tracheids.
Interxylary (included) phloem see below.
ANOMALOUS STRUCTURE
Anomalous structure of the foraminate type (corpus lignosum foramina-
latum) noted by Pfeiffer (1712) in Endodesmia.
ROOT
Secretory canals, similar to those of the stem, present in the cortex alone
phloem in certain species of Calophyllum, Clusia, Gar-
or in the cortex and
dnia, Mammea, Rheedia, Xanthochymus.
TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
The anatomy of the leaf and young stem is fairly uniform throughou^:
the
Guttiferae. In spite of the fact that some botanists regard the Kielmeyeroideae
4594 N
178 GUTTIFERAE
as rather distinct from the remainder of the family on the basis of their
external morphological characters, there does not appear to be any corre-
sponding demarcation in the structure of the leaf and shoot so far as these
have been investigated. Similarly the Hypericaceae have many points in
common with the Guttiferae and it seems to be largely a matter of opinion
whether the Hypericaceae should be treated as a distinct family or as a tribe
of the Guttiferae. The Theaceae are distinguished from the Guttiferae in
lacking secretory canals, but it is interesting to note that canals occur in the
Dipterocarpaceae.

(ii) FROM WOOD STRUCTURE

Twoextreme groups can be distinguished, the Clusieae and the Calo-
phylloideae, the former having distinctly more primitive vessel and ray
characters. These two groups are more sharply distinguished than are many
families.
The Garcinieae and Moronoboideae resemble each other in most characters
and have many features in common with the Calophylloideae, though lacking
the characteristic vessel pattern and vasicentric tracheids of this group.
Though they have been placed with the Calophylloideae, it should be borne
in mind that they also have characters in common with the Clusieae and that
at least some of the difference might be accounted for as the result of the

higher level of specialization in the Garcinieae and Moronoboideae.

The Kielmeyeroideae appears to be a mixed group. Mahurea and Marila
have many features in common with the Clusieae, whereas Caraipa and
Haploclathera closely resemble the Calophylloideae.
Mammea differs from all the other genera of the Calophylloideae in having
diffuse parenchyma.
ECONOMIC USES
The timbers are not generally of great commercial importance, though some
are very well known locally. The most important genus is Calophyllum, the
woods of which are characteristically strong and durable. The best-known
species are C. brasiliense Camb., which is widely used in South America for
general construction, shipbuilding, furniture, and other purposes, C. calaba
Jacq., the Santa Maria of Central America, C. inophyllum Linn., the Palo
Maria of the Philippines and Poon of India, and C. tomentosum Wight, Poon
or Poonspar from India, which is used for masts and spars and for a variety
of other purposes. The timber of Mesua ferrea Linn, is very strong and
durable and has a high reputation in India for railway sleepers, bridging, and
heavy construction. General utility timbers are provided by some of the other
genera, such as Kayea and Symphonia.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Allanblackia, Calophyllum,* Caraipa, Chrysochlamys, Clusia,* Garcinia,*
Haploclathra, Havetia, Havetiopsis, Kayea, Kielmeyera, Mahurea, Mammea,*
Marila, Mesua, Montrouziera, Moronobea, Oedematopus, Pentadesma,
Pilosperma, Platonia, Poeciloneuron, Quapoya, Rheedia,* Symphonia, Tovo-
mita, Tsimatimia, Xanthochymus.
* Kew slide
Represented in the collection.
 GUTTIFERAE 179

(ii) FOR WOOD STRUCTURE

Allanblackia, Calophyllum, (Caraipa), (Chrysochlamys), Clusia, Garcinia,
(Haploclathra), Havetiopsis, Kayea, Mahurea, Mammea, Marila, Mesua,
(Montrouziera), Moronobea, Pentadesma, (Pentaphalangium), Platonia,
Poeciloneuron, Rheedia, Symphonia, Tovomita, Tovomitopsis.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Brandza 262, Buttrick 325, De Cordemoy 469, Engler 634, Kienholz
1236, Vestal 2329.

(ii) On Wood Structure

Beekman 167, Benoist 169, den Berger 180, Besson 186, Burger-
Bailey 78, Bausch 154,
stein 310,Chalk 364, Cooper 461, Dadswell 525, Dadswell and Record 533, Demougeot
563, Desch 574, Foxworthy 705, Garratt 748, Gonggrijp 794, Greene 809, Janssonius
1147, 1154, Jentsch 1171, 1178, Jones 1191, Kanehira 1206, 1209, Kribs 1283, Lecomte
1334, M&iiaud 1491, Pearson and Brown 1679, Pereira 1687, Pfeiffer, H. 1712, PfeifTer
J. Ph. 1713, Record 1793, 1801, 1843, 1851, Record and Hess 1886, Record and Mell
1894, Stone 2202, 2206, 2207, Tang 2231, Torres 2269, Vestal 2329, Webber 2377.

49. QUIINACEAE
(FiG. 39 on p. 166; FIG. 43 on p. 176)

SUMMARY
(i) GENERAL ANATOMY
A small family of trees,
shrubs, and climbers from tropical America. The
family is
usually regarded as having affinities with the Guttiferae, but differs
from them in lacking schizogenous canals with resinous contents, and in
possessing lysigenous intercellular spaces filled with mucilage, notably
enclosed within the vascular strand of the midrib and petiole. The venation
of the leaf has a feathery appearance in herbarium specimens, and this is
caused by the sheath of very thick- walled fibres by which the vascular bundles
are surrounded, some of the fibres extending into the mesophyll or lying
immediately below the epidermis parallel with the surface of the leaf.

(ii) WOOD
solitary, occasionally with a slight radial or oblique pattern,
Vessels mostly
perforations simple, intervascular pitting alternate and very small, pits to ray
cells similar, members moderately to very long. Parenchyma apotracheal,

diffuse; reported to be sometimes predominantly paratracheal; sometimes

septate. Rays up to 2-4, or rarely 5-6, cells wide, high, markedly hetero-
geneous. Fibres with distinctly bordered pits, moderately long. Vasicentric
tracheids occasionally present.

LEAF
Dorsiventral. Hairs infrequent. Cells of the upper epidermis polygonal,
straight sided, but with thickenings of the cell wall projecting into the lumen.
Anticlinal walls of the cells of the lower epidermis rather unevenly thickened.
Stomata confined to the lower surface, up to about 20 p, in longitudinal
diameter (surface view); rubiaceous. Mesophyll composed of 2 layers of
i8o QUIINACEAE
palisade cells in Q. rhytidopus TuL, but of only one layer in Q. guianensis
Aubl., together with a broader layer of spongy tissue consisting of small
cubical cells in both species. All vascular bundles of the veins, especially the
smaller ones, surrounded by sheaths of very thick- walled fibres with branches
from the latter extending into the mesophyll and sometimes lying beneath
the epidermis parallel with the surface of the leaf. Midrib with an annular
vascular strand, flattened or slightly concave towards the adaxial surface,
enclosing a collateral, medullary bundle. Petiole of Q. rhytidopus (Fig. 39 c),
in transverse sections through the distal end, exhibiting an interrupted ring
of collateral bundles accompanied by 2 additional strands in the cortical
region. There is a large secretory cavity enclosed within the main vascular
strand. Petiolar structure similar in Q. guianensis (Fig. 39 E) but with an
accessory arc of medullary vascular strands as well as i large and 2 small
secretory cavities, the larger one being on the concave side and the 2 smaller
ones in the convex side of the arc of medullary strands. Cluster crystals
present in the petiole and mesophyll of both species. Secretory cells with
amorphous contents observed in Q. rhytidopus. For secretory cavities see
'Petiole* above.

Axis
YOUNG STEM (Fig. 39 F)
Cortex containing occasional groups of thick-walled, pitted stone cells.
Pericycle including groups of thick-walled fibres. Phloem and xylem in
the form of closed cylinders, traversed by uniseriate medullary rays. Small
groups of thick-walled fibres scattered throughout the phloem. Vessels
solitary and in radial groups, up to about 50 ^ in radial diameter; perforations
simple, oblique. Groundwork of the wood composed of radial rows of very
thick- walled fibres, the inner part of the walls being clearly demarcated from
the outer portion. Pith composed of pitted, parenchymatous cells with
moderately thick walls. Secretory cells with amorphous contents scattered
throughout the parenchymatous tissues; more numerous in Q. guianensis
Aubl. than in Q. rhytidopus Tul. Cluster crystals also present; especially
abundant in the cortex of Q. guianensis.

WOOD (Fig. 43 A-C)

Vessels moderately small (mean tangential diameter 50-100 /-i); mostly
solitary, but with a few radial pairs; sometimes with a tendency towards a
radial or oblique pattern, e.g. in Touroulia guianensis Aubl. about 20 per sq.
;

mm. Perforations simple and moderately oblique. Intervascular pitting

alternate, very small to minute; sometimes with coalescent apertures (1886);
Solid deposits common.
pits to ray cells similar to the intervascular pitting.
Mean length 0-9-1-3 mm. Parenchyma apotracheal in all the material
examined, as isolated cells or groups of 2 or 3 scattered among the fibres
(diffuse), but Record and Hess (1886) describe the parenchyma as 'para-
tracheal, sometimes confluent; also diffuse .'.
. Strands usually of 2-4 cells,
.

the individual cells sometimes containing single septa; chambered solitary

crystalsnumerous in Touroulia guianensis and rare septate parenchyma con-
taining druses reported (2329) in Quiina cruegeriana Griseb. Rays up to
2-4 (occasionally 5-6) cells wide and more than i mm. high; uniseriates
 QUIINACEAE 181

numerous and composed entirely of upright cells; 17-20 rays per mm.;
markedly heterogeneous (Kribs's Type I), commonly with 8 or more marginal
rows of upright cells, which often link together 2 or more multiseriate parts.
Deposits of gum abundant. Fibres with distinctly bordered pits that are
equally numerous on both radial and tangential walls; cells in contact with
the vessels with very numerous small pits in Lacunaria, but not distinctly
shorter than the other fibres. Walls very thick. Mean length i -6-2-1 mm.
Vasicentric tracheids occasionally present in Quiina and Touroulia.

TAXONOMIC NOTES
The
characters recorded in the 'Summary' seem to provide sufficient
justification for separating the Quiinaceae from the Guttiferae. Vestal (2329)
considers the wood structure of this family to be at a similar level of develop-
ment to that of the Guttiferae, but as taking its origin from the Theaceae.
The length of the vessel members and the primitive type of ray suggest that
it might be regarded as intermediate in this respect between the Calophyllum

and Clusia types of the Guttiferae (see p. 173).

Although in type of parenchyma the Quiinaceae resembles the Theaceae
rather than the Guttiferae, and in ray type the Theaceae and the Clusia group
of Guttiferae rather than the Calophyllum group, yet the occasional presence
of vasicentric tracheids and of a loose oblique or radial vessel pattern suggest
affinity with the Calophyllum group of the Guttiferae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
The part dealing with general anatomy is based on a study of specimens of
Quiina guianensis AubL* and Q. rhytidopus Tul.* in the Kew Herbarium.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Lacunaria, Quiina, Touroulia.

LITERATURE
(i) On General Anatomy
Engler 634.

(ii) On Wood Structure

Record 1843, z^S 1 * Record and Hess 1886, Vestal

50. THEACEAE
(Fic. 44 on p. 182; FIG. 45 on p. 186; FIG. 46 on p. 192)

SUMMARY
(i) GENERAL
A family of trees and shrubs, often with leathery leaves, which occurs in
Tropical Asia and temperate regions in the Far East. Hairs infrequent, but
where present they are generally unicellular, acuminate? with thick walls.
Fasciculate and tufted hairs with unicellular rays have also been recorded.
i8z THEACE AE
The leaf is generally dorsiventral with 1-3 layers of palisade tissue. The
epidermis consists partly or wholly of mucilaginous cells in certain species.
Stomata are confined to the lower surface in many genera surrounded by
;

2-5 narrow, subsidiary cells, more or less distinctly differentiated from the

FIG. 44. SAURAUIACEAE, A; THEACEAE. B-D; GUTTIFERAE. E

A, Transverse section through part of the leaf (upper side) of Saurauja nepaulensts DC, with arm-
palisade-parenchyma. B, Part of a transverse section of the leaf of Frextera undulata Swartz, with a
mucilaginous epidermal cell and with rod-shaped idioblasts. C-D, Idioblasts of Cleyera grandiflora
Hook. fil. et Thorns. ; C, in a transverse section of the leaf; D, in a superficial section of the upper side
of the leaf. E, Secretory cavity in a mesh between the veins of Caraipa fasciculata Camb. : vessels
extend from the vascular bundles of the veins to the secretory cavity. By Solereder.

remaining epidermal cells. In other cases the stomata are ranunculaceous.

Sclerenchymatous idioblasts in the parenchymatous tissues of the leaf as well
as in the cortexand pith of the axis are especially characteristic of the family.
Variations in their size, shape, and frequency are helpful in distinguishing
genera and species. (For details see 'Leaf'.) The petiole in most genera is
characterized, in transverse sections, by a solitary, crescentic vascular strand,
 THE ACE AE 183
but in some instances small lateral bundles are present as well. Prismatic and
clustered crystals are common in both leaf and axis. The family may be
divided into two distinct groups so far as the seat of origin of the cork in
young stems is concerned. In one of these its origin is superficial, most
frequently sub-epidermal, whilst in the other it arises in the pericycle. Where
the cork arises superficially the cortex is relatively broad and durable, the
outer part being collenchymatous but in genera with a deeply seated pheilo-
;

gen it is often narrower, and soon cut off by the formation of cork in the peri-
cycle. The endodermis is usually ill defined. The pericycle includes
heterogeneous sclerenchyma, but becomes entirely sclerified in most of the
genera. The phloem and xylem appear, in transverse sections, as closed
rings traversed by narrow rays, and surround the central pith which is often
heterogeneous and may contain idioblasts and crystals.

(ii) WOOD
Vessels typically small, predominantly or exclusively solitary, semi-ring-
porous in some species, commonly with spiral thickening, perforation plates
very oblique and scalariform, often with many bars, intervascular pitting
scalariform or opposite, pits to parenchyma similar, sometimes simple.
Members very to extremely long. Parenchyma predominantly apotracheal,
diffuse or in short lines, vasicentric present in addition in some genera. Rays
sometimes of 2 distinct widths, 1-8 cells wide, commonly only up to 2 or
3 cells, moderately short to high, heterogeneous with 3-10 marginal rows of
upright cells. Fibres with conspicuous bordered pits, of medium length to
very long.

LEAF
Generally dorsiventral, and frequently covered by thick cuticle. Hairs
infrequent, but when present unicellular, acuminate, thick walled in Camellia,
Eurya, Gordonia, Nabiasodendron, Stewartia, Ternstroemiopsis. Fasciculate
hairs with unicellular rays recorded in Gordonia and Ternstroemiopsis. Tanni-
niferous cork warts present on the lower surface in certain species of
Adinandra, Anneslea, Camellia, Eurya, Ternstroemia. Marginal hydathodes,
consisting of cells elongated at right angles to the leaf surface, recorded by
Lepeschkin (1361) in the leaf teeth of Camellia japonica Linn. Epidermis
including mucilaginous cells in certain species of Archytaea, Camellia,
Cleyera, Eurya, Freziera (Fig. 446), Gordonia, Laplacea, Nabiasodendron.
Stomata almost or completely confined to the lower surface, surrounded by
2-5, narrow subsidiary cells in certain species of Adinandra, Anneslea,
Camellia, Cleyera, Freziera, Ternstroemia, Visnea; rubiaceous in Archytaea',
ranunculaceous in some genera. Guard and subsidiary cells lignified in
Camellia as shown by treatment with phloroglucin and hydrochloric acid
according to Heilbronn (932). Hypoderm present on the upper side of the
leaf in certain species of Archytaea. Mesophyll. Palisade tissue consisting
of 1-3 layers, the number of layers being of some value for the identification
of genera and species. Spongy parenchyma occupies half to two-thirds of
the mesophyll. Sclerenchymatous idioblasts of various types, sometimes
appearing as transparent dots in the leaves, present in the mesophyll of
Adinandra, Anneslea, Camellia, Cleyera (Fig. 44 C-D), Eurya, Franklinia,
184 THEACEAE
Freziera (Fig. 448), Gordonia (extending from the upper to the lower
epidermis in the last two genera), Laplacea, Nabiasodendron, Pyrenaria,
Schima, Ternstroemia, Visnea. Idioblasts varying in shape in different genera.

(i)
Much thickened and provided with more or less elongated branches
with pointed ends in Anneslea, Camellia, Franklinia, Laplacea,
Pyrenaria, Schima, Ternstroemia.
(ii) Branched, but with rounded endings to the branches in Gordonia.
(iii) Only slightly branched and thickened, solitary or in groups in
Adinandra, Eurya, Visnea.
(iv) Almostcircular or rectangular, slightly thickened and mostly solitary
in Archytaea and Ploiarium.
(v) In the form of spicular cells traversing the palisade tissue
or the whole
of the mesophyll in certain species of Cleyera, Freziera, Gordonia, and
Schima.
(vi) Absent from Stewartia (except the pedicel).

Midrib usually containing a V- or U-shaped vascular strand, surrounded

by thick- walled tissue. Petiole, in transverse sections through the distal end,
exhibiting a solitary, arc-, U-, or V-shaped vascular strand in Adinandra,
Anneslea, Camellia (Fig. 46 B) (pro parte, margins of strand frequently in-
curved), Cleyera, Eurya, Freziera (pro parte), Laplacea, Nabiasodendron,
Pyrenaria, Stewartia, Ternstroemia, Ternstroemiopsis, Visnea. Large median
strand accompanied by several smaller lateral ones in Anneslea (base of
petiole), Archytaea, Camellia (pro parte), Gordonia, Lacathea, and Schima.
Vascular strand of the petiole closed in certain species of Freziera. Idioblasts
present, especially in the 'cortex' of the petiole in Adinandra, Anneslea,
Camellia, Cleyera, Eurya, Franklinia, Freziera, Gordonia, Laplacea, Nabiaso-
dendron, Pyrenaria, Schima, Ternstroemia, Ternstroemiopsis, Visnea. The
development and structure of the idioblasts in the petiolar ground tissue of
Camellia japonica have recently been investigated by Foster (694). The idio-
blasts arise from parenchymatous cells with specially large nuclei, and remain
uninucleate throughout the life of the protoplast. The cells soon put out 'one
or more delicate tubular branches which extend between the walls of neigh-
bouring tissue-elements and may ultimately penetrate certain of the inter-
cellular air-spaces. The continued ramification of the sclereid is accompanied

by the origin and development of "spicules" which, like the major branches,
grow in between the walls of adjacent parenchyma cells.' A pitted secondary
wall is formed around the cell after the intercellular development is complete.
Foster suggests that the idioblasts develop by 'intrusive* growth, but con-
cludes that elongation of the major branches is not necessarily restricted to
their tips.
'Pericycle' sclerenchymatous in Anneslea, Lacathea, Pyrenaria, Schima,
Stewartia (pro parte), Ternstroemia (partly fibrous); collenchymatous in cer-
tain species of Camellia, Stewartia, and Ternstroemiopsis. Petiole not well
defined in Archytaea, owing to the decurrent leaf bases. Solitary prismatic
and/or clustered crystals of calcium oxalate present m
the mesophyll of
Anneslea, Cleyera, Freziera, Stewartia, Ternstroemia, Visnea. Sphaero crystal-
line masses recorded in the epidermis of certain species of Eurya. Crystals

present in the 'cortex* of the petiole in Cleyera, Eurya, Pyrenaria, Visnea.

 THE ACE AE 185
Axis
YOUNG STEM (Fig. 46 D-F and L)
Young stem usually circular, but rarely (Visnid) quadrangular in trans-
verse sections.Cork arising in the sub-epidermis of Adinandra, Anneslea,
Archytaea, Cleyera (Fig. 46 L), Eurya, Franklinia, Freziera, Gordonia (pro
parte), Schima, Ternstroemia, Visnea\ pericyclic in Camellia (Fig. 46 E),
Gordonia (pro parte), Laplacea, Nabiasodendron, Pyrenaria, Stewartia,
Tutcheria. Cork cells thin-walled in Anneslea,
Archytaea, Cleyera, Eurya,
Franklinia, Freziera, Schima, Stewartia, Ternstroemiopsis (majority of cells),
Visnea', with U-shaped thickening in species of Camellia, Laplacea, Nabia-
sodendron, Ternstroemia, Ternstroemiopsis. Cells containing tannic substances
in Anneslea and Ternstroemiopsis. Primary cortex consisting of up to about
12 layers of cells, outer part frequently collenchymatous, inner part spongy,
containing sclerenchymatous idioblasts in Adinandra, Anneslea, Cleyera
(Fig.46 D and F), Eurya, Freziera, Laplacea, Nabiasodendron, Schima, Tern-
stroemia, Visnea. Idioblasts rounded in Archytaea; absent from Stewartia
except from the pedicel, and sometimes from young stems of Camellia.
Endodermis indistinct. Pericycle varying somewhat according to the
age of the shoot always heterogeneous but entirely sclerified in Adinandra,
;

Archytaea, Camellia (pro parte), Cleyera, Eurya, Franklinia, Freziera, Gor-

donia, Laplaceae, Nabiasodendron, Pyrenaria (fibre groups becoming separated
owing to the action of sub-adjacent phellogen), Schima, Stewartia, Tern-
stroemiopsis. Pericycle containing isolated groups of fibres in Anneslea,
Camellia (pro parte), Ternstroemia, Visnea. Phloem and xylem appearing,
in transverse sections, as continuous cylinders traversed by narrow rays.
Phloem containing a variable proportion of fibres in different genera. Phloem
fibres especially large and thick walled in Adinandra, Anneslea, Gordonia,
Laplacea, Pyrenaria, Ternstroemia', usually absent from Eurya, Stewartia
(very young stem), and Camellia (very young stem).
Xylem. Vessels usually solitary, up to about 40 p, in diameter; perforation
plates scalariform, often with many bars. Perimedullary region frequently
sclerified. Pith composed exclusively of isodiametric cells in Pyrenaria, but
more or less heterogeneous in other genera, this character being especially
well developed in Camellia and Stewartia. Pith, when heterogeneous, con-
sisting of a mixture of large thin- walled cells embedded in a network
of smaller
ones with thicker and sometimes pitted walls. Sclerenchymatous idioblasts
sometimes occur in the pith, having been recorded in certain species of
Adinandra, Camellia, Eurya, Franklinia, Laplacea, Nabiasodendron, Schima,
Visnea. Idioblasts form diaphragms in the pith in certain species of Cleyera,
Eurya, Freziera, Ternstroemia, and Visnea. Prismatic or clustered crystals
present in the cortex of Anneslea, Cleyera, Eurya, Franklinia, Freziera, Gor-
donia, Stewartia, Ternstroemia, Ternstroemiopsis, Visnea. Crystals also occur
in the phloem of Eurya and Stewartia.

WOOD (Fig. 45 A-G, K-L, and N)

Vessels typically small (less than 100 /x mean tangential diameter, some-
times very small (25-50 /i), e.g. in species of Camellia, Pyrenaria, and
Sakakia, slightly more than 100 /x in some species of Schima and Ternstroemia,
e.g. S. noronhae Reinw.; predominantly or exclusively solitary; very variable
 N
Fio. 45. THEACEAE, A-G, K-L, and N; ACTINIDIACEAE, H-I; BONNETIACEAE, J and M
villas* Choby. B, Gordonia concentricatrix Burk.
D
A, Adinandra
Freziera sp.
v
E, Camellia japonica Linn. F, Temstroemta penangtana Choisy. r, cmaw
y. H, Actimdia callosa Lindl. I P ^4. cnWosfl Lindl. J, Bonnetia crtusa Gleason. K, CameUia
L, Ploiarium sp. M, Bouwrtw eroiffl Gleason. N,
Ploumim ap.
 THE ACE AE 187
in number even within single genera, mostly 30-140 per sq. mm,, fewest
(16-40) in some species of Adinandra, Gordonia, Schima, and Ternstroemia,
more than 100 per sq. mm. in some species of Adinandra, Camellia, and
Sakakia, exceptionally numerous in some species of Camellia, up to 250
observed by the author and up to 400 reported by Kanehira (1206) semi-ring-
;

porous in some species of Camellia, Laplacea, and Pyrenaria\ spiral thickening

present in Adinandra, Camellia, Eurya, Franklinia (1861), Gordonia (1861),
Sakakia, Schima, Stewartia, and Ternstroemia, the spirals sometimes limited
to the tips of the elements and sometimes surrounding and linking together
the pit borders, e.g. in Camellia. Perforation plates very oblique, scalariform
and often with numerous bars (up to 100). Intervascular pitting often difficult
to observe owing to the absence of vessel groups, typically scalariform to
opposite, distinctly opposite in Adinandra, Anneslea, Cleyera, Eurya, Sakakia,
and Ternstroemia', pits to ray cells similar to the intervascular pitting in the
above genera with opposite pitting, elongated, scalariform; with wide aper-
tures and narrow borders or simple in the other genera. Occasionally with
solid contents or a few tyloses. Mean member length i-o-i-6 mm., but about
0*7-1 in Camellia and up to 2-4 in Adinandra. Parenchyma predominantly
apotracheal, diffuse, occasionally in short tangential lines (Fig. 45 A and E);
occasional narrow sheaths of paratracheal parenchyma also present in some
genera (Fig. 45 F). Crystalliferous strands of chambered parenchyma and
strands of barrel-shaped crystalliferous cells commonly present in Schima.
Rays sometimes of two distinct widths; the larger rays most commonly 2-3
cells wide, but 4-8 cells wide in some species of Anneslea, Camellia, Eurya,
Gordonia, Pyrenaria (1154), and Ternstroemia', almost exclusively uniseriate in
some species of Adinandra and Schima, e.g. A. dumosa Jack, and S. noronhae
Reinw., and also, according to Record (1861), in Franklinia and Gordonia
lasianthus Linn.; multiseriate rays usually less than i mm. high, but over
2 mm. and often very high in Adinandra, Anneslea, Cleyera, Eurya, and
composed of upright cells
Ternstroemia', uniseriate rays of multiseriate species
only, varying from moderately numerous and high, to few and low, e.g. in
Camellia and Stewartia', 9-17 rays per mm.; heterogeneous (mostly Kribs's
Types I and II A), typically with uniseriate marginal rows of 4-10 upright
cells, but with only 2 or 3 rows in Camellia and Stewartia sheath cells common
;

in Ternstroemia and, according to Janssonius (1154), occasionally present in

Camellia, Eurya, and Ternstroemia. Swollen ray cells containing solitary
crystals present in Camellia (Fig. 45 K) and, according
to Vestal (2329), druse

crystals present in Camellia, Gordonia, and Schima. Fibres with conspicuous

bordered pits (smaller in Stewartia) on all walls, sometimes
more numerous
on either the radial or the tangential walls (Janssonius 1154); walls often very
thick. Mean length 1-2-3-1 mm., up to 2 mm. or more in some species of
Adinandra, Cleyera, Eurya, Gordonia, and Schima.

ROOT
Root most fully examined by Beauvisage (163) in Camellia sasanqua Thunb.
where it is characterized by a rapidly caducous piliferous layer; a cortex of
about 12 layers of parenchyma; a slightly suberized endodermis; a central
cylinder containing 2 groups of xylem and 2 of phloem. Structure very
similar in Eurya sp.
i88 THEACEAE
ANOMALOUS GENERA
The anatomy of the following genera has been described separately because
their taxonomic position is rather uncertain.

I. CLEMATOCLETHRA Maxim.
Chinese shrubs. Description based on Beauvisage's (163) account of C.
scandens (Franch.) Maxim.
LEAF
Mesophyll consisting of 2 layers of palisade tissue, together with very
lacunar spongy tissue. Arm-palisade cells present. Large, multicellular and
uniseriate hairs occur on the petiole. Petiole, in transverse sections,
exhibiting a U-shaped vascular strand with outwardly directed arms in C.
scandens, but vascular strands tending to form a closed ring in C. tiliacea Kom.
Mesophyll containing raphides situated in elongated tubes or sacs similar ;

crystals also present in the phloem of the veins. Raphides sometimes replaced
by crystal-sand.

Axis
YOUNG STEM
Cork arising superficially; consisting of about 5 layers of cells with thin
walls. Pericycle generally provided with a composite, continuous ring of
sclerenchyma, although isolated groups of fibres also recorded. Xylem with
isolated, infrequent vessels, 50-70 n in radial diameter, and numerous,
narrow medullary rays. Pith becoming hollow. Raphides recorded in cortex
and in the phloem.

II. SLADENIA
Shrubs from the Himalayan region belonging to the single species S.
celastrifolia Kurz, the following description is based on that of Beauvisage

LEAF
Dorsiventral. Mesophyll consisting of a single layer of tall palisade cells,
and very lacunar spongy parenchyma. Stomata confined to the lower surface ;
ranunculaceous. Large clustered crystals present, particularly in the petiole.
Vascular system of the petiole appearing, in transverse sections, as a single
U-shaped strand.

Axis
YOUNG STEM
Cork originating in the epidermis. Pericycle containing isolated groups
of fibres when young, but becoming converted to a continuous composite
ring; the latter stated to consist of 3-4 layers of cells. Phloem containing
stone cells, either solitary or in groups; often accompanied by solitary crystals
in the adjoining cells. Xylem with vessels in radial rows; the latter 35-65 /x
in diameter; provided with bordered pits and scalariform perforation plates.
Wood fibres in radial rows, elements with wide lumina. Rays 1-2 cells wide.
Pith consisting of thickened pitted cells. Clustered crystals present in the
cortex, often arranged in axial rows. Raphides and crystal-sand absent.
 THE ACEAE 189

III. TREMATANTHERA F. VOH Mxill.

Shrubs belonging to the single species T. dufauri F. von M. The following
description is based on Beauvisage's (163) account.

LEAF
Dorsiventral. Asingle layer of aqueous hypoderm present. Mesophyll
consisting of 3 layers of palisade tissue interspersed with cells containing
raphides, and a broad region of lacunar spongy tissue. Vascular system of
the petiole appearing, in transverse sections, in the form of a closed ring
surrounding an accessory, collateral bundle. Two accessory bundles also
present in the cortical region of the petiole.

Axis
YOUNG STEM
Cork arising in the sub-epidermis. Outer part of the cortex containing
chlorophyll; middle part collenchymatous, but containing isolated or small
groups of sclereids; inner part consisting of thin-walled parenchyma. Endo-
dermis consisting of broad cells containing starch. Pericycle with a com-
posite ring of sclerenchyma. Xylem containing mostly isolated vessels, each
surrounded by a sheath of vasicentric parenchyma. Rays numerous, 1-2 cells
wide. Pith sclerenchymatous in the perimedullary region; the remainder
consisting of cells of very unequal sizes. Elongated cells containing raphides
and other forms of crystals also occur in the middle and inner part of the
cortex.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
The genera placed by Bentham and Hooker in the Ternstroemiaceae do
not constitute a homogeneous group. This was fully appreciated by Solereder
who was unable to find any anatomical characters which are common to all
of them. The genera here described under Theaceae are fewer in number,
and approximate to those regarded as members of this family by Melchior
(1481). Bonnetia and Pelliciera have also been excluded and described under
separate families. From a perusal of the account of the anatomical structure
of the genera described above (excluding those which are anomalous), it is
clear that they constitute a far more homogeneous group than those in the
Ternstroemiaceae of Bentham and Hooker. It is interesting to note, however,
that certain species of Freziera differ from the rest of the family in having, in
the petiole, a solitary vascular strand which appears as a closed ring in sections
through the distal end. Archytaea differs from the other genera in having
decurrent leaf bases, and rubiaceous stomata. Stewartia appears also to be
somewhat aberrant in the absence of idioblasts from the leaf. The most note-
worthy variation in stem structure is between those genera in which the
phellogen arises in the sub-epidermis and those in which its origin is peri-
cyclic. It is doubtful, however, whether much significance should be attached
to this variation, because similar divergences are known to occur within other
i 9o THE ACE AE
families. The tribes Camellieae and Ternstroemieae do not appear to be very
clearly demarcated on anatomical grounds.
The taxonomic position of the genera described above as anomalous has
never been well established, since they possess characters in common with the
Dilleniaceae and Theaceae. In this connexion it is interesting to note that
Hutchinson (1113) has pointed out the probable existence of affinities between
the Dilleniales and Theales. Sladenia seems to differ somewhat from most of
the Dilleniaceae, notably in the absence of raphides, and in the distribution
of the vessels in radial groups.

(ii) FROM WOOD STRUCTURE

Excluding Asteropeia, Bonnetia, Tetramerista and possibly Ploiarum, leaves
genera within the family that are much more uniform in structure.

ECONOMIC USES
Camellias are cultivated in gardens as ornamental shrubs. The roots and
bark have sometimes been used as a source of tannin. Tea consists of the
dried tips and upper leaflets of Camellia sinensis (L.) O. Ktze. The micro-
scopical characters of tea include the following. Upper epidermis consisting
of cells 50 IJL in diameter with slightly sinuous anticlinal walls, devoid of
stomata or hairs. Lower epidermis consisting of cells about 70 /z in diameter
with more sinuous anticlinal walls. Stomata confined to the lower surface,
surrounded by 3-4 narrow, subsidiary cells. Amount and distribution of
cutin in the guard cells as seen in transverse section believed by Rehfous
(1905) to be of diagnostic value in the recognition of different varieties of tea,
*
and also as a means for distinguishing Camellia* from tea-leaves. Palisade
cells circular in surface view. Spongy parenchyma consisting of star-shaped
cells. Idioblasts in the mesophyll very variable in form and size, up to 150 JJL

long, broadened at the ends and provided with simple or forked branches.
Rosette crystals abundant. Vascular bundles and idioblasts in tea-leaves are
coloured pink when treated with strong hydrochloric acid owing to the
presence of phloroglucin. Stracke (2211) claims to have shown that this
reaction is not given by substitutes for tea such as Camellia japonica Linn.
Recent investigations into the structure and physiology of the tea plant have
been made by Bond (227, 228).
The timbers of this family, which are of the general utility class, are not
much used. Campano, Laplacea brenesii Stand!., however, is reported (1886)
to be 'one of the best known local timbers on the market in the Cartago region
of Costa Rica*. Pearson and Brown (1679) include Needle Wood, Schima
wallichii Choisy, among the commercial timbers of India.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Adinandra, Anneslea, Archytaea, Camellia,* Clematoclethra, Cleyera,*
Eurya,* Franklinia, Freziera, Gordonia, Laplacea, Nabiasodendron, Ploi-
arum, Pyrenaria, Schima, Sladenia, Stewartia,* Ternstroemia, Tern-
stroemiopsis, Trematanthera, Tutcheria,* Visnea.
* Kew
Represented in the slide collection.
 THE ACE AE 191

(ii)
FOR WOOD STRUCTURE
Adinandra, Anneslea, Camellia, Cleyera, Eurya, (Franklinia), Gordonia,
Haemocharis, Laplacea, Pyrenaria, Sakakia, Schima, Stewartia, Ternstroemia.

LITERATURE
(i) On
General Anatomy
Beauvisage 163, Bond, T. E. T. 227, 328, Foster 694, Heilbronn 932, Hutchinson 1113,
Kobuski 1259, Lepeschkin 1361, Melchior 1481, Pekelharing 1681, Rehfous 1905,
Stracke 2211.

(ii) On Wood Structure

Beekman 167, den Berger 179, 182, Janssonius 1154, Kanehira 1206, 1209, Pearson and
Brown 1679, Record 1743, 1843, 1851, 1861, 1864, Record and Hess 1886, Tang 2231,
Vestal 2329, Williams 2430, Yamabayashi 2478.

51. ACTINIDIACEAE
(Fio. 45 on p. 186; FIG. 46 on p. 192)

SUMMARY
Trailing and climbing shrubs from the Far East belonging to the single
genus Actinidia. The most significant anatomical character is the occurrence
of raphides. The wood exhibits the following characters. Vessels small or
of two types, small and large; with spiral thickening; perforation plates simple
or scalariform; members moderately short. Parenchyma apotracheal,
diffuse or in irregular bands. Rays up to 6 cells wide, markedly hetero-
geneous. Fibres with bordered pits.

LEAF
Dorsiventral. Hairs including simple, uniseriate, and multicellular glan-
dular types. Stomata confined to the lower surface; ranunculaceous.
Mesophyll. Palisade tissue in A. callosa Lindl. consisting of one layer.
Arm palisade cells stated sometimes to occur. Spongy parenchyma in the same
species with abundant intercellular spaces. Petiole, in transverse sections
through the distal end, exhibiting a crescentic strand with incurved ends in
A. kolomikta Maxim, and a dorsally flattened almost or completely cylindrical
strand in A. callosa Lindl. (Fig. 46 c); accessory strands present in the wings
in both species. Raphides present in elongated sacs or tubes, situated in the
mesophyll and also in the cortical region of the petiole.

Axis
YOUNG STEM (Fig. 46 j)
Cork arising superficially at an early stage; consisting of thin- walled cells
svith brown contents. Later formed cork relatively deep-seated in origin.

Pericycle containing a continuous but very narrow ring of sclerenchyma.

Phloem devoid of sclerenchymatous elements. Xylem with relatively in-
frequent, mostly solitary vessels, the latter varying in diameter in different
species. Vessels of the primary xylem tending to be in radial rows. Perfora-
tion plates mostly scalariform, with numerous bars, but simple perforations
also occur. Wood fibres in radial rows separated by narrow medullary rays.
FIG. 46. STACHYURACEAE, A; THEACEAE, B, D-F, and L; ACTINIDIACEAE, C and J;
BONNETIACEAE, G; SAURAUIACEAE, H; MARCGRAVIACEAE, I and K
A, Stachyurus chinensis Franch. Petiole Xi5. B, Camellia cuspidate Veitch. Petiole XH. C, Acti-
mdia callosa Lindl. Petiole X 15. D, Cleyera ochnacea DC. Idioblast from cortex X 300. E, Camellia
cuspidata Veitch. Stem Xi$. F, Cleyera ochnacea DC. Idioblast from cortex Xsoo. G, Bometia
anceps Mart, et Zucc. Stem X 5. H, Sauratda subspinosa Anthony. Petiole x ip. 1^ Marcgravia
wnbtUata Linn. Petiole X 13. J, Actimdia callosa Lindl. Stem X 12. K, Marcgravia umbellata Linn.
Stem X 13. L, Cleyera ochnacea DC. Stem x 15.
 ACTINIDIACEAE 193
Pith septate in A. kolomikta Maxim, becoming hollow in A. callosa Lindl.
Very long raphides, situated in elongated idioblasts or sacs, present in the
primary cortex, phloem, and pith. These crystals sometimes resemble a
gummy or resinous material on superficial examination, possibly owing to the
presence of mucilage. The true nature of the crystals is most clearly revealed
in longitudinal sections. Crystals exhibiting forms transitional between
raphides and crystal-sand also occur, particularly in the phloem.
WOOD (Fig. 45 H-I)
Vessels generally of two types, (a) large, tending to be 'locally zonate' (533),
and with simple perforations, and (b) small, mostly solitary and with simple
or scalariform perforation plates, the latter with several to many bars; with
spiral thickening. With only small vessels in the material examined of A.
kolomikta Maxim. Intervascular pitting small, often with coalescent apertures ;

pits to ray and wood parenchyma similar. Mean member

length o-y-O'S mm.
Parenchyma apotracheal, scattered and in irregular lines (Fig. 45 i). Strands
commonly of 8 cells. Rays tending to be of two sizes, uniseriate and composed
entirely of square or upright cells, or multiseriate, up to 6 cells wide, composed
of procumbent cells with several marginal rows of square or upright cells
(Kribs's Heterogeneous Type I). Fibres with numerous, distinctly bordered
pits on both radial and tangential walls.

ROOT
Raphides present.
TAXONOMIC NOTES
Actinidia probably has affinities with the Dilleniaceae and Theaceae.

GENUS DESCRIBED
Actinidia.*
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Cr6te* 498, Gilg and Werdemann 782.
(ii) On Wood Structure
den Berger 182, Dadswell and Record 533, Record 1843, 1851, Vestal 2329.

52. BONNETIACEAE
(Fic. 45 on p. 186; FIG. 46 on p. 192)

SUMMARY
Bonnetia, which is a Brazilian genus, has the general features of the
Theaceae but the leaf bases are decurrent, whilst the stomafa are rubiaceous.
The wood exhibits the following characters. Vessels solitary, perforations
simple, intervascular pitting alternate, pits to ray cells large, members of
medium length to moderately short. Parenchyma varying from pre-
dominantly diffuse to predominantly paratracheal (aliform). Rays up to
2 or 3 cells wide, with numerous uniseriates, heterogeneous. Fibres with
4594 O
i 94 BONNETIACEAE
conspicuous bordered pits on all walls, of medium length to moderately short.
Vasicentric tracheids present.
LEAF
Lamina thick, dorsiventral; the upper surface covered with a thick smooth
cuticle. Cells of the epidermis of two very distinct sizes, the larger ones

being very tall and filled with a mucilaginous secretion. The size and arrange-
ment of the mucilaginous cells vary in different species, and in some the cells
appear to form a hypoderm. Stomata confined to the lower surface;
rubiaceous. Mesophyll consisting of 2-5 layers of short palisade cells, and
a broader region of spongy tissue. Midrib containing an annular vascular
strand, surrounded by a sclerified pericycle; usually including some muci-
laginous cells on the lower side. Petiole, in transverse sections, exhibiting
a median vascular strand which is usually crescent-shaped or almost cylindrical
through the ends being strongly incurved. In some sections of B. anceps
Mart, it was completely closed. The median vascular strand is
accompanied
on either side by a few small bundles.

Axis
YOUNG STEM (Fig. 46 G)
Epidermis covered with a thick, smooth cuticle. Cork superficial. Cortex
somewhat spongy; containing a few unbranched, thick-walled, sclerenchy-
matous idioblasts and also cluster crystals. Pericycle including a composite,
continuous, or slightly interrupted ring of sclerenchyma. Phloem and xylem
in the form of continuous cylinders traversed by narrow rays. Strands of
fibres noted in the phloem of B. anceps Mart. Vessels of B. anceps mostly
isolated, except in the primary xylem where they are in radial rows and more
oval in shape, up to about 90 p, in radial diameter; perforations not easily
seen, but simple and scalariform types with few bars apparently present.
(See also 'Wood'.) Wood fibres in the same species provided with abundant
bordered pits. Perimedullary region composed of a narrow zone of lignified
elements. Pith spongy, composed of somewhat elongated, sclerosed, pitted
cellsand usually containing cluster crystals. Crystals, see 'Cortex* and
Tith' above.

WOOD (Fig. 45 j and M)

Vessels moderately small (mean tangential diameter 50-100 p) to medium-
sized (100-200 fc) exclusively solitary 9-25 per sq. mm. Perforations simple,
; ;

slightly oblique (see also 'Young Stem'). Intervascular pitting alternate,

moderately large, sometimes tending to be scalariform near the primary wood;
pits to ray and wood parenchyma commonly large and almost simple. Solid
deposits and tyloses sometimes abundant. Mean member length o-3-o-8 mm.
Parenchyma varying from predominantly paratracheal with a little diffuse
to predominantly diffuse, with a few cells associated with the vessels; the
paratracheal parenchyma commonly as small aliform clusters of cells at the
sides of the vessels; sometimes, e.g. in Bonnetia tristyla Gleason, tending to
surround the vessels except on the abaxial sides (Fig. 45 j). Sometimes con-
taining chambered crystals. Strands commonly of 2-4 cells. Rays up to 2 or
3 cells wide; less than i mm. high; uniseriates numerous, composed of high
 BONNETIACEAE 195

upright cells and commonly only i or 2 cells high; 8-12 rays per mm.;
distinctly heterogeneous (Kribs's Type II A), usually with 1-4 marginal rows
of square to upright cells, but with more in some species. Solid contents
abundant. Fibres with very numerous, conspicuous bordered pits on both
radial and tangential walls. Walls very thick. Mean length o-8-i-i mm.
Vasicentric tracheids present.

TAXONOMIC NOTES
Bonnetia possesses characters in common with the Theaceae, although it
differs from that family in a few respects. It seems to be a matter of opinion
whether the genus should be placed in a distinct family, or regarded as a tribe
or sub-family within the Theaceae.
Vestal (2329) considers that the anatomical evidence supports the sugges-
tion that this group stands as a connecting-link between the Theaceae and the
Guttiferae.
is a very close resemblance between the wood
There anatomy of some
species of Bonnetia, e.g. B. tristyla Gleason, and some members of the Kiel-
meyeroideae, e.g. Caraipa richardiana Camb., the points of difference being
mainly the exclusively uniseriate rays of the latter and the occurrence of the
parenchyma (otherwise similar) on the abaxial and not on the axial sides of
the vessels.
GENUS DESCRIBED
(i) FOR GENERAL ANATOMY
B. anceps Mart.,* B. roraimae Oliv.*
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Bonnetia.
LITERATURE
(i) On General Anatomy
Beauvisage 163.

(ii) On Wood Structure

Record 1861, Record and Hess 1886, Vestal 2329, Williams 2430.

53 CARYOCARACEAE
(FiG. 47 on p. 202)

SUMMARY
A
family of trees and shrubs from Latin American countries, comprising
the two genera Anthodiscus and Caryocar. The wood exhibits the following
characters. Vessels moderately small in Anthodiscus, large in Caryocar; per-
forations simple; intervascular pitting alternate, medium-sized and with large
simple pits to ray cells in Caryocar, minute and with pits to ray cells similar
in Anthodiscus; members of medium length to moderately long. Parenchyma
predominantly apotracheal (diffuse) but with some paratracheal in Caryocar,
paratracheal only (vasicentric to slightly aliform) in Anthodiscus; crystal druses
sometimes present. Rays with small biseriate parts and numerous marginal
196 CARYOCARACEAE
rows of square to upright cells; druse crystals sometimes present. Fibres
with simple pits; occasionally septate in Caryocar, of medium length to
moderately long. Anomalous structure has been recorded in the root of a
Caryocar.
LEAF
Generally dorsiventral, but sometimes tending to be centric, e.g. in Caryocar
glabrum Pers, and certain other species of Caryocar. Hairs infrequent or
absent; short, bent, pointed, unicellular trichomes recorded on the petiole of
Anthodiscus. Extra-floral-nectaries present at the margin of the young leaf
and on the stipules. Paired hydathodes also present near the apices of the
stipules. Cork warts recorded on the lower surface of Caryocar. Cuticle on
both surfaces thick. Stomata mostly confined to the lower surface, but a few
in depressions beside the veins on the upper surface in Caryocar nuciferum
Linn, observed by Blank (203), who also counted up to 435 per sq. mm. on
the lower surface. Stomata ranunculaceous in Caryocar. A single layer of
aqueous hypoderm recorded in Anthodiscus. Mesophyll including 2 layers
of palisade cells in Caryocar nuciferum according to Blank (203); spongy
portion with but few intercellular spaces in the same species. Branched,
sclerenchymatous idioblasts present in the mesophyll (in C. nuciferum
especially beside the veins and at the leaf margin) and also in the ground
tissue of the petiole in both Anthodiscus and Caryocar. Vascular bundles of
the veins accompanied above and below but not surrounded by strands
of cellulose fibres in C. nuciferum. Vascular structure of the midrib of
Caryocar similar to that of the petiole (see below). Petiole in Caryocar some-
times becoming covered with 2-4 layers of cork. Petiolar vascular system in
Caryocar described by Blank (203) as consisting of bundles which unite at
the distal end to form a sinuous ring, the latter being accompanied by a few
cortical and more numerous (up to 10 in C. brasiliana St. Hil.) medullary
strands. Petiolar vascular strands said to be separate at the base and at the
distal end, but united to form a cylinder in the middle portion of the petiole
in Anthodiscus. Crystals numerous in the palisade tissue of Caryocar
nuciferum. Prismatic crystals also recorded in the petiolar ground tissue of
Anthodiscus. Dark spots in the lamina of Caryocar nuciferum, visible by
transmitted light, represent tanniniferous cells, according to Blank (203).

Axis
YOUNG STEM
Surface in Caryocar said to be covered with abundant short hairs when
very young, but these soon become detached. Cork arising in the sub-
epidermis or outer part of the cortex consisting, in Caryocar, of a mixture of
;

cells with thin walls and others with U-shaped thickenings. Primary cortex
about 10 cells wide; including nodular, sclerenchymatous idioblasts. Endo-
dermis in Caryocar nuciferum Linn, said by Blank (203) to be clearly defined
but to be devoid of casparian thickenings. Pericycle in Caryocar bounded
externally by a ring of fibres locally interrupted by parenchymatous cells.
Phloem and xylem in the form of continuous cylinders traversed by narrow
rays. Occasional fibres present in the phloem. Vessels tending to be in radial
multiples of 2-3 in Caryocar generally solitary in Anthodiscus up to 60 /z in
; ;
 CARYOCARACEAE 197
radial diameter in Anthodiscus, rather larger in Caryocar; perforations trans-
verse to very oblique in both genera but always simple. Pith in Anthodiscus
and containing rounded, sclerenchymatous idioblasts. Only the peri-
sclerified

pheral part of the pith in Caryocar lignified when young, but becoming more
completely lignified when older, or with the central portion of the pith some-
times becoming disorganized. Branched, sclerenchymatous idioblasts present
in the pith of the same genus. Knee-shaped crystals and less numerous
druses accompany the pericyclic sclerenchyma of Caryocar according to
Blank (203); cluster crystals also recorded in the secondary phloem. Secre-
tory canals observed by Blank (203) in the hypocotyl and base of the stem
of Caryocar nuciferum.

WOOD
Caryocar (Fig. 47 E and G)
Vessels medium-sized (mean tangential diameter 100-200 ju) to large
(more than 200 /u.); solitary and in radial multiples of 2 or 3 cells; 1-2 per
sq. mrn. Perforations typically simple, but Williams (2429, 2430) notes a
tendency to multiple plates. Intervascular pitting alternate, moderately large ;

pits to ray cells often large, irregularly elongated and simple. Tyloses present
and often abundant. Mean member length about 0-8 mm. (Kribs 1283,
Vestal 2329). Parenchyma predominantly apotracheal, as scattered cells
and short tangential lines, but with some paratracheal parenchyma (vasicentric
to slightly aliform) in addition (Fig. 47 G). Chambered cells containing
solitary crystals moderately common, sometimes containing druses (2329).
Strands commonly of 8 cells. Rays up to 2 cells wide and often more than
i mm.
high; the biseriate parts seldom more than 7 cells high, little wider
than the uniseriate cells and often more than one per ray (Fig. 47 E) uni- ;

seriates not very common(most of the rays being biseriate in part), composed
entirely of square to upright cells; 11-18 rays per mm.; markedly hetero-
geneous (Kribs's Type II A), with several marginal rows of square to upright
cells. Crystals sometimes present in subdivided marginal cells and sometimes

consisting of druses (2329). Fibres with small simple pits, mostly on the
radial walls. Septa rare to common (Williams 2429). Walls very thick and
often with a gelatinous inner layer. Mean fibre length about 1*8-2*0 mm.
(Kribs 1283, Williams 2430).

Anthodiscus (Fig. 47 F and H)

Vessels moderately small (mean tangential diameter 50-100 ^), solitary
and in short radial multiples of 2 or 3 cells; about 15 per sq. mm. Perforations
typically simple, but Vestal (2329) refers to occasional scalariform plates.
Intervascular pitting alternate, minute pits to ray cells similar. Solid deposits
;

common; ty loses absent. Member length about 0*7 mm. Parenchyma para-
tracheal, partially surrounding the vessels and sometimes slightly aliform
(Fig. 47 H). Chambered cells containing crystals present in some species, but
apparently absent from others (1886). Strands commonly of up to 6-8 cells.
Rays similar to those of Caryocar except for the presence of abundant gum-
like deposits and the absence of crystals. Fibres similar to those of Caryocar

except that gelatinous layers and septa are absent, mean length about
1*5 mm.
198 CARYOCARACEAE
ANOMALOUS STRUCTURE, see 'Root* below
ROOT
The root structure of Caryocar nuciferum Linn, has recently been investi-
gated by Blank (203) from whose account the following particulars are taken.

(i)
ADVENTITIOUS AND SUBSIDIARY ROOTS
Mostof the cells of the sub-epidermal layer provided with strongly
thickened and suberized inner tangential walls, but occasional unthickened
cells also present. Endodermis also suberized, but casparian thickenings
visible only when young. Primary vascular structure diarch. Phloem well

developed when roots are sufficiently mature. Paired, monoclinic crystals of

calcium oxalate present.

(ii) PRINCIPAL ROOTS ARISING FROM THE SWOLLEN BASE OF THE HYPOCOTYL
Several layers of cells with suberized walls and tanniniferous contents
present in the hypodermal region. Endodermis with conspicuous casparian
thickenings. Primary vascular structure triarch, tetrarch, pentarch, hexarch,
or octarch. Phloem including fibres. Xylem with more numerous vessels
than in the adventitious and subsidiary roots. Wood fibres with lignified
primary and unlignified secondary walls. Medullary rays mostly 2-6 cells
wide. Anomalous structure. Xylem and phloem of the normal ring show
unequal development at different points on their circumference. A second
xylem body, devoid of vessels, develops in the large pith, partly by lignifica-
tion of the pith cells and partly from 2 to 6 small groups of meristematic cells.
The latter cut off xylem on the outside and phloem on the inside, but the
phloem becomes surrounded by xylem and is thus interxylary. The inter-
xylary phloem ends blindly in the tuberous base of the hypocotyl at one end
and towards the root apex at the other. This anomalous structure is of the
corpus lignosum foraminulatum type. More towards the apex of the root, cork
cells are differentiated from the abnormal phloem tissue, and a phellogen may
also arise between them and the phloem. The abnormal xylem locally traverses
the almost closed inner phloem ring and so unites with the inner ring of cork.
All of the abnormal cork cells contain tannin, or their end walls sometimes
break down to form larger tanniniferous cavities. Secretory cavities present
in the normal protophloem in the portion of the root nearest to the swollen
base of the hypocotyl. Knee-shaped crystals present in both the normal and
the abnormal phloem.

TAXONOMIC NOTES
The
presence of idioblasts, the nature of the hairy covering, and the
sub-epidermal origin of the cork are suggestive of affinities between
the Caryocaraceae and Theaceae. The vascular system of the petiole of the
Caryocaraceae is more complex, however, and since considerable taxonomic
significance can probably be attached to this difference, it seems probable
that the Caryocaraceae should be regarded as a distinct family. Blank (203)
believes that the secretory cavities in the root, stem base and hypocotyl of
Caryocar are indicative of affinities with the Guttiferales.
Vestal (2329) suggests a close affinity between Caryocar and Tetramerista,
 CARYOCARACEAE 199
but against this must be noted marked differences in ray type, intervascular
and ray- vessel pitting and fibre pits.

ECONOMIC USES
'Butter Nuts* are the fruits of Caryocar nudferum Linn., the seeds of which
have a high content of edible fat. The structure of the fruit and seed has
been investigated in considerable detail by Blank (203).
The
timbers of some species of Caryocar are well known locally and are
used for such purposes as shipbuilding, warehouse flooring, wheel hubs, and
felloes.

GENERA DESCRIBED
Anthodiscus, Caryocar.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Blank 203, Pilger 1721.

(ii) On Wood Structure

Howard 1088, Kribs 1283, Pfeiffer, J. Ph. 1713, Record 1843,
Benoist 170, Blank 203,
1851, Record and Hess 1886, Vestal 2329, Williams 2429, 2430.

54. MARCGRAVIACEAE
(Fie. 46 on p. 192; FIG. 47 on p. 202)

SUMMARY
(i) GENERAL
A
family of climbing shrubs, and epiphytes and occasional erect species
which occur in Tropical America. The branch system of Marcgravia is
dimorphic, consisting of (i) sterile portions which are attached to the sub-
stratum and bear two rows of leaves, and (ii) fertile, pendulous portions with
spirally arranged leaves and a terminal inflorescence. The leaves of the sterile
and fertile shoots respectively also differ in structure (see 'Leaf'). Raphides
and sclerenchymatous idioblasts are two of the most characteristic
anatomical features.

(ii) WOOD
Marcgravia and Souroubea
Vessels medium-sized to large, perforation plates simple or simple and
scalariform with few bars, intervascular pitting alternate, pits to ray cells
similar, members of medium length. Parenchyma paratracheal, vasicentric.
Rays up to 10 or more cells wide, composed almost entirely of square and
upright cells, containing raphides. Fibres septate, with small bordered pits,
of medium length to moderately short.

Norantea
Vessels moderately small, multiples of 2-6 cells common, perforation
plates scalariform with many fine bars, intervascular pitting opposite, pits to

ray cells commonly unilaterally compound, members very long. Parenchyma

200 MARCGRAVIACEAE
apotracheal, diffuse. Rays up to 10-12 cells wide, markedly heterogeneous,
chambered crystalliferous cells common. Fibres with bordered pits, non-
septate, sometimes with spiral thickening, very long.

LEAF
Dorsiventral. Hairs Structures resembling water-pores, but of
absent.
unknown physiological significance, present at the apices of the leaves. Extra
floral nectaries and cork warts present on the lower surface of the leaf of
certain species of Marcgravia and Norantea. Their physiological function is
not definitely known, but they may serve for the secretion of resinous
material. Cells of the upper epidermis provided with a thick cuticle. A
single layer of aqueous hypoderm present beneath the upper epidermis.
Stomata mostly confined to the lower surface, but also occurring in small
numbers on the upper surface in Marcgravia. Mesophyll consisting of i or
2 layers of palisade tissue, accompanied by a broader, lacunar spongy tissue.
Sclerenchymatous idioblasts present in the mesophyll, and in the cortical
parenchyma of the petiole in all genera; exhibiting a variety of shapes and
frequently much branched, especially in Norantea. Petiole, in transverse
sections, exhibiting a single crescent-shaped vascular strand in Marcgravia
coriacea VahL a main vascular bundle in the form of an almost or completely
;

closed ring, sometimes accompanied by a small, accessory, collateral bundle

immediately external to the ring in Marcgravia umbellata Linn. (Fig. 46 i),
Norantea, Ruyschia, Souroubea. Secretory canals present in the midrib of
Marcgravia, and cells containing oleo-resin in the petiole of Souroubea.
Starch not observed in any member of the family; inulin thought by
Melchior (1482) to be the main product of carbon assimilation at least in
certain species of Marcgravia. Bundles of raphides occur in special cells in
the mesophyll of Marcgravia, Norantea, Souroubea. Similar cells filled with
mucilage or oil drops also recorded.
Leaves on the sterile differ from those on the fertile shoots in the following
respects: (i) The rougher upper surface caused by the papillose epidermis,
(ii)
The presence of stomata on both surfaces, (iii) The absence of an aqueous
hypoderm below the upper epidermis, (iv) The much greater infrequency of
sclerenchymatous idioblasts in the mesophyll. (v) The chlorophyll granules
being much larger than those of the fertile shoots.

Axis
YOUNG STEM (Fig. 46 K)
Cork superficial in origin in Marcgravia, Ruyschia, Souroubea', cells with
U-shaped thickenings in Marcgravia', mixed thin- walled cells and others
with U-shaped thickenings or completely sclerosed in Souroubea. Cortex
frequently collenchymatous in the outer part, but somewhat spongy towards
the inside; sometimes including much branched sclerenchymatous idioblasts
in all genera.Pericycle with a continuous ring of-sclerenchyma, consisting
of fibres when young but becoming composite when older. Phloem and

xylem appearing, in transverse sections, as closed rings traversed by narrow

medullary rays. Vessels in Marcgravia umbellata Linn, mostly somewhat
angular in transverse section, up to about 45 /z in diameter; solitary or
tending to be in short radial groups; lateral walls with large, circular, oval,
 MARCGRAVIACEAE 201

or horizontally elongated bordered pits with wide borders; perforations very

oblique, simple. (See also under 'Wood* below.) Pith consisting of thin-
walled cells in all genera, but with branched idioblasts interspersed amongst
them in Marcgravia, Norantea, Ruyschia. Raphides present in the cortex
and sometimes the pith. Secretory cells containing oleo-resin recorded in
the cortex of Souroubea.

WOOD (Fig. 47 A-D)

Marcgravia and Souroubea
1

Vessels typically medium-sized (mean tangential diameter 100-200 p\

occasionally large (more than 200 JJL) in some species of Marcgravia (2329);
solitary and in radial pairs and threes; about 6 per sq. mm. Perforations
probably typically simple in Marcgravia, though referred to in some of the
literature (e.g. 959, 2430) as simple and scalariform; perforation plates
scalariform, with few bars, in Souroubea and occasionally foraminate or
reticulate. (See also under 'Young Stem' above.) Intervascular pitting alter-
nate, of medium size; commonly with coalescent apertures; pits to ray cells
similar, but without coalescent apertures. Mean member length about 0-6 mm.
(Marcgravia). Parenchyma paratracheal, vasicentric, forming a narrow
sheath i or 2 cells wide (Fig. 47 B). Strands commonly of 2-4 cells in Marc-
gravia. Rays tending to be of 2 distinct sizes, the larger up to 10 or more
cells wide and several millimetres high; uniseriates numerous; both uniseriate
and multiseriate rays formed predominantly of square or upright cells about ;

12 rays per mm. Large cells containing raphides present in both uniseriate
and multiseriate rays. Fibres with small bordered pits that are equally
numerous on both radial and tangential walls. Septate. Walls thin to
moderately thick. Sometimes with conspicuous intercellular spaces in Marc-
gravia. Mean length about 0*9 mm. (Marcgravia).

Norantea
Vessels moderately small (mean tangential diameter 50-100 /u,); solitary
and in numerous multiples of 2-6 cells; about 15 per sq. mm. Perforation
plates scalariform, with many fine bars, in material examined by the author;
Record and Hess (1886), however, describe the perforations as simple. Inter-
vascular pitting opposite, of medium size; pits to ray cells similar in shape
and size to the intervascular pitting, but often unilaterally compound, one
elongated ray pit subtending 2-4 circular vessel pits. Mean member length
about i -7 mm. Parenchyma apotracheal, diffuse and in short tangential
lines (Fig. 47 D). Strands mostly of 8 cells. Rays of 2 distinct sizes, the larger

up to 10-12 cells wide and 3 mm. high; about n

rays per mm.; uniseriates
numerous, composed of high upright cells and seldom more than i mm. high;
markedly heterogeneous (Kribs's Type I), often with 10 or more marginal
rows of high upright' cells and with some sheath cells. Upright and pro-
cumbent cells often chambered and containing crystals. Fibres with rather
few, large, bordered pits, which occur mostly in the radial walls. Walls rather
thin. Sometimes with faint spiral thickening. Mean length about 2-5 mm.

1
No material of Souroubea was examined by the author ; its description here is based mainly
on the data given by Hess (959) and Vestal (2329).
 FIG. 47- MARCGRAVIACEAE, A-D; CARYOCARACEAE, E-H; SAURAUIACEAE, I-J
A, Marcgravia rectiflora Tr. et PI. B, Af. recttflora Tr. ct PI. C, Norantea sttbsessiKs Donn. D, AT.
$wfc!7w Donn. E, Caryocar villosum (Aubl.) Pers. F> Anthodiscus trifoliatus G.F.W. Mey. G, Caryo-
carglabrum Pers. H, Anthodiscus trifoliatus G.F.W. Mey. I, Saurauia griffitkii Dyer. J, S. griffithii
Dyer.
 MARCGRAVIACEAE 203
ROOT
Cortex containing raphides. Vascular cylinder consisting of 5 groups
of xylem and 5 of phloem in Marcgravia.

TAXONOMIC NOTES
The
anatomical characters do not sharply differentiate the Marcgraviaceae
from the Theaceae, the presence of sclerenchymatous idioblasts being
especially characteristic of both families. On the other hand, the more
complex character of the vascular strand in the petiole, and the presence of
raphides, serve to distinguish the Marcgraviaceae from the Theaceae.
Vestal (2329) considers that in the wood structure 'the genus Norantea
possesses anatomical characters that would seem to link this family very
closely to the Theaceae'.
Marcgravia and Norantea differ in almost every character of their wood
anatomy and there appears to be no reason for associating them together.
Souroubea appears from descriptions to be similar to Marcgravia. Marcgravia
has a more highly specialized type of wood than Norantea.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Marcgravia,* Norantea, Ruyschia, Souroubea.
* Kew
Represented in the slide collection,

(ii)
FOR WOOD STRUCTURE
Marcgravia, Norantea (Souroubea).

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gilg and Werdemann 783, Melchior 1482, Richter 1933.

(ii) On Wood Structure

Dadswell and Record 533, Hess 959, Record 1843, 1851, Record and Hess 1886, Vestal
2329, Williams 2430.

55. MEDUSAGYNACEAE
SUMMARY
A
small family from the Seychelles, consisting of shrubs belonging to the
single genus Medusagyne. The species most fully
described anatomically is
M. oppositifolia Baker which exhibits the characters recorded below.

LEAF
Cuticle on the upper surface thick. Cells of the upper
Dorsiventral.
epidermis much taller than those of the lower. Hypoderm of i or 2 layers
of mucilaginous cells present beneath the upper epidermis. Stomata con-
fined to the lower surface; ranunculaceous. Mesophyll consisting of about
3 layers of palisade cells and an extensive, very lacunar, spongy mesophyll.
204 MEDUSAGYNACEAE
Vascular strand of the midrib in the form of a flattened ellipse, composed of
numerous small vascular bundles. Petiole, in transverse sections, exhibiting
an arc of about 6-9, variously orientated and more or less U-shaped vascular
bundles. Mucilage cells present in the mesophylL Crystals all clustered ;

occurring in the petiole and around the vascular strand of the midrib.

Axis
YOUNG STEM
Cork arising in the sub-epidermis; consisting of about 6 layers of cells, the
outermost ones with thin walls, but those towards the phellogen provided
with U-shaped thickenings. Cortex fairly broad, parenchymatous, but
including a proportion of sclerosed cells and some which contain cluster
crystals. Small scattered strands consisting mainly of fibres, but sometimes
including a few vessels, recorded by Engler and Melchior (647) in the primary
cortex. Pericycle containing a closed ring of sclerenchyma when very young,
but the ring becomes interrupted by parenchyma and stone cells when older.
Phloem stratified into alternating concentric rings
of sieve tissue and strongly
thickened cells. ring interrupted by primary rays which are mostly
Xylem
5-8 cells wide, the secondary rays being 2-3 cells wide. Vessels mostly
solitary, 15-20 \L in diameter; perforations simple. Pith consisting of
thickened, pitted elements containing prisms of calcium oxalate. Crystals,
see 'Cortex' and Tith' above.

TAXONOMIC NOTES
The taxonomicposition of Medusagyne is somewhat uncertain, as it has
been variously ascribed to the Guttiferae and Theaceae. The genus differs
from the Guttiferae and Marcgraviaceae in the absence of resin-canals and
from the Theaceae in the absence of the sclerenchymatous idioblasts which
are a characteristic feature of the parenchymatous tissues of nearly all members
of that family. It resembles the Ochnaceae in having cortical bundles.

GENUS DESCRIBED
Medusagyne.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Beauvisage 163, Engler and Melchior 647.

56. PELLICIERACEAE
SUMMARY
The sole representative of this family is Pelliciera rhizophorae Trian. et
Planch., a tree growing in watery places in Latin American countries. In
habit it resembles a Rhizophora. Unlike most of the true Theaceae the plant
has decurrent leaf-bases. A notable anatomical feature is the occurrence of
raphides in special cells situated in the parenchymatous tissues.
 PELLICIERA CEAB 205
LEAF
Thick and leathery with the midrib not projecting above the surface.
Dorsiventral. Hairs. None observed. Upper epidermis composed of small,
polygonal cells with highly cutinized outer walls and slightly sinuous anti-
clinal walls. Cells of the lower epidermis somewhat larger and more thickly
cutinized. Stomata confined to the lower surface, each surrounded by about
5 concentric subsidiary cells. A single layer of thin-walled, aqueous hypo-
derm present beneath the upper epidermis, the anticlinal walls of which
become considerably contracted in herbarium material. A single layer of
smaller aqueous cells also occurs on the inside of the lower epidermis.
Mesophyll composed of 2 layers of palisade cells, interspersed with sacs of
raphides, and a highly lacunar spongy tissue. Sinuous fibres, mostly lying
parallel to the leaf surface, scattered throughout the mesophyll; groups of
stone cells also present in the spongy tissue. Petiole collenchymatous in the
outer part of the cortical region, but more spongy and including sinuous
fibres nearer the almost annular but somewhat interrupted vascular strand;
occasional raphide sacs occur in the cortical region and small prismatic
crystals chiefly in the phloem.

Axis
YOUNG STEM
Cork
arising superficially in very young stems; composed of thin- walled
cells. clearly differentiated into an outer collenchymatous and an
Cortex
inner spongy region. Pericycle bounded externally by a composite, con-
tinuous ring of sclerenchyma, with small prismatic crystals in some of the
component elements. Phloem and xylem in the form of closed cylinders
traversed by uniseriate rays. Vessels in radial rows of about 3-10, up to about
45 /LL
in radial diameter; perforations simple, oblique. Wood fibres consisting
of radial rows of moderately thick-walled elements, with inconspicuously
bordered pits. Pith consisting of spongy, parenchymatous, comparatively
thin-walled but pitted cells. Branched sclerenchymatous idioblasts and
elongated thick-walled fibres present in the inner, spongy portion of the
cortex as well as in the pith. Crystals, see under Tericycle* above.

TAXONOMIC NOTES
Although was treated as a tribe of the Theaceae by Melchior
Pelliriera

(1481) it
conspicuously from the other members of this family in the
differs

frequent occurrence of raphides, and in the almost annular structure of the

vascular strand of the petiole. There are also other important differences in
the floral and external morphological characters. Beauvisage (163), suggested
that Pelliciera should form the basis of a family closely related to but distinct
from the Theaceae, and intermediate between them and the Marcgraviaceae.

GENUS DESCRIBED
Pelliciera,*
*
Represented in the Kew slide collection.

LITERATURE
On General Anatomy
Beauvisage 163, Melchior 1481,
 (206)

57. PENTAPHYLACACEAE
(Fio. 48 on p. 206)

SUMMARY
The Pentaphylacaceae are represented by the single genus Pentaphylax,
which consists of shrubs with leathery leaves occurring in China and Malaya.
The anatomical characters which have been recorded about the general
anatomy refer only to P. euryoides G. et C. The wood exhibits the following
features. Vessels solitary, perforation plates and intervascular pitting scalari-
form, members extremely long. Parenchyma diffuse. Rays up to 5 cells
wide, heterogeneous. Fibres with distinctly bordered pits, extremely long.

FIG. 48. PENTAPHYLACACEAE, A-B; STACHYURACEAEf C-D

A, Pentaphylax arborea Ridley. B, P. arborea Ridley. C, Stachyurus chinensis Franch. D, 5.
praecox S. et Z.

LEAF
Dorsiventral. Epidermis mucilaginous. Stomata confined to the lower
epidermis; figured by Beauvisage (163) as rubiaceous in most instances.
Mesophyll consisting of one layer of palisade tissue together with a region
of very lacunar spongy parenchyma, the latter occupying two-thirds of the
thickness of the lamina. Vascular bundles of the veins provided with a sheath
of sclerenchyma. Petiole, in transverse sections, exhibiting a solitary, slightly
U-shaped vascular strand; pericycle unlignified except at the distal end.
Solitary crystals present in the mesophyll.

Axis
YOUNG STEM
Corkarising in the sub-epidermis; consisting of small elements with thin
walls. Primary cortex slightly collenchymatous in the outer part. Endo-
dermis not clearly defined. Pericycle including a composite and continuous
 PENTAPHYLACACEAE 207

ring of sclerenchyma. Xylem with small isolated vessels 15-30 /u, in radial
diameter; perforation plates scalariform; wood fibres with very thick walls;
rays 1-2 cells wide, frequently contiguous to the vessels. Pith consisting of
cells with thickened, pitted walls. Secretory cells with mucilaginous con-
tents present in the cortex. Prismatic crystals occur in the phloem, and a
solitary crystal in each cell of the endodermis.

WOOD (Fig. 48 A-B)

Vessels very small (25-50 /z mean tangential diameter) and angular;
almost entirely solitary, except for apparent tangential pairs due to over-
lapping ends; about 30 per sq. mm. Heimsch (938) refers to spiral thickening
as abundant in a twig of Pentaphylax euroides G. et C. but as present only in
a few vessel tips in mature material of P. arborea RidL Perforation plates
scalariform, usually with about 15 bars but sometimes with more than 25
(938). Intervascular pitting scalariform to transitional; pits to ray and wood
parenchyma similar in outline to the intervascular pitting, but almost simple.
Mean member length about 2-0 mm. Parenchyma apotracheal, as isolated
among the fibres, with a slight tendency to form short uniseriate
cells scattered
lines. Strands usually of 6-8 very high cells. Rays up to 4 or 5 cells wide ;

uniseriates moderately numerous and composed of square to upright cells;

about 8 rays per mm. markedly heterogeneous (Kribs's Type I-II A), with
;

1-8 marginal rows of square to upright cells. Dark, gum-like deposits

abundant. Fibres with distinctly bordered pits, which are more abundant on
the radial than on the tangential walls. Walls thick. Mean length 3-3 mm.

TAXONOMIC NOTES
Pentaphylax was treated by Bentham and Hooker as a member of the
Ternstroemiaceae-Ternstroemieae. It is retained in the Theaceae by Hutchin-
son (i 113), but in the Englerian system it is treated as a separate family having
affinities with the Cyrillaceae and Celastraceae. Beauvisage (163) also regards
the genus as related to the Cyrillaceae. Heimsch (938) considers that, on the
basis of the wood anatomy, the genus would be better placed near the
Celastraceae or the Theaceae than in the Terebinthales.

GENUS DESCRIBED
(i)
FOR GENERAL ANATOMY
Pentaphylax. The above description is based mainly on Beauvisage's (163)
account of the structure of P. euryoides G. et C.

(ii) FOR WOOD STRUCTURE

Pentaphylax (a single sample of P. arborea RidL).

LITERATURE
(i) On General Anatomy
Beauvisage 163, Hutchinson 1113.

(ii) On Wood Structure

Heimsch 938.
 FIG. 49. SAURAUIACEAE, A, C, D; GUTTIFERAE> B

A, Branched trichome of Saurauia napaulensis DC. B, Lower epidermis of the leaf of Caraipa
glabrata Mart., with stellate hairs. C, Clustered hair of Saurauia spadicea Bl. D, Shaggy hair of
Saurauia napaulensis DC. By Solereder,

FIG. 50. CHLAENACEAE

A, Transverse section through one of the small depressions (pits) on the lower surface of the leaf of
Schizolaena rosea Thouars. B, Small scale of Sarcolaena multiflora Thouars in surface-view,- By
Solereder.

FIG. 51. DIPTEROCARPACEAE FIG. 52. ANCISTROCLADACEAE

Lower epidermis of the leaf of Epidermis of lower side of leaf in Ancistro-
Anisoptera lanceolata Waip. By cladus heyneanus Wall By Solereder.
Solereder.
 (209)

58. SAURAUIACEAE
(Fio. 46 on p. 192; FIG. 47 on p. 202 ;
FIG. 49 on p. 208)

SUMMARY
Trees and shrubs belonging to the single genus Saurauia which occurs in
tropical and sub-tropical regions in Asia and America. One of the most
significant anatomical characters is the occurrence of raphides. The wood
exhibits the following characters. Vessels exclusively solitary, perforation
plates scalariform with many bars, intervascular pitting scalariform to opposite
and pits to parenchyma members very long. Parenchyma diffuse,
similar,
containing raphides. Rays of 2 sizes, the larger up to 6 cells wide and
markedly heterogeneous. Fibres with distinctly bordered pits, very long.

LEAF
Dorsiventral. Clustered hairs (Fig. 49 c), visible to the naked eye, and
branched trichomes (Fig. 49 A) present, particularly on the lower surface.
Shaggy hairs (Fig. 49 D) also recorded. Two layers of hypoderm stated to
occur beneath the upper epidermis in S. napaulensis DC. Stomata generally
confined to the lower surface; some tending to be rubiaceous, but mostly
ranunculaceous. MesophylL Arm-palisade cells recorded in S. napaulensis.
Petiole, in transverse sections through the distal end, generally exhibiting
a single U-shaped vascular strand with incurved ends, but tending to be
cylindrical in certain species. About 3 medullary bundles were observed
within the main U-shaped strand in S. subspinosa Anthony (Fig. 46 H).
Raphides or styloids present in the spongy mesophyll and in the parenchy-
matous tissues and phloem of the petiole. Secretory cells observed in the
cortical region of the petiole in S. subspinosa.

Axis
YOUNG STEM
The following features were observed in S. subspinosa Anthony grown at
Kew.
Cork arising superficially. Pericycle with a composite, continuous ring
of sclerenchyma. Xylem forming a continuous cylinder, traversed by mostly
narrow and occasional broader rays; including vessels with scalariform per-
many bars. Pith broad and somewhat spongy. Raphides
foration plates with
abundant, especially in elongated sacs in the phloem.

WOOD (Fig. 47 i-j)

Vessels moderately small to medium-sized (slightly smaller or larger than
100 p mean tangential diameter); solitary, but commonly appearing as tan-
gential pairs owing members; angular; 8-15 per sq. mm.;
to overlapping
spiral thickening reported (1851) on the tips of the members. Perforation
plates typically scalariform, occasionally reticulate, oblique, with more than
20 and up to 50 (1154) bars. Intervascular pitting rare owing to the lack of
vessel groups; when present, opposite to scalariform; pits to ray and wood
parenchyma similar, often simple. Mean member length 1-5-1 -8 mm.
4594 P
2io SAURAUIACEAE
Parenchyma apotracheal, sparse to moderately abundant, scattered and
tending to form short uniseriate bands from ray to ray (Fig. 47 j) with ;

enlarged mucilaginous cells containing raphides in some species, the crystals

sometimes up to 610 p long according to Hess (959). Rays of 2 distinct sizes,
the larger up to 5-10 cells wide and more than 2 mm. high; uniseriates
numerous and composed entirely of high upright cells about 1 3 rays per mm. ; ;

heterogeneous (Kribs's Type I), commonly with 10 or more marginal rows of

upright cells; sheath cells often present; with conspicuous solid contents.
Record and Hess (1886) refer to raphides as present in the rays of some
species, but it seems to be more typical for raphides, when present, to occur
in the parenchyma only (959, 1154). Fibres with large bordered pits on both
radial and tangential walls. Walls moderately thin to moderately thick. Mean

length about 2*5 mm.

TAXONOMIC NOTES
Saurauia probably has affinities with the Theaceae and Dilleniaceae.

GENUS DESCRIBED
Saurauia,*f
*
Represented in the Kew slide collection.
t The description of the wood was based on Saurauia griffithii Dyer, the only species
examined by the author.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Buscallioni and Muscatello 321.

(ii) On Wood Structure

Hess 959, Janssonius 1154, Kanehira 1206, Record 1833, 1851, Record and Hess 1886.

59. STACHYURACEAE
(Fic. 46 on p. 192; FIG. 48 on p. 206)

SUMMARY
Shrubs or small trees belonging to the single genus Stachyurus which occurs
in the Far East. The wood is characterized by the following features. Vessels
small, solitary, commonly with spiral thickening, perforation plates scalari-
form with many bars, members very long. Parenchyma diffuse. Rays up
to 2-4 cells wide, markedly heterogeneous. Fibres with distinctly bordered
pits, sometimes with spiral thickening, long.

LEAF
Lamina dorsiventral, but with a thin mesophylL Stomata confined to the
lower surface; ranunculaceous. MesophylL Palisade tissue consisting of a
single layer of cells in S. praecox Sieb. et Zucc. Spongy tissue occupying
two-thirds of the thickness of the lamina. Midrib projecting considerably
from the lower surface; supplied by a U-shaped vascular strand. Petiole,
in transverse sections, exhibiting a large, median, arc-shaped vascular strand,
accompanied by two smaller lateral ones (Fig. 46 A). No definite ring of
 STACHYURACEAE 211

sclerenchyma in the pericyclic region of the petiole. Cluster crystals present

in the spongy mesophyll and in the parenchymatous tissues of the petiole.
A few of the parenchymatous cells of the petiole contain drops of an oily-
looking substance.

Axis
YOUNG STEM
Epidermis composed of dome-shaped cells with a thick cuticle. Cork
originating in the epidermis; composed of thin- walled, tanniniferous cells.
Cortex up to about 18 cells wide; outer part composed of smaller, more com-
pact cells than the inner part; including cells stained very darkly with haema-
toxylin. Pericycle marked by a composite, continuous ring of sclerenchyma.
Phloem and xylem appearing, in transverse sections, in the form of con-
tinuous rings, interrupted only by very numerous, uniseriate rays. Vessels
mostly tending to be in radial rows, but some isolated; somewhat angular in
transverse section, up to about 45 p in radial diameter (rather larger and more
frequently isolated in S. praecox Sieb. et Zucc. than in S. chinensis Franch.);
perforation plates scalariform, with numerous bars. Perimedullary region
consisting of a narrow zone of thick-walled cells. Pith composed of very
thin- walled parenchyma. Large cluster crystals present in the cortex and
pith,

WOOD (Fig. 48 C-D)

Vessels very small (mean tangential diameter 25-50 ft) exclusively solitary,
;

but commonly appearing as tangential pairs owing to overlapping members;

angular; sometimes with spiral thickening (1206, 2261). Perforation plates
scalariform and oblique, with about 5 bars; sometimes locally reticulate.
Intervascular pitting rare, small; opposite to transitional; pits to parenchyma
similar. length about 1-2 mm. Parenchyma apotracheal,
Mean member
scattered among the fibres; strands usually of 6-8 cells. Rays usually up to
4 cells wide, but up to 7 cells wide in Stachyurus chinensis Franch. and not
more than 2 cells wide in 5. himalaicus Hook. f. et Thorns. (1206); uniseriates
numerous and composed entirely of upright cells; heterogeneous (Kribs's
Type I), with several marginal rows of square or upright cells. Fibres with
bordered pits on all walls; sometimes with spiral thickening, e.g. in S.
chinensis i
mean length about 1*7 mm. (1206).

TAXONOMIC NOTES
A small family of uncertain taxonomic position. from most of the
It differs
Theaceae in the absence of sclerenchymatous and by the fact that
idioblasts,
the cork arises in the epidermis. It is included by Hutchinson (1113) in the
Hamamelidales, a suggestion which Tippo (2261) believes to be supported
from the evidence of wood anatomy. A comparison of the stem of Stachyurus
with those of a selection of the Hamamelidaceae certainly reveals a very
strong similarity in the general topography and in some details of structure.
A comparison of Stachyurus with some of the Flacourtiaceae indicates possible
affinities with this family also. Plants which are characterized by tannini-
ferous tissues, a somewhat spongy cortex in the young stem, xylem with small,
somewhat angular vessels with scalariform perforations, and by the presence
212 STACHYURACEAE
of large cluster crystals in the parenchymatous tissues, occur in both the
Flacourtiaceae and the Hamamelidaceae. In the circumstances it is difficult to
decide on anatomical grounds alone to which of these families Stachyurus is
most closely related.
GENUS DESCRIBED
Stachyurus.*f
* Kew
Represented in the slide collection.
f The account of the secondary xylem is based entirely on published descriptions.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gilg 774, Hutchinson 1113.

(ii) On Wood Structure

Kanehira 1206, Record 1837, 1843, 1851, Tippo 2261.

60. DIPTEROCARPACEAE
(FlG. 51 on p. 208; FIG. 53 on p. 214; FIG. 54 on p. 216; FIG. 55 on p. 218)

SUMMARY
(i) GENERAL
A
family of large resinous trees, with leathery dorsiventral leaves mainly
native of tropical Asia, the Monotoideae alone being African. The most
outstanding anatomical characteristic is the occurrence throughout the family,
except in the Monotoideae, of a branched system of resin canals, usually
lined with a delicate epithelium. In the young stem the resin canals are
always present in the pith, especially in the perimedullary region, but they
also occur in the phloem and/or cortex. Their size and distribution show a
considerable range in different genera and species, so that they are of definite
taxonomic value, but allowance must be made for the fact that transverse
sections taken at different levels in a single internode of a given species often
exhibit considerable differences in the number and distribution of the canals.
It therefore, necessary to examine sections from the corresponding region
is,
of internodes of twigs of the same age when comparing genera or species.
Even then it must be appreciated that there may be variations between
different twigs from a single species. Before the taxonomic significance of the
resin canals in the young stem can be fully assessed a thorough reinvestigation
is most desirable. The resin canal system extends from the axis via the petiole

and passes out into the lamina of the leaf beside the vascular bundles of the
veins. For the occurrence of resin canals in the secondary xylem see below.
Also important are the usually triangular phloem strands in the axis, with
the apex towards the exterior. Each phloem strand is stratified into alter-
nating fibrous and unlignified zones. The portions of the rays traversing
the phloem are also triangular with their apices towards the interior. This
structure recalls that of the Malvales. Cortical vascular bundles are also
especially characteristic of the young axis. The hairs are simple, unicellular,
sometimes tufted, or of various glandular types. Stomata, confined to the
lower surface of the leaf, are sometimes accompanied by subsidiary cells, but
 DIPTEROCARPACEAE 213
these are not always well defined. In a few instances they are rubiaceous.
The smaller veins of the leaf are vertically transcurrent. Although three
separate vascular bundles enter the base of the petiole, transverse sections
through the distal end exhibit a closed or very slightly open ring of vascular
bundles surrounding a central medullary region with accessory bundles
embedded in it. The petiolar vascular structure is always complex, and
exhibits a range of structure which might prove to be of considerable diagnos-
tic value after further investigation. Other features of specific
diagnostic
value are :
(i) The presence or absence of mucilage cells in the leaf epidermis,
(ii) The occurrence of fibres in the mesophyll.

(ii) WOOD
Vessels usually medium-sized, exclusively solitary or with some radial
multiples of 2 or 3 cells, often with a slight oblique pattern; perforations
simple, intervascular pitting alternate and moderate-sized to rather large,
pits to ray cells often large, elongated and simple members usually of medium
;

length, sometimes moderately long. Parenchyma usually abundant and

most commonly including both paratracheal (usually aliform) and apotracheal
(diffuse) types, with either predominant sometimes almost exclusively para-
;

tracheal, and then sometimes very sparse. Rays most commonly up to 4-8
cells wide, exclusively uniseriate in Marquesia and Monotes often more than
;

i mm. high; uniseriates few to numerous; rays

heterogeneous to almost
homogeneous. Fibres with simple or bordered pits, of medium length to
moderately long. Vasicentric tracheids present in many genera. Vertical
intercellular canals characteristic of all the genera except Marquesia and
Monotes radial canals occasionally present in addition.
;

LEAF
Dorsiventral. Hairs,Simple, unicellular trichomes with thick walls
(i)
and narrow lumina. (ii)
Tufted
hairs, (iii) Peltate glands, of various types
but resembling those of the Oleaceae, recorded in certain species ofAnisoptera
(Fig. 51), Doona, Hopea, Monoporandra, Parashorea, Penlacme, Shorea,
Valeria, (iv) Stalked glands with more or less lobed, unicellular heads also
present in certain species of Anisoptera, Dryobalanops, and Hopea. (v) Glan-
dular hairs, with multicellular, peltate heads in Valeria sp. Disk-shaped
extra-floral nectaries recorded on the adaxial surface of the foliage leaves
and on the abaxial surface of the stipules of Shorea sp. Cells of the epidermis
small, polygonal in surface view; palisade-like in Valeria sp.; mucilaginous
in Balanocarpus heimii King and certain species of Diplerocarpus, Doona,
Hopea, Shorea. Stomata confined to the lower surface surrounded by more
;

or less distinct subsidiary cells, the latter being particularly well defined and
parallel to the pore (rubiaceous) in some species of Balanocarpus, Shorea, and
Valeria. Medium-sized veins usually, if not always, vertically transcurrent.
Vascular system of the petiole (Fig. 53 A-D) always exhibiting a very complex
structure in transverse sections through the distal end, and including a
number of resin canals accompanying the vascular strands. Variations in the
vascular structure and in the number and distribution of resin canals are
apparently highly distinctive in different genera and species, some indication
of the range being shown in Fig. 53. (In so far as there is a generalized type
 FIG. 53. DIPTEROCARPACEAE
A, Anisoptera oblonga Dyer. Petiole x 18. B, Isoptera sumatrana v. SI. Petiole X20. C, Diptero-
carpus turbtnatus Gaertn. Petiole X 15. D, Valeria indica Linn. Petiole X 18. E, Hopea odorata Roxb.
Stem xi 3. F, Anisoptera oblonga Dyer. Stem X 13.
m.c. Mucilage cells, s.c. Secretory canals.
 DIPTEROCARPACEAE 215
it isusually characterized by an outer, usually somewhat interrupted, ring or
arc of bundles surrounding an elaborate conducting system in the medullary
region. A
thorough reinvestigation of the petiolar structure of the Diptero-
carpaceae, based on accurately named material, should yield data of the highest
diagnostic value. Meanwhile the synopsis given by Solereder (pp. 140-3)
may be consulted.) Mucilage cells sometimes in rows, situated in the
ground tissue of the petiole and/or midrib and lateral veins in Balanocarpus
heimti and certain species of&ipterocarpus, Doona, Hopea, Parashorea, Shorea,
Valeria^ and Vatica. Resin canals often found associated with the outer
part of the xylem of the larger veins and in the petiole. Crystal-idioblasts,
faith mucilaginous inner membranes and containing solitary crystals, situated

immediately below the upper epidermis in Doona, but next to the lower
epidermis in Hopea. Cluster and, more rarely, solitary crystals commonly
present and frequently abundant in the parenchymatous tissues of the
mesophyll and petiole.

Axis
YOUNG STEM (Fig. 53 E-F)
Cork, in the limited number of species investigated, originating in the
epidermis itself or the outermost layer of the cortex. Cortex sometimes con-
taining stone cells. Phloem in triangular strands (Fig. 53 E) (not always well
defined in herbarium material) with the narrow portion outwardly directed,
each group consisting of alternating bands of fibres and parenchyma when
sufficiently mature (cf. Tilia). Xylem, in transverse sections, appearing as
a more or less continuous ring. Vessels with simple perforations. Cortical
vascular bundles (Fig. 53 F), either concentric or half- moon shaped, each
accompanied by fibres in the phloem and a resin canal associated with the
xylem, are especially characteristic of the family. Mucilage cells and
cavities have been recorded in the cortex and pith of Balanocarpus heimii
King and certain species of Dipterocarpus, Doona, and Shorea, Resin canals
always present in the pith, but varying in number and position in different
genera and species, (i) A single resin canal or a group of canals produced by
the division of a single one in Dryobalanops. (ii) Confined to the perimedullary
region, being always closely associated with the primary xylem groups in the
remainder of the family. About 100 present in Dipterocarpus, and 3 to 30 in
the remaining genera. For a synopsis of the number and distribution of resin
canals in the axis in individual genera see the details given by Solereder
(pp. 140-3).

WOOD (Figs. 54-5)

Vessels typically medium-sized (100-200 mean tangential diameter) to
JJL

large (more than 200 /*),

but small (less than 100 /*) in some species of
Cotylelobium, Hopea, Monotes, and Vatica almost exclusively solitary in
;

Anisoptera, Cotylelobium, Dipterocarpus, Dryobalanops, Marquesia, Monotes,

Upuna, and Vatica; mostly solitary, but with a few pairs and short radial
multiples, in the other genera; often with an oblique pattern, particularly in
Vateria, Vatica, and in a few species of Hopea, e.g. H. nervosa King, Para-
skorea, e.g. P. lucida (Miq.) Kurz., and Shorea, e.g. S. bentongensis Foxw. and
S. bracteolata Dyer. Usually about 3-7 per sq. mm., but distinctly more
2l6 DIPTEROCARPACEAE
numerous (15-26) in Monotes and Vatica. Perforations simple, horizontal or
slightly oblique. Intervascular pitting alternate, typically moderate-sized to
rather large, but small in Balanocarpus heimii King and Hopea\ the apertures
often elongated; vestured; typically with some large, round to elongated,

FIG. 54. DIPTEROCARPACEAE

A, ffopea semicuneata Sym. B, Balanocarpus heimii King. C, Monotes africana A. DC. D, M.
kerstingii Gilg. E, Dipterocarpus crinitus Dyer. Showing small intercellular canals. F, D. rotundifolius
Foxw. G, Hopea odor at a Roxb. H, Dipterocarpus obtusifolius (Miq.) Teysm. I, D. turbinatus Gaertn.
i.e. Intercellular canal.

simple pits to raycells ; pits to wood parenchyma cells occasionally scalariform ;

large pits rare in Marquesia and Monotes. Tyloses usually very abundant,
except in Dipterocarpus, and often pitted. Mean member length usually
O'4-o*7 mm.; longest (up to i-i mm.) in Dipterocarpus.
Parenchyma typically with both scattered (apotracheal) and aliform (para-
tracheal) types and usually abundant. Both types abundant and neither
predominant in Anisoptera, Dipterocarpus p.p., e.g. Z). alatus Roxb. and D.
 DIPTEROCARPACEAE 217

grandiflorus Blanco, Hopea p.p., e.g. H. odorata Roxb., and Pentacme stamen-
sis (Miq,) Kurz. (see Fig. 55 H); the paratracheal parenchyma typically
aliform in these genera, but mostly vasicentric in several species of Diptero-
carpus. Predominantly apotracheal, as scattered cells and short tangential
lines in Anisoptera p.p., e.g. A. laevis Ridl. and A. scaphula (Roxb.) Pierre,

Cotylelobium, Marquesia, most specimens of Monotes (Fig. 540) (pre-

dominantly aliform and abaxial in some specimens), a few species of Shorea,
e.g. S. talua Roxb., and Upuna (573). Predominantly paratracheal (aliform),
(a) with moderately numerous scattered cells or short apotracheal bands
(Figs. 54 A and 55 D) in Anisoptera p.p., Balanocarpus heimii King, Doona,
Dryobalanops, Hopea p.p. (the scattered parenchyma particularly charac-
teristic ofthe Euhopea group, e.g. H. helferi (Dyer) Brandis, H. odorata
Roxb. and H. semecuneata Sym., but also present in the Pierrea group),
Shorea p.p., e.g. S, robusta Gaertn. f. and S. palembanica Miq., and Vatica;
and (b) with little apotracheal parenchyma (Fig. 55 A) in
Hopea p.p., parti-
cularly species of the Bracteataand Dryobalanoides groups, Parashorea and
Shorea p.p., e.g. S. assamica Dyer forma globifera (Ridl.) Sym., S. benton-
gensis Foxw. and S. dealbata Foxw. Very sparse and almost entirely para-
tracheal, apart from the parenchyma associated with the intercellular canals
(Figs. 54 E and 55 c), in some species of Shorea, e.g. S. dasyphylla Foxw,,
S. leprosula Miq., and S. ovalis (Korth.) Bl., and Dipterocarpus, e.g. D.
crinitus Dyer. Distinctly banded in a few species of Shorea, e.g. S. max-
welliana King and S. sericeifolia Fischer et Hutch. Chambered crystals
present in at least some species of Hopea (Dryobalanoides section), Para-
shorea, Pentacme, Shorea p.p. (particularly the 'Balau* group), and Valeria.
Druses present in Vateria. Strands most commonly of 4 cells, but sometimes
up to 8 cells in some species of Anisoptera, Cotylelobium, and Vatica and
occasionally in Dipterocarpus\ strands of 2 cells common in some species of
Hopea. Parenchyma also occurs surrounding the intercellular canals, often in
continuous bands, and occasionally as apparently terminal bands. Rays
exclusively uniseriate in Marquesia and Monotes, typically up to 4-8 cells wide
in the other genera, though occasionally slightly more than 8 cells wide in
some species of Anisoptera and Shorea and seldom more than 3 cells wide in
some species of Dipterocarpus and Doona; very variable in height even within
a genus, often more than i mm. high and sometimes up to 6 mm. uniseriates ;

moderately numerous and composed mostly of square to upright cells in

Anisoptera, Cotylelobium, Dipterocarpus, Doona, Shorea p.p., Vateria, and
Vatica, rather few and composed of square to upright and procumbent cells in
the other genera; usually between 5 and 9 rays per mm., but more numerous
(12-17) m
Marquesia and Monotes; most commonly heterogeneous (Kribs's
Types II A and B), with 1-4 marginal rows of square to upright cells, but
almost homogeneous (Kribs's Type I) in a few species of Parashorea and
Shorea, and homogeneous (Kribs's Type III) in Marquesia and Monotes ;
sheath cells sometimes present in Anisoptera, Balanocarpus, Dipterocarpus,
Hopea, Valeria^ and Vatica] with upright cells interspersed among the pro-
cumbent cells in some species of Hopea (Euhopea and Pierrea groups) and
Balanocarpus heimii (see Fig. 54 B). Solitary crystals common in the ray
cells of some species of Cotylelobium, Hopea, Pentacme, Vateria, and Vatica",
druses present in Vateria. Silica reported (794) in some species of Anisoptera,
218 DIPTEROCARPACEAE
Cotylelobium, Dtpterocarpus, and Dryobalanops and by Desch (573) as charac-
teristic of the 'Meranti Pa'ang* (Anthoshorea) group of Shorea, e.g. 5.
sericeiflora Fischer et Hutch., and all species of Anisoptera, Cotylelobium,
Dipterocarpus, and Dryobalanops] Besson records high percentages of silica

FIG. 55. DIPTEROCARPACEAE, A-H

A, Shorea multiflora (Burck.) Sym. T.S. B S. laevis Ridl. Ray. C, S. leprosula Miq. T.S. D, S.
robusta Gaertn. f. T.S. E, S. macrantha Brandis. Ray. F, S. faguetiana Heim. Ray. G, Anisoptera
laevis Ridl. Ray. H, Pentacme (Shorea) siamensis (Miq.) Kurz. T.S.
i.e. Intercellular canal.

in Dipterocarptis sp. and Anisoptera cochinchmensis Pierre. Fibres with

moderately conspicuous bordered pits in both radial and tangential walls in
Anisoptera, Cotylelobium, Dipterocarpus, Dryobalanops, Marquesia, Monotes,
Upuna, and Vatica; pits simple or indistinctly bordered and tending to be
1
mostly in the radial walls in the other genera. Walls typically thick to very
thick, except in some species of Shorea. Mean length i'Z-2-o mm.; longest
1
Janssonius (1154) describes the pits in Dipterocarpus, Hopea, Shorea, and Vatica as more
numerous on the tangential walls.
 DIPTEROCARPACEAE 219
in Dipterocarpus. Vasicentric tracheids present in Ealanocarpus heimii,
Doona, Hopea, Parashorea, Pentacme, and Shorea, and occasionally in
Dipterocarpus, the cells irregular in shape and with conspicuous bordered
pits. Cells intermediate between vasicentric tracheids and fibre-tracheids
occur round the vessels in some of the other genera, particularly Anisoptera,
Dryobalanops, and Marquesia. Intercellular canals of the vertical type
present in all the genera except Marquesia and Monotes; scattered through
the wood singly or in tangential series of 2-7 in Anisoptera, Cotylelobium,
Dipterocarpus (Fig. 54 E and i), Valeria, and Vatica, and occasionally in
tangential lines particularly in Anisoptera and Dipterocarpus; typically in
tangential lines in the other genera. The canals typically smaller than the
vessels, but occasionally as large or larger, e.g. in Anisoptera laevis Ridl.,
Dipterocarpus concavus Foxw., Shorea balanocarpoides Sym. (573), and some
species of Vatica. Radial canals occur in the rays of Ealanocarpus heimii and
some species of Shorea and Valeria (Fig. 55 F); according to Desch (573),
they are characteristic of the 'meranti damar hitam* group of Shorea, e.g. S.
balanocarpoides and a few of the species of the 'red meranti' group, e.g. S.
,

leprosula. Growth rings. The seasonal development of the growth rings in

Shorea robusta Gaertn. f. has been described by Chowdhury (414).

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
If the genera Ancistrocladus and Lophira, which were included by Bentham
and Hooker in the Dipterocarpaceae, be omitted, the remainder of the family
is anatomically very well defined. The peculiar structure of the petiole, the
presence of resin canals and cortical bundles, and the triangular phloem
groups stratified into sclerified and soft portions are especially characteristic
features. It is interesting to note the similarity of the phloem, and to a lesser
extent the structure of the petiole, to those of the Malvales.

(ii) FROM WOOD STRUCTURE

in his work on the Malayan Dipterocarpaceae has distinguished
Desch (573),
fifteengroups; these he combines into four tribes, which include the following
genera: I. Anisoptera, Cotylelobium, Upuna, and Vatica. II. Dipterocarpus.
III. Dryobalanops, and IV. Ealanocarpus, Hopea, Parashorea, Pentacme, and
Shorea. He suggests that the Euhopea section of Hopea, together with Hopea
ferrea Lannesan and Ealanocarpus heimii King may possibly represent a
further separate tribe. He regards tribes I and II as distinctly more primitive
than IV.
The different species of Shorea exhibit a wide range of variation, and Desch,
commenting that 'unlike most other genera of the Dipterocarpaceae, Shorea
is not a natural genus' distinguishes six groups, which he treats as if they
were separate genera.
Monotes and Marquesia are distinguished from the rest of the family by the
absence of intercellular canals and by their exclusively uniseriate rays.
Vestal (2329), in a study of the Guttiferae and their allies, has suggested
that the family Dipterocarpaceae 'stems in the Exalbuminosae of the Ochna-
ceae, with the connecting genus probably Lophira'. He also holds the view
that 'although the Dipterocarpaceae have occasionally been allied to the
220 DIPTEROCARPACEAE
Guttiferae due to the presence of the secretory canals*, the latter should be
regarded as a parallel development.

ECONOMIC USES
of the members of this family are medium-sized to large trees and
Most
their timbers are very widely used for general purposes in the countries in
which they grow. The timbers vary from soft and light, e.g. the Malayan
Red Meranti (Shorea leprosula Miq. et al.\ to hard, heavy, and durable woods
such as are furnished by some species of Hopea, some forms of 'Balau'
(Shorea spp.), and Chengal (Balanocarpus heimii King). few of them are A
moderately well known on world markets, e.g. Meranti (Shorea spp.) and
Keruing (Dipterocarpus spp.) from Malaya, Apitong (Dipterocarpus spp.)
from the Philippines, the Serayas and Lauans from Borneo and the Philippines
(Shorea and Parashorea spp.); and Gurjun (Dipterocarpus griffithii Miq. and
D. turbinatus Gaertn. f.) from India; but on the whole the woods tend to lack
distinctive character and none of the species furnish outstanding special-
purpose timbers. The large size and good form of the logs from most species
and a tendency to gregariousness help to increase the popularity of these
woods as general-purpose timbers.
Dammars and resins are obtained from the trunks of species of Balano-
carpus, Hopea, Shorea, and Vateria, and oil and fat from the kernels or nuts
of Isoptera and Shorea. Camphor occurs as a natural deposit in the wood of
Dryobalanops.
GENERA DESCRIBED
(i) GENERAL ANATOMY
Anisoptera,* Balanocarpus,* Dipterocarpus,* Doona,* Dryobalanops,*
Hopea,* Isoptera,* Monoporandra,* Parashorea,* Pentacme,* Shorea,*
Stemonoporus,* Vateria,* Vatica.*
* Kew
Represented in the slide collection.

(ii) WOOD STRUCTURE

Anisoptera, Balanocarpus, Cotylelobium, Dipterocarpus, Doona, Dryo-
balanops, Hopea, Marquesia, Monotes, Parashorea, Pentacme, (Upuna),
Vateria, Vatica.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gilg 771.

(ii) On Wood Structure

Beekman 167, den Berger 179,
Bailey 78, Bancroft 125, 126, 127, 128, Bausch 154,
180, 182, 183, Besson 186, Bianchi 194, Chowdhury 411, 4x4, 417, Clarke 428, Desch
569, 570, 571, 573, 574, 575, Foxworthy 704, 705, Gonggrijp 794, Gupta 848, 849,
Hale 870, Howard 1088, Jamsonius 1154, Jones 1191, Kanehira 1206, 1209, Lecomte
1334, M&iiaud 1491, Pearson and Brown 1679, Record 1780, 1783, 1801, 1809, 1843,
1851, Reyes 1927, Schneider 2044, Stone 2206, Vestal 2329.
 (221)

61. ANCISTROCLADACEAE
(Fic. 52 on p. 208)

SUMMARY
A
family of scandent shrubs, consisting of the single genus Ancistrocladus.
They occur for the most pan in tropical Asia, although at least one species is
known from the African tropics.

LEAF
Dorsiventral. Peltate glands (Fig. 52), visible to the naked eye and which
secrete wax, occur in depressions in the surface of the leaf, the heads of the
glands being close to the leaf surface conceal the pits in which they are situated.
Stomata (Fig. 52) confined to the lower surface; actinocytic. Hypoderm
sometimes present. Mesophyll including more or less distinct palisade
tissue. Veins with vascular bundles vertically transcurrent. Three separate
bundles enter the base of the leaf, but unite to form a closed or more or less
open tube throughout the midrib; generally accompanied by 5-19 accessory,
inversely orientated bundles situated in the outer part of the zone of fibres
around the main vascular strand.

Axis
YOUNG STEM
Cork arising at various levels in the cortex or outer part of the pericycle.
Pericycle at first parenchymatous, but provided with a continuous or
interrupted ring of stone cells when older. Phloem not stratified into fibrous
and parenchymatous portions as in the Dipterocarpaceae, but containing
stone cells, often branched and sometimes vertically elongated. Groundwork
of xylem composed of fibre-tracheids. Vessels mostly solitary, tending to be
in radial or oblique rows in some species. Wood parenchyma well developed,

usually in tangential bands. Solitary and clustered crystals present in the

primary cortex and pith. Van Tiegem's record of secretory canals in the
pericycle was an error. Isolated thick-walled secretory cells also occur
in the primary cortex.

TAXONOMIC NOTES
The genus Ancistrocladus was included by Bentham and Hooker in the

Dipterocarpaceae, but it is now generally placed in a special family.

The
anatomical structure confirms that it differs somewhat from the Diptero-
carpaceae. The position of the family is not well established.

GENUS DESCRIBED
Ancistrocladus.

LITERATURE
On General Anatomy
Gilg 778.
 (222)

62. CHLAENACEAE
(FiG. 50 on p. 208)

SUMMARY
Shrubs or trees, confined to the Mascarene Islands. The wood exhibits
the following characters. Vessels exclusively solitary, perforations simple,
intervascular pitting alternate. Parenchyma apotracheal, diffuse. Rays
exclusively uniseriate. Fibres very to extremely short.

LEAF
Leaf dorsiventral. Hairs (Fig. 50). (i) Simple, unicellular, thick-walled,
(ii)Unicellular, thin- walled, twisted at the apex, (iii) Two-armed, unicellular,
(iv) Peltate, (v) Multicellular, glandular, of various types. Cells of the
epidermis polygonal as seen in surface view. Stomata surrounded by a
rather large number of ordinary epidermal cells; sometimes in depressions,
e.g. in Schizoclaena (Fig. 50 A); confined to the upper surface according to
Gerard (757) and to the lower surface according to Solereder. Hypoderm
of 1-2 layers sometimes present well developed in Schizochlaena. Mesophyll
;

including sclereids according to Beauvisage (163). Vascular bundles of

the veins always accompanied by sclerenchyma; vertically transcurrent in
Rhodochlaena, Sarcochlaena, and Schizochlaena. Petiole examined only in
Eremochlaena by Beauvisage (163), according to whose account the vascular
strand appears, in transverse sections, in the form of a closed ring surrounding
a fairly large, central, medullary strand. Mucilage cells present in the
mesophyll; those in Eremochlaena described by Beauvisage as very large.
Cluster crystals recorded by the same author in the mesophyll and cortical
region of the petiole in Eremochlaena.

Axis
YOUNG STEM
Primary cortex including sclerotic cells in Eremochlaena and Schizochlaena.
Pericycle containing groups of fibres, fairly widely separated in most genera,
but almost continuous in Sarcochlaena. Xylem and phloem forming closed
cylinders, traversed by narrow rays. Primary phloem containing 1-4 layers of
fibres. Secondary phloem stratified into fibrous and parenchymatous portions.
Vessels isolated or in small groups, up to about 40 /z in diameter, circular or
oval perforations simple. Paratracheal parenchyma sometimes well developed.
;

Wood fibres provided with thick walls and bordered pits. Perimedullary
region sclerosed in Eremochlaena. Pith containing sclerified cells of varying
abundance, the sclereids especially numerous in Sarcochlaena. Pith almost
entirely collenchymatous in Xerochlamys. Cluster crystals always present in
the soft tissues, especially numerous in Xerochlamys. Mucilage cells, vary-
ing in frequency in different species, generally occur in the primary cortex
and pith. Mucilage cells tend to disappear as the stems grow old, and are
sometimes absent from Schizochlaena.
 CHLAENACEAE 223
WOOD 1

Vessels of medium size (100-200 ^ mean tangential diameter), exclusively

solitary, 8-16 per sq. mm., perforations simple, intervascular pitting alternate.
Parenchyma apotracheal, as isolated cells scattered among the fibres. Rays
exclusively uniseriate, less than i mm.
high, 16-18 per mm. Fibres with
moderately thick walls; mean length about 0*5 mm.

TAXONOMIC NOTES
The
presence of mucilage cells in the cortex, and the complex structure of
the vascular system of the petiole suggest that this family may have affinities
with the Dipterocarpaceae. It differs from the Dipterocarpaceae, however,
in the absence of resin canals. Gerard (757) points out that the mucilage cells
of the pith recall those which occur in certain of the Malvaceae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Although they are not all mentioned by name in the text, the above
description of leaf and young stem is based mainly on Gerard's (757) study
of the following genera: Eremochlaena (especially E. humboltiana Baill.),
Leptochlaena, Rhodochlaena, Sarcochlaena, Schizochlaena, Xerochlamys,
Xylochlaena.

(ii) FOR WOOD STRUCTURE

(Leptochlaena), (Rhodochlaena).

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gerard 757.

(ii) On Wood Structure

Lecomte 1333, Perrot 1694.

63. MALVACEAE
(Fio. 56 on p. 224; FIG. 57 on p. 228; FIG. 60 on p. 244)

SUMMARY
(i) GENERAL
Herbs and shrubs, widely distributed throughout the world. The stellate
hairs are especially characteristic, but simple unicellular or uniseriate, peltate
and glandular trichomes also occur. Mucilage cells are very common in
the parenchymatous tissues, and should not be confused with the secretory
glands with brown/contents which also occur, but less frequently. The cells
of the epidermis, especially in the leaf, sometimes have mucilaginous inner
walls and appear as transparent dots. The leaf is. dorsiventral in most
instances although exceptions occur. Stomata are present on both surfaces.
The petiole usually contains a circle of about 6, but sometimes more,
separate, collateral vascular strands; but in a few instances the xylem and
1
Based entirely on the descriptions given by Lecomte (1333) and Perrot (1694).
224 MALVACEAE
phloem appear, in transverse sections, as a continuous ring. Cork in the axis
arises in the epidermis itself, or in the outermost part of the cortex. Stone
cells are rarely present in the cortex. The pericycle of the young stem

usually contains strands of fibres opposite the phloem groups, although in a

few instances there is a tendency for the fibres to form a closed cylinder. The
phloem, in most instances, is especially characteristic, consisting of triangular
strands as seen in transverse sections with the bases towards the xylem, and
stratified tangentially into alternating fibrous and non-fibrous bands. The

FIG. 56. MALVACEAE, A-E; STERCULIACEAE, F

A, Stellate hair of Althaea offidnalis L. B, Stellate hair of Malachra radiata L. C, Stellate hair of
Malvastrum asperrimum Garcke. D, Peltate hair of Durio lanceolatus Mast. E, Hairy covering of
Malvastrum capense Garcke. A, After Vogl; B-C and E, After Kuntze; D, After CX Bachmann.
F, Peltate hair of Heritiera macrophylla Wall. After CX Bachmann.

primary medullary rays, where passing through the phloem, are also triangular
but with the apices towards the xylem. The triangular shape of the phloem
groups is not very well defined in a few species. The xylem of the young
stem usually forms a practically continuous ring, except where broken by the
rather broad primary rays. The pith generally consists of thin-walled
parenchyma, but in a few genera there are fibres in the perimedullary region.
Medullary bundles recorded in one species of Abutilon.

(ii)
WOOD
Vessels small to medium-sized; semi-ring-porous and withspiral thicken-
ing in some genera; perforations simple, intervascular pitting small and
alternate, pits to parenchyma sometimes simple and larger than the inter-
vascular pitting; members of medium length to very short. Parenchyma
 MALVACEAE 225
(a) in the Malveae and Ureneae, predominantly paratracheal, vasicentric to
confluent, (b) in the Hibisceae, predominantly apotracheal, in numerous,
often short bands 1-2 cells wide, accompanied by some paratracheal paren-
chyma; occasionally intermediate between paratracheal and apotracheal;
terminal parenchyma sometimes present; fusiform cells often present; usually
storied. Rays. Often tending to be of 2 distinct sizes; multiseriate rays
usuallyup to 4-9 and occasionally up to 23 cells wide; low to high; uniseriates
numerous to few; typically heterogeneous and often composed of mingled
upright and procumbent cells, but occasionally almost homogeneous; com-
monly with sheath cells druses occasionally present storied when short and
; ;

sufficientlynumerous. Fibres with simple pits, sometimes storied, of medium

length to moderately long. Vascular tracheids reported in 3 genera. Inter-
cellular canals of the vertical traumatic type present in a few species with
;

'gossypol cavities' in some species of Gossypium and possibly in Thespesia.

LEAF
Generally dorsiventral, but centric structure recorded in Malva parviflora
Linn., occasionally isobilateral (see 'Mesophyll' below). Covered externally
with characteristic stellate and tufted hairs (Fig. 56 A-D and 60 A), but small
peltate scales, bristles and glandular hairs are to be found, several of these
types sometimes occurring together in a single species. Peltate hairs recorded
in Hibiscus and Thespesia. X-shaped hairs recorded on the lower surface of
Malvaviscus arboreus Cav. by Gehrig (753). Multicellular, capitate glandular
hairs of various sizes and shapes occur in species of Gossypium Hibiscus,
,

Kokia, Lagunaria, Thespesia smaller glandular hairs, composed of fewer cells,

;

in species of Althaea, Hibiscus, Malva, Pavonia; pitcher- or flask-shaped

glandular hairs frequent in Malva, Malvastrum, Wissadula; depressed glands
recorded in Althaea officinalis Linn. Extra-floral nectaries, consisting of
groups of secretory trichomes, situated either on the surface or in concavities
or depressions, recorded on the lower side of the midrib or three principal
'veins in certain species of Cienfuegosia, Decaschistia, Dicellostyles, Gossypium
(not in G. tomentosum Nutt. ex Seem.), Hibiscus, Ingenhouzia, Julostylis,
Kydia, Thespesia, Urena. Their shape and distribution stated by Janda(i 143),
Koernicke ( 1 265), and Stanford and Viehoever (21 84) to be of limited taxonomic
value owing to the variation occurring within a single species. Small, inflated,
refractive, spherical, multicellular glands were observed especially near the
veins on the lower surface of the leaves of Hibiscus manihot Linn, growing at
Kew; the appearance being very striking and beautiful as seen under the
binocular microscope. (These are probably similar to the 'intumescences' on
the leaves of H. vitifolius Linn, described by Dale (534). Dale's experiments
led to the conclusion that the intumescences are pathological structures
formed only in a sufficiently humid atmosphere and when the temperature
and light conditions are appropriate.) Glandular hairs stated by Solereder
sometimes to serve as hydathodes. The hairs on the leaf of Abutilon sp.
reported by Luckan (1399) to be capable of absorbing water. For further
details of hair structure see articles by Roiisseau (1963), Sabnis (1977),
Ulbrich (2308), Youngman and Panda (2499, 2500). Cells of the epidermis
with straight or undulating anticlinal walls; some of them, especially on the
upper surface, specialized for the secretion of mucilage, the large cells
4594
Q
226 MALVACEAE
sometimes penetrating into the mesophyll, e.g. in a few species of Althaea,
MalvaviscuSy and Pavonia. Mucilage cells in the epidermis inconspicuous
or lacking in Erioxylum, most species of Gossypium (but fairly numerous
in G. kirkii Mast, and G. klotzschianum Anderss.), Kokia, fairly numerous
in species of Hibiscus, Lagunaria, Thespesia. Scattered epidermal cells with
purplish or reddish cell sap present in most species of Gossypium^ more
widely distributed in plants with reddish than with green leaves. Stomata
ranunculaceous; always present on the lower and frequently on the upper
surface as well; present on both surfaces in most species of Gossypium,
although always more numerous on the lower side, the relative frequency
on the upper and lower surface respectively varying from species to species.
Hypoderm said to be well developed in Lagunaria patersonii G. Don. Water-
storage cells stated by Alexandrov (12) to occur in Gossypium. Mesophyll.
Palisade tissue recorded as present on both surfaces in certain species of
Althaea and Gossypium. Hibiscus cannabinus Linn, from Java is stated by
Magitt (1414) to be provided with palisade tissue towards both surfaces, but
palisade tissue observed only towards the upper side of leaves of the same
species from Persia. Isolated sclerenchymatous elements recorded in the
mesophyll of certain species of Goethea and Pavonia. Vascular bundles of the
larger veins seldom accompanied by sclerenchyma except in the Ureneae
and in Hibiscus\ smaller veins vertically transcurrent or embedded in the
mesophyll. Petiole, in transverse sections through the distal end, generally
exhibiting isolated collateral bundles arranged in a circle, e.g. in Abutilon
(strands rather close together), Althaea (Fig. 60 c), Malva, Pavonia, Sida,
Sidalcea] but the xylem forms a closed ring in Abutilon (pro parte), Anoda,
Hibiscus (Fig. 60 B), Lavatera, Sphaeralcea. A solitary arc-shaped bundle
present in Hoheria sexstylosa Colenso. The vascular system, in transverse
sections through the middle of the petiole of Gossypium, has the form of a
closed ring, an arc, or a circle of isolated bundles. The number of bundles,
where these are separate, varies in different species, but there are often about
6; sometimes of 2 distinct sizes in Lavatera and Malva. Mucilage re-
ceptacles occur in the mesophyll, e.g. in a few species of Hoheria and
Plagianthus. Lysigenous 'secretory glands' (cavities) with brown contents,
not to be confused with the mucilage cells just described, sometimes appear-
ing as transparent or dark opaque spots in the leaves, present in Gossypium.
(When the glands are present in parts of the plant exposed to the light, the
surrounding envelope of cells is believed to contain anthocyan, partly or
wholly in the form of the glucosides quercimeritrin or isoquercitrin, or in
ethereal oils, resins, or tannins. Gossypol occurs in the glands when not
exposed to light.) Similar glands, but fewer in number, also recorded by
Stanford and Viehoever (2184) in Cienfuegosia, Erioxylum, Ingenhouzia,
Pavonia, Sida, and Thespesia. Crystals, mostly clustered but sometimes soli-

tary, generally present; druses often very abundant although none observed
atKew in a few species of Malva. H-shaped crystals, in addition to ordinary
druses, recorded in the Ureneae by Gehrig (753).
Exposure of seeds or very young seedlings of cotton to X-radiation under
suitable conditions is stated by Singh, Choudri, and Kapoor (2106) to induce
the petioles during subsequent development to become grooved or dorsiven-
tral, and the vascular bundles to assume the form of a closed ring instead
 MALVACEAE 227
of the usual circle of separate bundles. Variegation, puckering, and the
development of tumours and variations in the vascular system of the lamina
have also been induced by the same treatment. For detailed structure of the
leaf in different species of Lavatera and Malva see Rousseau's
(1963) article
and for the leaf of the family in general Gehrig's (753) thesis.

Axis
YOUNG STEM (Fig. 60 F~G)
Cork arising in the epidermis or the outermost part of the cortex. Cortex
generally with collenchymatous and parenchymatous zones. Stone cells
present in the cortex of certain species of Goethea, Pavonia, and Sphaeralcea.
Pericycle always containing fibres, usually in groups external to the phloem,
but sometimes tending to form a ring. Phloem in triangular strands with the
narrow portions outwardly directed; more or less distinctly stratified owing
to the presence of tangential bands of fibres. The triangular shape of the
phloem groups is more apparent in some genera than in others. The peri-
cycle and phloem of many Malvaceae yield commercial fibres (see 'Economic
Uses'). The primary medullary rays also triangular where traversing the
phloem but with the base towards the exterior. Xylem forming a practically
continuous ring in most instances, except where traversed by the broad,
primary, medullary rays, e.g. in Malva sylvestris Linn. (Fig. 60 G). Vessels
with simple perforations. Pith generally consisting of thin-walled paren-
chyma, but containing sclerenchymatous fibres situated on the inside of the
xylem in certain species of Hoheria, Matisia, and Plagianthus. Mucilage
cells and cavities present in the cortex and pith; especially numerous in
species of Althaea, Anoda, Cristaria, Hoheria, Kitaibelia, Lavatera, Malope,
Malva, Napaea, Palava, Sida; less numerous in species of Abutilon, Fugosia,
Goethea, Gossypium, Hibiscus, Kydia, Lagunaria, Malachra, Malvaviscus,
Pavonia, Sidalcea, Sphaeralcea, Thespesia, Urena, Wissadula. Secretory
glands (cavities) with brown contents, similar to those in the leaf, occur in
the cortex of Cienfuegosia, Erioxylum, Gossypium, Ingenhouzia, Pavonia, Sida,
Thespesia. (For further particulars see 'Leaf '.) Mucilage canals recorded in
the pith of a few species of Hibiscus, Kokia, Thespesia. Cluster crystals very
common and frequently abundant in the cortex, phloem, rays, and pith, but
the frequency varies in different species; those in the phloem sometimes
smaller than those in the other tissues; probably occurring in most genera,
and definitely observed in species of Abutilon, AltJiaea, Gossypium, Hibiscus,
Hoheria, Lagunaria, Malva, Pavonia, Sida, Sidalcea, Sphaeralcea. Large
solitary crystals sometimes present as well as the clusters although less
frequently, e.g. in species of Hoheria and Lagunaria. Crystals sometimes more
abundant in very young than in slightly older stems, e.g. in the pith of very
young stems of Abutilon insigne Planch. Small spherical outgrowths or
papillae, with walls at first containing pectic substances but subsequently
becoming cutinized, described by Blackwell (202) as arising from the pith of
Althaea rosea Cav. when wounded. Callus formation in wounded stems of
Hibiscus rosa-sinensis Linn, stated by Sharpies and Gunnery (2084) to
originate from the exposed medullary ray cells, the cambium playing a part
only after the initial formation of a callus cushion.
 FIG. 57. MALVACEAE
A, Kydia calydna Roxb. B, Cephalohibiscus peekelii Ulbricht. C, Hibiscus elatus Sw. D, H. elatus
Sw, E, Gossypwm thurberi Tod. F, G. obtusi/olium Roxb.
G, Abutihn niveum Griseb. H, Hoheria
populnea A. Cunn I, Mo/vawcm mo//w DC. J, Thespesia populnea
.
(L.) Soland. K, Gossypium morilli
Cook et Hak. L, Thespesia populnea (L.) Soland.
 MALVACEAE 229
WOOD (Fig. 57)
Vessels small (less than 100 //, mean tangential diameter) or medium-
sized (100-200 p), the latter mostly less than 150 /x in diameter; extremely
small (less than 25 /x) in Sphaeralcea (2373), very small (25-50 p) in some
species of Abutilon, Alyogyne, Althaea (2373), Cienfuegosia (2373), Erioxylum,
Hoheria, Lavatera, Malvastrum (2373), Pavonia (2373), Plagianthus, Shantzia
(2373), Tetrasida, and Wissadula (2373), moderately small (50-100 p) in
some species of Abutilon, Bastardiopsis, Gossypium, Hibiscus, Hoheria (2373),
Lagunaria, Malvastrum, Pavonia, Shantzia, Sida, and Urena, largest (about
150 fji) in some species of Hibiscus, Kydia, and Paritium, and, according to
Solereder, in Plagianthus sidoides Hook. solitary, in irregular clusters and in
;

radial multiples of 2 or 3 multiples commonly of 4 or more cells present in

;

some species of Abutilon, Hibiscus, Lavatera, Malvastrum, Malvaviscus,

Pavonia, Sphaeralcea, and Urena and sometimes producing a radial pattern
in Lavatera and Malvastrum; sometimes with a rather vague tangential
pattern in Abutilon and Lavatera, entirely limited to the tangential arcs of
parenchyma separating the fibres into groups in Hoheria (Fig, 57 H) and
Plagianthus and intermediate in Sphaeralcea; except in the woods with pro-
nounced pattern or multiples, seldom more than 20 per sq. mm.; 5-10 per
sq. mm. in some species of Bombycidendron, Erioxylum, Gossypium, Hibiscus,
Paritium, and Thespesia, fewer than 5 per sq. mm. in Cephalohibiscus, Gossy-
pium, Hibiscus, Kokia (2373), Kydia, Thespesia (2373) and Wercklea, up to
30 per sq. mm. in Tetrasida; ring-porous or semi-ring-porous in some species
of Gossypium, Hibiscus, Malvastrum, Paritium, Sida (2430), Sphaeralcea, and
Thurbergia; spiral thickening observed in the small vessels of some species
of Abutilon, Hibistus (rare), Hoheria, Malvastrum, Plagianthus, Sida (1851),
Sphaeralcea, Thurberia, and Wissadula (2373). Perforations simple. Inter-
vascular pitting alternate, sometimes with some much elongated slit-like pits,
e.g. in Malvastrum p.p., and sometimes striated owing to coalescent apertures,
e.g. Gossypium dryparioides Seem.; Webber (2373) describes the pitting as
alternate, opposite or occasionally scalariform in part; small to minute; pits
to ray and wood parenchyma typically similar to the intervascular pitting but
sometimes simple, and with some horizontally or obliquely elongated pits in
some species of Abutilon, Cephalohibiscus, Hoheria, Lavatera, Malvaviscus,
and Wercklea and particularly in Pavonia p.p., Plagianthus, and Sphaeralcea,
Most of the genera with occasional deposits of gum (2373); tyloses observed
only in Bombycidendron and Hibiscus. Mean member length 0-2-07 mm.,
mostly, according to Webber (2373), 0-33-0-45 mm. Parenchyma rather
scanty to abundant; predominantly paratracheal in the Malveae and Ureneae:
vasicentric to slightly aliform in Abutilon p.p., Malvastrum p.p., Malvaviscus,
Pavonia, Sida p.p., and Urena, aliform to confluent in Abutilon p.p., Bastar-
diopsis,Lavatera, Sida p.p., and Tetrasida, confluent in Hoheria, Malvastrum
p.p., Plagianthus and Sphaeralcea', with numerous biseriate apotracheal bands
predominating in Kydia (Fig. 57 A); in Lavatera, in addition to the para-
tracheal parenchyma, there are occasional broad bands of very thin-walled
parenchyma cells, sometimes numerous locally or linking up to form aground-
tissue in which, in cross-section, the groups of fibres and vessels appear as
small islands or bands in Urena there are numerous small groups of thin-
;

walled cells (apparently composed of fusiform parenchyma cells) scattered

230 MALVACEAE
over the cross-section in a manner similar to that of the strands of included
phloem in Strychnos\ single specimens of Afalvaviscus mollis DC. and Pavonia
spicata Cav. were observed to have predominantly apotracheal parenchyma
of the type described below for the Hibisceae and to differ in this respect from
other species of these genera. Apotracheal parenchyma always present and
often predominant in most of the Hibisceae: in numerous, short bands
1-2 cells wide in Alyogyne, Cephalohibiscus, Dicellostyles (2373), Erioxylum,
Gossypium (rare in some species), Hibiscus, Juliostylis (2373), Paritium,
L), and Wercklea, and with more regular and more
Shantzia, Tfiespesia(ig. 57
continuous bands in some specimens of Gossypium and Hibiscus the para-
;

tracheal parenchyma more prominent and tending to be aliform in some

of the species of Hibiscus, Paritium, and Thurberia\ particularly in the
late wood, in broader bands, intermediate between metatracheal and con-
fluent in Lagunaria and possibly in Kokia (2373) vasicentric or aliform and
;

without apotracheal parenchyma in Bombycidendron. Terminal parenchyma

present in some genera. According to Webber (2373), frequently with
crystals in Althaea, Bombycidendron, Hibiscus, Kokia, and Thespesia, and
with yellow, reddish, or brown gum in Althaea, Gossypium, Hibiscus, Kokia,
Kydia, Malvaviscus, and Thespesia. Strands most commonly of 2-4 cells,
tending to be mostly of 2 cells in the Malveae, though Webber states that they
are up to 8 cells in Kydia. Fusiform cells moderately common in many of the
genera. Storied in nearly all the species, except in very small stems. Rays
very variable in type, ranging from (a) high multiseriate rays composed mainly
of narrow upright cells, together with numerous uniseriate rays, to (b) large
homogeneous rays with few uniseriates, or (c) short, heterogeneous, storied
rays. In the Ureneae Malvaviscus, Pavonia, and Urena the multiseriate
rays are high, 2-5 cells wide, with relatively few procumbent cells, which are
scattered among the upright cells as described by Chattaway (272) for her
Tterospermum* type of tile cell (Fig. and the uniseriate rays are
57 i),

numerous, composed entirely of upright and moderately high (Kribs's

cells,
Het. Type I). In the Hibisceae similar types, though with wider multiseriate
rays, occur in a few species of Gossypium (Fig. 57 K) and Hibiscus and in
Wercklea, but in most of the woods of this sub-family that have high rays, the
procumbent cells are more numerous, the upright cells are shorter and some-
times almost square (particularly in Lagunaria) and the uniseriate rays are
few, often composed of procumbent cells and commonly only 2 or 3 cells
high (Kribs's Het. Type II B), e.g. in Erioxylum, Gossypium p.p., Hibiscus,
Lagunaria, Shantzia, Sphaeralcea, and Thurberia; in other genera or species,
e.g. Bombycidendron, Cephalohibiscus, Hibiscus p.p., and Paritium, the rays
are low, 2-3 (occasionally 4) cells wide, heterogeneous with i or 2 marginal
rows of upright cells, with a central part composed entirely of procumbent
cells, and storied (see Fig. 57 B and c); Thespesia is intermediate, with most
of its rays of the latter type, but with some high rays, in which procumbent
and upright cells alternate (see Fig. 57 j). In the Malveae the tendency is
towards more homogeneous rays. High multiseriate rays and relatively few
uniseriates, both with alternating procumbent and upright cells, occur in
Abutilon, Lavatera, Malvastrum, Sphaeralcea, but the upright cells are barely
more than square in Abutilon and Lavatera', in Hoheria and Plagianthus
almost all the rays are very large and composed of small procumbent cells
 MALVACEAE 231
surrounded by sheath cells, small rays being scarce and usually uniseriate;
in Bastardiopsis and Kydia the large rays are shorter and with less conspicuous
sheath cells, and small rays up to 3 cells wide are numerous, storied and
heterogeneous; in Sida all the rays are homogeneous and the storied small
rays are nearly all uniseriate in Tetrasida most of the rays are only i story
;

high, 2-4 cells wide, homogeneous and storied, and uniseriates are rare. Rays
2-3 cells wide in some species of Bombycidendron, Cephalohibiscus, Gossypium,
Hibiscus, Malvavisctts, Paritium, and Pavonia; up to 10-14 ce ^ s wide m some
species of Bastardiopsis, Hibiscus, Lagunaria, and Malvastrum; up to 17 cells
in Hoheria and 23 cells in Plagianthus up to 4-9 cells wide in the other genera
;

and in some species of Gossypium, Hibiscus, Paritium, and Pavonia; commonly

with a tendency to be of 2 distinct sizes in the woods with larger rays; exceed-
ing i mm. in height except in some species of Abutilon, Bastardiopsis,
Bombycidendron, Cephalohibiscus, Kydia, Paritium, Shantzia, Tetrasida, and
Thespesia; fewer than 5 per mm. in some species of Hibiscus, Hoheria,
Lagunaria, and Plagianthus, 12-15 per mm. in Abutilon, Malvaviscus, Pavonia,
Tetrasida, and Urena, 6-n per mm. in the remainder; homogeneous in Sida
and almost homogeneous in Bastardiopsis and Tetrasida', markedly hetero-
geneous in most of the other genera; with square or upright cells intermingled
with procumbent cells and tending to alternate in groups (see Fig. 57 F, i, j,
and K), as in Chattaway's (272) Tterospermum* type of tile cell, in Abutilon
p.p., Alyogyne, Erioxylum, Gossypium, Hibiscus p.p. (a few specimens only),
Lavatera, Malvaviscus, Pavonia, Shantzia, Sphaeralcea, Thespesia, Thurbera,
Urena, and Wercklea; with square or upright cells (other than sheath cells)
limited to i or 2 marginal rows in Abutilon p.p., Bastardiopsis, Bombyciden-
dron, Cephalohibiscus, Gossypium p.p. (a few specimens only), Hibiscus p.p.,
Hoheria, Kydia, Paritium, and Plagianthus', with sheath cells in Abutilon p.p.,
Alyogyne, Erioxylum, Gossypium p.p., Hibiscus p.p., Hoheria, Malvastrum,
Thespesia, Thurbera, and Urena', very commonly containing crystals, with
druses in some species of Althaea (2373), Lavatera, Paritium, Thurbera, and
Sphaeralcea and sometimes with gummy contents. Webber (2373) records
the occurrence of gossypol cavities in some species of Gossypium. All the rays
storied in Abutilon p.p. (according to Webber, but not in any of the 6 species
examined), Bombycidendron, Hibiscus p.p., Paritium, Tetrasida, and Thespesia
(2373); with the small rays storied in Abutilon (rare), Bastardiopsis, Dicello-
styles (2373), Hibiscus p.p., Juliostylis (2373), Lagunaria, Sida, and Thespesia
p.p. Fibres typically with small simple pits, but according to Webber (2373),
fibre-tracheids form a considerable portion of the wood in Pavonia and
Wercklea ;
numerous in the radial walls of Bastardiopsis and Sida.
pits very
With thin walls insome species of Cephalohibiscus, Gossypium, Hibiscus,
Lavatera, Malvaviscus, Pavonia, and Urena, and with very thin walls in
Wercklea\ with thick walls in species of Abutilon, Bombycidendron, Hoheria,
Kydia, Plagianthus, Sida, Tetrasida, and Thurbera. Commonly storied in
woods in which the parenchyma is distinctly storied. Mean length 0-36-
2-33 mm. (2373); usually of medium length (0-9-1-6) but sometimes longer,
e.g. in Bombycidendron, Hibiscus p.p., and Hoheria. Vascular tracheids
reported by Webber (2373) in Malvastrum, Plagianthus, and Wissadula.
Intercellular canals of the traumatic vertical type reported in species of
Hibiscus and Urena by Webber (2373) who suggests that the secretory
232 MALVACEAE
cavities observed by Hohnel in Thespesia populnea (L.) Soland. may be of this

type or may possibly be 'gossypol cavities'. Growth ring formation in Hibiscus

has been investigated by Coster (481) in Java.

ROOT
Most completely described by Gore and Taubenhaus (796) for Gossypium
as follows. Secondary xylem constituting a large proportion of the root;
vessels solitary or in groups of 2-4, frequently containing conspicuous tyloses ;

wood fibres with thick walls, tapering ends, and bordered pits rays 4 cells
;

wide and many rows of cells tall; secretory glands, similar to those of the
stem and leaf, present in the portions of the rays situated in the phloem.
Concentric layers of cork are formed around a single vessel or group of
vessels in the xylem of Althaea officinalis Linn. Fibres occur in the phloem
of all the investigated genera. This character is stated by Solereder to be rare
in other families with the exception of the Sterculiaceae and Tiliaceae. The
root of Althaea officinalis is described under 'Economic Uses', and some
particulars concerning the roots of Gossypium and Hibiscus given in the same
section.

ANOMALOUS STRUCTURE
Medullary vascular bundles recorded by Sabnis (1977) in the pith of
Abutilon fruticosum Guill et Perr.

TAXONOMIC NOTES
(i)
FROM GENERAL ANATOMY
The Malvaceae constitute a homogeneous group from the anatomical stand-
point. There is evidence that they have very close affinities with the Bom-
bacaceae, and also, but to a less pronounced extent, with the Tiliaceae and
Sterculiaceae. It will be recalled that certain genera described under
Flacourtiaceae (see p. 126) are believed to have affinities with the Tiliaceae.

(ii)
FROM WOOD STRUCTURE
The tribe Hibisceae (Fig. 57 D and L) appears tobe clearly separated by its
parenchyma from the Ureneae and Malveae (Fig. 57 G and H), but Webber
(2373) states categorically that the distinction is not valid, at any rate on the
basis suggested by Dumont. Webber has shown that the vessel characters
indicate that the family is a fairly advanced group, and has drawn attention to
the unusually wide range of variation within the genera as at present delimited
and even within some species.
There are no true tile cells, but the rays of the Ureneae might represent the
early stages of development leading to true tile cells, such as occur in the
Durioneae of the Bombacaceae and the Buettnerieae.

ECONOMIC USES
The outstanding economic importance of this family is due to the fibres
and hairs which they yield. Of these, by far the best known is cotton, which
consists of the hairs attached to the seeds of several species of Gossypium.
For modern descriptions of the anatomical structure and variations within
the genus Gossypium see Webber's (2376) and Hector's (929) accounts. Other
 MALVACEAE 233

important fibres are obtained from the pericycle and phloem of many members
of the family, some of which are as follows. Queensland Hemp (Sida rhombi-
folia Linn.), Chinese or Indian Hemp (Abutilon avicennae Gaertn.), Aramina
fibre from Brazil (Urena lobata Linn.), Rozelle Hemp (Hibiscus sabdariffa
Linn.), Deccan Hemp (Hibiscus cannabinus Linn.), Cuba Bast (Hibiscus
elatus Sw.), To lorn Pom' fibre (Thespesia macrophylla}$\\imt\ (Dantzer 540),
Gambo Hemp (Hibiscus sp.), (Horst 1085). For details of the anatomical
structure of some of these fibres see Wiesner (2423). The family also includes
several well-known ornamental plants such as the mallows (Malva spp.),
Hollyhocks (Althaea spp.), and Hibiscus spp. The leaves and roots of Althaea
officinalis Linn, are used for medicinal purposes. The root of Althaea
following anatomical characters. Groups of 3-25 phloem
officinalis exhibits the
fibres, unlignified except for the middle lamella; numerous starch grains,
mostly simple and ovoid; occasional cluster crystals; numerous mucilage
cells, easily stained by ruthenium red.

Secondary thickening in the roots of cotton infected by Fitsarium sp. is

stated by Dhannarajulu (581) to be initiated much earlier than in healthy
roots, whilst peripheral cork is especially well developed, and cells in the
region of the invading mycelium become suberized in cotton plants infected
by the same fungus. Twisting and hypertrophy of the vascular system,
enlargement of the cortical cells and medullary rays and the more abundant
deposition of crystals are induced in roots of Hibiscus esculentus Linn, when
invaded by the eelworm Heterodora radicicola according to Saran (1995).
Roots of Gossypium grown in acid soils exhibit, according to Gore and
Taubenhaus (796), proliferationswhich originate from the ray cells, and from
a stimulation of the cambium and phloem the original phloem sometimes
;

being killed, but subsequently replaced by new phloem containing the usual
elements but in varying proportions.
The timbers are usually light, though sometimes moderately heavy, are
not very durable and are of no importance. The wood of Thespesia populnea
(L.) Soland. is reputed to be durable under water (Desch 574) Hibiscus elatus
;

Sw. furnishes a timber that is highly valued where it occurs, e.g. in Jamaica.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Abutilon,* Althaea,* Anoda, Cienfuegosia, Cristaria, Decaschistia, Dicello-
styles, Fugosia, Goethea, Gossypium,* Hibiscus,* Hoheria,* Ingenhouzia,
Julostylis, Kitaibelia, Kokia, Kydia, Lagunaria,* Lavatera, Malachra, Malope,
Malva,* Malvastrum, Malvaviscus, Napaea, Palava, Pavonia,* Plagianthus,
Senra, Sida,* Sidalcea, Sphaeralcea,* Thespesia, Urena, Wissadula.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Abutilon, Alyogyne, Bastardiopsis, Bombycidendron, Cephalohibiscus,
(Cienfuegosia), Erioxylum, Gossypium, Hibiscus, Hoheria, (Kokia), Kydia,
Lagunaria, Lavatera, Malvastrum, Malvaviscus, Paritium, Pavonia, Plagi-
anthus, Shantzia, Sida, Sphaeralcea, Tetrasida, Thespesia, Thurberia, Urena,
Wercklea, Wissadula.
234 MALVACEAE
LITERATURE
(i) On General Anatomy
Alexandrov 12, D' Almeida 536, Blackwell 202, Dale 534, Dantzer 540, Dharmarajulu
581, Farr 672, Gehrig 753, Gore and Taubenhaus 796, Hector 929, Hess 957, Horst
1085, Janda 1143, Kienholz 1236, Koermcke 1265, Luckan 1399, Magitt 1414, Rousseau
1963, Sabnis 1977, Saran 1995 , Sharpies and Gunnery 2084, Singh, Choudri and Kapoor
2106, Spieth 2169, Stanford and Viehoever 2184, Ulbrich 2308, Webber 2376, Wiesner
2423, Youngman and Panda 2499, 2500.
(ii) On Wood Structure
Beekman 167, den Berger 179, 182, Brown, F. B. H. 282, Burgerstein 310, 312, Chatta-
way 272, Coster 481, Cozzo 494, Dadswell and Record 533, Desch 574, Edlin 622, Forsaith
690, Garratt 744, Greguss 2522, Hyde 11x7, Janssonius 1154, Kanehira 1206, 1209,
Lecomte 1334, Pearson and Brown 1679, Record 1781, 1787, 1801, 1809, 1843, 1848,
1851, 1885, Record and Hess 1886, Record and Mell 1894, Riera 1937, Webber 2373,
Williams 2426, 2430.
 64. BOMBACACEAE
(Fic, 58 on p. 238; FIG. 59 on p. 240; FIG. 60 on p. 244)

SUMMARY
(i) GENERAL
A tropical family of trees with many anatomical features in common with the
Malvaceae. Such differences as there are between them appear to be mainly
correlated with the more definitely arboreal habit of the Bombacaceae. Some
members of the family attain an enormous size (e.g. Ceibapentandra Gaertn.),
while others such as the baobab (Adansonia digitata B. Juss.) possess -barrel-
shaped trunks, which, although short, reach a considerable diameter. The
timber of most species is very light and soft, the most outstanding example
being Balsa (Ochroma lagopus Sw.) (see 'Economic Products' below). On the
relatively young branches and trunks of some of the species there are large,
deciduous, woody spines* The Bombacaceae tend to be deciduous, the
leaves falling in dry seasons. Some of the larger trees are supported by but-
tress roots. Peltate and stellate hairs are very common, whilst mucilage,
secreted either in cells or cavities, is very characteristic of the family. The
vascular structure of the petiole tends to be complex. The phloem in young
stems appears as triangular strands in transverse sections, the apices of the
triangles being towards the exterior. It is stratified into fibrous and non-
fibrous zones. The portions of the medullary rays which traverse the phloem
ire also triangular -but with their apices inwardly directed. Cluster crystals
ire common in the parenchymatous tissues.

[ii) WOOD
Vessels typically medium-sized to large perforations simple, intervascular
;

3itting alternate, small in the Durioneae and in a few genera of the Matisieae,
arge in the other genera pits to parenchyma often multiples of the intervas-
;

:ular pits or unilaterallycompound, except in some of the genera with small

ntervascular pitting; members of medium length to moderately short. Paren-
chyma abundant and sometimes forming the ground tissue of the wood;
ypically in numerous fine, metatracheal lines and in narrow sheaths round
,he vessels (vasicentric) with strands of 8 cells and not storied in the
;

Durioneae; typically of 4 cells and storied in the other tribes. Rays of

:he Adansonieae and Matisieae typically 4-10 cells wide, 1-5 to several mm.

ligh, 2-6 per mm.heterogeneous, with 1-2 marginal rows of upright cells in
.he large rays and rather few uniseriates, which are usually storied if suffi-
ciently numerous. Rays of the Durioneae 3-10 cells wide, typically i to
several mm. high, 7-12 per mm., heterogeneous, with 4 or more marginal
ows of upright cells and numerous uniseriate rays; with tile cells, except in
Maxwellia] uniseriate rays not storied. Fibres with simple or bordered pits,
septate in a few genera; scanty and occupying less space than the parenchyma
n several genera; commonly storied except in the Durioneae; moderately
ihort to moderately long. Intercellular canals of the vertical traumatic

ype occur in some genera.

236 BOMB AC'ACE AE
LEAF
Generally dorsiventral (see also 'Mesophyll'). Hairs sometimes absent, but
peltate kinds very characteristic, recorded especially in Bombax, Boschia,
Camptostemon, Coelostegia, Cullenia, Cumingia, Dialycarpa, Durio, Neesia,
and Quararibea. Extra-floral nectaries, similar to but differing somewhat
in structure from those of the Malvaceae, situated on the lower side of the
midrib and on the dorsal surface of petiole in certain species of Adansonia,
Ceiba, Chorisia, and Pachira. Cuticle often thick, occurring as high ridges in
certain species of Carolinea, Ceiba, Chorisia, and Pachira. Epidermis many
layered in certain species of Adansonia (Gehrig 753), Bombax, and Durio;
some of the epidermal cells divided horizontally in Chorisia and Quararibea
sp. Hypoderm present in Durio. Stomata confined to the lower surface.
Mesophyll of the cotton tree (Ceiba pentandra Gaertn.) found by Magitt
(1414) to be dorsiventral when grown in America, but, when grown in the
Old World, possessing palisade tissue on both sides with a small amount of
spongy tissue at the centre. Leaves of hybrids between plants from both
sources, as well as seedling plants and greenhouse specimens of the same
species stated to have the palisade tissue confined to the upper side. Isolated
sclerenchymatous elements in the mesophyll recorded in Bombax. Petiole,
in transverse sections, exhibiting a cylindrical vascular strand (cf. Hibiscus in
Malvaceae) in many of the Bombacaceae, but more complex types such as a
closed but lobed vascular strand or a closed strand surrounding two central
strands present in certain species of Bombax, Coelostegia, and Durio. Muci-
lage cells situated in the epidermis, especially on the upper side, sometimes
extending into the mesophyll in Adansonia, Boschia, Durio, and Quararibea.

Axis
YOUNG STEM (Fig. 60 H)
Cork superficial in origin. Cortex of most genera including stone cells.
Other scattered cells in the cortex, readily stained with haematoxylin, are

presumably tanniniferous. Pericycle rather indefinitely marked by strands

of fibres, mostly situated at the outer periphery of the phloem groups, but
extending laterally in a few instances. Phloem strands, and portions of the
primary medullary rays between them, appearing triangular in transverse
sections. Phloem usually stratified into fibrous and non-fibrous tangential
zones. Xylem tending to be in the form of a closed cylinder, but usually
somewhat interrupted by rather broad, often lignified primary medullary
rays. Vessels with simple perforations. Pith sometimes very broad and con-
sisting of thin-walled spongy tissue, e.g. in Bombax malabaricum DC. and
Pachira macrocarpa Walp.; at others more compact and composed of thick-
walled, pitted cells, e.g. in Camptostemon philippinensis Becc. Cortical
bundles observed in Bombax malabaricum and Camptostemon philippinensis.
Fruit stalks of Adansonia polystelic. Cluster crystals usually abundant in
the parenchymatous tissues; solitary crystals also occur but rather less
frequently. Mucilage cells and cavities, situated in the parenchymatous
tissue of the cortex and pith, are abundant in Adansonia, Bombax, Ceiba,
Chorisia, and Pachira\ rare in Hampea, Ochroma, Quararibea, and Scleronema;
very rare in Boschia, Coelostegia, Durio. Mucilage cells sometimes arranged
 BOMBACACEAE 237
in longitudinal rows. Secretory cavities, often somewhat elongated, present
in the pith and in the portions of the medullary rays passing through the
phloem in Pachira and Durio.

WOOD (Figs. 58-9)

Vessels typically medium-sized (100-200 /x mean tangential diameter) to
large (more than 200 ^); moderately small (50-100 fi) in some species of
Camptostemon, Matisia, Maxwellia, and Quararibea about 200 ft or more in
1

diameter in at least some species of Adansonia, Bernoullia, Bombacopsis^

Bombax, Cavanallesia, Ceiba, Chorisia, Coelostegia, Durio, Gossampinus,
Huberodendron, Matisia, Neesia, Ochroma, Pachira, and Scleronema} mostly
solitary, with a few multiples of 2 or 3 cells, with a tendency to the formation
locally of longer multiples, irregular clusters and tangential groups mostly
;

'5~5 P er scl- nun., slightly exceeding this maximum in some individual

specimens, about 12 per sq. mm. in Maxwellia. According to Record and
Hess (1886) the pores are more numerous in the outer part of the growth ring
in some specimens of Bombacopsis, Bombax, Cavanillesia, Ceiba, Chorisia,
Ochroma, and Pachira. Perforations simple; intervascular pitting alternate,
minute in Camptostemon, Matisia, and Quararibea, very small in Boschia,
Coelostegia, Cullenia, Durio, Gyranthera, Hampea, Maxwellia, Neesia, and
Scleronema, large in the other genera; sometimes with striations due to
coalescent apertures, e.g. in Boschia and Scleronema; pits to ray and wood
parenchyma cells often large, simple and unilaterally compound, less com-
monly so in genera with very small intervascular pitting and not observed in
Boschia p.p., Camptostemon, Coelostegia, Durio, Hampea, Huberodendron,
Matisia, Maxwellia, Montezuma, Quararibea, and Scleronema, Small amounts
of gummy deposits present in some species, tyloses present and sometimes
abundant in some species of Aguiaria (1857), Bernoullia, Bombacopsis, Bombax
(2436), Cavanillesia, Ceiba, Chorisia, and Scleronema (1857); deposits of
calcium carbonate have been observed by Record (1818) in callus tissue and
normal heart wood of Quararibea spp. Mean member
in dark streaks in the
length 0-3-0-8 mm. Parenchyma abundant (reported by Solereder to be
scanty in Pavonia), sometimes forming the ground tissue, predominantly
apotracheal, but vasicentric always present as well; in closely spaced (6 or
more per mm.), regular, uniseriate (occasionally biseriate) bands, separated
by 1-3 fibres, in Adansonia, Bernoullia, Bombacopsis, Bombax, Chorisia,
Gossampinus, Gyranthera, Hampea, Matisia, Montezuma, Pachira, and
Quararibea (Fig. 58 E and G); in more widely spaced and less regular bands,
which sometimes merge into diffuse parenchyma, in the Durioneae, e.g.
Boschia, Camptostemon, Coelostegia, Cullenia, Durio, Maxwellia, and Neesia,
and also in Huberodendron (Fig. 58 A and D); forming a continuous ground
tissue, in which the fibres are interspersed in groups, in Cavanillesia, Ceiba 9

Chorisia p.p., and Ochroma\ in widely spaced bands 2-10 cells wide (possibly
terminal) in Scleronema, together with diffuse, vasicentric and a little aliform;
in broad 5~2o-celled bands, separated by narrower bands of fibres, in Aguiaria
(1857) an d Catostemma (Fig. 59 c). A few genera with chambered crystals
and others with dark gummy contents; with druses in Adansonia and accord-
ing to Janssonius (1154) in Ceiba\ Solereder refers to silica bodies present in
Coelostegia borneensis Becc. and C. griffithii Benth. (794). Strands, in the
 238 BOMBACACEAE
apotracheal parenchyma, usually
of 4 cells, strands of 2 cells common in

Camptostemon, Hampea, Afontezuma, Ochroma,

and Scleronema\ strands
seldom of less than 8 cells in the Durioneae, e.g. Boschia, Coelostegia, Cuttenia,
Durio, Maxwellia, and Neesia, and in some species of Matisia, Pachira,
and

FIG. 58. BQMBACACEAE

A, Camptosteman philippinensis Becc. B, Cullenia exceka Wight. C, Coehstegia grifjitkii Benth.
I> C. gnffithii Benth, E, Bombast malabaricum DC. F, Ochroma iagopus Sw. G, Bernoutlia flvmmea
Oliv.

strands of the
Quamribea; fusiform cells common in Jiampea and Ochroma\
parenchyma usually containing more cells than those
vasicentric of the
in the Durioneae, but not
apotracheal. Typically storied except distinctly
storied in Chorisia p.p., Matisia, and Quamribea\ storied in Camptostemon of
the Durioneae; sometimes with secondary storying due to regular subdivision
of the strands, e.g. in Bombax. Cells on cross-section comiiionly much wider
than the fibres and irregular in shape, except in the Durioneae. Rays
uniseriate in Camptostemon, rarely more
typically 4-10 cells wide; exclusively
 BOMBACACEAE 239
than 3 wide in Maxwellia, 10-15 ce U $ w*de in some species of Aguiaria
cells

(1886), Bombacopsis, Ceiba (1885), Chorisia (1885), Gyranthera, Hampea,

Matisia, Quararibea, and Scleronema\ often described as of two distinct
sizes, e.g. by Record and Hess (1886), but intermediate sizes are usually
common; uniseriate rays numerous in the Durioneae, not common in the
other sub-families and sometimes rare, e.g. in Adansonia, Bernoullia,
Cavanillesia, Gyranthera, Huberodendron, Montezuma, Ochroma, and Quara-
ribea p.p.; multiseriate rays typically high, often 4 mm. or more, e.g. in
Bombax, Catostemma, Pachira, and Quararibea, and up to io.mm. in Matisia,
less than i mm. in some species of Boschia, Huberodendron, an& Montezuma',

typically with well-marked sheath-cells except in the Durioneae, but these

cells are less pronounced in some species of Bombax, Huberodendron, and
Pachira 7-12 per mm. in the Durioneae, 2-11 per mm. in the other
i

tribes, than 4 per mm. in some species of Adansonia, Bernoullia,

less

Bombacopsis, Cavanillesia, Gyranthera, Huberodendron, Matisia, and Quara-

ribea, seldom exceeding 6 per mm. in the other genera except in Matisia
cordata H.B. et K. heterogeneous, the larger rays seldom with more than i or 2
;

marginal rows of upright cells (Kribs's Type II B) except in the Durioneae,

in which the rays commonly have 4 or more marginal rows and in some
species 10 or more (Kribs's Types I and HA?); homogeneous (Kribs's
Type III) in Camptostemon crystals occur in chambered or ordinary cells in
\

only a few species but are sometimes abundant, e.g. in Camptostemon and
Matisia; in the form of druses in Adansonia and Bernoullia; dark gum-like
contents common. Large rays not storied, but small rays storied where
sufficiently numerous, except in the Durioneae other than Camptostemon all ;

the rays short and storied in the latter and in one specimen of Montezuma
cubensis Urb. Tile cells of the 'Durio' type (Chattaway 372) occur in all the
Durioneae examined except Camptostemon and Maxwellia, i.e. in Boschia,
Coelostegia, Cullenia, Durio, and Neesia; an intermediate form, in which the
upright cells are less narrow radially and distinctly higher axially than the
procumbent cells (Chattaway's 'Pterospermum' type), occurs iq Montezuma
and Ochroma, both of the Matisieae (Fig. 58 F). In the 'Durio' type the
uniseriate rays present an unusual appearance in tangential section, the pro-
cumbent cells being round and the interspersed tile cells consequently with
concave walls where they abut on the procumbent cells; the small rays of
Montezuma and Ochroma are also composed of alternating upright and
procumbent cells, but as the upright cells are twice as wide tangentially as the
procumbent cells the latter are always biseriate and in consequence wholly
uniseriate rays are rare. Fibres with simple or indistinctly bordered pits
equally numerous in both radial and tangential walls; pits more distinctly
bordered and mostly in the radial walls in the Durioneae, e.g. in Boschia,
Coelostegia, Cullenia, Durio, Maxwellia, and Neesia. Septate in Adansonia
and Bombacopsis and, according to Record (1857), in Bombax, Hampea, and
Pachira. Walls very thin, e.g. in Ochroma, to thick in some species of Bom-
bacopsis, Matisia, Montezuma, and Pachira, and extremely thick in Cato-
stemma the light weight of many of the woods, e.g. Cavanillesia, is due to the
;

small proportion of fibres rather than to the thinness of their walls; fibres in
narrow, often uniseriate lines or small groups and occupying less space
than the parenchyma in Adansonia, Bombax, Cavanillesia, Ceiba, Chorisia,
 BOMBACACEAE
Gossampinus, Hampea, Ochroma, and Pachira. Commonly storied. Mean
length o*7-2'O mm. Intercellular canals of the vertical traumatic type re-
ported by Record (1857) in Bombacopsis, Bombax, Catostemma, Cavanillesia,
Ochroma, Pachira, and Scleronema, and by Desch (574) in Durio. Growth
rings. Studies of seasonal growth have been made by Chowdhury (414) in
Bombax and by Coster (481) in Ceiba and Gossampinus. Besson (186) notes
a high ash content in the woods of Bombax and Ceiba.

FIG. 59. BOMBACACEAE

A, Matisia cor data H.B. et K. B, Durio stibethinus Murr. C, Catostemma fragrans Rodrig.
D, Montezuma speciosissima Sesse et Moq. E, Cavanillesia platanifolia H.B. et K.

ROOT
Navez comparing meteorological data with the arrangement
(1582), after
of the so-called buttress roots in Ceiba pentandra Gaertn., observed that the
buttresses attain their maximum development on the side exposed to the
prevailing wind. It was, therefore, concluded that they support the trees by
acting as resistance cables rather than buttresses.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
The anatomical characters indicate quite clearly that the Bombacaceae and
Malvaceae are very closely related. It is interesting to note the more complex
vascular structure of the petiole which seems to be associated with the
arboreal habit of the Bombacaceae. The shrubby genus Hibiscus of the
 BOMBACACEAE 241
Malvaceae, with its solitary cylindrical vascular strand, is exactly comparable
with the least complex type to be found in the Bombacaceae. The anatomical
structure also confirms that there are affinities with the Sterculiaceae and
Tiliaceae.

(ii) FROM WOOD STRUCTURE

The tribe Durioneae is clearlydistinguished from the others by its tile
parenchyma type, and numerous uni seriate rays. Maxwellia, however,
cells,

though agreeing in other respects, lacks tile cells. Camptostemon appears from
its wood structure to be out of place in this tribe.
The Matisieae are nearer to the Durioneae than are the Adansonieae.
Catostemma stands apart on account of its
parenchyma and fibres,

ECONOMIC USES
The Baobab
(Adansonia digitata B. Juss.) is the source of a bast fibre. The
Silk Cotton trees, belonging to the genera Bombax and Ceiba t yield a floss
from around the seeds which arises from the wall of the fruit, and is used in
the manufacture of life-saving and other equipment where buoyancy is impor-
tant, as well as for upholstery purposes. The best-known floss is Kapok,
derived from Ceiba pentandra Gaertn. For details concerning the structure
of Ceiba pentandra Calvet's (329) Thesis may be consulted. Balsa wood is the
product of Ochroma lagopus Sw. According to Pierce (1717) most of the
so-called distinct species of Ochroma which yield Balsa wood are in reality
ontogenetic stages or ecological varieties of the one species Ochroma lagopus.
The nature of the indumentum on the leaf of the plant varies with age, since
the indumentum is deciduous. The Durian (Durio zibethinus Murr.) yields
edible fruits.
The timbers of this family vary from light and soft, e.g. in the Bombacineae,
to heavy and hard, e.g. in the Catostemmataceae. Some of the woods are
exceptionally light, particularly those derived from the genera Aguiaria,
Cavanillesia, Ceiba, and Ochroma^ the best known of which is the Balsa,
Ochroma lagopus Sw. The timber of Bombax malabaricum DC. is of some
importance in the East for the match industry, and this and other species
are used for tea and rubber boxes and packing-cases. In tropical America
Bombacopsis and Ceiba supply timber of local utility and occasional export
(1886).

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Adansonia, Bombax,* Boschia, Camptostemon,* Carolinea, Ceiba, Chorisia,
Coelostegia, Cullenia, Cumingea, Dialycarpa, Durio,* Eriodendron, Hampea,
Matisia, Neesia, Ochroma, Pachira,* Quararibea, Scleronema.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Adansonia, (Aguiaria), Bernoullia, Bombacopsis, Bombax, Boschia, Camp-
tostemon, Catostemma, Cavanillesia, Ceiba, Chorisia, Coelostegia, Cullenia,
Durio, Gossampinus, Gyranthera, Hampea, Huberodendron, Matisia, Max-
wellia, Montezuma, Neesia, Ochroma, Pachira, Quararibea, Scleronema.
242 BOMBACACEAE
LITERATURE
(i) On
General Anatomy
Braun 265, 266, Calvet 329, Gehrig 753, Little 1379, Magitt 1414, Navez 1582, Pierce
17*7-

(ii) On Wood Structure

Benoist 169, den Berger 179, 182, Besson 186, Br. Hond. F. D. 274, Chattaway 372,
Chowdhury 414, Cooper 461, Coster 481, Cozzo 494, Dadswell and Record 533, Desch 574,
Ewald 666, Foxworthy 705, Gonggrijp 794, Howard 1088, Hyde 1117, Janssonius 1154,
Jentsch 1172, Kanehira 1206, 1209, 1214, Kribs 1283, Lecomte 1334, Panshin 1649,
Pearson and Brown 1679, Pereira 1687, Pfeiffer, J. Ph. 1713, Record 1780, 1781, 1787,
1801, 1809, 1818, 1834, 1843, 1848, 1851, 1857, 1^85, Record and Hess 1886, Record
and Mell ^894, Rowlee 1964, Stone 2206, Webber 2372, Williams 2430.

65. STERCULIACEAE
(Fic. 56 on p. 224; FIG. 60 on p. 244; FIG. 61 on p. 246; FIG, 62 on p. 250)

SUMMARY
(i)
GENERAL
Trees or shrubs, more rarely herbs. The family occurs in tropical and
sub-tropical regions. Anatomical features which recall those of the Bomba-
caceae and Malvaceae include: (i) The predominantly stellate hairs, often
accompanied by simple unicellular, uniseriate, glandular, and peltate types,
(ii) The usually triangular and stratified phloem strands of the young stem,
(iii)
The occurrence of mucilage receptacles which may be in the form
of cells, cavities, or canals. The leaf is generally dorsiventral, but in a few
species the mesophyll is composed entirely of palisade cells. The stomata
are usually ranunculaceous. The main vascular strand of the petiole usually
appears, in transverse sections, as a closed cylinder of xylem and phloem, or
as a cylinder formed by a number of separate but closely situated vascular
bundles. In either of these cases there are often several accessory bundles,
of very varying shapes in different species, situated in the pijh of the petiole.
In the young stem cork arises from the outermost layer of the cortex; the
pericycle contains well-developed fibre strands usually forming caps at the
exterior of the phloem (see above), the xylem constitutes a closed cylinder,
but is traversed by the relatively broad primary medullary rays. Crystals are
quite frequent and secreted in the form of large clusters solitary types also ;

occur but are less common. Scattered cells, stained deeply with haematoxylin
and presumably tanniniferous, occur in parenchymatous tissues of the stem
and leaf. They have been observed in species of Cola, Dombeya, Fremontia,
Guazuma, Heritiera, Theobroma, Trochetia, and probably occur in other
genera as well. Spines have been recorded in Buettneria and Eriolaena.

(ii) WOOD
(a) Excluding Buettnerioideae
Vessels medium-sized to large, few (1-5 per mm.); perforations simple,
intervascular pitting alternate, small; pits to parenchyma sometimes larger
and elongated members typically of medium length, occasionally moderately
;

short. Parenchyma principally of 2 types: (a) diffuse, with a little vasi-

centric, and (V) broad apotracheal to confluent bands, also with some
 STERCULIACEAE 243
vasicentric parenchyma; terminal bands sometimes present; storied. Rays
often of 2 distinct sizes, the larger up to 3-20 cells wide, 1-4 mm. high in
most genera, uniseriate rays scarce in some genera, heterogeneous, with
sheath cells; the small rays sometimes storied. Fibres with simple pits,
storied in some genera, of medium length to very long. Intercellular canals
of the vertical traumatic type present in a few specimens.

(b) Buettnerioideae
Vessels mostly medium- sized, with multiples of 4 or more cells in some
genera, occasionally in large groups with a radial or oblique pattern mostly
;

5-20 per mm., sometimes very numerous (more than 50 per mm.); with a
tendency to be ring-porous in several genera occasionally with spiral thicken-
;

ing; perforations simple, intervascular pitting alternate, small, pits to paren-

chyma similar in size and shape members of medium length to very short.
;

Parenchyma of 2 alternative types: (a) diffuse, with a little vasicentric, and

(b) exclusively paratracheal, vasicentric, aliform, or locally confluent; storied
in most of the genera. Rays often of 2 distinct sizes, the larger mostly
4-10 cells wide and commonly exceeding i mm. in height, typically hetero-
geneous, but almost homogeneous in a few genera, commonly with sheath
cells the small rays often storied and the rays predominantly storied in some
;

genera; tile cells present in several genera. Fibres with small simple pits,
storied in some genera, of medium length.

LEAF
Generally dorsiventral, but palisade and spongy mesophyll not always
clearly differentiated ; mesophyll consisting wholly of palisade tissue in Mel-
hania spp. according to Sabnis (1977) and in Reevesia, but composed mainly
of spongy tissue with only a little palisade in Heritiera littoralis Ait. according
to Kienholz (1236). Hairs, like those of the Malvaceae, consisting of simple
unicellular, uniseriate, glandular, tufted, stellate, and peltate types. Peltate
hairs recorded particularly in species of Cheirolaena, Dombeya, Heritiera
(Fig. 56 F) (whole of the lower surface covered with overlapping scales in
H . Trochetta. One of each pair of stipules in Pterosper-
littoralis), Tarrietia,
mum javanicum Jungh. becoming hollow and lined with short-stalked glan-
dular bodies so as to have the function of extra-floral nectaries. Cuticle
sometimes thick in Sterculia. Cells of the epidermis commonly mucilaginous,
especially in Sterculia. One layer of hypoderm recorded in Ungeria by Zebe
(2504) and 3 layers in Heritiera littoralis by Kienholz (1236). Stomata similar
to those of the Malvaceae, generally ranunculaceous, but said by Zebe (2504)
to be rubiaceous in Reevesia confined to the lower surface in species of Co/a,
;

Guazuma, Sterculia, and Theobroma examined by Gehrig (753). Midrib

including sclerenchyma in the species of Cola, Guazuma, Sterculia, and
Theobroma examined by Gehrig (753). Petiole supplied by a vascular system
appearing, in transverse sections, as a closed ring of xylem and phloem
(Fig. 60 D), or a circle of closely placed but separate bundles, in certain
species of Abroma, Cheiranthodendron, Dombeya, Guazuma, Heritiera, Her-
mannia, Kleinhovia, Lasiopetalum, Pterospermum, Sterculia, Theobroma; vas-
cular structure as above but often enclosing one to several accessory bundles
embedded in the pith in Cola, Heritiera (Fig. 60 p), Sterculia, and Theobroma
FIG. 60. MALVACEAE, A-C and F-G; STERCULIACEAE, D and N-P; ELAEOCARPACEAE,
E and L; BOMBACACEAE, H; TILIACEAE, I-K and M
A, Althaea qfficinalis Linn. Typical hair X 90. B, Hibiscus syriacus Linn. Petiole X 15. C, Althaea
officinaKs Linn. Petiole X 12. D, Guazuma tomentosa H.B. et K. Petiole X 13. E, Aristotelia macqui
L'Hdrit. Petiole x 13. F, Hibiscus syriacus Linn. Stem x 10. G, Malva sylvestris Linn. Stem xj.
H, Durio zibethinus Murr. Stem X 12. I, TiKa vulgaris Hayne. Petiole x 13. J, Corchorus olitorius
Linn. Petiole x 13. K, C. olitorius Linn. Stem X 12. L, Aristotelia macqui L*H&rit. Stem x 13.
M, Tilia vulgaris Hayne. Stem x 13. N, Theobroma cacao Linn. Petiole x 10. O, T. cacao Linn.
Stem x 10. P, Heritiera macrophylla Wall. Petiole x 10.
c Mucilage cavity.
.
 STERCULIACEAE 245

(Fig. 60 N), the medullary bundles sometimes forming an almost complete

inner ring in Heritiera: yascular strand in the form of an arc with strongly
incurved ends in Fremontia mexicana Macbride.
The above observations and records concerning the petiolar vascular struc-
ture of the Sterculiaceae have recently been considerably amplified by the
investigations of Dehay (558, 559). This author found transverse sections
through the middle rather than the distal end of the petiole to be the most
useful for diagnostic purposes, and, in species with a complex system of
medullary bundles, there are considerable variations in structure depending
on the level at which the sections are taken.

Dehay's central type of petiolar structure for the Sterculioideae exhibits a

vascular strand consisting of an adaxial and an abaxial arc almost united to
form a ring, and enclosing a central concave, collateral medullary strand with
adaxial xylem, e.g. in Sterculia tragacantha Lindl. (This type also occurs in
certain members of the Tiliaceae.) In Sterculia acerifolia A. Cum. the
medullary system reduced, and from S. foetida Linn, it is absent. The
is

medullary system is, on the other hand, progressively more complex in

the series Sterculia carthaginensis Jacq,, Cola altissima Engl., and Pterygota
aubrevillei Pellegr.

Dehay also recognizes a distinct central type for the Buettnerioideae, con-
sisting of a deeply concave vascular crescent with the ends very much incurved,
e.g. in Commersonia echinata var. platiphylla Forst. This type also occurs
in Mansonia altissima A. Chev. One modification of this central type is
represented by petioles in which adaxial portions of the strand appear, in
transverse sections, as partly independent strands, e.g. in Commersonia bar-
tramia Merr. In other species the adaxial portions of the vascular system
appear, again in transverse sections, to be completely independent of the
main abaxial strand. The abaxial xylem then appears circular and is
accompanied by 2 smaller circles of xylem towards the adaxial side. All
3 circles of xylem are embedded in phloem. This type occurs in Reevesia
pubescerts Mast. In Scaphopetalum blackii Mast., an adaxially concave vas-
cular cylinder is accompanied by 2 cortical strands towards the wings and
encloses a solitary, centric strand with central phloem. In Theobroma grandi-
florum Sch. and other species of Theobroma the main abaxial arc is accom-
panied by a complex adaxial system.
The Sterculioideae and Buettnerioideae are not, however, sharply demar-
cated by their petiolar structure since certain species of Dombeya (e.g. Z).
malacoxylon Sch.), Hua, Pterospermum (e.g. P. diversifolium BL), and Theo-
broma exhibit vascular systems resembling those of the Sterculioideae.
Mucilage (see also 'Stem') as in the Malvaceae, commonly present in
special cells, cavities, or canals situated in the ground tissue of the petiole,
but cells and cavities more widely distributed than canals. Clustered and
solitary crystals generally present in the mesophyll, the former type being
especially large in Rulingia sp. Tannin abundant.

Axis
YOUNG STEM 60 o)
(Fig.
Cork arising in the outermost layer of the cortex. Cortex sometimes
containing stone cells in Co/a, Tarrietia, and Ungeria. Pericycle including
 FIG. 61. STERCUUACEAE
'

A, Sterculia blancoi Rolfe. B, Pterocymbium javanicum R.Br.' C, *Scaphium macropodium Miq.'

D, Tarrietia simplicifotia Mast. E, Cola cordifolia Sim. F, Brachychiton acerifolius . Muell.
G, Tarrietia utitis Sprague. H, Heritiera littoralis (Dryand.) Ait. I, 'Scaphium macropodium Miq.*
J, Stfrculia carthaginensis Cav.
 STERCULIACEAE 247
strands of thick-walled fibres, mostly developed as caps to the exterior of the
phloem groups. Primary medullary rays usually triangular where traversing
the phloem, with their broad portions outwardly directed; this character being
particularly well defined in certain species of Fremontia, Guazuma, Heritiera,
and Theobroma. Phloem strands also triangular in transverse sections, but
with the narrowest portions towards the exterior; more or less distinctly
stratified into fibrous and non-fibrous bands in old material. Xylem forming
a closed cylinder except where interrupted by the relatively broad primary
medullary rays. Vessels with simple perforations. Medullary bundles, see
*
Anomalous Structure'. Mucilage cavities and cells (see also 'Leaf*) occur
in members of all the tribes recognized by Bentham and Hooker, but not in
all of the genera and species; situated in the primary cortex and phloem.

Mucilage canals, partly lysigenous and partly schizogenous in origin, vary-

ing in number at different levels in the same plant, arranged in i or 2 more
or less regular circles in the primary cortex and/or pith. Fruit stalks poly-
stelic in certain species of Helicteres, Kleinhovia, and Sterculia.

WOOD'
(a) Excluding Buettnerioideae (Fig. 61)
Vessels mostly medium-sized (100-200 mean tangential diameter), large /u,

(more than 200 jn) in some species of Pterocymbium, Pterygota, Scaphium,

Sterculia, and Tarrietia, very small in Octolobus\ solitary and in small radial
multiples, the percentage of solitary vessels highest (50-6 per cent.) in the
2
woods with the largest vessels, e.g. Pterocymbium, Sterculia, and Tarrietia\
multiples of 4 or more cells common locally in some woods, e.g. Heritiera spp. ;
1-5 per sq. mm., except in Octolobus (n per sq. mm.); ring-porous in
occasional species of Firmiana and Sterculia (1851); spiral thickening
reported in Heritiera macrophylla Wall. (2158) and Sterculia p.p. (1851).
Perforations simple. Intervascular pitting alternate, never large, small
in Argyrodendron, Heritiera^ and Octolobus, often with coalescent apertures;

pits to ray or wood parenchyma cells similar to the intervascular pitting except
in Brachychiton, Firmiana, and Sterculia, in which there are some elongated,
simple or slightly bordered pits, with the longer axes often oblique or vertical,
and some unilaterally compound pits, e.g. in Firmiana. Usually without
contents but solid deposits sometimes present in small amounts; tyloses
observed only in a single specimen of Heritiera macrophylla. Chattaway
arranges the genera in the following order according to vessel member length:
Pterocymbium (mean length 540 /u,), Scaphium, Brachychiton, Sterculia A,
Firmiana, Tarrietia, Argyrodendron, Cola, Pterygota, Sterculia B, Heritiera
1
Based largely on Chattaway (375).
*
Chattaway (375) divides the species of Sterculia into the following groups according to
their wood anatomy. Sterculia A. Metatracheal parenchyma predominantly in lines i cell
wide S. angustifolia Roxb. ; *9. cariboea R. Br. S. carthaginensis Cav. S. Columbians Sprague
: ; ; ;

iS. crassiramea Merril S. foetida L. S. harmanda Pierre S. hypochra Pierre S. javanica R,

; ; ; ;

Br. S. macrophylla Vent. S. montana Merril S. oblongata R. Br. S, ornata Wall. 5. parvi-
; ; ; ; ;

flora Roxb.; S. philippinensis Merril; S. recordiana Standl. (Standley 1935); S. rubiginosa

Vent. S. spangleri R. Br. S. tragacantha Lindl. S: urceolata Smith S. villosa Roxb.
; ; ; ;

Sterculia B. Metatracheal parenchyma and paratracheal parenchyma often indistinguish-

able, predominantly in broad bands 3 or 4 cells wide S. appendiculata K. Schum. 5. blancoi
: ;

Rolfe S. blumei G. Don. S. dnerea A. Rich. S. coccinea Roxb. S. elegantiflora Hutch,

; ; ; ;

and Dalz. S. oblonga Mast.; S. pollens Wall.; S. quinqueloba K. Schum.; S. rhinopetala

;

K. Schum. S. urens Roxb.;

248 STERCULIACEAE
(370 fjC),
and single specimens of Octolobus and Eribroma at about 250 ft.
Parenchyma abundant; as scattered cells and narrow, uniseriate lines,
together with distinct vasicentric sheaths in Brachychiton (Fig. 61 F), Heritiera,
Pterocymbium, Sterculia A, and Tarrietia (Fig. 61 D); in broad bands, 4 or
more cells wide, in Argyrodendron, Cola (Fig. 61 E), Eribroma, Firmiana,
Octolobus, Ptergygota, and Sterculia B (Fig. 61 A), the bands sometimes
regular and apparently apotracheal together with distinct vasicentric sheaths,
sometimes irregular, anastomosing and enclosing all the vessels, particularly
in some species of Firmiana and Pterygota entirely paratracheal (vasicentric
;

or slightly aliform) in Scaphium, the wings sometimes confluent between

adjacent vessels or, more rarely, connected with narrow bands; terminal
bands present in several species, often rather sporadic in occurrence. Crystals
regularly present in Brachychiton, Eribroma, and Pterygota, present in most
species of Argyrodendron, Heritiera, Sterculia, and Tarrietia, occurring
sporadically in Cola and not observed in Firmiana, Octolobus, Pterocymbium,
and Scaphium silica present in species of Heritiera and Tarrietia (794) some-
; ;

times with dark, gum-like contents. Strands usually of 2 cells in the scattered
cells and uniseriate bands, and with some fusiform cells, particularly in some

species of Pterocymbium and Sterculia A, but of 4, occasionally 8, cells in the

vasicentric sheaths strands usually of 4, sometimes 8, cells in the broad bands,
;

though occasionally of only 2 cells on the margins of the bands. Chattaway

(375) has shown that broad bands and strands of 4 cells are characteristic of
the genera with short vessel members, and narrow bands and strands of 2 cells
of the genera with longer vessel members. Distinctly storied. Rays often of
2 distinct sizes, e.g. in most species of Agyrodendron, Brachychiton, Cola,
Eribroma, Pterocymbium, Pterygota, Scaphium, and Sterculia', the larger rays
5-10 cells wide in most species of Heritiera and Octolobus and more than
10 cells wide in most species of Argyrodendron, Brachychiton, Eribroma,
Pterocymbium, Pterygota, Scaphium, Sterculia, and Tarrietia and up to 20 or
more cells in some species of Brachychiton and Sterculia A; varying above and
below 10 cells wide in Cola, Firmiana, and Tarrietia', the larger rays more than
i story high and commonly 1-4 mm. high except in Heritiera and some
species
of Tarrietia', uniseriate fays scarce in Cola p.p., Eribroma, Octolobus, Ptero-
cymbium p.p., Pterygota, and Sterculia B; uniseriate rays numerous and
storied and multiseriate rays relatively few in Scaphium', typically 2-4 rays
per mm., but 5-8 per mm. in some species of Cola, Firmiana, Heritiera,
Octolobus, and Scaphium markedly heterogeneous with 2 to several marginal
;

rows of upright cells (Kribs's Type II A and B) and with sheath cells, except
in Heritiera and Octolobus, in which sheath cells are lacking and in which the
multiseriate rays of some species are almost homogeneous and the uniseriate
rays contain many procumbent cells. Sometimes with dark gum-like contents ;

crystals present in at least some

species of Argyrodendron, Brachychiton, Cola,
Eribroma, Heritiera, Pterygota, Sterculia, and Tarrietia', silica present in some
species of Heritiera, Scaphium, and Tarrietia (794); the small rays storied
where sufficiently numerous, e.g. in Scaphium, Fibres with simple pits that
tend to be limited to the middle portion when the fibres are storied; storied
in Argyrodendron, Brachychiton, Firmiana, Pterocymbium, Pterygota, Scaphium,
Sterculia A, and Tarrietia. Crystal-bearing fibres are recorded by Chattaway
(37S) i*1 Eribroma and Sterculia spp., but Milanez (1526) questions whether
 STERCULIACEAE 249
these elements are properly described as fibres. Chattaway (375) discusses
the lengths of the fibres in relation to vessel member length the mean lengths
;

for the genera vary from 1-2 to 2-5 mm. (mostly 1-7-2-0 mm.) and the ratios
of fibre length to vessel member length from 3 to 6-5, the higher ratios being
typical of the genera with the shortest vessel member lengths. Intercellular
canals of the vertical traumatic type observed in occasional specimens of
Brachychiton and Heritiera and also reported by Record (1787) in Ster-
culiaand Tarrietia and by den Berger (182) in Firmiana, Pterocymbium, and
Scaphium. Anomalous structure. In Brachychiton rupestris K. Schum.,
the 'Australian bottle tree', the xylem is separated into thin layers by wide
sheaths of parenchyma containing large empty cavities, which are possibly
connected with the storage of water.
Besson (186) shows a rather high percentage of silica in the ash of Tarrietia
cochinchinensis Pierre and a very high percentage (65*85) in Cola attiensis
Aubrev. et Pellegr.

(b) Buettnerioideae (Fig. 62)

Vessels mostly medium-sized (100-200 /x mean tangential diameter),
slightly larger in some species of Pterospermum and in Triptochiton, moderately
small (50-100 jn) in some species of Reevesia, Scaphopetalum, and Theobroma,
very small (25-50 fji) in Helicteres, Leptonychia, Mansonia gagei Drummond, and
Waltheria; solitary and in small multiples and clusters, multiples of 2 or 3 cells
very common in someof the genera, multiples of 4 or more cells common in
Dombeya p.p., Eriolaena p.p., Helicteres, Kleinhovia p.p., and Leptonychia and
tending to be abundant in localized zones in several other genera; in large
groups of very small vessels in Fremontia (Fig. 62 j) and most species of
Reevesia* the groups tending to produce an oblique or tangential pattern;
mostly 5-20 per sq. mm., 1-5 per sq. mm. in Cheirostemon, Guazuma,
Pterospermum Theobroma p.p., and Triplochiton, 50-75 per sq. mm. in
,

Helicteres, Mansonia gagei, and Waltheria, and too numerous to count in

Fremontia and Reevesia] 1 semi-ring-porous or ring-porous in some species of
Fremontia, Helicteres, Kleinhovia, and Reevesia; with spiral thickening in
1

Fremontia and Reevesia. Perforations simple. Intervascular pitting alternate,

1

rather small, sometimes minute, e.g. in Dombeya, Helicteres, Mansonia, and

Pterospermum, moderately large and occasionally elliptical in Cheirostemon
and Theobroma\ pits to ray and wood parenchyma similar to the intervascular
pitting. Usually without contents, but solid deposits present in small amounts
in Mansonia gagei and Scaphopetalum-, tyloses observed only in Eriolaena
spectabilis Planch, and Triplochiton. Mean member length o*2-o4 mm.
Parenchyma often abundant; most typically as scattered cells, or numerous
short uniseriate lines alternating with 1-3 rows of fibres (Fig. 62 A and D),
together with distinct sheaths round the vessels, except in Reevesia ;* entirely
or predominantly paratracheal, as vasicentric, aliform, or locally confluent
sheaths round the vessels (Fig. 62 c and G), in Dombeya, Eriolaena, Helicteres
p.p., Melochia, and Waltheria. Crystals, either in chambered or ordinary
cells, and gummy deposits present in a few species, but usually absent from the

parenchyma cells even though abundant in the rays. Strands typically of

4 cells; strands of 2 cells common or predominant in Dombeya p.p., Eriolaena t

1
Except Reevesia wallichii R. Br,
 H I
FIG. 62. STERCUL1ACEAE-BUETTNERIOWEAE

J, Fremontia califomica Ton.

 STERCULIACEAE 251
Fremontia, and Triplochiton^ and of 6-8 cells in Leptonychia', fusiform cells
common in some species of Dombeya, Eriolaena, and Triplochiton. Distinctly
storied, except in Commersonia, Fremontia, Helicteres, Leptonychia, Melochia,
Pterospermum and Waltheria. Rays of 2 distinct sizes in Cheirostemon^
p.p.,
Leptonychia, Pterospermum p.p., Reevesia p.p., Scaphopetalum, and Theo-
broma\ the larger rays mostly 4-7 cells wide; 2-3 cells wide in Eriolaena,
Helicteres,Mansonia, and Pterospermum p.p., 8-10 cells wide in Cheirostemon,
Fremontia, Reevesia, and Theobroma p.p. and up to 20 cells wide in some
1

species of Theobroma the larger rays typically more than i story high, except
;

in Mansonia and a few species of Pterospermum, more than i mm. high

in Cheirostemon, Commersonia, Guazuma, Leptonychia, Melochia p.p., Ptero-
spermum p.p., Reevesia, and Scaphopetalum, and often showing evidence of
1

dissection into smaller units ;

uniseriate rays numerous, except in Eriolaena
p.p., Guazuma, and Mansonia, and very low, often of only i or 2 cells, in
Fremontia', most commonly 4-12 rays per mm., but 13-20 per mm. in
Helicteres, Kleinhovia p.p., Leptonychia, Pterospermum
Scaphopetalum, p.p.,
a few species of Theobroma, and Waltheria typically distinctly heterogeneous,
;

with 2-4 marginal rows of upright cells and heterogeneous uniseriate rays
(Kribs's Type II B), with more numerous marginal rows and uniseriates of
upright cells only (Kribs's Type II A) in Scaphopetalum, verging on homo-
geneous in some species of Dombeya and Mansonia', with sheath cells in
Cheirostemon, Commersonia, Fremontia, Leptonychia, Melochia p.p., Reevesia
wallichii R. Br., Scaphopetalum, and Theobroma p.p. Sometimes with dark
gum-like contents; crystals observed in at least some species of Commersonia,
Fremontia, Helicteres, Kleinhovia, Leptonychia, Mansonia, Pterospermum, Sca-
phopetalum, Theobroma, and Triplochiton, rays nearly all short and prominently
storied in Dombeya p.p., Kleinhovia, Mansonia, and Pterospermum p.p., with
only the smaller rays storied in some species of Dombeya, Eriolaena, Helicteres,
Pterospermum, Reevesia, and Theobroma. Tile cells of the *Durio' type (372)
present in Guazuma, Kleinhovia, Leptonychia, and Scaphopetalum, and of the
'Pterospermum' type in Melochia p.p. and Pterospermum, and intermediate
between the 2 types in Reevesia and Triplochiton. Fibres with small simple
pits, sometimes markedly more numerous on the radial than on the tangential
walls and varying from few to numerous. With plugs of gum and very
occasional septa in Eriolaena. Walls usually rather thin, very thin in some
species of Melochia and Theobroma, thick in Eriolaena, Fremontia, Hibiscus,
and Mansonia. Moderately distinctly storied in Cheirostemon, Guazuma,
Kleinhovia, and Mansonia and rather vaguely storied in some other genera.
Mean length io-i*7 mm. Intercellular canals of the vertical traumatic
type reported by Record (1787) in Theobroma.

ROOT
Root generally containing mucilage cells and cavities (but not canals),
situated in the primary cortex as well as in the rays where passing through
the phloem. Mucilage receptacles stated to be absent from certain species
of Cheiranthodendron, Dombeya, Heritiera, Pterospermum, and Theobroma.
Buttress roots with very excentric structure, recorded and described by
Francis (707) in a variety of Tarrietia argyrodendron Benth. Peg-like pneu-
1
Except Reevesia wallichii R. Br.
252 STERCULIACEAE
matophores, arising as localized wings from the horizontal roots and often
coalescing with the buttress roots, described by Groom and Wilson (827) for
Heritiera. The wood of the pneumatophores contains fewer vessels and fibres
than normal roots, the tissues being largely parenchymatous and often filled
with abundant starch. Lenticels present on the surface of the pneumato-
phores; intercellular spaces also well developed.

ANOMALOUS STRUCTURE
Medullary bundles, with centrally directed phloem, recorded by Solereder
and by Pfeiffer (1712) in Leptonychia sp.

TAXONOMIC NOTES
(i)
BASED ON GENERAL ANATOMY
The resemblance in anatomical characters between the Bombacaceae,
Malvaceae, and Sterculiaceae indicates that these families are all closely
related to one another. The arc-shaped vascular strand, with incurved ends,
which occurs in the petiole of Fremontia mexicana Macbride is a very much
simpler type than is to be found in most members of the Sterculiaceae. This
suggests that Fremontia may be rather remotely related to the other genera.
Dehay's (558, 559) investigations concerning the petiolar vascular structure
raise problems of considerable interest to phylogenists. He shows that the
tribes cannot be arranged in a phylogenetic sequence based on petiolar struc-
ture, nor does increasing complexity of petiolar structure run parallel to
corresponding stages in floral evolution. Dehay's work fully confirms the
close affinities between the Sterculiaceae and Tiliaceae.

(ii)
BASED ON WOOD STRUCTURE
Chattaway (375) states that the anatomical evidence supports the conclusion
of Edlin (622) that the family name Sterculiaceae should be restricted to the
Sterculieae. Discussing the taxonomic position of the genera within the
family, Chattaway makes the following suggestions: Sterculia pattens Wall,
to be transferred to Ftrmiana, Brachychiton and Eribroma to be again sunk
in Sterculia^ and the genus Sterculia to be subdivided into 2 sub-genera.
The woods which have here been described under Sterculiaceae-
Buettnerioideae, and which some botanists treat as a distinct family the
Buettneriaceae, fall into 2 fairly distinct groups according to their parenchyma,
as follows :

(a) Parenchyma predominantly apotracheal (diffuse) in Edlin's Buett-

nerieae, Helictereae, and Mansonieae.
Parenchyma exlusively paratracheal (vasicentric to aliform) in Edlin's
(b)
Dombeyeae, Eriolaeneae, and Hermannieae.
Tile cells follow much the same grouping as above, being commonly present
in Edlin's Buettnerieae, but absent from Dombeyeae and Eriolaeneae, and
rare in Hermannieae.
Reevesia wallichii R. Br. differs markedly from R. cavaleriei Leveille et
Variot and JR. pubescensMzst., though the differences might conceivably be due
to habitat. Cistanthera is very similar to Mansonia, and would fit easily in the
Buettnerioideae. Cheirostemon would fit better in the Bombacoideae (but not
in the Durioneae). Fremontia is exceptional, with ulmiform vessels, spiral
 STERCULIACEAE 253
thickening, and almost homogeneous, wide and high rays. This type of vessel
does occur in Reevesia p.p. but is more common in the Malvaceae, e.g. Malva-
viscus. The parenchyma would fit the Bombacaceae but other features less
well. Edlin's grouping of Fremontia with Cheirostemon at first sight seems
improbable, but the chief differences are of the same nature as those that
distinguish the different species of Reevesia.

ECONOMIC USES
Fibre obtained from the inner bark of various members of the family.
is

Gums are obtained from several species of Sterculia. Kola nuts, used as
substitutes for tea and coffee, are the dried seeds of Cola vera K. Schum.
Cocoa consists of the specially prepared seeds of Theobroma cacao Linn.
The timbers of this family vary from soft, coarse, and perishable to hard,
heavy, and durable. None of the timbers derived from the Sterculiaceae
sensu stricto is of first-class importance, though two of the genera, Heritiera
and Tarrietia, are of some importance locally; the wood of Heritiera fames
Buch. affords an example of the heavy, durable type of wood and is extensively
used in India for boat-building, wheel spokes and felloes, tool-handles, &c.,
and species of Tarrietia from Australia (the 'Tulip Oaks') and West Africa
(Niangon) are the source of timbers suitable for cabinet-work, panelling, &c. ;

some of the lighter woods, such as Sterculia spp., furnish packing-case

timbers. The Buettnerioideae furnish two timbers of more than local impor-
tance, both of them from West Africa Obeche, Ayous, or Samba (Triplochiton
;

scleroxylon K. Schum.) has been widely used for plywood and other purposes,
and Mansonia or Pruno (Mansonia altissima A. Chev.) is a walnut-like joinery
timber.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Abroma, Cheiranthodendron, Cheirolaena, Cola,* Commersonia, Dom-
beya,* Fremontia,* Guazuma,* Helicteres, Heritiera,* Hermannia, Hua,
Kleinhovia, Lasiopetalum, Leptonychia, Lysiosepalum, Melhania, Ptero-
spermum, Pterygota, Reevesia, Rulingia, Scaphopetalum, Sterculia, Tar-
rietia, Theobroma,* Trochetia,* Ungeria.
* Kew slide
Represented in the collection.

(ii) FOR WOOD STRUCTURE

A. Excluding Buettnerioideae
Argyrodendron, Brachychiton, Cola, Eribroma, Firmiana, Heritiera, Octo-
lobus, Pterocymbium, Pterygota, Scaphium, Sterculia, Tarrietia.

B. Buettnerioideae

Cheirostemon, Commersonia, Dombeya, Eriolaena, Fremontia, Guazuma,

Helicteres, Kleinhovia, Leptonychia, Mansonia, Melochia, Pterospermum,
Reevesia, Scaphopetalum, Theobroma, Triplochiton, Waltheria.

LITERATURE
(i) OnGeneral Anatomy
Dehay 558, 559, Francis 707, Gehrig 753, Groom and Wilson 827, Kienholz 1236,
Pfeiffer 1712, Sabnis 1977, Shelton 2086, Zebe 2504.
254 STERCULIACEAE
(ii) On Wood Structure
A. Excluding Buettnerioideae. Aubreville 51, 52, Baker 104, Benoist 169, den Berger 182,
Besson 186, Chattaway 370, 371, 372, 375, Cooper and Record 461, Coster 481, Dadswell
525, Dadswell and Record 533, Dixon 592, Edlin 622, Foxworthy 705, Giordano 786,
Gonggrijp 794, Howard 1088, I sen berg 1124, Janssonius 1154, Jentsch 1174, Jolly 1188,
Jones 1191, Kanehira 1206, 1209, Kribs 1283, Lecomte 1334, Leon 1359, Link 1377,
Mniaud 1492, Milanez 1526, Panshin 1649, Pearson and Brown 1679, Record 1781, 1783,
1801, 1809, 1824, *843, 1851, 1871, 1885, Record and Hess 1886, Record and Mell 1894,
Ridley 1935, Swain 2224, Tang 2231, Torres 2269, Tupper 2295, Williams 2430.
B. Buettnerioideae. AubreVille 52, Beekman 167, den Berger 179, 182, Besson 186,
Chattaway 371, 372, 375, Cooper and Record 461, Cozzo 494, Dadswell and Record 533,
Edlin 622, Howard 1088, Janssonius 1154, Jentsch 1172, Kanehira 1206, Lecomte 1334,
Me*niaud 1491, Metcalfe 1496, Ortega 1641, Pearson and Brown 1679, Record 1781, 1787,
1801, 1809, 1818, 1823, 1829, I ^43i 1851, 1871, 1872, 1873, 1874, Record and Hess
1886, Record and Mell 1894, Stone 2206, Tang 2231, Tupper 2295, Williams 2430.

66. TILIACEAE
(Fic. 60 on p. 244; FIG. 63 on p. 256)

SUMMARY
(i) GENERAL
A world-wide family consisting mainly of trees and shrubs, but including a
few herbs. The family has many anatomical features in common with the
Bombacaceae, Malvaceae, and Sterculiaceae. Mucilage is present in cells,
cavities, or, more rarely, canals in the cortex and/or pith of the stem as well
as in the cortex of the petiole and sometimes in the leaf lamina. The stomata
are ranunculaceous. Veins are embedded in the mesophyll sometimes ver- ;

tically transcurrent. The

vascular structure of the petiole is very variable,
ranging from a single arc-shaped strand to cylindrical types enclosing i or
more medullary bundles. The cork arises in the sub-epidermis of the young
stem; the cortex often contains an outer collenchymatous and an inner
parenchymatous zone. In the pericycle there are usually well-developed
strands of fibres, situated externally to the phloem. Transverse sections
nearly always show the phloem as triangular strands with the apices out-
wardly directed, and stratified into fibrous and non-fibrous portions.

(ii) WOOD
Vessels small to medium-sized, with radial multiples of 4, or more, cells
in some genera, semi-ring-porous and with spiral thickening in a few species;
perforations simple, intervascular pitting minute to moderately large, pits to
ray and wood parenchyma similar to the intervascular pitting or elongated
and simple members of medium length to moderately short. Parenchyma
;

predominantly apotracheal, in numerous, short, irregular, uniseriate bands,

but predominantly paratracheal vasicentric to aliform in some genera and
intermediate between apotracheal and paratracheal in others; often storied.
Rays (a) in the Brownlowieae, 2-3 cells wide, with few uniseriates, short and
storied, (A) in the other genera, typically, but with some exceptions, of 2 sizes,
with numerous uniseriates that tend to be storied, and high, unstoried rays
4-15 cells wide; sheath cells and tile cells present in several genera. Fibres
with numerous simple pits, frequently storied very short to moderately long,
;

but mostly of medium length.

 TILIACEAE 255
LEAF
Generally dorsiventral, but consisting wholly of palisade tissue in certain
species of Apeiba, Berrya, Corchorus, Diplodiscus, Grewia, as well as in
other genera especially amongst the Brownlowieae. Hairs unicellular, uni-
seriate, stellate, tufted, peltate, and glandular. ^Specially long, unicellular
hairs recorded by Gehrig (753) in Sparmannia africana Linn. Peltate scales
recorded in species of Brownlowia, Diplodiscus, Mollia, Pentace, but probably
present in other genera as well. Stomata generally confined to the lower
surface (recorded on the upper surface in species of Corchorus); usually
ranunculaceous. A xerophytic form of Tilia glabra Vent, described by
Starr (2188) as differing from a mesophytic form of the same species in having
(i) the upper epidermal cells less tall, but larger in surface area and divided

periclinally (ii) palisade cells taller, sometimes tending to be in 2 layers.

;

Midrib of Tilia always including some sclerenchyma according to Gehrig

(753). Vascular structure of the petiole very variable, its transverse appear-
ance ranging from a single arc-shaped strand to cylindrical types enclosing
i or more
subsidiary bundles in the pith. Petiolar vascular strand arc-shaped
in species of Corchorus (Fig* 60 j) and Grewia vernicosa Schinz; arc-shaped
but with accessory bundles within the arc in Glyphaea grewioidts Hook, f in .
;

the form of an almost or completely closed ring in Berrya ammonilla Roxb.

and Grewia sp. ;
form of a closed ring surrounding a complex of medul-
in the
lary bundles in TiKa spp. (Fig. 60 i). Dehay's (560) recent investigations
on the petiolar structure of the Tiliaceae confirm and extend the fore-
going records based on material examined at Kew, Dehay records, in trans-
verse sections through the distal end, a simple crescentic vascular strand in
Corchorus, Sparmannia, and Triumfetta, but with vestiges of a more complex
structure in certain species of Sparmannia and Triumfetta an almost closed
\

vascular ring surrounding a medullary strand in Berrya and Tilia a vascular ;

ring with the adaxial part very strongly invaginated, and dissected into
separate strands situated close to the bundles of the medullary system in
Brownlowia and Columbia, whilst actual fusion between the adaxial part of the
ring and the medullary bundles was observed in Columbia scabra A. DC.
Mucilage cells recorded in the epidermis of Desplatzia subericarpa Bocq.
and Glyphaea gremoides Hook. f. Mucilage cells or spaces common in the
mesophyll. Mucilage canals recorded by Coverlid (489) in the abaxial side
of the midrib and near the upper surface of the leaf in Grewia polygama Willd.
Petiole also containing mucilage cells or cavities, the latter often considerably
elongated, situated in the 'cortex* and/or 'pith* of most genera. Crystals
common; mostly clustered, but sometimes solitary; in some species most
numerous around the vascular bundles.

Axis
YOUNG STEM (Fig. 60 K and M)
Cork sub-epidermal in origin; cells with fairly thin walls and strongly
compressed in a radial direction in Tilia\ cells sclerotic in Apeiba and Vasi-
vaea. Cortex containing parenchymatous and collenchymatous zones. Peri-
cycle generally including groups of fibres situated externally to the phloem.
Phloem most commonly as triangular strands with the narrowest portions
 FIG. 63. TILIACEAE
A, Cistanthera papaverifera A. Chev. B, Columbia floribunda Kurz. C, Pentace burmanica Kurz.
D, Grewia excelsa Vahl. E, G. mollis A. Juss. F, Tilia vulgaris Hayne, G, Christiana africana DC.
H, Heliocarpus popayanensis H. B. et K. I, Cistanthera papaverifera A. Chev. J, Lueheopsis flavescens
(Britt.) Burrct. K, Tilia cordata Mill.
 TILIACEAE 257
towards the exterior, separated by the wide distal ends of the primary
medullary rays with their broadest portions towards the exterior (cf. Bom-
bacaceae, Malvaceae, and Sterculiaceae). Xylem in the form of a more or
less closed cylinder, but locally traversed by rather broad medullary rays.

Interruptions in the xylem ring much broader and more conspicuous in some
genera and species than in the others. Vessels with simple perforations.
Mucilage usually present in cells or cavities situated in the cortex phloem
and/or pith. Scattered, presumably tanniniferous cells, with contents readily
stained with haematoxylin, common in the parenchymatous tissues. Solitary
and clustered crystals of calcium oxalate common. Large, elongated pris-
matic crystals resembling styloids abundant in the bark of Tilia. 'Adventi-
tious stemlets' have been described by Richter (1934) arising from a stem
of Tilia cordata Mill, which had been damaged by lightning.

WOOD (Fig. 63)

Vessels mostly medium-sized (100-200 ft tangential diameter), moderately
small (50-100 ft), in some species of Berrya, Brownlowia, Carpodiptera,
Christiana Cistanthera, Desplatzia, Entelea (1295), Glyphaea, Microcos,
,

Pityranthe, Schoutenia (1154), Sparmannia, Tilia, Triumfetta, Vallea, and

Vinticena (1295); with multiples of four or more cells few to moderately
common in Brownlowia, Carpodiptera, Christiana, Cistanthera, Heliocarpus,
Luehea, Microcos, Pentace, and Pityranthe', irregular clusters sometimes
locally abundant in Brownlowia, Columbia, and Heliocarpus', commonly 6-20
per sq. mm., fewer than 5 per sq. mm. in some species of Althoffia (1295),
Apeiba, Belotia, Brownlowia, Entelea (1295), Goethalsia, Heliocarpus, Luehea,
Lueheopsis, Microcos, Mortoniodendron, Pentace, and Trichospermum, 20-40 per
sq. mm. in Cistanthera, Pityranthe, Sparmannia, and Triumfetta p.p. 40-70 ;

per sq. mm. in Tilia and Triumfetta p.p. semi-ring-porous in some species of
;

Erinocarpus and Grewia with spiral thickening in Luehea seemmannii PL et Tr.

;

and Tilia, and Chartocalyx (1295). Perforations simple.

in the smaller vessels of
Intervascular pitting commonlysmall to minute, sometimes extremely small,
e.g. in Diplodiscus, Glyphaea, Mollia, and Mortoniodendron (1886), tending to
be less small in the Greivieae, e.g. Columbia, Desplatzia, Duboscia, Erinocarpus,
and Triumfetta, and moderately large to large in Belotia, Heliocarpus, Spar-
mannia, and Trichospermum alternate, rather rarely with coalescent apertures
; ;

pits to ray and wood parenchyma usually similar to the intervascular pitting
but with some simple, larger, and elongated pits, and, according to Kukachka
and Rees (1295), often unilaterally compound in most of the Brownlowieae,
e.g. Berrya, Brownlowia, Carpodiptera, Diptodiscus, and Pentace, and also in
Erinocarpus, Heliocarpus, Sparmannia, Triumfetta, and Trichospermum', also,
according to Kukachka and Rees (1295), * n Glyphaea\ simple, without being
larger, in a few genera, e.g. Belotia. Sometimes with abundant gummy
deposits, e.g. Actinophora, Carpodiptera, Cistanthera, Duboscia, and Pentace;
tyloses observed in Berrya, Carpodiptera, Christiana, Grewia, Heliocarpus,
and Pentace. Mean member length 0*26-0-7 mm., mostly 0-35-0-45 mm.;
shortest in Carpodiptera, Chartocalyx, Grewia p.p., and Vinticena (1295).
Parenchyma usually moderately abundant, (a) predominantly apotracheal,
in irregular, short, numerous, uniseriate bands (Fig. 63 i and K) in Actino-
phora, Apeiba, Brownlowia, Cistanthera, Columbia, Desplatzia, Diplodiscus^
4594 S
258 TILIACEAE
Duboscia, Goethahia, Heliocarpus p.p., Luehea, Luehopsis p.p., Microcos,
Pentace, Schoutenia p.p. (1154), Tilia, and Trichospermum, often with some
vasicentric parenchyma in addition, particularly in Apeiba, Diplodiscus,
Grewia, Heliocarpus , Luehea, Pentace, and Trichospermum', in more regular
bands 1-2 cells wide in some species of Luehopsis (Fig. 63 j) (b) predominantly
;

paratracheal vasicentric to aliform with or without a little diffuse paren-

chyma in Belotia (diffuse abundant in some species, 1295), Erinocarpus,
Grewia (Fig. 63 E), Heliocarpus p.p., Mollia, Pityranthe, Schoutenia p.p.
(1154), Sparmannia, Triumfetta, and Vinticena (1295). Intermediate forms
occur in some genera, e.g. aliform mixed with irregular apotracheal bands and
scattered cells in Carpodiptera and Christiana (Fig. 63 G) and aliform with
occasional broad, possibly confluent bands in Berrya and Pityranthe. Terminal
bands usually present. The woods of Apeiba, Entelea, Heliocarpus, Spar-
mannia, and Triumfetta are characterized by the occurrence of an unusual
form of pith-like parenchyma, with very thin walls and few contents, in
addition to the common types. In cross-section this parenchyma appears as
islands (comparable with included phloem of the 'Strychnos' type), as bands
of varying width (up to ^ in. in Apeiba) or as both. The cells are elongated
radially in Apeiba and, to a less extent, in Entelea (1295). Chambered crystals
present in a few genera, e.g. Cistanthera, Erinocarpus, Grewia, Heliocarpus,
and Lueheopsis ; dark gummy contents abundant in some species. Silica bodies
have been observed in Brownlowia sp. (2158). Strands usually predominantly
of 4 or 8 cells, strands of 2 cells common in only a few species with some;

fusiform cells in Erinocarpus and Grewia celtidifolia Juss. (1154). Storied in

Apeiba, Belotia, Berrya, Brownlowia, Carpodiptera, Christiana, Cistanthera,
Columbia, Diplodiscus, Erinocarpus, Goethalsia, Grewia, Heliocarpus, Luehea,
Lueheopsis, Mollia, Pent ace, Pityranthe, Schoutenia (1295), Tilia, and Vinticena
(1295). Rays (a) in the Brownlowieae (Fig. 63 c), e.g. Berrya, Brownlowia,
Carpodiptera, Christiana, Diplodiscus, Pentace, and Pityranthe typically 2-3
cells wide, but up to 4-5 cells in some species of Pentace and Pityranthe',
largest rays less than i mm. high and seldom more than i story high; uni-
seriate rays scarce; 6-12 rays per mm.; heterogeneous (Kribs's Type II B),
with i or 2 marginal rows of square or upright cells to almost homogeneous
(Type I); frequently containing dark gummy contents and crystals in the
upright or procumbent cells; all the rays regularly storied. In Diplodiscus the
rays are similar in width, but differ in several other respects, e.g. the uniseriate
rays are common, composed almost entirely of procumbent cells, often more
than i story high and only vaguely storied the multiseriate rays are high (up
;

to 2 mm.), not storied and almost homogeneous, (b) In the other tribes
commonly of 2 distinct sizes, with numerous uniseriate rays; the larger rays
4-9 cells wide in most of the genera, 10 or more cells wide in Colombia,
Duboscia, and Sparmannia, and 3 cells wide in Cistanthera] typically more
than i story and more than i mm. high, though slightly less than i mm. in
some species of Desplatzia and Microcos and showing evidence of dissection
into smaller units in Columbia (Fig. 63 B) and Heliocarpus p.p. rays all short
;

and storied in Cistanthera and Luehea most commonly 11-20 per mm. but
;

sometimes fewer (8-10 per mm.) in Columbia, Erinocarpus, Heliocarpus,

Lueheopsis, Microcos, Sparmannia, Tilia, Triumfetta, and Trichospermum, and
4-7 per mm. in Mortoniodendron and Tilia, p.p.; typically heterogeneous
 TILIACEAE 259

(Kribs's Type II A and B) with 4 or more marginal rows of square or upright

;

cells in Althoffia (1295), Apeiba, Belotia, Desplatzia p.p., Erinocarpus,

Glyphaea, Heliocarpus p.p. (Fig. 63 H), Lueheopsisp.p., Microcos, Mollia, and
Sparmannia, and with 10 or more rows in Duboscia, Heliocarpus p.p., Tricho-
spermum, and Triumfetta; sometimes homogeneous (Kribs's Type I) in Tilia
and some specimens of Cistanthera papavenfera A. Chev. sheath cells present;

on Althoffia (1295), Entelea (1295), Erinocarpus, Goethalsia, Grewia, Helio-

carpus, Luehea, Lueheopsis, Mortoniodendron, Trichospermum, Triumfetta, and
Vinticena (1295); tl ^ e ce ^ s f ^
'Durio* type (372) present in Columbia,
Desplatzia, Grewiopsis (1295), Luehea, Lueheopsis, Mollia, Mortoniodendron,
and Vinticena (1295), and of the Tterospermum' type (372) in Belotia, 1

Duboscia, Grewia (Fig. 63 D), Mortoniodendron, and Trichospermum, and

possibly also in Sparmannia and Triumfetta, in which the large rays are com-
posed almost entirely of upright cells but with occasional procumbent cells
scattered among them. Crystals present in most of the species, sometimes
abundant gummy contents common in the procumbent cells. Nearly all the
;

rays storied in Cistanthera and Luehea p.p. and, according to Record (1894),
in Diplodiscus; the smaller rays storied in Apeiba, Belotia, Carpodiptera,
Columbia, Diplodiscus, Duboscia, Goethalsia, Grewia, Heliocarpus, Lueheopsis,
Mottia, Schoutenia (1154), Tilia p.p., Trichospermum p.p., and Vinticena
(1295). Fibres typically with numerous slit-like simple pits on the radial
walls and relatively few on the tangential walls less numerous in a few genera,
;

e.g. Pentace and Tilia, and with small borders in Tilia] often with trumpet-
shaped canals, the inner aperture sometimes resembling a small border when
seen in surface view. With gum-plates in Berry a. Walls thick in Actinophora,
Brownlowia, Carpodiptera, Christiana, Erinocarpus, Grewia p.p., Lueheopsis,
Microcos p.p., Pentace p.p., and Pityranthe, and thin in Apeiba, Belotia p.p.,
Duboscia, Goethalsia, Heliocarpus, Microcos p.p., Trichospermum, and Trium-
fetta. Storied, though sometimes rather indistinctly, in the species with
storied parenchyma. Deposits of calcium carbonate are reported by Record
(1818) in callus tissue of Tilia americana L. Mean length 0-8-1*6 mm. (1295).
Hyde (1117) refers to the occurrence of a few short fibre-tracheids around the
vessels in Heliocarpus, and Kukachka and Rees (1295) to vascular tracheids
in Erinocarpus and Vinticena. Large, widely separated radial channels occur
in Heliocarpus and, according to Record and Hess (1886), are present in all
species.

GENUS OF UNCERTAIN AFFINITIES. NESOGORDONIA

(The following information is based on Beauvisage's (163) description of
N. bernieri Baill.)
SUMMARY
A tree with thick -leaves, native of Madagascar.

LEAF
Dorsiventral. Cuticle thick on the upper surface. Mesophyll composed
of about 3 layers of short palisade cells and a broad region of spongy tissue.
Outer part of the cortical region of the petiole collenchymatous, inner part
1
Kukachka and Rees (1295) describe the tile cells in Belotia as of the *Durio' type.
260 TILIACEAE
composed of somewhat spongy parenchyma. Pericyclic region containing
bundles offibres. Petiole supplied by a closed vascular strand with an acces-

sory medullary bundle except at the distal end where the almost annular
strand is adaxially concave with inwardly directed ends. Mucilage cells
present in the epidermis, mesophyll and cortical region of the petiole. Cluster
crystals also occur.

Axis
YOUNG STEM
Cortex collenchymatous opposite the bundles of fibres in the pericycle.
Pericycle containing a ring composed of alternating groups of fibres and
parenchyma. Phloem strands and distal endings of the primary medullary
rays triangular, the phloem being stratified into fibrous and non-fibrous
portions. Xylem with vessels isolated or in radial rows of 2-5, 20-45 /* * n
diameter; perforations simple, oblique. Wood fibres composed of radial rows
of very thick-walled elements. Rays numerous, 1-3 cells wide. Pith com-
posed of lignified pitted cells, not much thickened. Elongated mucilage
cells present in the cortex and solitary and cluster crystals in the cortex and
pith.

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The Tiliaceae constitute an anatomically homogeneous family having un-
doubted affinities with the Bombacaceae, Malvaceae, and Sterculiaceae.
Genera like Aristotelia, Elaeocarpus, and Tricuspidaria are sufficiently distinct
from the Tiliaceae to be treated as a separate family, the Elaeocarpaceae.
Hasseltia, Plagiopteron, and Prockia, although included in the Tiliaceae in the
Bentham and Hooker system, are here treated as members of the Flacour-
tiaceae, to which they seem to be anatomically more similar. This is also the
position to which they have been assigned in the system of Engler and Prantl.
Nesogordonia, whose taxonomic affinities have been much disputed, exhibits
many characters in common with the Tiliaceae, as has already been pointed
out by Beauvisage (163). Dehay (560) has drawn attention to the similarity
of the petiolar vascular system of the Tiliaceae to that of the Sterculiaceae.

(ii)
FROM WOOD STRUCTURE
Kukachka and Rees (1295) have made a detailed study of the systematic
anatomy of the woods of this family. They propose the following 9 taxonomic
groups and phylogenetic sequence, the sequence being based on several
characters, but chiefly on vessel member length and the relative amount of
elongation of the fibres ('fibre-vessel length ratio'). Group I: Althoffia,
Belotia, and Trichospermum. Group II Apeiba, Entelea, Erinocarpus, Hetio-
:

carpus, Honckenya, and Triumfetta. Group III: Colona, Goethalsia, Luehea,

and Microcos. Group IV: Desplatzia, Duboscia, Glyphea, and Grewiopsis.
Group V: Lueheopsis and Mollia. Group VI: Grewia and Vinticena. Group
VII: Chartocalyx, Schoutema, and Titia. Group VIII: Brownlowia, Diplodis-
cus,and Pentace. Group IX: Berrya, Carpodiptera, and Christiana.
This arrangement is very similar to that proposed by Burret (315), the chief
points of difference in the constitution of the groups being the association in
 TILIACEAE 261

Group III of Microcos and Leuhea with Colona and Goethalsia and the
inclusion of Glyphaea with Grewiopsis and Desplatzia in Group IV.
Group I, the Trichospermae of Burret, has the most unspecialized wood
characters, with Althoffia as the most primitive genus, and Groups VIII and
IX, which are substantially Burret's Brownlowieae, the most highly specialized.
The high position assigned by Burret to his 3 tribes Triumfetteae, Spar-
mannieae, and Apeibeae is not held to be justified it is not clear, however,
;

that the authors have made sufficient allowance for the possible influence of
the shrubby habit on the characters, such as ray type, by which specialization
of the wood is judged. The group is characterized by an unusual type of
pith-like parenchyma.
The 4African genera in Group IV are regarded as an offshoot of Microcos ;

Grewiopsis is regarded as a distinct genus and not a synonym of Desplatzia.

The conclusion reached by Chattaway (373) that Microcos is anatomically
distinct from Grewia is not only upheld but further evidence is provided.
The status of Vinticena is regarded as uncertain.
The anatomy of the wood is held to substantiate Burret's transfer of
Chartocalyx from the Brownlowieae, but not his reduction of the genus to
synonymy with Schoutenia.
Groups VII and IX are substantially Burret's Brownlowieae and have
many characters in common, particularly vessel-parenchyma pitting and ray
type; they stand out as distinct from the other groups.
Record (1845) disagrees on anatomical grounds with the suggestion of
Gleason that Goethalsia should be transferred to the Flacourtiaceae.
The anatomical evidence supporting the separation of the Elaeocarpaceae
is discussed under that family.

ECONOMIC USES
two well-known timbers, the European Lime or Linden and
Tilia furnishes
the American Basswood. Burma Mahogany or Thitka, Pentace burmanica
Kurz is used in India for high-class furniture, mathematical instruments,
boat-building, &c., and Trincomalee Wood, Berry a ammonilla Roxb., and
Dhaman, Grewia tiliaefolia Vahl., are used in Burma and India respectively
for shafts, spokes, and the bent parts in carriage and cart construction. Danta,
Cistanthera papaverifera A. Chev., is sometimes exported from West Africa.
The woods of Heliocarpus and Belotia are very light and soft.
Valuable fibres are obtained from the phloem of several members of the
family, the best known being Jute, which is derived from Corchorus capsularis
Linn., C. olitorius Linn., and other species of Corchorus. Less familiar fibres
are derived from Grewia, Honckenya, and Triumfetta. Jute fibres are poly-
gonal in transverse section, but with rounded or oval lumina, the latter vary-
ing considerably in width throughout the length of the individual elements ;
the transverse section is stained yellow throughout when treated with iodine
solution and sulphuric acid; individual elements several mm. long and 17-
23 fi wide.
The fibres of Triumfetta rhomboidea Jacq. are, according to Pasqualet (1658),
in groups of 15-25 cells; individual elements 90 p long and 4-7 /LC broad;
thickness of the wall slightly undulating; cell endings pointed, rounded, or
262 TILIACEAE
spoon-shaped; fibres in stems 9-12 months old become violet-coloured when
treated with ammoniacal fuchsin.
Decoctions of the leaves of Grewia polygama Willd. are stated by Coverlid
(489) to be used in Australia in cases of dysentery.

GENERA DESCRIBED
(i) FOR GENERAL
ANATOMY
The above description is actually based on other genera besides those
mentioned by name in the text. Those mentioned in the text are: Apeiba,
Berrya,* Brownlowia, Columbia, Corchorus,* Desplatzia, Diplodiscus,
Glyphaea,* Grewia,* Mollia, Nesogordonia, Pentace, Sparmannia, Tilia,*
Triumfetta, Vasivaea.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Actinophora, (Althoffia), Apeiba, Belotia, Berrya, Brownlowia, Carpo-
diptera, Christiana, Cistanthera, Columbia, Desplatzia, Diplodiscus,
Duboscia, Entelea, Erinocarpus, Glyphaea, Goethalsia, Grewia, Heliocarpus,
Luehea, Lueheopsis, Microcos, Mollia, Mortoniodendron, Pentace, Pityranthe,
(Schoutenia), Sparmannia, Tilia, Trichospermum, Triumfetta, (Vinticena).

LITERATURE
(i) On
General Anatomy
Beauvisage 163, Burret 315, Coverlid 489, Damla 538, Dehay 560, Gehrig 753, Kundu
1302, Mullan 1571, Pasqualet 1658, Richter 1934, Sabnis 1977, Starr 2188, Zebe 2504.
(ii) On Wood Structure
Baehni 62, Bausch 154, Benoist 169, den Berger 179, 182, Besson 186, Bienfait 197,
Brown, H. P. 228, 229, Burgerstein 310, Chalk 364, 365, Chattaway 371, 372, 373,
Coster 481, Cozzo 494, Edlin 622, Greguss 2522, Hale 870, Horn 1084, Howard 1088,
Hyde 1117, Janssonius 1154, Jayawardana 1159, Jolly 1188, Jones 1191, Kanehira 1206,
1209, Kribs 1283, Kukachka and Rees 1295, Lecornte 1334, Nicoloff 1593, Pearson and
Brown 1679, Pereira 1687, Record 1781, 1783, 1808, 1809, 1818, 1843, 1845, 1851,
1871, 1872, 1873-4, 1885, Record and Hess 1886, Record and Mell 1894, Stone 2203,
Tang 2231, Williams 2426, 2430, Yamabayashi 2478.

67. ELAEOCARPACEAE
'
(FiG. 60 on p. 244; FIG. 64 on p. 264; FIG. 65 on p. 270)

SUMMARY
(i) GENERAL
A
small tropical and sub-tropical family allied to and at one time included
in the Tiliaceae. Although possessing characters in common with the
Tiliaceae, the Elaeocarpaceae are distinguished by the lack of mucilage
cavities and canals, although mucilage cells are known to occur. The
members of the Elaeocarpaceae which have been examined anatomically do
not exhibit triangular stratified phloem strands in transverse sections of the
stem, nor are the distal ends of the rays triangular. Large cluster crystals
are common, as in the Tiliaceae and other related families, but Gehrig (753)
found solitary crystals to be more frequent than druses in the leaf.
 ELAEOCARPACEAE 263

(ii) WOOD
(a) Excluding Dicraspidiaand Muntingia
Vessels usually rather small, with pronounced radial multiples in some
genera, perforations simple, sometimes accompanied by a few vestigial
scalariform plates; intervascular pitting large, usually opposite, occasionally
alternate; elongated, simple pits to ray cells common; members moderately
short to moderately long. Parenchyma paratracheal, very sparse; some-
times terminal. Rays commonly of 2 sizes, the larger 4-9 cells wide, more
than i mm. high, and markedly heterogeneous. Fibres with small bordered
pits, septate in nearly all the genera, moderately short to moderately long.
Intercellular canals of the vertical traumatic type sometimes present.

(b) Dicraspidia and Muntingia

These genera differ from the others mainly in having solitary vessels, small,
alternate intervascular pitting and similar pits to ray cells, parenchyma
apotracheal and storied, and few uniseriate rays.

LEAF
Usually dorsiventral. Simple unicellular and glandular hairs recorded in
Sloanea. Cells of the epidermis frequently mucilaginous. Stomata, in
species of Elaeocarpus and Sloanea examined by Gehrig (753), confined to the
lower surface. Hydathodes stated to be present in Aristotelia macqui L'Herit,
Vascular bundles of the veins, in species of Elaeocarpus and Sloanea examined
by Gehrig (753), surrounded by 3 or 4 layers of sclerenchyma. Petiole, in
transverse sections through the distal end, exhibiting a slightly interrupted
vascular ring in Aristotelia macqui (Fig. 60 E) and Elaeocarpus serratus Linn,
(Fig. 65 A), the ring being accompanied by 2 strands in A. macqui and
several accessory strands towards the wings in Elaeocarpus serratus. Petiole
of Tricuspidaria dependent Ruiz, et Pav. as illustrated in Fig. 65 c. Crystals
mostly solitary in species of Elaeocarpus and Sloanea examined by Gehrig
(753),but clusters also present beside the vascular bundles, e.g. in Sloanea
dentata Linn.

Axis
YOUNG STEM
Cork arising in the sub-epidermis. Cortex frequently differentiated into
an outer collenchymatous and an inner parenchymatous but somewhat spongy
zone. Pericycle including a somewhat interrupted to almost continuous
ring of fibres in certain species of Aristotelia^ Elaeocarpus^ and Tricuspidaria.
The triangular shape of the phloem strands and distal ends of the rays, which
are characteristic of the Tiliaceae, have not been observed in species of
Aristotelia^ Elaeocarpus, and Tricuspidaria available for examination. Phloem
and xylem in the form of closed cylinders traversed by narrow rays. Vessels
with mostly simple perforations and occasional scalariform plates. Pith
usually somewhat heterogeneous,
264 ELAEOCARPACEAE
WOOD
(a) Excluding Dicraspidia and Muntingia (Fig. 64 AHF)
Vessels usually very to moderately small (25-100 //, mean tangential
diameter), medium- sized (100-200 pJ) in some species of Elaeocarpus com- \

monly with radial multiples of 4 or more in Aristotelia, Crinodendron (Fig. 64 E),

G
ELAEOCARPACEAE
FIG. 64.
A, Sloanea australis F. v. M. B, Vallea stipularis Linn. C, Elaeocarpus grandis F. v. M. D, E*
floribundus Bl. E Crinodendron tucumanum Lillo. F, Sloanea australis F. v. M. G, Dicraspidia
f

donnell-smithii Standl. H, D. donnell-smithii Standl.

many species of Elaeocarpus, Tricuspidaria, and Vallea. Fewest, about 5-20

per sq. mm., in Elaeocarpus and Sloanea p.p., most numerous, 40 or more per
sq. mm., in Aristotelia, Crinodendron^ and Vallea\ spiral thickening present
in some species of Elaeocarpus. Perforation plates exclusively simple in most
genera and oblique, but some vestigial scalariform plates occur in Aristotelia
and Vallea (1295). Intervascular pitting large; typically opposite, but
alternate or transitional to alternate in Crinodendron and Vallea p.p. (1295);
pits to ray and wood parenchyma frequently simple and often elongated
horizontally (less commonly so in Elaeocarpus) or unilaterally compound.
 ELAEOCARPACEAE 265
Solid deposits and tyloses present in some species. Mean member length
0-4-1-0 mm. Parenchyma paratracheal and in narrow terminal bands
(Fig. 64 D); the paratracheal parenchyma usually limited to a few cells round
the vessels and sometimes extremely sparse or absent, e.g. in Aristotelia and
Vallea\ in narrow sheaths round the vessels, confluent, and diffuse in
Echinocarpus (1295). Rays up to 2-3 cells wide in Aristotelia, Crinodendron,
Tricuspidaria, and Vallea; 4-10 cells wide in Echinocarpus, Elaeocarpus, and
Sloanea; of 2 distinct sizes in Aristotelia, Elaeocarpus, Sloanea, and Tn-
cuspidaria; the larger rays more than i mm. high, except in Tricuspidaria and
Vallea; uniseriates numerous and typically composed entirely of upright
cells, but with some procumbent cells in Vallea; about 10-20 rays per mm.;

heterogeneous (Kribs's Types I and II A), with wide margins of 10 or more

square to upright cells; sheath cells often present. Sometimes containing
dark deposits crystals in chambered upright cells prominent in most species
;

of Elaeocarpus and occasionally present in Crinodendron; silica present in

some species of Elaeocarpus (794). Fibres with small bordered pits that are
more numerous in the radial than in the tangential walls, the borders some-
times very inconspicuous. Septate in Aristotelia, Crinodendron, Elaeocarpus,
Sloanea, Tricuspidaria, and Vallea, the septa sometimes limited to the fibres
adjoining the vessels. Walls never very thick and usually thin, Janssonius
(1154) refers to the presence of occasional solitary crystals in the septate
fibres of Sloanea sigun K. Schum. Mean length 0*7-1-9 mm. Intercellular
canals of the vertical traumatic type reported (1801) in Elaeocarpus and
Sloanea.

(b) Dicraspidia and Muntingia (Fig. 64 G-H)

These 2 genera resemble each other very closely, but differ from the rest of
the family, particularly in the following features.
Vessels mostly solitary; semi-ring-porous in Muntingia (1851); inter-
vascular pitting small and alternate and pits to ray cells similar; mean member
length in Muntingia 0-3 mm. (100). Parenchyma as (a) uniseriate sheaths
round the vessels, and (b) in numerous short uniseriate bands and scattered
cells (diffuse) (Fig. 64 H); storied. Rays homogeneous in Muntingia (1886),
composed almost entirely of square and upright cells in Dicraspidia. Fibres
sometimes septate in Dicraspidia (376); with simple pits; storied.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY

The anatomical structure of the members of this family is sufficiently

distinct from that of the Tiliaceae to justify treating the Elaeocarpaceae
separately.

(ii) FROM WOOD STRUCTURE

Dicraspidia and Muntingia differ considerably from the other genera in
theirwood anatomy. Except for these 2 genera, which have many points in
common with the Tiliaceae, and possibly Echinocarpus, the wood anatomy
supports the establishment of the Elaeocarpaceae as a separate family.
Kukachka and Rees (1295) note that 'practically all the characters of the
266 ELAEOCARPACEAE
Elaeocarpaceae point to a primitive type of structure* and regard the family
as distinctly less highly specialized than the Tiliaceae, with Echinocarpus as
a possible link. These authors, however, do not favour the removal of this
genus from the Elaeocarpaceae. Their arrangement of the genera within the
family is essentially the same as that of Engler and Prantl, with the tribe
Elaeocarpeae in the most primitive position, except that Sloanea is placed
before Elaeocarpus and that Echinocarpus is regarded as a genus and not a
section under Sloanea.
Bausch (154) has suggested that Eucryphia may have affinities with this
family, but, in the author's opinion, the wood anatomy lends little support to
such a view.

ECONOMIC PRODUCTS
The fruits of the Macqui (Aristotelia macqui L'H&it.) are eaten in Chile.
None of the timbers are of more than local importance, but some species of
Elaeocarpus and Sloanea are used in Australia for veneers, core-stocks, and
cabinet-work.

GENERA DESCRIBED
(i)
GENERAL ANATOMY
Aristotelia,* Elaeocarpus,* Muntingia, Tricuspidaria.*
* Kew
Represented in the slide collection.

(ii)
WOOD STRUCTURE
A. (Aristotelia), Crinodendron, (Echinocarpus), Elaeocarpus, Sloanea,
Tricuspidaria, Vallea.
B. Dicraspidia, (Muntingia).

LITERATURE
(i) On General Anatomy
Gehrig 753.

(ii) On Wood Structure

Tupper 100, Bausch 154, Beekman 167, den Berger 179, 182, Brown, F. B. H.
Bailey and
280, Cozzo 494, Dadswell 525, Desch 574, Descole 576, Garratt 744, Gonggrijp 794,
Howard 1088, Janssonius 1154, Kanehira 1206, 1209, Kribs 1283, Kukachka and Rees
1295, Pearson and Brown 1679, Record 1801, 1809, 1843, 1851, Record and Hess 1866,
Record and Mell 1894, Tang 2231, Williams 2430.

68. SCYTOPETALACEAE
(Fie. 66 on p. 276)

SUMMARY
A small family of trees from west tropical Africa. The wood exhibits the
following characters. Vessels with simple perforations and a few scalariform
perforation plates, intervascular pitting alternate, pits to parenchyma elongated
and almost simple, members of medium length. Parenchyma apotracheal,
in numerous uniseriate bands. Rays up to 6 cells wide, with rather few
uniseriates, heterogeneous. Fibres with simple pits and of medium length.
 SCYTOPETALACEAE 267
LEAF
Usually dorsiventral, but palisade tissue not always distinct. Hairs, where
present, simple and unicellular. Cells of the epidermis sometimes elongated
and divided by tangential walls in Oubanguia and Scytopetalum inner walls ;

convexly arched and some of the cells filled with red contents in Brazzeia.
Stomata present on both surfaces or confined to the lower side; cruciferous.
All subsidiary cells filled with violet contents in Oubanguia and Rhaptopetalum,
but similar contents confined to the smallest subsidiary cell in Brazzeia, and
absent altogether from Scytopetalum. Mesophyll containing idioblasts
connected with the sclerenchyma of the veins and spreading out beneath the
epidermis, in all 4 genera, but not in all of the species of each genus. An
arc-shaped vascular strand passes out to the leaf from the axis, but the
petiole is stated, in transverse sections, to exhibit a large median and 2 smaller
lateral strands.

Axis
YOUNG STEM
Cork arising in the sub-epidermis or outer part of the cortex in Oubanguia,
and from the epidermis itself in Brazzeia, Rhaptopetalum, and Scytopetalum ;

cork cells with U-shaped thickenings in Rhaptopetalum. Pericycle contain-

ing isolated bundles of fibres. Phloem stratified into lignified and unlignified
portions. Xylem traversed by rays 1-3 cells wide and including abundant
parenchyma. Vessels of Scytopetalum klaineanum Pierre exhibiting a wide
range of diameter and provided with scalariform and reticulate lateral thicken-
ing as well as steep scalariform perforation plates. Two cortical vascular
bundles commonly present. 'Cristarque cells' (i.e. cells with U-shaped
thickenings containing crystals, of. Ochnaceae) present in all parts of the
primary cortex of Oubanguia and Scytopetalum, but not recorded for
Rhaptopetalum .

WOOD (Fig. 66 J-K)

Vessels medium-sized (100-200 /u mean tangential diameter); solitary and
in multiples of 2 or 3 cells; about 3 per sq. mm. Perforations mostly simple,
but some scalariform plates, with up to 12 bars, usually present. Intervascular
pitting alternate and rather small; pits to ray and wood parenchyma cells
often large and simple and a/ranged in scalariform groups with the long axes
of the pits horizontal, vertical, or oblique. Mean member length about 0*5 mm.
Parenchyma apotracheal, in very numerous uniseriate bands and scattered
among the fibres. Strands usually of 8 cells. Rays up to 6 cells wide; rays
less than i mm. high uniseriates few and composed of both procumbent and
;

upright cells; about 9 rays per mm. heterogeneous (Kribs's Type II B), with
;

i or 2
marginal rows of square to slightly upright cells. Fibres with simple
pits, equally numerous on both radial and tangential walls. Walls very thick.
Mean length i-i mm.

TAXONOMIC NOTES
This small family is included in the Malvales in the Engler system and in
the Tiliales by Hutchinson (i 113). Rhaptopetalum was treated in the Bentham
268 SCYTOPETALACEAE
and Hooker system anomalous genus under Olacineae. Of the rather
as an
limited facts recorded about the anatomy of the family, the occurrence of
phloem stratified into lignified and unlignified portions is somewhat suggestive
of affinities with the Tiliaceae; the wood anatomy is also not inconsistent with
such a view, but is of a less highly specialized type. On the other hand,
'cristarque cells' have not been recorded in the Tiliaceae although they are a
characteristic feature in some of the Scytopetalaceae. It is interesting to note
that these rather uncommon elements also occur in the Ochnaceae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Brazzeia, Oubanguia, Rhaptopetalum, Scytopetalum.

(ii) FOR WOOD STRUCTURE

Scytopetalum.

LITERATURE
(i) On General Anatomy
Hutchinson 1113.

(ii) On Wood Structure

Cooper and Record 461, Record 1851.

7
69. LINACEAE
(Fic. 65 on p. 270; FIG. 66 on p. 276)

SUMMARY
(i) GENERAL

Most of the Linaceae are shrubs and small trees from tropical regions,
although a few herbaceous genera such as Linum and Radiola flourish in
cooler climates. The family also includes woody climbers which are sup-
ported by tendrils resembling coiled watch springs which arise from the
secondary branches. The anatomical structure of the leaves and young shoots
of the woody species is known only very incompletely. The most familiar
feature in Linum is the well-defined masses of fibres in the pericycle of the
stem, where they are sometimes sufficiently close together to form an almost
continuous ring. The cells of the epidermis are often mucilaginous in
Hugonia and Linum, Stomata are usually rubiaceous. Crystals mostly
solitary except in Reinwardtia sp., where clustered crystals occur in the cortex
of the stem. The xylem and phloem are in the form of continuous cylinders
in Linum except in very young stems, where the bundles are individually
distinct but not very widely separated.

(ii) WOOD
Vessels exclusively solitary except in Hugonia, perforation plates scalari-
form or simple, pits to ray cells small and bordered or large and simple;
members moderately to very long. Parenchyma very varied in different
genera, mostly aliform or confluent and sometimes abaxial; in broad, blunt-
 LINACEAE 269
ended bands dissociated from the vessels in some genera. Rays up to 2-5 cells
wide, markedly heterogeneous. Fibres with numerous distinctly bordered
pits, grading to tracheids adjoining the vessels, of medium length to moderately
long.

LEAF
Generally dorsiventral, but mesophyll not clearly differentiated into
palisade and spongy regions in Linum usitatissimum Linn. Hairs usually
consisting of narrow, unicellular (more rarely multicellular) trichomes of
varying length. Glandular shaggy hairs recorded on the leaf margin of Linum
viscosum Linn. tufted hairs apparently confined to Ctenolophon. Cells of the
;

epidermis with straight or sinuous anticlinal walls sometimes mucilaginous

;

in certain species of Hugonia, Linum, and Roucheria. Stomata usually con-

fined to the lower surface, but present on both sides in Linum usitatissimum
and in at least some of the other species of Linum; rubiaceous in Linum,
Radiola and Reinwardtia, A single layer of hypoderm recorded in Ixonanthes
y

sp, Mesophyll including idioblasts in Hugonia and Ochthocosmus sp. Spiral

tracheae, similar to those of Nepenthes, occur in the mesophyll of Ochthocosmus
roraimae Benth. Vascular bundles of the veins accompanied by well-developed
sclerenchyma. Three bundles pass out from the axis to the base of the leaf
in Ctenolophon, Hugonia, Ixonanthes, Roucheria. Petiole of Reinwardtia
trigyna Planch. (Fig. 65 B) with a shallow, crescentic vascular strand in
transverse sections through the distal end.

Axis
STEM (Fig. 65 H, j, and L)
Cortex not exhibiting any noteworthy features in Linum. Pericycle
characterized by specially large, conspicuous fibre strands in Linum (Fig. 65 j
and L), the best known being the flax fibre of commerce derived from L.
usitatissimum Linn, (see 'Economic Uses* on p. 272). Pericyclic fibres absent,
e.g. from Reinwardtia tngyna Planch. (Fig. 65 H). Xylem and phloem con-
stituting closed cylinders traversed by narrow medullary rays in mature stems
of Linum, but bundles individually distinct near the stem apex in the same
genus. Vessels of Linum mostly in radial rows, but in some species tending
to be solitary, or, more rarely, in tangential pairs; perforations simple (see
also 'Wood'). Pith becoming hollow in Linum.

WOOD (Fig. 66 A-D and G)

Vessels medium-sized (100-200 mean tangential diameter), largest in
ju,

Hugonia; exclusively solitary, except in Hugonia and Lepidobotrys, in which

a few radial multiples of 2 or 3 cells occur; 3-30 per sq. mm., most numerous
in Ctenolophon. Spiral thickening reported by Solereder in Indorouchera
griffithiana (Planch,) Hallier. Perforation plates scalariform in Ctenolophon
and Indorouchera (938), usually with less than 25, rather thick bars, but the
bars more numerous and often anastomosing in C. englerianus Mildbr,;
exclusively or predominantly simple in the other genera. Intervascular
pitting very rare except in Hugonia, in which the pits are moderately large
and with round borders and
alternate, horizontal, frequently coalescing
apertures, and Lepidobotrys, with minute alternate pitting. Pits to ray cells
270 LI N ACE A E
large, simple, and elongated axially, obliquely, or horizontally in Hebepetalum,
Ixonanthes, and Ochthocosmus, and sometimes with scalariform pits to paren-
chyma cells in Lepidobotrys\ small and bordered in Ctenolophon and Hugoma,

FIG. 65. ELAEOCARPACEAE, A and C; LINACEAE, B, H, J, and L; ZYGOPHYLLACEAE,

D-E; MALPIGHIACEAE, F-G, 1, andM; ERYTHROXYLACEAE, K
A, Elaeocarpus serratus Linn. Petiole x 15. B, Reinwardtia trigyna Planch. Petiole Xzo. C,
Tricuspidaria dependent Ruiz, et Pav. Petiole X32. D, Guaiacum officinale Linn. Petiole Xis,
E, G. officinale Linn. Stem X 10. F, Galphimia gracilis Bartl. Petiole x 22. G, Heteropteris chryso-
phylla H. B. et K. Petiole XQ. H, Reinwardtia tetragyna Planch. Stem X9. I, Hiptage madablota
Gaertn. Petiole X 9. J, Linum usitatisstmwn Linn. Group of fibres. X 60. K, Erythroxylum coca Lam.
Stem X 13. L, Linum perenne Linn. Stem Xi3. M, Galphimia gracilis Bartl. Stem x 15. ,
v
m.c. Mucilage cells.

alternate in the latter, in horizontal rows in Ctenolophon and occasionally

unilaterally in irregular groups similar in size and shape to the
compound;
simple pits in Ixonanthes. Tyloses absent from most species but moderately
abundant in some species of Ixonanthes \ patches of sclerosed tyloses occa-
 LINACEAE 271

sionally present in Ochthocosmus africanus Hook. f. ; solid deposits rare. Mean

member length 07 (Ixonanthes) to 1-25 mm. (Ctenolophon). Parenchyma
often difficult to classify as either paratracheal or apotracheal; in Ctenolophon
and Roucheria calophylla Planch. (Heimsch 938) forming uniseriate rows
along the abaxial sides of the vessels and extending tangentially as narrow
wings that spring usually from the tops of the vessels but sometimes from the
sides; in Ixonanthes and Ochthocosmus and, according to Heimsch (938), in
Phyllocosmus in bands 2-3 cells wide that start and end blindly, as in confluent
parenchyma, but independently of the vessels; aliform in Hebepetalum',
aliform and confluent in Hugonia; with scanty paratracheal parenchyma,
according to Heimsch, in Linum, Reinwardtia, and Roucheria, vasicentric in
the latter and in Tirpitzia\ abundant, as scattered cells, in Lepidobotrys;
Heimsch describes theparenchyma in Indorouchera as 'abundant diffuse-in-
aggregate with a strong tendency to form narrow apotracheal bands'. Cells
usually filled with a dark gum-like substance; crystals in chambered cells
present in some species, particularly of Ctenolophon, Lepidobotrys, and
Ochthocosmus and very abundant in Hugonia. Heimsch has observed sclerotic
parenchyma cells in Hebepetalum. Strands usually of 8 cells, often up to 1 6
cells in Ctenolophon. Rays multiseriate, up to 2 or 3 cells wide in Ctenolophon,

Hebepetalum, Ixonanthes (4 cells wide in /. reticulate Jack.), and Ochthocosmus

and up to 5 cells wide in Hugonia; rays almost exclusively uniseriate in
Lepidobolrys and, according to Heimsch, in some species of Ixonanthes and
Ochthocosmus commonly about 1-5 mm. high in Ctenolophon and Hugonia,
;

less than i mm. in the other genera; uniseriate rays numerous, composed

entirely of upright cells in Ctenolophon and Hebepetalum, mostly of procum-

bent cells in the other genera; 9-12 rays per mm.; heterogeneous (Kribs's
Types I-II B and occasionally III), with uniseriate margins of 4 or more
rows of square or upright cells; often with more than 10 rows in Ctenolophon
and Hugonia', almost homogeneous in Lepidobotrys', the multiseriate part of
the ray sometimes only a few cells high and not much wider than the uni-
seriate margins. Cells with conspicuous gum-like contents; with chambered

crystals in Ctenolophon englerianus and Hebepetalum. Fibres with numerous

bordered pits on all walls except in Ixonanthes and Lepidobotrys, in which the
pits are more numerous on the tangential walls and with only minute borders
in the latter. Walls moderately thick to thick. Cells near the vessels often
with thinner walls, wider lumina, and more numerous pits, with all gradations
from the long fibre-tracheids of the ground tissue to short vasicentric
tracheids. Mean length i-o (Ixonanthes) to 2-0 mm. (Ctenolophon).

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The anatomical structure differs considerably from that of the Erythroxy-
laceae which was included in this family by Bentham and Hooker. The
Erythroxylaceae have, therefore, been described separately. It has recently
been suggested that Ctenolophon may have affinities with the Humiriaceae
(see 'Humiriaceae').

(ii)FROM WOOD STRUCTURE

The marked differences between genera strongly suggest that this family is
272 LIN ACE AE
not a sound natural group. Gtenolophon, which has a much more primitive
structure than the other genera, except possibly Indorouchera, bears a close
resemblance to the Humiriaceae.
Heimsch (938) appears to regard the differences between the other genera
of the Linaceae and those of the Humiriaceae as compatible with the obviously
greater degree of specialization in the former. Indeed, he states that 'no
evidence from xylem structure militates against a union of these families as
is done by Hallier and Winkler', but he does not approve of Hallier's proposal

to transfer Ctenolophon to the Celastraceae.

The rather unusual type of parenchyma band found in Ixonanthes, Ochtho-
cosmus, and Phylfacosmus, together with the tendency to an oblique vessel
pattern and vasicentric tracheids, is suggestive of affinity with the Calo-
phylloideae group of the Guttiferae and Vismia of the Hypericaceae.
Lepidobotrys, as judged by a single twig of L. staudtii EngL, is exceptional
in many respects, but almost all its features, taken singly, can be matched in
some other genus, suggesting that, while it has affinities with the Linaceae,
it stands somewhat apart from the other genera.

ECONOMIC USES
The most important economic plant in this family is flax, the commercial
types of which are derived from varieties of Linum usitatissimum Linn. Lin-
seed oil is expressed from the seeds of the same species. The residue left
after expressing the oil is used to manufacture cattle cake. Best-quality flax
is obtained from specially selected varieties grown close together so as to

induce long, slender, unbranched stems. According to Djakonov (593) the

quantity of fibre present in the stem appears to depend on the moisture
content of the soil. Individual fibre elements are mostly 1 2-25 /x in diameter
(greater in fibres from the base of the stem and root) and from about 20 to 40 or
more mm. long, the longest fibres being in the upper part of the stem. In rare
instances the fibres may be as much as 120 mm. long while the diameter has
been recorded as ranging from 9-6 to 201*6 /x. The secondary walls are
divided into 2-5 major layers, composed mainly of cellulose but becoming
lignified to a greater or less extent during development, the amount of lignin
varying in different parts of the plant. The middle lamella is at first composed
of pectic substances, but later tends to become lignified. Fibres sharply
polygonal in transverse section, the lumen usually very small, but broader in
fibres from the base of the stem or from the root. Fibres from the root very
similar to those of Hemp (Cannabis sativa), but stated by Korn (1268) to be
distinguishable by differential staining with basic dyes, by treatment with
ammoniacal copper oxide, or according to Sonntag (2166) by an intensive
study of differences in the striation of the cell wall. Cell endings usually
pointed, but sometimes rounded, this difference depending on the position
within the plant from which they have been derived. For further particulars
of fibre structure see Wiesner (2423), Hayward (927), Kremer (1280), and,
for the comparative stem structure of closely related species and varieties
of Linum, the articles by Melkinov (1488, 1489) and Nestler (1587). The
developmental anatomy of the flax plant has been summarized by Hector (929),
Hayward (927), and Esau (656, 657). For further microscopical differences
between the fibre of flax and hemp see 'Cannabinaceae Economic Uses'.
 LI N ACE A E 273
GENERA DESCRIBED
(i) GENERAL ANATOMY
Ctenolophon, Hugonia, Ixonanthes, Linum>* Ochthocosmus, Radiola,
Reinwardtia,* Roucheria.
* Kew slide
Represented in the collection.

(ii)^WooD STRUCTURE
Ctenolophon, Hebepetalum, Hugonia, (Indorouchera), Ixonanthes, Lepi-
dobotrys (twig only), (Linum), Ochthocosmus, (Phyllocosmus), (Reinwardtia),
(Roucheria), (Tirpitzia).
LITERATURE
(i) On General Anatomy
Crooks 508, Djakpnov 593, Esau 656, 657, Hayward 927, Hector 929, Korn 1268,
Krerner 1280, Melnikov 1488, 1489, Nestler 1587, Sonntag 2166, Tever 2244, Tobler
2266, Wiesner 2423, Winkler 2441.
(ii) On Wood Structure
Bausch 154, den Berger 182, Burgerstein 312, Heimsch 938, Kanehira 1206, Lecomte
*334 Record 1843, 1851, Record and Hess 1886.

-f
70. ERYTHROXYLACEAE
(Fic. 65 on p. 270; FIG. 66 on p. 276)

SUMMARY
(i) GENERAL
A family of trees and shrubs wholly confined to the tropics. The more
noteworthy anatomical features are the usually abundant, prismatic, solitary
crystals present in the parenchymatous tissues, especially the epidermal
cells the presence of sclerenchymatous idioblasts in the mesophyll of the
;

leaf; the superficial origin of thecork in the stem, and the existence of
cortical bundles in the young stem. The xylem and phloem in young
stems constitute continuous cylinders traversed by narrow rays.

(ii) WOOD
Vessels occasionally tending to form long radial multiples, perforations
simple, intervascular pitting alternate and very small, pits to parenchyma
elongated and almost simple, members of medium length. Parenchyma
predominantly paratracheal with a little diffuse, the paratracheal as vasicentric,
aliform or confluent. Rays 2-5 cells wide, heterogeneous. Fibres with
distinctly bordered to simple pits, of medium length.

LEAF
Generally dorsiventral; approximately centric in a few species of Erythroxy-
lum. Cells of the epidermis mucilaginous in certain species of Aneulophus
and Erythroxylum. Lower epidermis frequently papillose in Erythroxylum
spp. and in Nectaropetalum kaessneri Engl. but not in N. capense O. Stapf.
Stomata confined to the lower surface rubiaceous in the species of Erythroxy-
;

lum used as the source of cocaine (see Economic Uses*), and in Nectaro-
petalum. Mesophyll commonly including sclerenchymatous idioblasts in
Erythroxylum spp. Vascular bundles of the veins in Nectaropetalum capense
4594 T
z 74 ERYTHROXYLACEAE
embedded in the mesophyll, but accompanied by sclerenchyma. Three
bundles enter the base of the petiole, the two lateral ones giving off branches
to the stipules, but then uniting with the main arc-shaped vascular strand in
the distal end. Minor differences occur in the arrangement of the vascular
system of the petiole of Aneulophus and Erythroxylum respectively. Scattered
solitary crystals occur in the palisade tissue and in the parenchymatous
tissues of the petiole of Erythroxylum coca Lam.
The structure of the epidermis, the form of the palisade cells, the structure
of the midrib, the occurrence and distribution of papillae and sclerenchyma
provide characters which have enabled a table for the identification of different
species of Erythroxylum to be drawn up (Ballard 116).

Axis
YOUNG STEM (Fig. 65 K)
Cork arising in the epidermis of Aneulophus and in the sub-epidermis of
Erythroxylum, Pericycle containing isolated groups of fibres in Erythroxy-
lum and Nectaropetalum, but a composite, continuous ring of sclerenchyma
present in Aneulophus. Phloem constituting a continuous cylinder, contain-
ing scattered groups of fibres or stone cells in different species of Erythroxylum,
but provided with a practically continuous ring of sclerenchyma in Aneulophus.
Secondary phloem stratified into lignified and unlignified portions in
Nectaropetalum. Xylem also in the form of a closed cylinder traversed by
narrow rays. Vessels rather unevenly distributed, tending to be in radial rows
of about 2-6; perforations simple. See also 'Wood*. Cortical bundles
(Fig. 65 K) usually present in stems which are sufficiently young.
The following statement by Boodle [see Stapf and Boodle (2187)] con-
cerning the cortical bundles of Nectaropetalum kaessneri Engl. is quoted
verbatim.
'In Nectaropetalum two vascular bundles separate from the stele in the upper
part of the internode and pass upwards, as cortical bundles, through the node and
the next internode. They reach the second node, where apparently each of them
forks into two, one branch going to the leaf trace and one to the stipule. It follows
that there are two cortical bundles in the lower part of an internode, and four in the
upper part. Thus the behaviour of the cortical bundles is similar to that of the
cortical bundles in Erythroxylum, except that in that genus, according to Van

Tieghem (2318) the cortical bundles originate in the lower, instead of the upper,
portion of the internode, and stop at the next node, instead of the next but one.
In Erythroxylum emarginatum Schum. et Th., the two bundles originate practically
at the node, but one or both may separate from the stele just before or just after the
two cortical bundles from the node below pass out of the stem. Hence, as regards
the cortical bundles, this species forms a connecting link between Nectaropetalum
kaessneri and the species of Erythroxylum investigated by Van Tieghem/

Secretory cells with gum-like contents numerous in the cortex of

Erythroxylum coca Lam. Solitary, octahedral crystals occur in scattered cells
in the cortex, phloem, and pith of Erythroxylum coca, being most numerous
in the cortex.

WOOD (Fig, 66 E-F, H-I, L)

Vessels small (50-100 p mean tangential diameter) to medium-sized
(100-200/x); solitary and in multiples, some of the multiples of 4 or more
 ERYTHROXYLACEAE 275
cells inmost species; with a tendency to form oblique rows in E. mannii Oliv. ;

usually 12-40 per sq. mm. Perforations typically simple, and Heimsch (938)
states that they are so without exception; Williams (2430), however, refers to
a few scalariform plates. Intervascular pitting alternate and very small pits
;

to ray and wood parenchyma often elongated and almost simple, and tending
to form scalariform groups with the long axes of the pits horizontal, vertical,
or oblique. Tyloses sometimes present. Mean member length 0-4-0*8 mm.
Parenchyma variable and often difficult to classify, suggesting an inter-
mediate type between apotracheal and paratracheal. Predominantly para-
tracheal in most of the material examined, e.g. in E. cuneatum Kurz., E.
mannii (Fig. 66 E), and E. monogynum Roxb. as complete to partial sheaths,
sometimes limited to the abaxial sides of the vessels, aliform or confluent, and
usually with some scattered cells in addition, the latter abundant in E.
cuneatum. Chambered crystals usually abundant. Strands mostly of 6-8 cells.
Rays up to 2-5 cells wide; typically less than i mm. high; uniseriates
moderately to very numerous, composed of mixed upright and procumbent
cells;6-12 rays per mm. heterogeneous (Kribs's Type II B), with up to 6-8
;

marginal rows of square to upright cells. Fibres, Heimsch (938) states that
the fibrous elements consist of tracheids or fibre-tracheids; the borders,
however, are sometimes very small, e.g. in E. burmanicum Griff, and E.
mannii. Walls thick. Mean length 1-1-2-8 mm.
BARK
The smooth periderm becomes perforated by the development of wart-like
some species, in longitudinal rows. Cork
lenticels, usually scattered, but, in
exceptionally well developed in certain species of Erythroxylum.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY

The genera in this family were, in the Bentham and Hooker system,
included in the Linaceae, but they are now generally treated as a separate but
related family.
The existence of cortical bundles in Nectaropetalum capense O. Stapf was
believed by Boodle to lend support to Stapf's (Stapf and Boodle 2187)
opinion that this species, which had previously been known as Peglera
capensis Bolus and tentatively assigned to the Rhizophoraceae, is, in reality,
congeneric with Nectaropetalum belonging to the Erythroxylaceae.

(ii) FROM WOOD STRUCTURE

Heimsch(938) considers that, on the basis of the wood anatomy, the closest
of the family lie with the Linaceae rather than with any other family,
affinities
but that these two families are distinct.

ECONOMIC USES
Cocaine is derived from the leaves of Erythroxylum spp. The most impor-
tant commercial types are Huanuco or Bolivian Coca (E. coca Lam.) and
Truxillo or Peruvian Coca (E, truxillense Rusby). The shortly petiolate, oval
to lanceolate or occasionally ovate, somewhat leathery, glabrous, bitter leaves
of E. coca exhibit the following diagnostic microscopical features. The upper
 N
FIG. 66. LINACEAE, A-D and G; ERYTHRQXYLACEAE, E-F, H-I, and L;
SCYTOPETALACEAE, J-K; HUMIRIACEAE, M-N
A, Ixonanthes reticulata Jack. B, Ochthocosmos africanus Hook. f. C, Ixonanthes reticulata Jack.
D, Ctenolophon grandifolius Oliv. E, Erythroxylum mannii Oliv. F, E. burmamcum Griff. G, Ctenolo-
phon grandifolius Oliv. H, Erythroxylum cuneatum Kurz. I, E, burmamcum Griff. J, Scytopetalum
tieghemii Hutch, et Dalz. K, S. tieghemii Hutch, et Dalz. L, Erythroxylum mannii Oliv. M, Sacco-
glottis gabonensis Urb. N, S. gabonensis Urb.
 ERYTHROXYLACEAE 277
epidermis composed of cells which appear quadratic in transverse sections;
the lower epidermis consisting of papillose cells the rubiaceous stomata con-
;

fined to the lower surface; the single layer of palisade cells, some of which are
divided and contain solitary crystals; the small vascular bundles embedded
in the spongy parenchyma where they are accompanied by a few fibres*
According to Levin (1366), the vein islet number of Erythroxylum coca
Lam. is said to vary from 8 to 12, as compared with 15 to 26 in E. truxillense
Rusby, thus serving to distinguish these two important commercial species.
The timber of Erythroxylum is hard, strong, and durable, is used locally in
tropical America and West Africa, and has been exported to a limited extent.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aneulophus, Erythroxylum,* Nectaropetalum.
* Kew
Represented in the slide collection.
i

(ii) FOR WOOD STRUCTURE

Erythroxylum.
LITERATURE
(i) On General Anatomy
Ballard 116, Levin 1366, Schulz 2050, 2054, Stapf and Boodle 2187, van Tieghem 2318.
(ii) On Wood Structure
Besson 186, Desch 574, Heimsch 938, Howard 1088, Link 1377, Metcalfe 1497,
Normand 1614, Record 1843, 1851, Record and Hess 1886, Record and Mell 1894,
Williams 2430.

v/71. HUMIRIACEAE
(FiG. 66 on p. 276)

SUMMARY
(i) GENERAL
Trees or shrubs which occur in tropical America and Africa. The wood
exhibits .the following characters. Vessels solitary, perforation plates
exclusively scalariform; members extremely long. Parenchyma usually
predominantly diffuse with a little vasicentric, but ranging from this to
confluent, with or without a little diffuse, the paratracheal parenchyma often
abaxial. Rays 2 or occasionally 3 cells wide, with conspicuous marginal rows
of upright or square cells and filled with gum-like substance. Fibres with
numerous bordered pits, very long.

LEAF
Leathery and dorsiventral. Hairs rare, but simple unicellular or uniseriate
trichomes recorded in Vantanea sp., and short, conical, unicellular ones in
certain species of Saccoglottis. External glands, with a secretory epidermis
of palisade-like cells, occur at the leaf margins of Humiria, and appear like
dots. Cells of the epidermis larger on the upper than on the lower surface
in Humiria and Vantanea, but of equal size on both surfaces in Saccoglottis.
278 HUMIRIACEAE
Stomata ranunculaceous in Humiria and Saccoglottis \ rubiaceous, accom-
panied by subsidiary cells parallel to the pore, but divided by a wall per-
pendicular to the latter in Vantanea sp. MesophylL Palisade cells much
more elongated in Humiria than in Saccoglottis and Vantanea. Sclerenchy-
matous idioblasts extend from one epidermis to the other in certain species of
Saccoglottis. Clustered crystals present in the mesophyll in Humiria sp. and
solitary ones in Saccoglottis sp.

Axis
YOUNG STEM
Cork sub-epidermal in Humiria and Saccoglottis ; cells thin-walled and
somewhat compressed in a radial direction. Primary cortex containing stone
Humiria and Saccoglottis. Pericycle including a
cells in certain species of

composite and continuous ring of sclerenchyma in certain species of Aubrya,

Humiria, and Vantanea. Secondary phloem containing fibres. Distal ends
of the medullary rays, where traversing the phloem, sclerosed in certain
species of Humiria and Saccoglottis. Xylem in the form of a continuous
cylinder traversed by narrow rays vessels with scalariform perforation plates.
;

Solitary crystals present in cells adjoining the sclerenchyma of the phloem,

and clustered ones in the soft tissue in the same region.

WOOD (Fig. 66 M-N)

Vessels medium-sized (mean tangential diameter 100-200 /*); exclusively
solitary or nearly so; 6-10 per sq. mm. Perforation plates scalariform with
15-25 bars, which are usually rather thick. Intervascular pitting rare; when
present, opposite to alternate and moderately large pits to ray cells and
;

parenchyma similar to the intervascular pitting, round to oval with horizontal

apertures in Humiria and Saccoglottis in Vantanea these pits are simple or
;

with transitional borders (2158) and often oblong or scalariform (938). Mean
member length i '8-2-0 mm. Parenchyma typically both apotracheal and
paratracheal most commonly with scattered cells predominating and tending
;

to form short uniseriate lines, e.g. in Saccoglottis gabonensis Urb. (Fig. 66 N)

and Humiria the paratracheal parenchyma tending to be abaxial, and with
;

distinct wings in Saccoglottis obovata EngL and Vantanea p.p. Chambered

crystal cells present in woods with well-developed paratracheal parenchyma.

Rays typically up to 2 or 3 cells wide ; occasionally up to 4 cells in some species
(938); almost uniseriate in Saccoglottis obovata ; rays often slightly more than
i mm. high; uniseriates numerous, and composed of procumbent and square

to upright cells; about 12 rays per mm.; heterogeneous (Kribs's Type II A?),
with more than 4 marginal rows of square to upright cells (often more than
10 rows in Saccoglottis)^ which are not much narrower than the biseriate
parts. Cells often filled with dark contents. Fibres with numerous, distinctly
bordered pits on both radial and tangential walls; walls thick. Mean length
about 2*2 mm.

TAXONOMIC NOTES
(i)
FROM GENERAL ANATOMY
The idioblasts in the mesophyll of Saccoglottis recall those which occur in
members of the Theaceae.
 HUMIRIACEAE 279

(ii)
FROM WOOD STRUCTURE
Heimsch (938) considers that this family forms a homogeneous group,
distinct from the Linaceae, but more nearly related to it than to any other
family; the greatest resemblance is to Ctenolophon, which he believes has
closer affinities with the Humiriaceae than with the Linaceae. The woods of
the Humiriaceae are more unspecialized than those of the Linaceae, except
that of Ctenolophon.

ECONOMIC USES
The timbers of this family are dense they are of ;
little importance, though
some species of Humiria are used in tropical America for heavy construction,
wheels, &c.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aubrya, Humiria, Saccoglottis, Vantanea.

(ii) FOR WOOD STRUCTURE

Humiria, Saccoglottis, (Vantanea).

LITERATURE
On Wood Structure
Benoist 169, Cooper 461, Heimsch 938, Howard 1088, Pereira 1687, Pfeiffer, J. Ph.
1713, Record 1843, 1851, Record and Hess 1886, Record and Mell 1894, Stone 2202,
2206, 2207.

^72. MALPIGHIACEAE
(FiG. 65 on p. 270; FIG. 67 on p. 280; FIG. 69 on p. 290; FIG, 91 on p. 402)

SUMMARY
(i) GENERAL
A
mainly tropical family of trees, shrubs, or climbers, characterized
especially by hairs with i or 2 more or less horizontal arms, attached to the
plant by a long or short vertical stalk. True stellate hairs have been recorded
only in Thryallis. Large complex glands, visible with the naked eye, are
often present on the lower side of the petiole, and/or the lower side and margin
of the leaf. The stomata are rubiaceous. The mesophyll of the leaf is

dorsiventral or isobilateral, and often contains water-storage cells. The

structure of the mesophyll and the distribution of the water-storage cells
were found by Dubard and Dop (612) and by Niedenzu (1594) to be of value
for the identification of genera and species. Other features confined to the
leaves of certain species include: (i) A
many-layered epidermis, (ii) Muci-
laginous epidermal cells, (in) Fibre-like cells in the mesophyll. The petiole,
in transverse sections, exhibits a central,, arc-shaped, vascular strand, which
is sometimes accompanied by small accessory ones in the latero-superior

position. Crystals are sometimes solitary, but more frequently clustered;

styloids are rare. The cork of the young stem is usually superficial in origin,
the cells being either thin-walled, or with their inner tangential and, to a less
z8o MALPIGHIACEAE
extent, their lateral walls strongly thickened. Secretory cells, whose con-
tents are deeply stained with haematoxylin, are common in the parenchyma-
tous tissues of both leaf and stem. The pericycle contains small, isolated
groups of fibres, whilst the xylem and phloem have the form of continuous
cylinders traversed by narrow rays in the limited number of species which
have been examined. Various types of anomalous thickening occur in

FIG. 67. MALPIGHIACEAE

Hairs of: A, Malpighia wrens L. B-C, Peixotoa macrophylla Griseb. D, Mascagnia cordifolia (Juss.)
Griseb. E, Camarea hirsuta St. Hil. F, Hiraea chrysophylla Juss. G, Thryallis brachy stocky s Lindl.
H, A piece of the margin of a calyx-gland of Lophopterys splendem Juss. After Niedenzu.

certain members of the family. For instance, the xylem is sometimes furrowed
(Heteropteris, Peixotoa, Tetrapteris), or the furrows may become so extensive
that the xylem mass is divided into separate portions (Banisteria y Mascagnia,
Mezia Tetrapteris). Sometimes a secondary cambium arises in the parenchy-
y

matous tissue between the xylem groups (Stigmaphylbn). Interxylary

phloem is always present in Dicella and sometimes in Stigmaphyllon.

(ii) WOOD
Vessels mostly medium-sized, commonly in radial multiples of 2-3 and
sometimes with a definite radial pattern; semi-ring-porous in one species;
perforations exclusively simple and intervascular pitting alternate; pits to ray
cells mostly small and bordered, but large and simple in a few genera members ;
 MALPIGHIACEAE 281

of medium length. Parenchyma usually predominantly paratracheal, scanty

to confluent, occasionally abundant and banded, diffuse parenchyma some-
times present and occasionally predominant. Rays mostly 2-3 cells wide,
markedly heterogeneous. Fibres with simple pits on the radial walls, often
septate; of medium length to moderately long. Anomalous structure, see
under (i) above.

LEAF
Usually dorsi ventral, less frequently isobilateral or centric (for further
details see mesophyll). Hairs (Fig, 67 A-H) very characteristic, mostly con-

sisting of unicellular trichomes, usually with 2 more or less horizontal arms

attached to the plant by a short or long vertical stalk. The 2-armed hairs
exhibit modifications in different genera and species, the relative lengths of
the arms and the degree of twisting, as well as the smoothness of the external
walls being variable characters. Other modifications of the 2-armed trichomes
include stinging hairs in certain species of Malpighia and bristle hairs in
Camarea. Stellate hairs confined to Thryallis. Large complex sunken or
cushion-shaped glands common on the petiole as well as on the lower side
of the margins of the lamina. Glands with more or less elongated stalks also
recorded on the leaf in species of Aspicarpa and Gaudichaudia. Epidermis
composed of cells with very small lumina in species of Hiraea and Tetrapteris',
cells sometimes elongated and palisade-like in Acridocarpus, Banisteria, Bur-
dachia, Byrsonima, or very large (even wider than the whole of the assimilatory
tissue) in certain species of Camarea. Very large epidermal cells distributed
amongst much smaller ones and sometimes penetrating deeply into the meso-
phyll occur in species of Aspidopterys, Flabellaria, Malpighia, Microsteira,
Pterandra, Ryssopterys, Spachea, Triaspis, Tristellateia. The enlarged
epidermal cells sometimes appear as translucent spots in the leaves. Epidermis
consisting wholly or locally of 2 or more layers in certain species of Acrido-
carpus, Banisteria, Blepharandra, Hiraea, and Tetrapteris; mucilaginous in
Triaspis sp.; lower epidermis papillose in certain species of Burdachia,

Heteropteris, and Stigmaphyllon. Stomata nearly always confined to the

lower surface except in Caucanthus edulis Forsk and Triaspis spp. rubiaceous
;

in allinvestigated genera except Diacidia; guard cells usually elongated to

elliptical in surface view.
Mesophyll, according to Solereder, Dubardand Dop (612), and Niedenzu
(1598), isobilateral with paKsade tissue towards both surfaces or consisting of
homogeneous palisade tissue in species of Acridocarpus, Banisteria, Byrsonima,
Dinemagonum, Echinopterys, Hiraea, Microsteira, Sphedamnocarpus, Stigma-
phyllon, Tetrapteris, Triaspis, Tricomaria, and other related genera. Palisade
cells specially tall in species of Galphimia, Heteropteris, Stigmaphyllon;

containing little chlorophyll and serving for the storage of water in species of
Diplopterys, Galphimia, Spachea. Water-storage cells also recorded at the
boundary between the palisade and spongy mesophyll in certain species of
Stigmaphyllon and Tetrapteris immediately on the inside of both upper
;

and lower epidermis in Banisteria, Janusia, Philgamia, and Tricomariopsis;

occurring only below the upper epidermis in the region of the veins in
Acridocarpus; situated at the centre of the lamina in species of Aspidopterys,
Caucanthus, Flabellaria, Microsteira, Sphedamnocarpus, Triaspis, where a few
282 MALPIGHIACEAE
or a large proportion of the mesophyll cells serve as water-storage organs.
Individual cells of the outermost layer of palisade tissue also serve for water
storage in species of Aspidopterys, Diploterys, Tetrapteris. Water-storage cells
are also common around the vascular bundles of the veins. Differences in the
structure of the epidermis and mesophyll, as well as in the distribution of
water-storage cells, are also stated by Niedenzu (1594) to be of value for the
identification of species. Leaf characters have also been used by Dubard and

Dop (612) to confirm the existence of affinities between members of the family
from America and Madagascar respectively. Vascular bundles of the veins
sometimes not supported by mechanical tissue, e.g. in species of Banisteria,
Heteropteris, Spachea, Stigmaphyllon accompanied by groups of mechanical
;

elements in the phloem in species of Diploterys and Malphigia\ surrounded

by a continuous ring of mechanical elements in species of Acmanthera,
Coleostachys, Dicella, Diploterys (pro parte), Glandonia, Lophopterys, Peixotoa,
Stigmaphyllon, Tetrapteris. A
bundle consisting almost entirely of mechani-
cal tissue follows the course of the leaf margin in species of Heteropteris y
Malpighia, and Tetrapteris. Cells over the midrib contain abundant tannin
in Janusia gracilis Gray, according to Clausen (431). Petiole, in transverse
sections, exhibiting a single arc-shaped vascular strand, sometimes accom-
panied by very small ones in the latero-superior position, in certain species of
Galphimia (Fig. 65 F), Heteropteris (Fig. 65 G), and Malpighia, ends of the
median vascular arc very much incurved, e.g. in Hiptage (Fig. 65 i); the
vascular strand accompanied by little or no sclerenchyma in certain species
of Banisteria, Galphimia, Heteropteris, Janusia, Malpighia, Philgamia, and
Tricomariopsis with, according to Dubard and Dop (612), sclerenchymatous
;

fibres on both sides in Acridocarpus. Cells containing a gum-like substance

present in the 'cortex' of the petiole in Heteropteris sp.

Calcium oxalate present in the form of clustered or solitary crystals, those
of Hiraea and certain species of Banisteria especially large; styloids occur in
the assimilatory tissue of Banisteria, Janusia, Peixotoa, Philgamia, and
Tricomariopsis. Solitary crystals more frequent in members of the family
from the Old World than in American species, but the exact form of the
crystals is stated by Niedenzu (1594) to be of very limited taxonomic value.

Axis
YOUNG STEM (Fig. 65 M)
Cork superficial in origin in certain species of Byrsonima, Galphimia,
Heteropteris, and Malpighia, but described by Clausen (431) as possibly
originating near the endodermis in Janusia gracilis Gray cells thin-walled in
;

Galphimia gracilis Bartl. but with strong thickenings on the inner tangential
and to a lesser extent on the lateral walls in Heteropteris chrysophylla H.B.
et K. and Malpighia sp. Cortex sometimes containing groups of stone cells
in Malpighia. Pericycle including isolated strands of fibres in the. few
species of Galphimia (Fig. 65 M), Heteropteris, and Malpighia actually
examined, but pericyclic sclerenchyma stated to be absent from a few species
of Byrsonima and Malpighia. Primary phloem constituting a narrow, closed
cylinder; with simple plates to the sieve tubes in Byrsonima sp. and Mal-
pighia sp.; secondary phloem, including concentric layers of thick-walled,
abundantly pitted fibres in certain species of Byrsonima and Malpighia.
 MALPIGHIACEAE 383

Xylem in the form of a closed cylinder, traversed by narrow

rays, in all the
species so far examined. Vessels with simple perforations. Pith said by
Clausen (431) to contain stone cells in Janusia gracilis. Clustered crystals
common in the cortex and phloem; solitary types recorded in Malpighia sp.
Crystals present in 2-chambered cells in the phloem of Hiraea and other
genera. Anomalous structure, see p. 284.

WOOD (Fig. 69 D-H)

size from very small (less than 50 /z mean tangential
Vessels varying in
diameter) in Malpighia, Ptilochaeta, Spachea, Tetrapodenia, and Thryallis to
large (over 200 ^) in some of the lianes, e.g. Banistariopsis and Stigma-
phyllon. Radial multiples common throughout, with 4 or more per group in
all but a few specimens, except for Heteropteris multiflora (DC.) Hochr., in
which the vessels are all solitary ; juxtaposition of the multiples gives a definite
radial pattern in Diacidia, Lophanthera, Ptilochaeta, and Thryallis, but in
Ptilochaeta the pattern may also be tangential locally; also with a tendency to
clusters insome species, particularly the small vessels of the lianes; mostly
between 20 and 40 per sq. mm., fewer in Bunchosia, Byrsonima, and Glandonia,
and more numerous in Malpighia; semi-ring-porous in Ptilochaeta and ring-
porous, according to Heimsch (938) in Galphimia glauca (Poir.) Cav.; spiral
thickening reported in Peixotoa (Solereder) and Tricomaria (25 1 7). Perforations
exclusively simple. Intervascular pitting alternate, typically very small to
minute, but moderately large in Banisteriopsis leptocarpa R. O. Williams pits ;

to ray cells generally similar to the intervascular pitting, except in Burdachia,

Glandonia, and Tetrapodenia, in which larger simple pits are common very ;

occasional large simple pits or unilaterally compound pits occur in most of

the genera; pits vestured (78). Tyloses present in only a few species; solid
deposits present in Banisteria, Malpighia, and Stigmaphyllon. Mean member
length 0-4-0*8 mm. Parenchyma predominantly paratracheal in most
genera, consisting often of only a few cells about the vessels (Fig. 69 F), but
sometimes aliform in Heteropteris and confluent in Bunchosia (Fig. 69 G),
Burdachia, and Ptilochaeta\ with some diffuse parenchyma in addition in
Burdachia, Lophanthera (1883), Ptilochaeta, Spachea (1883), and Thryallis
(1883); predominantly apotracheal (diffuse or banded) in Malpighia, Tetra-
podenia, and Triopteris ; in broad bands 4 or more cells wide in Stigmatophyllon ;

obscure or absent from Glandonia terminal in Diacidia. Typically filled with

;

dark contents and very commonly containing small to large chambered

crystals. Strands of 2-4 cells. Rays up to 2-3 (occasionally 4) cells wide;
uniseriate or partially biseriate, according to Williams (2430) in Spachea
tricarpa A. Juss. less than i mm. high, except in Burdachia, Thryallis, and
;

Triopteris uniseriates few to numerous, composed of square to upright cells;

;

usually 6-12 rays per mm., but sometimes more numerous; very distinctly
heterogeneous (Kribs's Type I and II A) with 2 or 3 marginal rows of large
upright cells and with 4 or more rows in Burdachia, Glandofiia, Heteropteris,
and Malpighia. Occasionally less markedly heterogeneous (Kribs's Type II B),
according to Heimsch, and sometimes only weakly heterogeneous in Bunchosia
(1886). Cells characteristically filled with a dark substance and often contain*
ing crystals in ordinary or chambered cells, and in idioblasts in Bunchosia and
Burdachia. Fibres with simple pits, more numerous on the radial than on the
284 MALPIGHIACEAE
tangential walls; septate in Banisteria, Byrsonima, Glandonia> Ptilochaeta^ and
Tetrapteris^ and, according to Solereder, in Hiraea and according to Heimsch
(938), with some septate fibres in Heteropteris and Spachea\ Heimsch refers
also to the common occurrence of gelatinous fibres; commonly containing
gum; walls moderately thick to thick. Mean length 1-0-1-8 mm.

ANOMALOUS STRUCTURE (Fig. 91 D)

Interxylary phloem present in all and in Stigma-
species of Dicella
phyllon sp.
In young stolons of Dicella nucifera Chod. the structure is normal. At a
later stage some of the cells cut off from the cambium towards the inside fail
to differentiate as xylem elements, but form blocks of parenchyma in which
sieve tubes develop. More xylem is then differentiated externally to the
abnormal phloem, so that the latter becomes isolated as islands in the xylem.
Successive islands of interxylary phloem are formed in the same way as the
stern grows older.
The same species exhibits the same type of anomalous structure
root of the
becomes locally swollen into ellipsoid or fusiform tubercles,
as the stem, but
5-10 cm. long X 3-4 cm. broad. The tubercles are composed mainly of
parenchyma, with groups of fibres embedded in it. They are stated by
Chodat and Vischer (399) to contain inulin and secretory cells with brown
contents.
The xylem is normal in structure and mode of development in
at first

StigmaphylloHj contains abundant parenchyma and ray tissue in

but it

which cambial layers are subsequently laid down, and give rise to new
bundles of xylem and phloem.
Certain lianes belonging to this family are characterized by a lobed mass of
xylem, or one which becomes lobed during development, e.g. in Heteropteris^
Peixotoa, Tetrapteris (Fig. 91 D); subsequent enlargement of the tissue
between the xylem radii sometimes causes the xylem to become split up into
groups in Banisteria, Mascagnia, Mezia, and Tetrapteris. For further parti-
culars see Solereder, Pfeiffer (1712), and Chodat and Vischer (399).
Lianes belonging to the genera Heteropteris (pro parte), Hiptage, Schwannia,
Tetrapteris (pro parte), and Thryallis are stated to be normal in structure.

TAXONOMIC NOTES
Heimsch (938) points out that the Malpighiaceae have a more highly
specialized wood
structure than the Linaceae, Humiriaceae, and Erythroxy-
laceae, with which they are often associated, and that this family and the
Vochysiaceae, though showing indications of relationship to these families
and to the Polygalaceae, Tremandraceae, Trigoniaceae, and Zygophyllaceae,
differ from all of them, particularly in having vestured vessel pits and simple-
pitted fibres.

ECONOMIC USES
According and Raymond-Hamet (1698), the anatomical structure
to Perrot
of a Brazilian plant which is capable of inducing a state of intoxication,
pleasant dreams, and other hallucinations, and of which the local names are
Yage, Ayahuasco, and Caapi, suggests that it is a species of Banisteria. The
 MALPIGHIACEAE 285
bark of certain species of Bynonima is used in tanning. Many of the woods
have an attractively variegated colour, but few of the trees are large enough
to be important sources of timber. Some species of Byrsonima are large trees
and produce timber that is used for various purposes.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acmanthera, Acridocarpus, Aspicarpa, Aspidopterys, Banisteria, Blepha-
randra, Burdachia, Byrsonima, Camarea, Caucanthus, Coleostachys,
Diacidia, Dicella, Dinemagonum, Diploterys, Echinopteris, Flabellaria,
Galphimia,* Gaudichaudia, Glandonia, Heteropteris,* Hiptage,* Hiraea,
Janusia, Lophopterys, Malpighia,* Mascagnia, Mezia, Microsteira, Peixotoa,
Philgamia, Pterandra, Ryssopterys, Schwannia, Spachea, Sphedamnocarpus,
Stigmaphyllon, Tetrapteris, Thryallis, Triaspis, Tricomaria, Trico-
mariopsis, Tristellateia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Banisteria, Bunchosia, Burdachia, Byrsonima, Diacidia, Glandonia,
Heteropteris, Lophanthera, Malpighia, (Peixotoa), Ptilochaeta, Spachea,
(Stigmatophyllon), Tetrapodenia, Tetrapteris, Thryallis, (Tricomaria),
Triopteris.

LITERATURE
(i) On
General Anatomy
Chodat and Vischer 399, Clausen 431, Dubard and Dop 612, Niedenzu 1594, 1598*
O'Donell and Lourteig 1631, Perrot and Raymond-Hamet 1698.

(ii) On Wood Structure

Bailey 78, Benoist 170, Chalk and Chattaway 358, Cozzo 2517, Heimsch 938, Kribs
1283, Pfeiffer, H. 1712, Pfeiffer, J. Ph. 1713, Record 1815, 1843, 1851, Record and Hess
1886, Record and Mell 1894, Stone 2202, 2207, Williams 2430.

s/73. ZYGOPHYLLACEAE
(Fic. 65 on p. 270; FIG. 68 on p. 286; FIG. 69 on p. 290)

SUMMARY
(i) GENERAL
A tropical and sub-tropical family of shrubs or somewhat woody herbs
with hairy, fleshy, or leathery leaves. Certain species of Kallstroemia, Tribulus,
and Zygophyllum are annuals. The Zygophyllaceae and Thymelaeaceae are
said to be the only families poisonous to camels. The hairs are usually
unicellular and
simple, but
more rarely unicellular and 2-armed. The rneso-
phyll of the and partly composed of water-storage
leaf is generally centric,
cells in certain species. vascular bundles of the leaf veins are sometimes
The
surrounded by a sheath of thin or thick-walled parenchymatous cells which
may contain chlorophyll. The petiole, in transverse sections, exhibits a
central vascular strand which is somewhat complex in structure, and therefore
variable in appearance in sections taken at different levels within a single
286 ZYGOPHYLLACEAE
petiole. In most instances the central strand has the form of a closed or
somewhat interrupted cylinder. Two or more subsidiary strands also occur
in the latero-superior position. The primary cortex of the axis is usually
composed of thin-walled parenchyma in which stone cells are sometimes
embedded. The pericycle is generally characterized by separate strands of
fibres, between which stone cells are sometimes present. The phloem and

FIG. 68. ZYGOPHYLLACEAE

A, Hair of Larrea divaricata Cav. B, Glandular hair of Fagonia arabica L. C, Crystal-cells from the
phloem of Guaiacum officinale L. D, Transverse section of the leaf of Tribulus alatus Del. A, By
Solereder; B and D after Volkens; C after MSller.

xylem in the internodes of the young stem constitute closed cylinders sur-
rounding the pith, or the xylem may be interrupted by broad primary
medullary rays. Crystals are mostly clustered, but sometimes solitary.
Styloids are especially characteristic of the phloem of certain species of
Guaiacum, Larrea, and Porlieria.

(ii) WOOD
The most striking characters of the woods are the storying of all the elements
and the high number of stories per mm. (8-17), the predominance of fusi-
form parenchyma cells, very small intervascular pitting, and the occurrence
of tracheids. With the exception of Balanites, which has large unstoried rays
and no crystals, other characteristic features are narrow, low rays and
abundance of crystals in the parenchyma and sometimes in the rays. Vessels
very small to moderately large, exclusively solitary or with oblique radial
 Z YGOPH YLLACEAE 287
pattern or in large clusters, perforations simple, intervascular pitting alter-
nate; members very to extremely short. Rays all narrow, short, and storied
or all large and not storied; homogeneous. Parenchyma usually typically
apotracheal, diffuse or in uniseriate bands. Fibres with bordered pits, the
borders moderately distinct to indistinct with thick walls, usually storied of
; ;

medium length in Balanites, very to extremely short in the other genera.

Vasicentric tracheids usually present.

LEAF
Usually dorsiventral, but centric in certain genera (see 'MesophylF below).
Hairs mostly simple and unicellular (Fig. 68 A), either with thick walls and
narrow lumina or with thin walls and relatively wide lumina; more rarely
unicellular but 2-armed, this type occurring especially in the Agrophyllum
section of Zygophyllum. A
dense clothing of procumbent unicellular hairs has
been recorded in species of Bulnesia, Chitonia, Kallstroemia, Sericodes,
Tribulus, Viscainoa. Glandular hairs common on small elevations of the leaf
surface in Fagonia (Fig. 68 B) and glands with a clavate or spherical head in
Peganum harmala Linn. The whole surface of the leaf is covered with
mucilage glands in Zygophyllum fabago Linn, according to Cunningham
(515). The resinous substances covering the surface of Neoschroetera
tridentata (DC.) Briq. (syn. Larrea mexicana Moric) are stated to be secreted
from the epidermis of the stipules and not from external glands. Cells of the
epidermis usually provided with a thick cuticle; polygonal in surface view
in certain species of Guaiacum, Larrea, Porlieria, and Zygophyllum. Every
epidermal cell contains a crystal in Guaiacum officinale Linn. Stomata
generally present on both surfaces, but more numerous on the lower side,
either sunk or raised above the level of the epidermis; ranunculaceous.
Sunken stomata recorded especially in the leathery leaves of species of Bul-
nesia, Guaiacum, Neoschroetera, Pintoa, Porlieria and in the fleshy leaves of
Zygophyllum. Small black dots, each consisting of a few large thin- walled cells
lying under the epidermis and accompanied by a few cells of the epidermis
itself with brownish contents, present in Pintoa chilensis Gay. Hypoderm
recorded beneath the upper epidermis in Peganum harmala. Mesophyll
centric in certain species of Guaiacum, Larrea, Nitraria, Porlieria, Seetzenia,
Tribulus (Fig. 68 D), and Zygophyllum', containing scattered mucilage cells in
certain species of Nitraria, and tanniniferous idioblasts in two species of the
same genus. Lowermost cell layer of the mesophyll consisting of colourless

water-storage cells in certain species of Tribulus; central portion of the

cylindrical leaf consisting of aqueous tissue in Zygophyllum simplex Linn.
Mesophyll containing spicular fibres branching off from the sclerenchyma of
the veins and sometimes extending to the epidermis in Nitraria sp. Vascular
bundles of the veins enveloped by sheaths of pitted cells with fairly thick
walls in Tribulus spp. (Fig. 68 D) and Zygophyllum sp. according to Sabnis
(1977) and Solereder, and by sheaths of thin-walled cells in Balanites rox-
burghii Planch, and Fagonia sp. Sheaths absent from around the veins in
Seetzenia sp. Petiole, in, transverse sections, exhibiting a central ring of
vascular bundles in Bulnesia, Fagonia, Guaiacum, Larrea, Porlieria, Tribulus,
but closed vascular cylinders also recorded or observed in Guaiacum (Fig. 65 D)
 288 Z YGOPH YLLA CEAE
and in Zygophyllum the central strand
; accompanied by 2 or more lateral
is

bundles in all of the species examined. The

course of the bundles is some-
what complex in the petiole owing to splitting and fusion, the degree of
separation of the strands in the central ring therefore varying in sections
taken at different levels. The main vascular strand of the petiole has the form
of a continuous cylinder accompanied by groups of pericyclic fibres similar
to those of the axis in Guaiacum and Zygophyllum when observed in trans-
verse sections through the distal end. Two subsidiary vascular strands also
present in a latero-superior position in the same 2 genera. Petiole of Balanites
aegyptiaca (L.) Delile with a circle of closely approximated but individually
distinct bundles surrounding a large, collateral, medullary strand. Leaves
of Zygophyllum stapfii Schinz become detached from the plant in periods of
drought according to Zemke (2505). Crystals usually clustered, the clusters
being very large in Balanites aegyptiaca, solitary ones more frequent in the
axis acicular crystals recorded in the leaf tissue of Nitraria sp. and Peganum
;

harmala Linn, and raphide sacs in Peganum crithmifolium Eichw. A single

crystal said to occur in every epidermal cell in Guaiacum officinale. Crystalline
masses recorded in the cells of Fagonia and in the intercellular spaces of
Nitraria and Zygophyllum, these being especially characteristic of species
growing on saline soils.

Axis
YOUNG STEM (Fig. 65 E)
Epidermis composed of cells with very thick cuticle on the outer walls in
Guaiacum officinale Linn, and Balanites aegyptiaca (L.) Delile. Stomata in
the last of these species situated in deep pits. Cork arising in the epidermis
or sub-epidermis in most species, but originating on the inside of the strands
of pericyclic fibres in Fagonia cretica Linn.; usually strongly developed.
Primary cortex consisting of aqueous tissue in Tribulus sp. and Zygophyllum ;

the broad cortex of Augea capensis Thunb. is composed of thin-walled

parenchyma according to Schonland (2047). Stone cells present in the cortex
of species of Balanites and Bulnesia, and stone cells and, according to Cunning-
ham (515), sclereids at the nodes of Zygophyllum fabago Linn. Bundles of
fibres were observed in the primary cortex as well as in the pericycle of Z.

fabago grown at Kew. Pericycle containing well-defined, isolated strands of

fibres in Augea, Balanites, Fagonia, Guaiacum (Fig. 65 E), Seetzenia, Tribulus,
and Zygophyllum the fibre strands stated by Cunningham (515) to be inter-
;

rupted at the nodes in Zygophyllum fabago. Pericycle containing a composite

and continuous ring of sclerenchyma in Balanites aegyptiaca and Peganum.
Xylem and phloem forming closed cylinders in the internodes of Guaiacum
and Zygophyllum, but the xylem is said to consist of a composite cylinder of
closely placed bundles in certain species of Balanites, Fagonia, Seetzenia, and
Tribulus. Vessels usually small (radial diameter seldom more than 30 \L in
Guaiacum officinale, but attaining 60 \i in Balanites aegyptiaca}, mostly
solitary, but sometimes tending to be in radial rows. Vessels with simple
perforations. Pith consisting of thin-walled cells in Fagonia, Tribulus, Zygo-
phyllum (very large and becoming hollow in Z. fabago)', containing stone cells
in Guaiacum. Crystals mostly solitary or clustered; styloids recorded in the

phloem of Guaiacum officinale (Fig. 68 c), Neoschroetera tridentata (DC.)

 ZYGOPHYLLACEAE 289
Briq. (syn. Larrea mexicana Moric); small acicular crystals present in the
phloem of Nitraria schoberi Linn, and in the epidermis, primary cortex,
phloem, and rays of Peganum harmala Linn. Large mucilage cavities
recorded in the primary cortex and secondary phloem of Nitraria sp.
crystals sometimes being deposited in them as well.

WOOD (Fig. 69 I-M)

Vessels medium-sized (mean tangential diameter 100-200 p) in Balanites
and Guaiacum, very small to moderately small (50-100 /*) in the other genera;
solitary in Guaiacum, Larrea, Nitraria (1493), and Porlieria, but with a few
multiples in Bulnesia and Sericodes (938) and in large clusters, mostly of
solitary vessels, in Balanites with a radial or oblique pattern in Bulnesia,
;

Nitraria (1493), Plectrocarpa (495), and Porleiria and sometimes in Balanites ;

varying in number from fewer than 5 per mm. in Balanites to more than
70 per mm. in Larrea\ tending to be ring-porous in some species of Bulnesia
and Plectrocarpa (495). Perforations exclusively simple. Intervascular pit-
ting alternate, very small and numerous; pits to ray sells similar to the inter-
vascular pitting (absent from Balanites as the vessels do not touch the rays).
Vessels commonly filled with gummy or resinous deposits, except in
Balanites. Mean member length 0-07-0- 2 mm. Parenchyma predominantly
apotracheal in Balanites, Larrea, Plectrocarpa, and Porlieria, diffuse and in
numerous, irregular, uniseriate bands; similar parenchyma present, and some-
times predominant, in Guaiacum but often tending to be associated with the
vessels abaxiaily and sometimes accompanied by narrow aliform wings
(Fig. 69 M); almost entirely paratracheal in Bulnesia and Nitraria (1493);
apparently terminal bands sometimes present. Storied; usually with 8-n
stories per mm., but according to Cozzo (495), commonly with 14, and up to
17 stories per mm. in Plectrocarpa. Fusiform cells very common, and some-
times almost the only type of parenchyma present apart from crystalliferous
strands; crystals present, except in Balanites', in idioblasts in Larrea and in
chambered cells in Bulnesia, Guaiacum, and Porlieria. Markedly disjunctive
and with conspicuously grouped pits. Rays (except in Balanites and Bulnesia
retama (Gill.) Gris. and Nitraria (1493)), small, 1-2 cells wide, exclusively
uniseriate in some species of Guaiacum, Porlieria, and Sericodes (938);
exceptionally low, not more than 8 cells high and commonly not more than
4-6; storied, 10-11 stories per mm.; according to Cozzo (495), the rays in
i
species of Plectrocarpa are up to about 3 cells wide, commonly more than
i story high and themselves not storied; homogeneous (Kribs's Types II and

III), except in Larrea, which has single marginal rows of square cells. In
Bulnesia retama and Nitraria the rays are reported (1493, 1886) to be up to
3 and 3-4 cells wide respectively and 30 cells high. In Balanites the rays are
all broad (uniseriate rays absent), up to 20 cells wide and 1-7 mm. high, and
with sheath cells. Fibres with distinctly bordered pits, which are more
numerous in the tangential than the radial walls, in Balanites ;.pit$ in the other
genera with moderately distinct (Larrea) to indistinct borders. Heimsch (938),
however, states that the fibrous elements are in most cases tracheids and
seldom fibre-tracheids. Storied except in Balanites. Walls thick. Mean
length about 1*0 mm, in Balanites, o*35~O'6 in the other genera. Vasicentric
tracheids typically present, but not observed in Porlieria.
 '

FIG. 69. OXALIDACEAE, A-C; MALPIGHJACEAE, D-H; ZYGOPHYLLACEAE, I-M

A, Connaropns griffitkii Planch. B, C, griffithii Planch. C, Averrhoa carambola Linn. D,Ptilochaeta
nudtpes Griscb. E Glandonia macrocarpa Griseb. F, Byrsonima coriacea (Sw.) Korth. var. B. G
Bunchona cornifolia H. B. et K. H, B. cornifolia H. B. et K. I, Balamtes aegyptiaca Del. J, B. aegyptiaca
Del. K, Bulnesia arborea Engi. L, Guaiacum officinale Linn. M, G. officinale Linn.
 ZYGOPHYLLACEAE 291
ROOT
Primary structure diarch in Fagonia, Tribulus, and Zygophyllum, but triarch
in Guaiacum and Porlieria.
Tubercles of two distinct sizes, somewhat resembling the bacterial nodules
of the Leguminosae, and infested both externally and internally by fungal
hyphae, recorded by Issatschenko (1125) in Tribulus terrestris Linn, and by
Sabet (1976 A) in Fagonia, Tribulus, and Zygophyllum. The root structure of
Tribulus cistoides Linn, is described under 'Economic Uses'.

TAXONOMIC NOTES
Balanites. Record (1786) states that the wood structure of Balanites bears no
resemblance to that of Bulnesia, Guaiacum, and Porlieria. Though strikingly
different owing to its large rays and clustered pores, it has many less obvious,
but not necessarily less significant, features in common
with these genera,
e.g. vasicentric tracheids, small vessel pitting, fibres with thick walls and
bordered pits, distribution of parenchyma, abundance of fusiform parenchyma
cells, storying of elements other than the rays and the number of stories

per mm.
Heimsch (938) considers that, apart from the rays, the wood anatomy of
Balanites suggests affinity with the Zygophyllaceae rather than with the
Simarubaceae. Elsewhere, however, he states that 'the high and wide rays of
Balanites may be evidence from a structural point of view for excluding this
genus from the Zygophyllaceae which otherwise have only low, and narrow
rays'.
The woods of the family form a very distinctive, natural group, and must
be regarded as highly specialized. They are remarkable for their combination
of several highly specialized features, such as a vessel member length of
O'i-c-2 mm,, storied structure, homogeneous rays and fusiform parenchyma
cells, with several other features, such as solitary vessels, diffuse parenchyma
and fibres with bordered pits, which are usually associated with an un-
specialized structure.

ECONOMIC USES
The only important timber from this family is the Lignum Vitae, Guaiacum
officinale Linn, from the western Indies and South America. This timber has
long been known to commerce one of its principle uses is for the bearings or
;

bushing blocks for lining the stern tubes of the propellor shafts of steamships,
for which the self-lubricating properties of the wood, owing to its resin con-
tent, make it especially suitable. Other uses include bowling balls, mallets,
and pulley sheaves. Guaiacum resin, used in medicine, is formed chiefly in
the cells of the medullary rays of the same species and of G. sanctum Linn.
The twigs of the Creosote Plant Neoschroetera tridentata (DC.) Briq. (syn.
Larrea mexicana Moric.) yield a resinous substance used locally in the treat-
*

ment of rheumatism.
The root of Tribulus cistoides Linn, possesses medicinal properties. Trans-
verse sections of the roots of this species i cm. thick exhibit, according to
Diepenbrock (587), a yellow central cylinder clearly demarcated from the
surrounding white tissues. Vessels tending to be in radial rows, and accom-
panied by tracheids and thick- walled parenchyma. Groundwork of the xylem
292 Z YGOPH YLLA CEAE
composed of fibres. Rays mostly uniseriate, but sometimes partly or wholly
2 or more cells wide. Wood surrounded by cambium and phloem, the latter
consisting of a cylinder of small, unthickened, starch-free cells. Large masses
of fibres are present externally to the phloem with extensions of the rays
between them. Cortex represented by a narrow band of starch-containing
cells, bounded externally by several layers of cork cells,

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Augea, Balanites,* Bulnesia, Chitonia, Fagonia, Guaiacum,* Kallstroemia,
Larrea, Neoschroetera, Nitraria, Peganurn, Pintoa, Porlieria, Seetzenia,
Sericodes, Tribulus, Viscainoa, Zygophyllum.*
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Balanites, Bulnesia, Guaiacum, Larrea, (Nitraria), (Plectrocarpa),. Porlieria,
(Sericodes).
LITERATURE
(i) On General Anatomy
Cunningham 515, Diepenbrock 587, Engler 639, Evenari (Schwarz) 665, Issatschenko
1125, Sabet 1976 A, Sabnis 1977, Schonland 2047, Zemke 2505.

(ii) On Wood Structure

Heimsch 938, Howard 1088, Jones 1191,
Burgerstein 312, Cozzo 495, Foxworthy 705,
Kanehira 1209, Messeri 1493, Record 1781, 1783, 1786, 1809, 1818, 1843, 1851, Record
and Hess 1886, Record and Mell 1894, Wallis 2348.

^/ 74. GERANIACEAE
(FiG. 70 on p. 296)

SUMMARY
(i) GENERAL
A
mainly herbaceous family which occurs chiefly in temperate regions. It
also includes succulent and shrubby species. The genera described are fairly
homogeneous in structure, the greatest differences being between herbs and
shrubs. The desert genus Sarcocaulon has a highly specialized structure in
relation to its habitat. The hairs consist either of simple, unicellular and
uniseriate types or they may be glandular. The oil secreted by the glandular
hairs is highly scented and sometimes distilled for use in perfumery. The
odour which is usually associated with Geranium is due to these oils. Tannic
acid is also common in the tissues, notably in Pelargonium as well as in species

of Erodium and Monsonia from the Egyptian desert. Crystals solitary or

clustered. The stomata are ranunculaceous. The pericycle in the axis of
Erodium, Geranium, and Pelargonium usually contains a well-defined, con-
tinuous ring of sclerenchyma, which is sometimes composite, especially in
Erodium. The vascular bundles in the stem of herbaceous species are
usually widely separated and individually distinct, arranged in i or sometimes
in 2 rings according to the species. The inner bundles tend to be medullary
 GERANIACEAE 293
in certain species of Geranium. In some species the phloem of the outer
bundles abuts directly on to the sclerenchyma of the pericycle, whilst in other
species, such as Geranium pratense Linn., inwardly directed sclerenchymatous
projections serve to connect the phloem groups of the individual bundles with
the main sclerenchymatous ring. The xylem and phloem form closed
cylinders in mature sterns of shrubby species owing to the development and
activity of interfascicular cambium. The vessels have simple perforations,
and the wood fibres are provided with simple pits. The root exhibits peculiar
thickening ridges in the sub-epidermis of many species of Er odium, Geranium,
and Pelargonium.

(ii)
WOOD
Vessels with simple or rare multiperforate plates and alternate to opposite
pitting. Parenchyma scanty paratracheal. Rays often absent. Fibres
commonly septate.

LEAF
Hairs simple, unicellular (Viviania marifolia Cav.) or uniseriate; glandular
types with uniseriate stalks of varying length and unicellular, spherical heads
in Erodium, Geranium, Monsonia, Pelargonium, Viviania^ and Wendtia\
glandular shaggy types with long multiseriate stalks and multicellular knob-
shaped heads in Biebersteinia. Glandular leaf teeth occur in Geranium
robertianum Linn. Glandular hairs, where present, secreting oils. Cells of
the upper epidermis described by Evenari (665) as having thin outer walls
and cuticle in Erodium glaucophyllum Ait. those of Sarcocaulon found by
;

Planchon (1728) and Zemke (2505) to be unequal in size and to contain

chlorophyll. Epidermis of Monsonia heliotropioides Boiss., according to
Sabnis (1977), papillose especially on the lower surface. Stomata present on
both surfaces or confined to the lower side according to the species; ranun-
culaceous. Stomata situated in grooves on either side of the principal vein on
the lower side of the leaf in Balbisia microphylla Phil.; present in equal
numbers on both surfaces in Pelargonium coronopifolium Jacq. although the leaf
is deeply furrowed only on one side owing to the margins being re volute.

Mesophyll isobilateral in Erodium glaucophyllum Ait. and Monsonia helio-

tropioides Boiss. Vascular bundles of the veins said by Legault (1342) to be
surrounded by an endodermis in Erodium and Geranium. Larger veins
vertically transcurrent in Monsonia heliotropioides Boiss. Petiole exhibiting
a circle of separate vascular bundles in transverse sections through the distal
end in Biebersteinia, Erodium, Geranium (Fig. 70 c), and Monsonia. An outer
circle of about 14 relatively small bundles and an inner circle of 5 larger ones
recorded in Geranium anemonifolium L'Herit. Xylem groups of the bundles
almost surrounded by phloem in Geranium phaeum Linn., but this character
is less pronounced in G. pratense Linn. Phloem groups bounded externally

by caps of fibres in the pericyclic region, the latter connected by interfascicular

sclerenchyma so as to form a ring in some species, e.g. in Geranium robertianum
Linn, and Pelargonium zonale (L.) Ait., but interfascicular sclerenchyma
poorly developed in other species, e.g. Geranium phaeum Linn., or absent,
e.g. in Geranium pyrenaicum Burm. f. and Erodium moschatum (L.) Hr.
Petiole structure of Pelargonium (Fig. 70 F) very similar to that of Geranium,
294 GERANIACEAE
but the ring of bundles encloses a large, central medullary strand in many
species. For details of petiolar structure in Pelargonium zonale (L.) Ait. var.
Meteor see Doyle's (609) article. A single arc-shaped strand flanked on either
side by a smaller bundle occurs in the distal end in Viviania petiolata Hook,
et Arn. and a solitary but well- developed strand in Rhyncotheca and Wendtia.

Crystals solitary or clustered, sometimes situated in idioblasts in Erodium\

styloids in Rhyncotheca', sphaerocrystalline masses in Erodium and Monsonia \

short rows of cells containing cluster crystals in Wendtia.

Axis
STEM (Fig. 70 A, D, j, and K)
Cork originating in the hypodermis, and consisting of cells with wide
lumina and thin walls inMonsonia. Cortex usually rather narrow, bounded
internally by a closed, but often composite (especially in Erodium (Fig. 70 D))
ring of mechanical tissue representing the pericycle. Vascular bundles of
Erodium moschatum (L.) Her. each provided with a cap of pericyclic fibres
which are smaller in diameter than those of the arcs of interfascicular scleren-
chyma which unite the fibre caps of the bundles so as to form a continuous
ring. The width of the ring of mechanical tissue varies in different species.
Pericycle containing isolated groups of fibres in certain species of Monsonia.
Endodermis conspicuous in young stems of Geranium pratense Linn. Vas-
cular bundles usually widely separated (Fig. 70 D and K), arranged in i or
sometimes in 2 more or less distinct rings according to the species, the inner
ones said by Solereder sometimes to be inversely orientated. The inner
bundles, where present, are larger than the outer ones. Bundles arranged in
a single ring in Geranium phaeum Linn. in 2 more or less distinct circles, but
;

connected by sclerenchymatous tissue to the mechanical ring, e.g. in the

pericycle of G. dissectum Linn., G. lucidum Linn., and G. maccorhizum Linn.;
in a single ring when young, but the inner bundles become embedded in the
pith with increase of age, e.g. in G. collinum Steph. and G. pratense Linn. ;

inner bundles free in the pith, outer ones connected with the sclerenchyma
ring in G. robertianum Linn, Stem structure in Erodium and the herbaceous
species of Pelargonium similar to that of Geranium, but the xylem and phloem
form closed cylinders in the shrubby species of Pelargonium (Fig. 70 j) owing
to the formation and activity of an interfascicular cambium. Vessels, except
in the protoxylem with spiral thickening, provided with horizontal bordered
pits perforations usually simple, but reticulate types recorded in Pelargonium
;

(Thompson 2254). Wood fibres with simple pits except in Viviania. Pith
consisting of thin- walled parenchyma; becoming hollow in certain species.

WOOD 1

Vessels solitary and in multiples and clusters, and often, especially in

Geranium, confined to radial strips; sometimes ring-porous; with spiral
thickening in Balbisia. Perforation plates typically simple, but occasional
multiperforate plates (ephedroid, reticulate, and scalariform) have been
observed (1408, 2254) in Pelargonium. Intervascular pitting predominantly
alternate, but usually with some transitional and opposite types; pits to ray
and wood parenchyma cells large and elongated. Parenchyma paratracheal,
1
Based mainly on the description given by Heimsch (938).
 GERANIACEAE 295

scanty. Rays, when present, heterogeneous and very variable in height and
Absent from some species of Balbisia, Momonia,- Viviana, and
size of cell.
Wendtia, and, according to Barghoorn (139), from the early formed wood of
Geranium tridens Hbd., though high-celled multiseriate rays develop later.
Fibres with simple or indistinctly bordered pitsj often septate; walls thin
except sometimes in Balbisia.
RHIZOME
Endodermis and pericycle not well defined in Geranium maculatum Linn,
according to Holm (1026) but with a single, somewhat excentric ring of
vascular bundles with interfascicular cambium between them.
ROOT
A characteristic network of thickening ridges recorded in Erodium,
Geranium, and Pelargonium by Solereder, Holm (1026), and Scott and Whit-
worth (2074). These thickenings appear to be lignified when tested with
phloroglucin and hydrochloric acid or with aniline sulphate, but respond to
cellulose reagents after treatment with Schultze's macerating fluid. Fatty
acids have been detected in them as well.

SARCOCAULON
The stem of Sarcocaulon, a frutescent plant from desert regions in South-
west Africa, which bears thorns formed from the petioles after the lamina
the end of the rainy season, is so specialized in structure that the
falls off at

following separate description, based on the accounts by Knuth (1255),

Planchon (1728), and Zemke (2505), is given.
Stem covered by a yellow, elastic inflammable layer of cork, easily detached
from the central tissues, and consisting of several layers of cells filled with
resinous contents. The cork serves as an hermetically sealed covering during
dry weather (cf. the similar cork in certain species of Kalanchoe (family
Crassulaceae) from Madagascar). Cortex represented by a zone of parenchy-
matous tissues containing chlorophyll and crystals of calcium oxalate, a special
form of starch (not stained readily by iodine) and resinous substances.
Pericycle demarcated by a few fibres situated externally to the phloem.
Phloem in radially arranged groups, separated by fairly broad, medullary
rays 4-6 cells wide. Ray cells filled with starch, a few cluster crystals of
calcium oxalate, and resirous fragments. Xylem containing numerous
vessels of small diameter. Pith similar in structure to the cortex.

TAXONOMIC NOTES
The family constitutes a homogeneous group except for the specialized
desert plant Sarcocaulon, in which the rather different structure has evidently
been evolved in response to the arid conditions in which the plant grows.
Heimsch (938) has pointed out the similarity between the woods of this
family and those of the Oxalidaceae, particularly the tendency to elimination
of rays, the scanty paratracheal parenchyma and the common occurrence of
septate fibres.
MacDuffie (1408) used the structure of the perforation plates of Pelargonium
argument against the hypothesis that the types of vessels in
to illustrate his
the Gnetales and the angiosperms are distinct in their mode of derivation.
 FIG. 70. GFRANIACEAE, A, OD, F, and J-K; BALSAMINACEAE, B and H;
TROPAEQLACEAE, E and L; OXALWACEAE, G and I
A, Geranium robertianum Linn, Stem X 15. B, Impatiens glandulifera Royle. Petiole X 10. C,
Geranium pyrenaicumBurm. f. Petiole x 12. D>Erodiummoschatum(L.)H&T, Stem xy. E,Tropaeolum
mains Linn. Petiole X7. F, Pelargonium zonale (L.) Ait. Petiole xiz. G, Oxalis corniculata Linn,
cult. var. Stem X 15. H, Impatiens glandulifera Royle. Stem x 10. I, Oxalis corniculata Linn.
Petiole X 15. J, Pelargonium sonale (L.) Ait. Stem X 7. K, Geranium pyrenaicum Burm. f. Stem X 7,
L, Tropaeolum majus Linn. Stem X 7.

e. Endodermis. k.p. Hollow pith. p.p. Parenchymatous pith.

 GERANIACEAE 297
ECONOMIC USES
Geranium oil is distilled chiefly from the leaves of selected varieties of
Pelargonium. Many species of Geranium and Pelargonium are cultivated for
ornamental purposes.
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Biebersteinia, Erodium,* Geranium,* Monsonia, Pelargonium,* Sarco-
caulon, Viviania, Wendtia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Balbisia), (Geranium), (Monsonia), (Pelargonium), (Viviana), (Wendtia),

LITERATURE
(i) On
General Anatomy
Doyle 609, Evenari (Schwarz) 665, Holm 1026, Knuth 1251, 1252, 1255, Legault 1342,
Planchon 1728, Sabnis 1977, Scott and Whitworth 2074, Thompson 2254, Zemke 2505.

(ij) On Wood Structure

Barghoorn 139, Heimsch 938, Macduffie 1408, Record 1843, 1851, Thompson, W. P.
2254-

y75. BALSAMINACEAE
(Fie. 70 on p. 296)

SUMMARY
A
small family of succulent herbs belonging to the genera Impatiens and
Hydrocera. The family is mainly tropical but some species occur in temperate
regions. The most characteristic anatomical feature is the occurrence of
raphides.
LEAF
Dorsi ventral. Extra-floral nectaries recorded on the petiole and stem of
certain species of Impatiens. Hydathodes present in the leaf teeth. Stomata
confined to the lower surface in /. sultani Hook, f., but observed on both
surfaces in other species of Impatiens examined at Kew mostly ranunculaceous
;

but some tending to be cruciferous. Petiole in transverse sections through

the distal end of Impatiens glandulifera Royle (Fig. 70 B) exhibiting an arc of
separate bundles, but with thin-walled tissue, resembling phloem, present
between the phloem groups of the individual bundles. Raphide-sacs
present in the leaf (and stem) of at least 62 species of Impatiem, sometimes
appearing as transparent dots; frequently containing mucilage as well as
raphides. Branched tannin-sacs present in the spongy parenchyma of
/. sultani.

Axis
STEM (Fig. 70 H)
cells. Cortex relatively narrow,
Epidermis composed of small, thin-walled
outer part consisting of small collenchymatous cells and the inner part of
298 BALSAMINACEAE
few species examined. Pericycle apparently
large, thin-walled cells in the
devoid of sclerenchyma, the rigidity of the stem being maintained by the
strongly turgescent ground tissue. Vascular bundles individually distinct
and arranged in a circle of about 12 as seen in transverse sections; consisting
mainly of xylem composed of a ground tissue of small, very thin-walled cells
with large vessels embedded in it, the latter being provided with very well-
developed spiral thickening. Phloem strands in each bundle small, those of
adjacent bundles being connected together by a small-celled tissue with thin
walls, rather suggesting phloem in appearance, the true nature of which needs
further investigation. Vessels with simple perforations. Interfascicular
cambium arising, in older stems, in the region of the small-celled tissue just
mentioned, but apparently giving rise only to thin-walled tissue on the inside,
comparable with the ground tissue of the xylem, but not including any
vessels. Pith becoming hollow at the centre. Isolated, annular, or spiral
vessels statedby Solereder to arise in the pith of certain species of Impatiens
before the development of the vascular bundles. Large cells with muci-
laginous contents scattered throughout the ground parenchyma. Bundles of
raphides present in the cortex.

TAXONOMIC NOTES
Impatiens was included by Bentham and Hooker in the Geraniaceae, from
which the genus differs, however, (i) in having an arc instead of a circle of
bundles in the petiole, (ii) in the absence of a ring of mechanical tissue from
the pericycle, (iii) in the presence of raphide sacs. These anatomical features
confirm the desirability of treating the Balsaminaceae as a separate family.

ECONOMIC USES
Several species of Impatiens are cultivated for ornamental purposes.

GENUS DESCRIBED
Impatiens.* The
description of the petiole and stem structure is based
mainly on an examination of /. glandulifera Royle grown at Kew.
* Kew
Represented in the slide collection.

76. LIMNANTHACEAE
SUMMARY
A small family of succulent marsh herbs occurring in North America. The
anatomy has been studied especially by Russell (1969) on whose paper the
following description is mainly based.

LEAF
Dorsiventral. Epidermis consisting of rounded, unthickened cells.
Stomata present on both surfaces, but more numerous on the lower than on
the upper side. Mesophyll consisting of i layer of palisade tissue interrupted
by intercellular spaces below the stomata, and a narrow zone of spongy tissue
below the palisade with small bundles embedded in it. A single hydathode
 LIMNANTHACEAE 299
is present at the tip of the leaf. Tannin cells occur on the lower side of the
leaf in Limnanthes, but not seen in Floerkea.

Axis
STEM
Weak and flaccid, remaining green and rarely branched in Floerkea; stouter
and more branched, especially at the base, in Limnanthes. Epidermis com-
posed of small rectangular slightly cutinized cells. Cortex 6-7 layered,
consisting of large round cells with thin walls. Endodermis indistinct.
Vascular bundles 8-10 in a ring, appearing individually distinct and
separate in transverse sections. Phloem poorly developed. Interfascicular
cambium absent.

ROOT
Root system consisting of a few short, fibrous, unbranched roots. Adventi-
tious roots also arise from the nodes in procumbent species. Epidermis
cutinized in mature roots. Cortex consisting of 2-4 layers of large round cells
with thin walls. Endodermis well defined, consisting of elliptical cells
thickened on the inner tangential and lateral walls. Stele consisting of a
simple diarch bundle.

TAXONOMIC NOTES
Limnanthes was included in the Geraniaceae in the system of Bentham and
Hooker, but more recently this and the anatomically similar genus Floerkea
have been placed in a separate family the Limnanthaceae. The anatomical
characters in themselves scarcely provide sufficient evidence to determine
whether they should constitute a separate family or not. It is interesting to
note, however, that the Limnanthaceae resembles the Geraniaceae in having
widely spaced vascular bundles in the stem, but there is, on the other hand,
no trace of the characteristic mechanical ring in the pericycle.

GENERA EXAMINED
Floerkea, Limnanthes.

LITERATURE
On General Anatomy
Russell 1969.

V/77. OXALIDACEAE
(FiG. 69 on p. 290; FIG. 70 on p. 296; FIG. 77 on p. 334)

SUMMARY
(i) GENERAL

Most members of the family are herbs, but a few shrubby species occur,
whilst a still smaller number resemble small trees. They mostly occur in
tropical and sub-tropical regions, but some species are temperate. There is
a great diversity of form even amongst the herbaceous species, since some are
rosette plants with no true stem, others have an underground rhizome, whilst
300 OXALIDACEAE
there are fleshy and woody species which serve as transitions to the shrubby
and arboreal types. Members of the tuberosae section of Oxalis bear small
root tubers, whilst in other sections of the genus there are species with con-
tractile roots or bulbs. Other interesting features include the conversion of

petioles into phyllodes after abscission of the leaflets in certain species of

Oxalts, and the possession of leaves which exhibit 'sensitive* movements. In
correlation with the range of habit there is a correspondingly wide diversity of
anatomical features. The fundamental vascular structure of both stem and
petiole consists of a circle of collateral bundles, often accompanied externally
by a sclerenchymatous ring in the pericycle. In some species of Oxalis the
xylem of the stem tends to be in the form of a closed cylinder in correlation
with a more woody habit, but this character has been developed even further
in genera like Averrhoa, where the xylem and phloem of the petiole as well as
of the stem appear, in transverse sections, as practically continuous closed
rings. Secretory cavities are characteristic of many species of Oxalis where
they occur in the leaf. As the name of the family implies, oxalic acid is very
common in the tissues, where it is believed to occur in the form of dissolved
potassium oxalate as well as being secreted as calcium oxalate, usually in the
form of small, solitary, cubical crystals. Crystal-sand is said to occur as well
but only very rarely.

(ii) WOOD
Vessels often in multiples of 4 or more cells, perforations simple, inter-
vascular pitting alternate and large, pits to parenchyma often simple, members
of medium length to very short. Parenchyma typically vasicentric and
scanty, but often with numerous diffuse crystalliferous strands. Rays uni-
seriate, heterogeneous to almost homogeneous. Fibres septate, with simple
pits, moderately to very short.

LEAF
Leaflets usually dorsiventral in Oxalis. Simple, unbranched, hairs of
variable length widely distributed in the family. Bladder-like hairs which
serve for water storage recorded in species of Oxalis. Glandular hairs, with
stalks of varying lengths and unicellular heads, stated to occur in Averrhoa,
Biophytum, and Oxalis. Cells of the epidermis of Oxalis often very large in
proportion to the total width of the lamina, sometimes arched outwards; in
some species tall, palisade-like, exceeding the remainder of the total thickness
of the leaf. Lower epidermis papillose in certain species of Eichleria and
Oxalis. Stomata rubiaceous with at least i and sometimes 2 subsidiary cells
parallel to the pore in Averrhoa, Biophytum, and Eichleria, Hydathodes in
depressions in the leaflets of certain species of Oxalis described by Vouk
(2342). Hypoderm present in Dapania scandens Stapf. Vascular bundles of
the veins provided with enlarged terminal tracheids in Averrhoa, Biophytum t

and Eichleria. Petiole, in transverse sections, exhibiting a circle of separate

bundles in many species of Oxalis. The phloem groups of the bundles in the
petiole are attached externally to a ring of sclerenchyma in some species,
e.g. O. corniculata Linn. (Fig. 70 i), but not in others, e.g. O. rosea Jacq.
Closed rings of xylem and phloem surrounding 2 medullary bundles recorded
by Solereder in the distal end of the petiole of Oxalis tetraphylla Cav. The
 OXALIDACEAE 301

xylem and phloem form a practically continuous cylinder around a fairly

extensive, thin-walled, parenchymatous pith in Averrhoa carambola Linn,
(Fig. 77 c). The vascular cylinder is also surrounded by a relatively broad,
almost continuous ring of thick-walled fibres in the same species. Secretory
cavities, variously interpreted as being schizogenous or lysigenous in origin,
containing either brown or red, and sometimes transparent and crystalline
substances, present in the pinnules of Oxalis spp., from both South Africa
and America. These cavities, which occur in different positions in the various
species, are confined to the margins of the pinnules in some, but cover the
whole surface in others, e.g. O. articulata Sav. and O. hirta Linn. Secretory
cavities absent from certain species of Oxalis with a well-developed stem,

e.g.O. cernua Thunb. and O. deppei Lodd.; generally situated on the dorsal
side of the vascular bundles when occurring on the bulb scales often elongated
;

and resembling canals. The distribution of secretory cavities and canals may
be of specific diagnostic value, but further investigation is necessary in order
to determine this point. Crystals mostly solitary, sometimes appearing as
transparent dots in certain species of Oxalis. Crystal cells also accompany
and form sheaths to the vascular bundles of the veins in Averrhoa, Biophytum,
and Eichleria.
In certain species of Oxalis the leaflets become detached from the petiole,
which then constitutes a phyllode. This is well seen, for instance, in O. herrerae
R. Knuth and O. bupleunfolia St. Hil., as described by Metcalfe (1495). In
the first of these species, the petiole becomes considerably swollen and club-
shaped after the leaflets have fallen, as they usually do when the plant is

growing in an exposed position. The increase in the bulk of the petiole is

caused by enlargement of the parenchymatous tissues of which the petiole
is mainly composed. The chlorophyll-containing tissue immediately below the

epidermis becomes more palisade-like the epidermal cells become larger, the
;

cuticle thicker, and the stomata more numerous. There is a circle of small,
individually distinct bundles in both swollen and unswollen petioles. In O.
bupleurifolia the petiole is flattened and leaf- like stomata are present on both
;

surfaces. The ground tissue of the petiole consists of loosely arranged paren-
chymatous cells containing chloroplasts. Vascular bundles interspersed with
fibre bundles are present towards the abaxial side. The bundles on the
adaxial side consist exclusively of fibres and probably represent reduced
fibro-vascular strands in a petiole which was originally radially symmetrical.
Leaves of certain members of the family exhibit 'sensitive* movements
comparable with those which are more familiar in certain members of the
Leguminosae. According to Steckbeck (2190) sensitive movements can be
Biophytum semivitum DC. and B. dendroides DC.
especially well observed in

Axis
STEM (Fig. 70 G and 77 K)
Epidermis consisting of cells of rather variable sizes in the few species of
Oxalis examined. Cork well developed in Averrhoa^ Connaropsis, Eichleria.
Cortex composed of large, loosely packed parenchymatous
relatively narrow,
cells inOxalis and of cells containing oil and large solitary crystals in
Averrhoa carambola Linn. Pericycle bounded by a composite and more or
less continuous ring of sclerenchyma in Averrhoa, Biophytum, Connaropsis,
302 OXALIDACEAE
Dapania, Eichleria, Hypseocharis, and Oxalis (pro parte). Vascular bundles
collateral, widely separated, arranged in a single ring in Oxalis corniculata
Linn. (Fig. 70 G) and O. rosea Jacq. Bundles individually distinct, arranged
in a single ring in young stems of Oxalis herrerae R. Knuth but with a closed
cylinder of xylem in older stems of the same species. Phloem and xylem
constituting closed rings in Averrhoa carambola Linn. (Fig. 77 K). Vessels
small (radial diameter seldom exceeding 40 /A in Averrhoa carambola Linn, or
30 p in Oxalis corniculata Linn,), solitary or in short radial rows; perforations
simple. Ground tissue of the xylem consisting of moderately thick-walled
prosenchymatous elements in Averrhoa carambola, locally replaced by cells
each of which contains a row of solitary crystals. Pith consisting of large,
thin-walled parenchyma in Oxalis corniculata and O. rosea Jacq.; some of the
cells with oily contents in Averrhoa carambola. Pith stated to contain fibres
in Dapania scandens Stapf.

WOOD (Fig. 69 A-C)

Vessels medium-sized (100-200 n mean tangential diameter) or slightly
less solitary, in multiples that often exceed 4 cells, and sometimes in irregular
;

clusters; about 5 per sq. mm.; sometimes with faint spiral thickening in
Sarcotheca. Perforations simple. Intervascular pitting alternate, large; pits
to ray and wood parenchyma commonly simple and sometimes larger than the
intervascular pitting, but tending to be round rather than elongated; occa-
sionally unilaterally compound. Mean length 0-3-0-7 mm. Parenchyma
consisting of a few cells about the vessels, with, in most species, numerous
crystalliferous strands scattered among the fibres (Fig. 69 B). The strands of
vasicentric parenchyma of 4-12 cells, the crystalliferous strands with 25-45
chambers. Rays typically exclusively uniseriate, but most species with a few
10-20 per mm.;
biseriate rays, particularly in Connaropsis griffithii Planch.;

distinctly heterogeneous with i or 2 marginal rows of rather high upright

cells, e.g. in Averrhoa carambola Linn, to almost homogeneous in Sarcotheca

subtriplinervis Hall. f. Containing crystals in some species of Averrhoa.

Heimsch (938) notes that the rays are very poorly developed in some species
of Oxalis. Fibres with numerous small, simple pits on the radial walls;
probably all septate in Averrhoa and Sarcotheca, but with septate fibres tend-
ing to be limited to the neighbourhood of the vessels in Connaropsis griffithii.
Walls thin to thick. Mean length 0-5-1*0 mm.

RHIZOME
Cork very thick, and wood very well developed in Hypseocharis according
to Chauvel (380).

ROOT
Certain species of Oxalis are provided with contractile roots bounded
externally by a few layers of cork, the bulk of the root being composed of
secondary phloem, consisting of radial rows of parenchymatous cells with
isolated sieve tubes scattered throughout. Cortex narrow, parenchymatous.
Stele tetrarch when young. The root is cast off from the plant by an abscis-
sion layer after having fulfilled its function. According to Duncan (614) and
 OXALIDACEAE 303
Rohde (1946) the contraction which serves to pull the plant into the ground
isachieved by the secondary phloem. The cells are vertically elongated at
first and with sap of high osmotic value. The osmotic value of the sap
filled
issubsequently reduced by chemical changes which are accompanied by the
formation of insoluble substances such as calcium oxalate. The cells then
contract longitudinally and, as the plant is securely anchored by the base of
the root, it tends to be pulled into the ground.

TAXONOMIC NOTES
The Oxalidaceae were included in the Geraniaceae in the system of Bentham
and Hooker. There are certain anatomical features common to both of these
families which confirm that they may be closely related to one another. The
basic vascular structure of the stem and petiole in both of them consists of a
ring of separate collateral bundles frequently associated with a characteristic
and well-developed sclerenchymatous ring in the pericyclic region. There
are aberrations from this type in both families, especially in species which
tend to be shrubby or arborescent in habit, where the xylem of adjacent
bundles becomes connected so as to form a continuous ring as the plant
grows older.
Heimsch (938) concludes that the wood anatomy justifies the placing of
Sarcotheca in this family rather than in the Linaceae.

ECONOMIC USES
tubers of certain species of Oxalis are edible. The fruits of the
The
Bilimbi (Averrhoa bilimbi Linn.) and the Carambola (A. carambola Linn.) are
preserved and eaten in India. Oxalic acid is especially abundant in Oxalis spp.
and can be extracted.
GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Averrhoa,* Biophytum, Connaropsis, Dapania, Eichleria, Hypseocharis,
Oxalis.*
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Averrhoa, Connaropsis, (Oxalis), Sarcotheca.

LITERATURE
(i) On General Anatomy
Chauvel 380, Duncan 614, Knuth 1253, 1254, Metcalfe 1495. Rohde 1946, Steckbeck
2190, Vouk 2342.

(ii) On Wood Structure

Heimsch 938, Janssonius 1154, Kanehira 1206, Lecomte 1334, Record 1851.
 (304)

y 78. TROPAEOLACEAE
(FiG. 70 on p. 296)

SUMMARY
Succulent, often scandent herbs from the Andes and other parts of Central
and South America. Some species are characterized by tuberous roots.
Owing to the small number of species which have been studied, the characters
are described under the organs in which they occur.

LEAF
Hairs infrequent, unbranched, consisting of several cells. Hill (969)
induced the formation of unicellular hairs on the usually glabrous petioles of
T< aduncum Smith by surgical manipulations such as removing the lamina.
Epidermis containing cells with mucilaginous contents in Tropaeolum majus
Linn. Stomata always present on the lower surface and in many species on
the upper side as well ranunculaceous. Water pores terminate the vascular
;

bundles in T. majus. Drops of liquid exuded from the leaf veins, petiole, and
young stem of T. majus when cut, are stated by Solereder to arise from
persistently juvenile vessel elements with nucleus and protoplasmic contents.
Petiole (Fig. 70 E) in transverse sections through the distal end of the few
species examined, exhibiting a single ring of separate, collateral bundles
embedded in thin- walled parenchyma; mechanical tissue absent. Spherical
secretory cells, with highly refractive contents, appearing as dots when
examined with a lens, recorded in T. pentaphyllum Lam. sphaero-crystalline
;

masses of the same substance said to occur in T. majus var. Myrosin present
but not apparently localized in special cells (cf. stem and root).

Axis
STEM (Fig. 70 L)
Epidermis consisting of comparatively small cells. Cork stated to arise
in the endodermis of 7\ peregrinum Linn. but apparently more superficial
;

in species examined at Kew. Cortex narrow, consisting of thin-walled

parenchyma. Endodermis fairly well defined and provided with casparian
thickenings in T. majus Linn. Vascular bundles (Fig. 70 L), at first separate,
somewhat variable in size, arranged in a single ring; thexylem groups of
the originally separate bundles later become connected by prosenchymatous
elements with simple pits, formed on the inside of an interfascicular cambium.
Interfascicular phloem is also formed by the activity of the cambium. Vessels
of T. aduncum Smith and T. majus up to about 80 \L in radial diameter
occupying the greater part of the transverse area of each of the bundles those ;

firstformed are provided with spiral thickening, but the later ones possess
small bordered pits; perforations usually simple, but reticulate types and
transitional forms have also been recorded by Thompson (2254). Pith occupy-
ing the greater part of the stem, composed of thin-walled, somewhat loosely
arranged parenchyma. Myrosin cells, capable of detection with Millon's
reagent, present in the sub-epidermal region as well as in the phloem. Con-
trary to Solereder's statement, no mechanical tissue was observed in the
pericycle.
 TROPAEOLACEAE 305
ROOT
Myrosin cells present in the primary cortex and phloem. The statement
that septate laticiferous tubes occur in the tuberous roots of a few species
requires confirmation.
TAXONOMIC NOTES
The genus Tropaeolum was included in the system of Bentham and Hooker
in the Geraniaceae, but since then it has generally been treated as a separate
family related to the Geraniaceae. The arrangement of the vascular bundles
in the stem and petiole is similar to that of the Geraniaceae, but the very
well-developed ring of mechanical tissue in the pericycle, which is a very
characteristic feature of the Geraniaceae, does not exist in the species of
Tropaeolum available for examination. Myrosin, which is commonly present
in Tropaeolum, has not apparently been found in the true Geraniaceae. These
facts favour the inclusion of Tropaeolum in a distinct family, which may,
however, have affinities with the Geraniaceae, although the anatomical
features are not very conclusive concerning this relationship. It is interesting
to note that myrosin also occurs in the Cruciferae, Resedaceae, and Cappari-
daceae, which are not generally regarded as close relatives of the Tropaeolaceae.

ECONOMIC USES
Several species of Tropaeolum are commonly cultivated in gardens for
ornamental purposes, the best-known example being the Nasturtium.

GENUS DESCRIBED
Tropaeolum.* Except where otherwise stated, the above description is
based on an examination of T. aduncum Smith and T. majus Linn, grown
at Kew.
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Farenholtz 671, Hill 969, Thompson 2254.

. RUTACEAE
(FiG.7i on p. 306; FIG. 72 on p. 306; FIG. 73 on p. 312; FIG. 77 on p. 334; FIG. 80 on p. 350)

SUMMARY
(i) GENERAL ANATOMY
A family consisting mainly of trees or shrubs and a very small proportion
of herbs all of which flourish in the tropics and sub-tropics. There is a con-
siderable range in the shape of the leaves in different sections of the family,
whilst thorns are common in some of the genera. The ground tissue of both
leaf and axis is nearly always characterized by the presence of secretory
cavities which, in the leaf, appear to the naked eye as transparent dots. The
secretory cavities are sometimes replaced or accompanied by secretory cells.
Resin cells are common in the pith, primary cortex, and rays of the young
4594 v
306 RUTACEAE
stem. The hairs are predominantly thick-walled and unicellular or some-
times uniseriate, but peltate, tufted, stellate, and multicellular glandular hairs
or warts also occur. The epidermis of the leaf is frequently composed of

FIG. 71. RUTACEAE

A-B, Glandular hair from the leaf of Toddalia aculeata Pers., A in section, B in surface-view.
C, Glandular shaggy hair from the floral region of Dictamnus albus Linn. D, Peltate hair of Phebalium
anceps DC. A-B by Solereder, C after Rauter, D after O- Bachmann.

FIG. 72
Transverse section through the leaf of Barosma serratifolia (Curt.) Willd. By Solereder.

cells ofwhich the inner walls are mucilaginous. Stomata are of various types.
The usually abundant crystals of calcium oxalate may be either solitary or
clustered, or a mixture of both of these types. Raphides and crystal-sand also
occur, although more infrequently. Dendritic crystals of diosmin have been
 RUTACEAE 307
recorded in a number of genera; berberin has been detected by Klein and
Bartosch (1245) in certain species of Evodia, Orixa, Toddalia. There is
usually a solitary, arc-shaped or cylindrical vascular strand in transverse
sections through the distal end of the petiole in most of the species which
have been examined. Anotable exception is Ruta graveolem Linn, with an
arc of separate bundles. Cork arises superficially in the stem of all of the
species in which its development has been studied, and in some species con-
sists of thin-walled cells, but in others the outer tangential walls are strongly
thickened. The pericycle of the stem usually contains well-defined strands
of fibres, although there is a tendency for the groups to coalesce in certain
species. The xylem and phloem constitute a closed cylinder in the stem in
nearly all species, although the primary medullary rays are moderately broad
in a few instances,

(ii) WOOD
Vessels small to medium-sized, often very small, typically in multiples and
sometimes with a distinct radial or oblique pattern, ring-porous or semi-ring-
porous in some species; perforations exclusively simple except for rare
multiperforate plates in a few species; intervascular pitting alternate, small to
minute, pits to rays cells similar members usually of medium length, some-
;

times moderately to very short. Parenchyma (a) terminal, (b) paratracheal,

usually vasicentric, but aliform and confluent in some genera, and (c) diffuse,
in some genera
only, and often consisting largely of chambered crystal cells.
Rays exclusively uniseriate or up to 2-4 cells wide; considerably larger in a
few genera; homogeneous or weakly heterogeneous and with few uniseriate
rays in most genera, but sometimes distinctly heterogeneous and with
numerous uniseriate rays; with distinct stories only in Chloroxylon, but in
echelon in a few other genera. Fibres with simple or slightly bordered pits,
with occasional septa in a few species; of medium length to moderately short.
Tracheids rare. Normal intercellular canals absent, but traumatic
vertical canals moderately common.

LEAF
Generally dorsiventral but sometimes centric. Hairs infrequent in many
members of the family, but both clothing and secretory types occur. Clothing
hairs most frequently thick- walled, unicellular; tufted types recorded in
Boronia, stellate forms in species of Asterolasia, Chorilaena, Correa, Crowea,
Diplolaena Eriostemon^ Flindersia (Fig. 80 B), Phebalium, Zieria, and scales in
y

Eriostemon, Flindersia, Nematolepis, Phebalium (Fig. 71 D). Adpressed hairs,

unicellular when young, but becoming septate and deciduous with increase
in age, recorded by Swingle (2225) in the drought resistant Eremocitrus glauca
Swingle (the Desert Kumquat) from Australia. External glands with clavate
or spherical multicellular heads recorded in certain species of Choisya,
Dictyoloma, Esenbeckia, Monnieria, Pilocarpus, Toddalia (Fig. 71 A-B), Zantho-
xylum, and Zieria. Sessile glands also occur on the lower surface of the leaf in
Boronia. Glandular shaggy hairs recorded in the floral region in Dictamnus
albus Linn. (Fig. 71 c). Cork warts sometimes present on the surface of old
Citrus leaves. Cells of the epidermis frequently have mucilaginous inner
walls in Acmadenia, Adenandra, Agathosma, Barosma, Coleonema, Diosma,
308 RUT ACE AE
Empleurum, Flindersta, Macrostylis, Phellodendron, Skimmia, Toddalia,
Zanthoxylum. Epidermis papillose in species of Boenninghausenia, Dictyoloma,
Eriostemon, Pilocarpus, and Spiranthera. Upper epidermis containing groups
of small cells with the palisade especially well developed below them in
Murraya exotica Linn. Stomata sometimes present on both surfaces
(Cneoridium dumosum Hook. f. and Rut a graveolens Linn.), but generally con-
fined to the lower side in most species; limited to a narrow median zone in
Acmadenia and Coleonema, and to two narrow strips on the lower surface
situated on either side of the midrib in Macrostylis; protected by scales in
Phebalium filifolium Turcz. Stomata provided with variously arranged sub-
sidiary cells in different genera and species.
Studies by Hirano (979) and Reed (1897) f t ^ie distribution of the stomata
on the leaves of a number of species of Citrus, collected and cultivated in one
place, showed the density to be related to the country of origin. Most tropical
species were found to have more than 500 stomata per sq. mm., but in the
hardier species and varieties from outside the tropics, with but few exceptions,
the density was lower. Stomatal density was also found to vary within the
single species Citrus sinensis Osbeck according to the stage of development
of the leaf, being greatest in very young leaves at the end of a period of
stomatal initiation. The size of the stomata was found to be only slightly
influenced by the number per unit area, but reduction of the light
intensity decreased their density. McLean (1468) found the greater resistance
of the Szinkum Mandarin (Citrus aurantium Linn. var. szinkum) to the canker
induced by infection with Pseudomonas citri Hasse compared with the lower
resistance of the Florida Grape-fruit (C. grandis Hassk.) to be associated with
differences in the structure of the openings to the outer chamber to the
stomata; those of the more resistant of these species being made narrower by
the development of larger ridges. The structure of Citrus stomata has also
been investigated by Turrell (2301). One or more layers of hypoderm,
sometimes locally distributed, occur in species of Gusparia, Eriostemon, Evodia,
Pagetia, Pilocarpus, Ruta. Mesophyll in the adpressed leaves of Agathosma
lediformis E. et Z. described by Solereder as having the palisade tissue
more strongly developed on the lower than on the upper side; isobilateral in
Eremocitrus glauca Swingle, the Australian Kumquat according to Swingle
(2225) and in Geijera parviflora Lindl. according to Wood (2457). Branched
sclerenchymatous cells are associated with the terminations of the veins in
several species of Boronia. Petiole, in transverse sections, usually exhibiting a
closed cylinder of xylem and phloem at the distal end, notably in species of
Calodendrum, Casimiroa, Choisya (slightly interrupted in C. ternata H. B. and
K. (Fig. 77 G)), Citrus, Correa, Dictamnus, Evodia, Murraya, Phellodendron,
Pilocarpus, Ptelea, Ravenia, Toddalia, Zanthoxylum ; with a closed cylindrical
bundle accompanied by medullary ones in Galipea; with a solitary arc-shaped
strand in species of Adenandra, Barosma (Fig. 77 E), Boronia, Orixa (phloem
present on part of the dorsal as well as on the ventral side of each strand),
Skimmia; with an arc of separate bundles in Ruta graveolens Linn. (Fig. 77 B).
Vascular strand of the petiole usually accompanied by a circle or groups of
pericyclic fibres except sometimes in species of Boronia, Calodendrum, Correa,
Orixa, Ptelea, Skimmia. Secretory cavities appearing as transparent dots
either in the uninjured leaf or on a cut surface, and variously interpreted as
 RUT ACE AE 309
schizogenous or lysigenous, almost always present in the mesophyll. Resin
cells have been recorded in Cneoridium. Tanniniferous idioblasts are stated
by Wood (2457) to occur in the mesophyll of Geijera parviflora LindL
Crystals very common, solitary ones being especially characteristic of certain
genera and clustered ones more frequent in others. Some of the palisade cells
transversely septate and containing clustered crystals in Pilocarpus spp.
Raphides and/or styloids recorded in Erythrochiton, Monnieria, Raputia, and
Rauia; crystal-sand in Cusparia and Orixa. Hesperidin (diosmin), usually
in the form of dendritic crystals, soluble in and giving a yellow colour to
solutions of caustic potash, recorded in the epidermis of certain species of
Agathosma, Barosma (Fig. 72), Calodendrum, Dictamnus, Empleurum, Pilo-
carpus, Ptelea, Skimmia, Toddalia, Zanthoxylum.

Axis
YOUNG STEM (Fig. 77 H)
Hairs of the following kinds have been recorded or observed, (i) Uni-
cellular, thick-walled in species of Agathosma, Calodendrum, Choisya', Dic-
tamnus, Evodia, Melicope, Murraya, Orixa, Ptelea, Ravenia, Skimmia,
Zanthoxylum. (ii) Uniseriate in species of Casimiroa and Evodia. (iii) Stellate
in species of Calodendrum and Correa spp. (iv) Glandular, club-shaped in
Agathosma and Evodia. Cork arising in the sub-epidermis in those species
so far investigated, notably in species of Agathosma, Barosma, Calodendrum,
Casimiroa, Citrus, Evodia, Melicope, Murraya, Orixa, Phellodendron, Ptelea,
Ruta, Skimmia, Zanthoxylum. Cork cells thin-walled in species of Casimiroa,
Cusparia, Galipea, Murraya, Ravenia, Ruta. Outer tangential walls of cork
thickened in Agathosma, Barosma, Orixa, Ptelea, Skimmia, and
cells strongly

Zanthoxylum. Primary cortex containing stone cells in certain species of

Almeidea, Casimiroa, Cusparia, Metrodorea, Ptelea, and Ravenia. Cortical
bundles sometimes present in Eriostemon and Pilocarpus. Pericycle usually
containing isolated strands or bands of sclerenchyma, notably in certain
species of Adenandra, Agathosma (groups small), Barosma (small and incon-
spicuous), Boronia, Casimiroa (very small), Citrus, Correa (not very widely
separated), Dictamnus, Eriostemon (very poorly developed), Evodia, Melicope,
Murraya (in older stems), Phellodendron, Pilocarpus (associated with large
stone cells), Poncirus, Ptelea (not well developed), Ruta (small), Toddalia,
Zanthoxylum. Pericyclic sclerenchyma less frequently tending to consti-
tute a continuous ring, but this character is sometimes to be seen in
Amyris, Calodendrum, Choisya, Correa, Evodia, Murraya, Orixa, Paramignya,
Zanthoxylum. Pericyclic fibres apparently absent from certain species of
Boenninghausenia, Coleonema, Crowea, Euchaetia, Ptelea, Ravenia, and
Skimmia. Phloem and xylem constituting closed cylinders traversed by
narrow rays in all of the species examined except in Choisya, Eriostemon, and
Pilocarpus, where relatively broad primary rays sometimes occur, especially
in very young stems. Vessels arranged in well-defined radial rows in certain
,

species of Adenandra, Agathosma, Barosma; Boronia (thin-walled), Caloden-

drum, Dictamnus, Melicope, Murraya, Ravenia, but with the radial arrange-
ment less pronounced in species of Citrus, Crowea, Eriostemon, Orixa, Ruta,
Zanthoxylum. Vessels not in well-defined radial rows in some species of
Casimiroa, Choisya, Citrus, Evodia, Phellodendron, Ptelea, Skimmia (flame-
3io RUTACEAE
shaped arrangement of small vessels), Toddalia (mostly solitary)* Vessels
usually with simple perforations. Pith homogeneous in species of Agathosma,
Barosma, Boronia, Calodendrum, Casimiroa, Choisya, Correa, Crowea, Die-
tamnus, Eriostemon, Orixa, Phellodendron, Ptelea, Ruta, Skimmia, Zanthoxy-
lum heterogeneous in species of Citrus, Evodia, Melicope, Murraya, Pilocarpus,
;

Toddalia. Medullary cork recorded by Holm (999) in the hypocotyl of

seedlings of Balsamocitrus dawei Stapf, but not in other closely related genera.
Large conspicuous secretory cavities, usually with oily or resinous con-
tents, very common in the parenchymatous tissues, especially the cortex,
notably in species of Adenandra, Agathosma, Barosma, Calodendrum,
Casimiroa, Choisya, Citrus, Dictamnus (relatively infrequent), Eriostemon
(specially large), Melicope, Murraya, Orixa, Phellodendron (sometimes in-
frequent), Pilocarpus, Poncirus, Ptelea (sometimes specially large), Ravenia,
Ruta, Skimmia, Toddalia, Zanthoxylum (sometimes infrequent). Secretory
cavities sometimes partly or wholly replaced by secretory cells in certain

species of Boronia, Citrus, Correa, Crowea, Melicope, Phellodendron, Ravenia

(contents deeply stained with safranin), Toddalia, Zanthoxylum (situated in
the inner part of the cortex or in the phloem). Intercellular secretory spaces
present in Evodia and secretory canals in Clausena sp. Crystals commonly
abundant in the cortex and/or pith and sometimes in the phloem as well,
e.g. in species of Adenandra, Agathosma, Barosma, Calodendrum, Casimiroa,
Choisya, Citrus, Eriostemon, Murraya, Pilocarpus, Poncirus, Ptelea, Ravenia,
Ruta, Skimmia, Toddalia, Zanthoxylum. Crystals predominantly solitary in
species of Casimiroa, Citrus, Murraya, Pilocarpus, Poncirus, Ravenia (very
large), Toddalia', usually clustered in certain species of Adenandra, Agathosma,
Barosma, Calodendrum, Choisya, Eriostemon, Pilocarpus, Ptelea, Ruta, Skim-
mia. Raphides and/or styloids present in certain species of Erythrochiton,
Galipea, Monnieria, Raputia, Ravenia, and Rauia, and sacs of crystal-sand in
Cusparia sp. and Orixa sp. Dendritic crystals of diosmin recorded in Barosma,
Eriostemon, Skimmia, Toddalia.

WOOD (Fig. 73)

Vessels mostly very to moderately small (25-100 [i mean tangential
diameter), sometimes extremely small (less than 25 \L), e.g. in Choisya and
Skimmia, medium-sized in about one-third of the genera and never more than
200 in mean diameter (excluding ring-porous species); never exclusively
fj,

solitary, radial multiples of 2-4, or more, common, the pores retaining a

characteristic rounded shape; clusters moderately common in Aeglopsis,
Balsamocitrus, Bosistoa, Erythrochiton, Feronia, Glycosmis, Halfordia, Limonia,
Nycticalanthus, Oricia, Phebalium, Ptelea, Skimmia, and Zieria\ multiples and
clusters sometimes forming a radial or oblique pattern, most marked in
Acradenia, Acronychia p.p., Amyris(Fig. 73 L), Balfourodendron, Calodendrum,
Choisya, Cneoridium (938), Evodia p.p., Orixa (938), Phebalium, Ptelea,
Raputia, Skimmia, and Vepris with tangential or ulmiform pattern towards
;

the outer part of the ring in Calodendrum (Fig. 73 K), Phellodendron, and
Vepris varying in number from 4 to 100 per sq. mm. but mostly between
;

8 and 40; ring-porous or semi-ring-porous in species of Acradenia, Bal-

fourodendron, Calodendrum, Choisya, Cneoridium, Erythrochiton (1889), Evodia,
Phebalium, Phellodendron, Ptelea, Thamno$ma> and Zanthoxylurn. Spiral
 RUTACEAE 311

thickening occurs in Acradenia, Asterolasia (2158), Calodendrum, Choisya,

Cneoridium, Eriostemon (2158), Evodia p.p., Orixa (938), Phebalium, Phello-
dendron, Philotheca (2158), Poncirus, Ptelea, Thamnosma, and Zieria (2158).
Perforations typically exclusively simple, but a few foraminate plates observed
in Acradenia and rare reticulate plates in 'Tetractomia'; Solereder records
occasional scalariform plates in some species of Acronychia, Adenandra,
Agathosma, Barosma, Boenninghausenia, Calodendrum, Clausena, Leptothyrsa,
and Paramignya\ modified scalariform plates are also reported (938, 1858) in
Adiscanthus and Platydesma. Intervascular pitting alternate, mostly small to
minute, moderately large in Aegle, Aeglopsis, Amyris, Balsamocitrus, Evodia
p.p., Fagara p.p., Murraya, Phebalium, Phellodendron, Ptelea, Skimmia,
l

Tetractomia\ and Zanthoxylum p.p.; Heimsch (938) notes some scalariform

and transitional pitting in Platydesma and Ravenia rosa Standl. coalescent
;

apertures common in many genera; pits to ray cells and parenchyma similar
to the intervascular pitting, occasionally unilaterally compound in Balfouro-
dendron, Citrus p.p., Evodia p.p. and Poncirus and simple in Orixa and
,

Skimmia (2158); Heimsch notes oblong pits in Erythrochiton, Orixa, and

Ravenia\ solid deposits common and sometimes abundant. Silica reported
in species of Evodia (794). Mean member length 0-2-0-6 mm.
Parenchyma absent from Nethadenia, present in the other genera as
(a) terminal bands, varying from i to 6 cells wide, typical of most species, but
not observed in Acronychia p.p., Araliopsis, Euxylophora, Hortia, Plethadenia,
*
and Tetractomid (b) paratracheal, predominant in all species, most commonly
;

scanty (Fig. 73 j and L), but aliform to confluent in Acronychia, Afraegle,

Bosistoa, Casimiroa, Citrus, Clausena excavata Durand (1154), Erythrochiton,
Evodia, Fagara p.p., Feronia, Flindenia p.p., Melicope, Nycticalanthus,
l

Oricia, Ravenia, Spiranthera, Tetractomia* and Zieria, and connecting the

,

vessels in woods with a pronounced vessel pattern, as in Acradenia and

Skimmia (c) diffuse, as single strands scattered among the fibres, often con-
;

sisting almost entirely of chambered crystalliferous cells, present in Aeglopsis,

Amyris, Araliopsis, Balfourodendron, Balsamocitrus, Bosistoa, Calodendrum^
Chloroxylon (Fig. 73 H), Citrus, Clausena, Evodia p.p., Feronia, Halfordia,
Heliettia, Limonia, Melicope, Merrillia, Murraya, Nycticalanthus, Oricia,
Platydesma, Poncirus, Raputia, and Vepris. Crystals in chambered cells some-
times abundant, observed in the following: Aeglopsis, Afraegle, Amyris,
Araliopsis, Atalantia, Balfourodendron, Balsamocitrus, Bosistoa, Casimiroa,
Chloroxylon, Citropsis, Citrus, Clausena, Esenbeckia, Evodia p.p., Fagara,
Flindersia, Halfordia, Helietta, Limonia, Merrillia, Murraya, Nycticalanthus ,
Oricia, Poncirus, Spiranthera, Teclea, Vepris, and Zanthoxylum p.p. in idio-
;

blasts in Citrus p.p. and Poncirus. Record and Hess (1889) have reported
large bundles of raphides in the diffuse parenchyma of Raputia magniftca
Engl. Strands typically of 2-4 cells; fusiform cells common in Plethadenia
(938). Storied in Chloroxylon and with a tendency to storied arrangement
(echelon) in Acronychia, Balfourodendron, and Feronia. Heimsch (938)
reports sclerotic cells in Flindersia ifflaiana F. v. Muell. Rays exclusively
uniseriate in Amyris, Choisya (occasionally biseriate), Cneoridium, Diosma,
Merrillia, Spiranthera, and Thamnosma\ mostly up to 2 or 3 cells wide; up to
6 or more cells wide in some species of Calodendrum, Clausena, Esenbeckia
(938), Evodia, Feronia, Flindersia, Geijera (938), Platydesma (938), and
 FIG. 73. RUTACEAE
A, Calodendrum capenris Thunb. B, Halfordia scleroxyla F. v. M. C, Chhroxyhn stvietenia DC.
D, Amyris balsamifera Linn. E, Citropns articulata Sw. et Kellerm. F, Plethadenia cubensis Urb.
G, Euxylophora paraensis Huber. H, Chloroxylon swietenia DC. I, Flindersia brayleyana F. v. M.
J, Tedea nmplicifolia (Engl.) Verdoorn. K, Calodendrum capensis Thunb. L, Amyris balsamifera
Linn.
M, Fagara angolensis Engl.
 RUTACEAE 313
Zanthoxylum (938); less than i mm. high; uniseriate rays usually scarce or
absent from woods with homogeneous or weakly heterogeneous multiseriate
rays; between 4 and 14, mostly 7-10, rays per mm.; homogeneous (Kribs's
Types I and II) in Aegle, Aeglopsis, Afraegle, Araliopsis, Balsamoritrus,
Bauerella (938), Calodendrum, Casimiroa, Citrus, Cneoridium (938), Diosma,
Euxylophora, Fagara p.p., Feronia, Flindersia, Halfordia, Helietta, Hortia,
Limonia, Merrillia, Murraya, Oricia, Phebalium, Phellodendron, Plethadenia,
Raputia, Sohnreyia, Spiranthera, Teclea, Vepris, and Zieria, and only weakly
heterogeneous in several other genera; typically homogeneous (Kribs's
Type III) in the woods with exclusively uniseriate rays, distinctly hetero-
geneous and sometimes with 4 or more rows of uniseriate marginal cells (not
all square or upright cells) in Acronychia
necessarily p.p., Atalantia, Choisya
(1889), Citropsis, Erythrochiton (1889), Esenbeckia, Evodia p.p., Glycosmis p.p.,
Melicope, Metrodorea
'
(1889), Micromelum, Nycticalanthus, Ravenia (1889),
Skimmia, Tetractomia' and Wenzelia (938); the most unspecialized rays
,

(Kribs's Het. II A), according to Heimsch, occur in the Rutoideae, e.g. in

some species of Erythrochiton, Evodia, Lunasia, and Ravenia, and in Rhabdo-
dendron', with a slight tendency to sheath cells in some species; chambered
crystalliferous cells present in Balfourodendron,
Bahamocitrus, Clausena,
Chloroxylon, Esenbeckia, Evodia p.p., Feronia, Limonia, and
Glycosmis,
Micromelum silica present in species of Achronychia and Evodia (794). Thin-
;

walled oil cells present in Euxylophora (Fig. 73 G). Storied in Chloroxylon

(about 4 per mm.) and in echelon in Fagara p.p., Phellodendron, Vepris, and
Zanthoxylum p.p. Fibres with simple or slightly bordered pits, rare on the
tangential walls, few to numerous on the radial walls; Heimsch (938) describes
the fibrous elements in Rhabdodendron as 'tracheids*; commonly containing
gum-like deposits, with numerous small crystals in Feronia lucida Scheff.;
often septate in Erythrochiton, Phellodendron wilsonii Hay. et Kanehira (1206)
and Raputia, and with occasional septa in some species of Araliopsis and
Evodia. Numerous gum plates, some of which are not easily distinguishable
from septa, occur in Flindersia. Walls moderately thick to very thick, some-
times mucilaginous. Mean length 0-6-1-3 mm. Tracheids occur mixed with
extremely narrow, often imperfect vessel members in Acradenia, Choisya, and
Skimmia. Intervascular canals of the traumatic vertical type observed or
reported (1889) in Afraegle, Atalantia, Balfourodendron, Citropsis, Citrus,
Esenbeckia, Euxylophora, Feronia, Fortunella, Helietta, Hesperethusa, Metro-
dorea, Micromelum, Pilocarpus, Ravenia, and Zanthoxylum. Webber (2377)
describes radial intercellular cavities in the rays of Citrus trees affected by the
physiological disease described as xyloporosis. Included (interxylary)
phloem reported (938, 1530, 1842) in Rhabdodendron amazonicum (Benth.)
Hub. ; of the 'concentric* type (c. I. circumvallatum), with successive develop-
ment of secondary groups of wood and bast.

ANOMALOUS STRUCTURE, see Preceding paragraph

ROOT
Large idioblasts with thin, yellow, unlignified cell walls and containing
resinous material recorded by Brandt (259) in the primary cortex of young
roots and in the secondary cortex of older roots of Ruta graveolens Linn,
3 i4 RVTACEAE
TAXONOMIC NOTES
The wood anatomy of this family is on the whole very uniform and repre-
sents a moderately high level of specialization. Heimsch (938) considers that
there is strong evidence for considering the Rutaceae, Simarubaceae,
Meliaceae, Sapindaceae, Burseraceae, and Anacardiaceae as forming a natural
group, with the Rutaceae standing rather apart, owing to its general lack of
septate fibres, and showing the closest resemblance to the Simarubaceae.
On the problem of whether Flindenia should be placed here or in the
Meliaceae, Dadswell (526), Harrar (906), and Record (footnote in Welch
2406) all agree that it is out of place in the Meliaceae owing to its homogeneous
rays and non-septate fibres. The genus, however, is variable and, though
most species show a greater resemblance to the Rutaceae than to the Meliaceae,
there are occasional species, e.g. F. brayleana F. v. M. that, except for the
presence of gum plates instead of septa in the fibres, are very similar to some
species of the Meliaceae, e.g. of Khaya. Record suggests that these differences
between the species may be of generic rank. Dadswell considers that the
genus is not absolutely typical of either the Rutaceae or the Meliaceae and
both he and Harrar support the suggestion of a separate family, the Flinder-
siaceae. The occurrence of secretory cavities and cells in other tissues besides
the secondary xylem favours the inclusion of Flindersia in the Rutaceae.
The wood anatomy of Rhabdodendron, with its bordered pits in the fibres
and anomalous structure, appears to be out of place in this family. Record
(1842) found little in common between the wood of this genus and that of the
Rosaceae, but found a marked affinity with certain genera of the Phyto-
laccaceae, and Record and Hess (1886) include it in this family.
Hess (950) has drawn attention to a marked dissimilarity between the
woods of Ravenia rosea Standl. and R. spectabilis (Lindl.) Planch.

ECONOMIC USES
The timbers are commonly dense and yellow-coloured and several of them,
e.g. Esenbeckia atata Pittier, have been used locally or suggested as boxwood
substitutes. The Satinwoods are of general commercial importance, the East
Indian Satinwood being furnished by Chloroxylon swietenia DC. and the
West Indian by Zanthoxylum flavum Vahl. The timbers of some species of
Flindersia are among the most important hardwoods of Australia; the wood
of F. brayleana F. v. M., for example, is used for cabinet work, veneers,
aeroplane construction, and rifle stocks, and that of E. ifflaiana F. v. M. is
reputed to be one of the most important structural hardwoods of north
Queensland (525). Pau Marfim, Balfourodendron riedelianum Engl., is
reported (1084) to be used in Brazil for laminated airscrews.
Apart from the timbers the most important economic products are the
Citrus fruits, such as oranges, limes, lemons, grape-fruits, &c. For particulars
of the density of stomata and oil glands in the peel of the Washington Naval
Orange see the article by Turrell and Klotz (2302); and for the structure of
the peel in relation to water spot the account by Scott and Baker (2077). Oil
of Rue is distilled from the leaves of Ruta graveolens Linn. Various products
of medicinal value are also obtained, including Cusparia Bark (Galipea offi-
cinalis Hancock); Buchu leaves (Barosma betulina Bart, and Wendl. and
other species of Barosma) Jaborandi leaves (Pilocarpus microphyllus Stapf and
;
 RUTACEAE 315
other species of Pilocarpus). Prickly Ash Bark or Toothache Bark is obtained
from Zanthoxylum americanum Mill, (northern type) and Z. clava-herculis
Linn, (southern type).

CUSPARIA BARK
The diagnostic anatomical characters are the cork with the outer tangential
:

walls of the cells strongly thickened; the fairly broad phelloderm consisting
of thin- walled cells; the primary cortex containing infrequent groups of
strongly thickened yellow fibres and stone cells; the secondary cortex con-
sisting mostly of thin-walled tissue arranged in alternating layers of paren-
chyma and disorganized sieve elements and traversed by rays 1-3 cells wide;
the secretory cells, raphides, other crystals, and starch. Copalchi Bark
(Croton niveus Jacq.) (see Euphorbiaceae) and Angusturo Bark (Esenbeckia
febrifuga A. Juss.) are sometimes used as substitutes. Angusturo bark may
be recognized by the 3 or 4 layers of colourless cells in the inner part of the
cork, which are capable of being stained bright blue by acids with oxidizing
properties, owing to the presence of the alkaloid evodin.

BUCHU LEAVES
The diagnostic anatomical characters are: the epidermis on both surfaces
composed of cells with straight anticlinal walls; abundant mucilage in the
epidermis and hypoderm dendritic crystals of diosmin, which occur especially
;

in the epidermis the large oil glands and cluster crystals of calcium oxalate in
;

the mesophyll the single layer of palisade tissue the ranunculaceous stomata,
; ;

confined to the lower surface.

The leaves of J3. betulina Bart, et Wendl. can frequently be distinguished from
those of other species of Barosma which are used as substitutes by a proportion
of the stomata being more than 38 /x in diameter. Levin (i 366) has shown how
differences in the vein-islet numbers may also be used to distinguish B. betulina
and B. crenulata (L.) Hook, from other species of Barosma (B. betulina 10-15
(average 127) and J5. crenulata 10-16-5 (average 13) per sq. mm., but varying
somewhat in the last of these species). The anatomy of an unidentified
adulterant of Buchu has been described by Small (2133).

JABORANDI LEAVES
*
The diagnostic anatomical characters are: the epidermis on both surfaces
composed of mostly pentagonal and hexagonal with practically straight
cells
anticlinal walls the stomata confined to the lower surface, each surrounded
;

by 4 or 5 concentric subsidiary cells; the single layer of palisade tissue; the

broader region of spongy tissue; the large secretory cavities, the scattered
secretory cells with reddish-brown contents, and the cluster crystals of cal-
cium oxalate, all of which occur in the mesophyll; the circular vascular strand
of the midrib surrounded by thick-walled fibres.

PRICKLY ASH BARK

The diagnostic anatomical features of the bark of Zanthoxylum americanum
Mill, are: the narrow, easily detached cork consisting of about 4 layers of brownish
thick- walled, tabular cells, stained golden orange when treated with iodine ;

the cortex composed of tangentially elongated, parenchymatous cells, in many

of which there are solitary, prismatic crystals; the broader phloem, traversed
316 RUT ACE AE
by narrow medullary rays; the thick- walled secretory cells with oily contents,
which are rounded or oval in transverse sections but somewhat elongated
longitudinally and occur in the cortex and, more abundantly, in the phloem;
the amorphous and oily material deposited chiefly in most of the ray cells and
some of the cortical cells; the sporadic small bundles of fibres and chains or
clusters of stone cells, both stained golden yellow by iodine, which occur in
the cortex; the abundant crystals attached to the inner surface of the bark.
It has been claimed by Wirth (2447) that the bark of Z. clava-herculis Linn,
can often be distinguished from that of Z. americanum by the absence of
crystals from the inner surface of the bark of the first of these species.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acmadenia, Adenandra,* Agathosma,* Almeidea, Amyris, Asterolasia,
Balsamocitrus, Barosma,* Boenninghausenia, Boronia,* Calodendrum,*
Casimiroa,* Choisya,* Chorilaena, Citrus,* Clausena, Cneoridium, Coleo-
nema,* Correa,* Crowea,* Cusparia, Dictamnus,* Dictyoloma, Diosma,
Diplolaena, Empleurum, Eremocitrus, Eriostemon,* Erythrochiton, Esen-
beckia, Euchaetis, Evodia,* Flindersia, Galipea, Macrostylis, Melicope,*
Metrodorea, Monnieria, Murraya,* Nematolepis, Orixa,* Pagetia, Para-
mignya, Peganum,* Phebalium, Phellodendron,* Pilocarpus,* Poncirus,*
Ptelea,*Raputia, Ravenia,* Ruta,* Skimmia,* Spiranthera, Toddalia,*
Zanthoxylum,* Zieria.
* Kew
Represented in the slide collection,

(ii)
FOR WOOD STRUCTURE
Acradenia, Acronychia, (Adiscanthus), Aegle, Aeglopsis, Afraegle, Araliop-
sis, Atalantia, Balsamocitrus, (Bauerella), Bosistoa, Calodendrum, Casimiroa,

Choisya, Citropsis, Citrus, Clausena, (Clausenopsis), Cneoridium, (Dictyo-

loma), Diosma, Erythrochiton, Esenbeckia, Euxylophora, Evodia, Fagara,
Feronia, Flindersia, (Fortunella), (Geijera), Glycosmis, Halfordia, Helietta,
(Hesperethusa), Hortia, Limonia, (Medicosma), Melicope, Merrillia, Metro-
dorea, (Microcitrus), Micromelum, Murraya, Nycticalanthus, Oricia, (Orixa),
(Paramignya), (Pelea), Phebalium, Phellodendron, (Pilocarpus), Platydesma,
Pleiococca, Plethadenia, Poncirus, Ptelea, Raputia, Ravenia, (Rhabdoden-
dron), Skimmia, Sohnreyia, Spiranthera, (Swinglea), Teclea, 'Tetractomia',
Thamnosma, Vepris, Zanthoxylum, Zieria.
LITERATURE
(i) On
General Anatomy
Brandt 259, Butler 324, Grossman 511, Elias 625, Engler 641, Hirano 979, Holm 999,
Klein and Bartosch 1245, Levin 1366, Livingstone 2526, McLean 1468, Reed 1897, Scon
2532, 2533, Scott and Baker 2077, Shelton 2086, Small 2133, Swingle 2225, Turrell
2301, Turrell and Klotz 2302, Wirth 2447, Wood, J. G. 2457.
(ii) On Wood Structure
Baker 104, den Berger 179, 182, Besson 186, Brit. Hond. F. D. 274, Brown, F. B. K.
382, Burgerstein 310, 312, Cockrell 435, Cooper 461, Coster 481, Dadswell 524, 525,
536, Foxworthy 705, Giordano 786, Gonggrijp 794, Greguss 2522, Harrar 906, Heimsch
938, Hess 950, Horn 1084, Howard 1088, Janssonius 1154, Jolly 1188, Kanehira 1206,
1209, Lecomte 1334, Metcalfe 1497, Milanez 1530, Pearson and Br. 1679, Pereira 1687,
Pfeiffer J. Ph. 1713, Record 1780, 1781, 1787, 1801, 1809, 1842, 1843, 1851, 1884,
Record and Hess 1886, 1889, Record and Mell 1894, Scott 2075, Stone 2202, Sudworth
22x8, Tang 2231, Webber 2377, Welch 2406, 2408, Williams 2430, Yamabayashi 2478.
 (3*7)

^80. SIMARUBACEAE
(FiG. 74 on p. 318; FIG. 75 on p. 322; FIG. 77 on p. 334)

SUMMARY
(i)
GENERAL
Trees or shrubs which occur chiefly in tropical countries. The hairs are
mostly simple, unicellular or uniseriate. Glandular hairs, sunken glands, and
extra-floral nectaries also occur in certain genera and species. The leaf is

usually dorsiventral, rarely isobilateral or sub-centric. The epidermis is

frequently mucilaginous and sometimes papillose. Stomata, mostly con-

fined to the lower surface, are usually ranunculaceous, except in a few genera
in which they are rubiaceous. Variously shaped, sclerenchymatous idioblasts
are common in the mesophyll. The petiole, in transverse sections through
the distal end, usually exhibits a cylindrical vascular strand which, in certain
genera, encloses medullary bundles, whose orientation and distribution have
been used in the identification of species. In the young stem the cork is
usually superficial in origin the pericycle is bounded by isolated strands of
;

fibres or acomposite and continuous ring of sclerenchyma the xylem and ;

phloem form continuous cylinders. Secretory canals occur in certain

genera especially in the peripheral region of the pith, whence they extend via
the petiole to the major leaf veins. Secretory cells containing oil, resin, or
mucilage, although less common, are to be found in the parenchymatous
tissues.

(ii) WOOD
This family includes a wide variety of anatomical features, but it has been
shown by Webber (2374) that this is largely due to dissimilarity between
some of the tribes.
Vessels evenly distributed in most genera, but with a radial or oblique
pattern in some a few species definitely or slightly ring-porous perforations
; ;

exclusively simple, with a few minor exceptions; intervascular pitting alter-

nate, varying from very small to large; pits to ray cells and parenchyma
similar to the intervascular pitting in most species but large and tending to be
simple in some; members of medium length to moderately short. Paren-
chyma extremely variable, ranging from extremely sparse or absent to
abundant, in broad bands; usually with at least some parenchyma associated
with the vessels diffuse parenchyma or fine metatracheal bands, predominant
;

in a few species and diffuse parenchyma sometimes present in woods with

predominantly paratracheal parenchyma; terminal parenchyma sometimes

present. Storied in several species. Rays multiseriate except in i genus,
usually between 2 and 4 cells wide, larger in Ailanthus\ commonly homo-
geneous, occasionally distinctly heterogeneous; often storied. Fibres with
small simple or bordered pits sometimes septate of medium length, Vasi-
; ;

centric tracheids present in a few species. Vertical intercellular canals,

probably traumatic, often present.
LEAF
Usually dorsiventral, but sometimes isobilateral or centric (see 'Mesophyll').
Hairs unicellular or uniseriate, mostly sclerenchymatous. Stalked glands
3*8 SIMARUBACEAE
(Fig. 74A-B) occur in Ailanthus, Alvaradoa, Brucea, Harrisonia, Hebonga,
Picramnia, Rigiostachys, Simaba, Simaruba, Soulamea, Suriana, but are
generally infrequent except in Suriana. Shaggy glands recorded in Eurycoma
(Fig. 74 c), and large, conspicuous, wart-like glands on the teeth at the base
of the leaflets of Ailanthus altissima Swingle (syn. A, glandulosa Desf,).
Sugar is stated by Petaj (1700) to be secreted from the so-called nectar
tissue of the glands of Ailanthus altissima directly through the epidermis, no
special secretory channels being present. Exudation, which does not appear
to be correlated with insect visits, begins as soon as the bud unfolds; is most
active in the early morning and slows down during the course of the day.
More recently the glands of Ailanthus have been studied by Davis (551).

FIG. 74. SIMARUBACEAE

Hairs of A, Picramnia sp. B, Rigiostachys bracteata Planch. C, Eurycoma longifolia Jack. By Solereder.

Extra-floral nectaries, in the form of sunken glands, also occur on various

parts of the leaf in numerous genera, (i) On the lower surface of the lamina
in Alvaradoa, Brucea, Eurycoma, Harrisonia, Picramia. (ii) On the upper
surface of the lamina in Cadellia, Hannoa, Odyendea, Simaba, Simaruba.
(iii) Confined to the apex and base of the lamina in certain species of Hannoa,

Mannia, Simaba, Simaruba, and Simarubopsis. (iv) On the petiole in Cadellia.

Extra-floral nectaries stated to be absent from the Irvingeae and from Castela,
Hebonga, Kirkia, Picrasma, Quassia, and Rigiostachys. Epidermis single-
layered in most genera, but cells sometimes becoming septate in Amaroria,
Castela, Hannoa, Soulamea. True hypoderm recorded by Solereder and by
Spiekerkoetter (2168) only in certain species of Ailanthus and Odyendea.
Lower epidermis papillose in certain species of Ailanthus, Alvaradoa,
Desbordesia, Eurycoma, Irvingella, Irvingia, Klainedoxa, Odyendea, Simaruba.
Epidermal cells mucilaginous in species of Ailanthus, Alvaradoa, Castela,
Hannoa, Harrisonia, Hebonga, Kirkia, Mannia, Odyendea, Picrasma, Simaruba,
Simarubopsis, as well as in all of the Irvingeae and Soulameae less frequently ;

mucilaginous in Brucea, Cadellia, Eurycoma, Guilfoylia, Picramnia, Picrasma,

Picrolemma, Quassia, Rigiostachys, Samadera, Simaba. Stomata very numer-
ous and almost exclusively confined to the lower surface, but present on the
upper surface as well in Suriana maritima Linn, and sporadically, especially
near the veins, in a few species of Ailanthus, nearly always at the same level
as the surrounding epidermal cells; ranunculaceous, usually surrounded by
4-10 ordinary epidermal cells, but rubiaceous types recorded in Castela,
Irvingia, Klainedoxa, and Picrodendron, Three to five epidermal cells surround
the stomata in Suriana, and sometimes exhibit a well-marked cruciferous
 SIMARUBACEAE 319

arrangement when each stoma is surrounded by 3 Stomata in groups in Castela

.

and Soulamea, and tending to be in parallel groups of about 5 in Irvingia and less

frequently in Klainedoxa. Peculiar processes are stated to be present at either end

of each stoma throughout most of the genus Picramnia. Mesophyll typically
dorsiventral, but somewhat variable centric in certain species of Ailanthus,
;

Castela, and Suriana\ isobilateral or sub-centric in Harrisonia abyssinica Oliv.

according to Spiekerkoetter (2168) and in Kirkia\ usually homogeneous and
palisade-like in the Irvingeae as well as in Cadellia and Picrodendron. Palisade
tissue consisting of 1-3 layers; cells described by Spiekerkoetter (2168) as
transversely septate in Amaroria, Harrisonia sp., Soulamea, and in certain of
the Irvingeae. Spongy parenchyma usually fairly lacunar; consisting of
hypha-like cells in Eurycoma, Simaba, Simaruba, Simarubopsis. Sclerenchy-
matous idioblasts, which exhibit a wide range of form and variations in the
thickness of the cell wall, occur in the mesophyll throughout Eurycoma,
Hannoa, Hyptiandra, Mannia, Qdyendea, Perrieria, Quassia, Simaba, Sima-
ruba, Simarubopsis. Those which are to be found in certain species of
Irvingella, Irvingia, Picramnia, Picrasma, and Samadera extend from the
fibres around the bundles and end blindly in the mesophyll. Small, in-
frequent idioblasts sometimes occur in other genera besides those listed above;
none recorded for Harrisonia and Klainedoxa. Veins very prominent in
Soulamea owing to the development of collenchyma on both sides of the leaf,
and in Castela owing to the presence of palisade tissue in the corresponding
position. Elsewhere, veins generally embedded in the mesophyll; provided
with well-developed parenchymatous sheaths in Cadellia, Kirkia, Picrasma,
Simaba, Simaruba, Simarubopsis, Soulamea (cells containing brown tannic
material), Suriana. Small veins vertically transcurrent in Irvingia, Klainedoxa,
and Picrodendron. Petiole, in transverse sections through the distal end,

exhibiting a closed, circular vascular strand surrounding a number of

variously orientated medullary bundles in many species. Medullary bundles
recorded in the petiole of Ailanthus (Fig. 77 L), Amaroria, Hebonga, Picrasma,
Picrolemma, Quassia, Samadera, Simaba, Simaruba, Soulamea, and possibly
other genera as well absent from at least certain species ofAeschrion (Fig. 77 1),
;

Alvaradoa, Brucea, Castela, Harrisonia, Irvingia, Klainedoxa, Picramnia,

Picrodendron. Orientation and number of medullary bundles stated by
Solereder to be of specific diagnostic value. Petiole containing a single arc-
shaped bundle in Suriana, and a closed or dissected annular bundle in
Rigiostachys. Secretory canals frequently accompany the vascular bundles
of the veins, especially in those genera where they occur in the axis also (see
'Young Stem'). Oil cells recorded by Engler (642) in the parenchymatous
portions of the leaf in Picrella trifoliata H. Baill. and in all species of Ailanthus,
Harrisonia, Hebonga, Simaba, Simaruba, and mucilage cells and cavities
throughout Desbordesia, Irvingia, and Klainedoxa. Crystals solitary or
clustered. Cluster crystals the commonest type, very variable in size, parti-
cularly large in Castela, Holacantha, Picramnia their size and distribution are
;

said to be of value in the identification of genera (for details see Boas's (208)
article). Solitary crystals especially frequent in the Irvingeae. Sphaerites
recorded in Picramnia and Suriana. Styloids confined to Alvaradoa. A sub-
stance resembling hesperidin recorded in certain species of Irvingella. Ash
containing 0-698 per cent, of copper in Odyendea gabunensis (Pierre) Engl.
3*o SIMARUBACEAE

Axis
YOUNG STEM 77 M)
(Fig.
Stomata deeply sunken Holacantha emoryi A. Gray, Cork
in the leafless
usually arising in the sub-epidermis except in the Irvingeae and in Amaroria,
Rigiostachys, Soulamea, and Suriana. Cork cells with U-shaped thickenings
recorded in certain species of Irvingella, Irvingia, and Klainedoxa. Cortex
sometimes containing stone cells, notably in species of Castela, Quassia,
Samadera, Simaruba. Inner part of the cortex strongly collenchymatous in
Aeschrion exceka (Sw.) Kunze (syn. Picraena excelsa Lindl.). Pericycle
sometimes containing isolated strands of fibres (e.g. with a broad but inter-
rupted ring in Aeschrion excelsa and Picrasma quassioides (Ham.) Benn.), but
with a composite ring of sclerenchyma in certain species of Ailanthus,
Alvaradoa, Cadellia, Castela, Picramnia, Quassia, slightly interrupted in Q.
amara Linn., Rigiostachys (ring becoming broken into groups with increase
in age), Samadera. Pericycle sometimes devoid of sclerenchyma in Hola-
cantha, but sub-epidermal groups of fibres separated by palisade-like
assimilatory tissue occur in this genus. Xylem and phloem constituting
closed cylinders, but primary rays fairly conspicuous, e.g. in Ailanthus
(Fig. 77 M). Vessels usually with simple perforations. Sclerenchymatous
elements often present in the primary and especially in the secondary phloem,
notably in species of Brucea, Harrisonia, Simaba, and Simaruba. Pith,
described by Spiekerkoetter (2168) as heterogeneous, consisting of small,
thick- walled cells distributed amongst larger cells with thin walls in Harrisonia,
Medullary secretory canals, often closely associated with the protoxylem,
and very variable in number and size, recorded in certain species of Ailanthus,
Amaroria, Brucea, Eurycoma, Hannoa, Mannia, Odyendea, Perrieria, Picrasma,
Picrella, Picrocardia, Picrolemma, Samadera, Simaba, Simaruba, Simarubop-
sis, Soulamea. Secretory canals apparently absent from species of Alvaradoa,

Cadellia, Castela, Eurycoma, Guilfoylia, Hannoa, Harrisonia, Holacantha,

Hyptiandra, Irvingia, Kirkia, Klainedoxa, Picramnia, Picrasma, Picrodendron,
Quassia, Rigiostachys, Samadera, Simaba, Spathelea, Suriana. The presence
or absence of medullary secretory canals is usually a character of generic
diagnostic value, although exceptions occur, notably in Simaba, where a group
of species with and another without canals was distinguished by Boas (208).
Canals recorded in the cortex in certain species of Harrisonia and Irvingia,
and also in the pericycle in Odyendea sp. Secretory cells also frequent in
the cortex and pith, either alone or accompanied by canals. Oil cells recorded
by Engler (642) in the cortex and pith of Picrella trifoliata H. Baill. and in the
same tissues throughout Ailanthus, Harrisonia, Hebonga, Simaba, Simaruba,
and mucilage cells and cavities in all species of Desbordesia, Irvingia, and
Klainedoxa. Secretory elements generally absent from the Alvaradoideae,
Castelineae, Eurycomineae, Kirkieae, Picramnioideae, and Surianoideae.
Simaruba is sharply differentiated from Simaba, according to Solereder, by
the presence of secretory cells in the axis, petiole, and lamina. Solitary and
clustered crystals common; small solitary ones occur in the cortex of
Ailanthus altissima Swingle and druses, sometimes very large, in the cortex
of Picrasma quassioides (Ham.) Benn.
 SIMARUBACEAE 321
BARK
Structure fully described for Ailanthus altissima Swingle by Miiller (1572)
as follows. Cork consisting of flattened, compact cells, with strongly thickened
inner walls. Phelloderm consisting of tangentially elongated cells. Cork and
phelloderm cells with homogeneous contents. Phelloderm also containing
small starch grains and large cluster crystals. Phloem including conspicuous
groups of fibres and solitary or, more frequently, grouped, tangentially
elongated cells with brown resinous contents. Medullary rays 1-3 cells wide,
consisting of thin-walled, radially stretched cells.

WOOD (Fig. 75)

Vessels mostly medium-sized (100-200 /x mean tangential diameter) in

diffuse-porous species, larger in some species of Ailanthus, Hannoa, Klaine-

doxa, Odyendea, and Simaruba, smallest in Alvaradoa, Brucea, Guilfoylia,
Picraena, Quassia, Samadera, and Suriana\ pore multiples not very common
in most species but multiples of 4 or more moderately common in Castela,
Guilfoylia (Fig. 75 i), Picramnia, and Suriana (2374); clusters present in the
foregoing genera and in Brucea and Picraena in clusters of very small, often
;

angular, vessels in the late wood of Ailanthus altissima Swingle, the clusters
often grouped into tangential lines (Fig. 75 j); tending to a flame-like or
oblique distribution in Alvaradoa, Castela, and Holacantha (938); varying in
number from i to 5 per mm. in the woods with larger vessels, not more than
20 per mm. in the remainder (excluding woods with marked groups of vessels) ;
ring-porous in some species of Ailanthus, Holacantha (938) and Picrasma
(938, 2374), semi-ring-porous in some species of Alvaradoa and Castela.
Spiral thickening present in the smaller vessels of Ailanthus altissima and in
Castela nicholsonii Hook. f. (2374), Holacantha emoryi A. Gray (2374), and
Picramnia (938). Perforations exclusively simple, except for occasional
multiple perforations in the late wood of Ailanthus altissima (2374). Inter-
vascular pitting alternate, often wider horizontally than vertically, including
a wide range of size, large (7-10 \i diam.) in Ailanthus, Guilfoylia, Hannoa,
Irvingia, Kirkia, Klainedoxa, Odyendea, Samadera, and Simaruba, small to
minute in Alvaradoa, Brucea, Castela, Picraena, Picramnia, and Quassia',
sometimes with coalescent apertures most pronounced in Hannoa, Picraena,
Picramnia, and Simaruba', sometimes transitional in Kirkia acuminata Oliv. ;

pits to ray cells similar to intervascular- pits where these are small, large,
unilaterally compound or simple in woods with large intervascular pitting,
e.g. in Hannoa, Irvingia, Klainedoxa, and Samadera. Tyloses abundant some-
;

gum and crystals in Kirkia (2374). Mean member length

times containing
O'3-O'7 mm. Parenchyma very sparse in or absent from Guilfoylia and
Picramnia, vasicentric in Brucea, Dictyoloma, Holacantha, and Kirkia, aliform
or confluent in Aeschrion, Ailanthus, Castela, Hannoa (Fig. 75 F), Irvingia,
Klainedoxa, Odyendea, Picrasma, Quassia, Simaba, and Simaruba, in con-
tinuous bands, probably but not obviously paratracheal in origin, in Castela,
Hannoa, Irvingia (Fig. 75 K), and Klainedoxa; also banded, according to
Heimsch, in Amaroria, Cadellia, Desbordesia, Eurycoma, and Soulamea;
diffuse or fine metatracheal bands predominating in Samadera, and present
in addition to paratracheal parenchyma in Cadellia (2374), Castela, Harrisonia
(938), Klainedoxa, Mannia (938), Picrodendron (938), and Suriana (2374);
4504 Y
 FIG. 75. SJMARUBACEAE
A, Kirkia acuminata Oliv. B, Irvingia gabonensis Baill. C, Brucea macrophylla Oliv. D, Odyendea
snmmermannn Bngl. E, ffannoa klaineana Pierre et Engl. F, H. klaineana Pierre et
EngK G, Ailanthus
alttsnma Swingle. H, Simaba multiflora Juss. I, Guilfoylia
monostylis (Benth.) F. Miill. J, Ailanthus
alttsstma Swingle. K, Irvingia gabonensis Baill.
 SIMARUBACEAE 323
terminal parenchyma present in some species, e.g. of Amaroria, Cadellia,
Holocantha.) and Simaba. Crystals present in the majority of species either
in the ordinary cells (large in Alvaradoa) or in chambered cells, not abun-
dant except in Hannoa undulata Pierre et Engl., Irvingia, Klainedoxa, and
Suriana (2374). Storied in Ailanthus p.p., Cadellia (2374), Castela, Picrasma,
Simaba, Simaruba, and Suriana (2374). Strands usually of 2-4 cells, but
commonly up to 8 cells and sometimes more in Irvingia, Kirkia, and Klaine-
doxa.Rays exclusively uniseriate in Aeschrion, Guilfoylia, and Quassia;
mostly 2-4 cells wide in the other genera, but occasionally, according to

Heimsch, up to 7 cells wide in Alvaradoa, Castela, Holacantha, Picrasma,

and Simaruba', larger and sometimes more than 10 cells wide in Ailanthus ;

more than i mm. high in Ailanthus, Brucea, Guilfoylia, and Samadera\

uniseriates typically few and composed of procumbent cells, but numerous
and composed of upright cells in Picramnia and Samadera, and of a few up-
right to procumbent cells in Alvaradoa and Brucea; rays varying in number
from about 4 per mm. in some species of Ailanthus, Kirkia, Odyendea,
and Picrasma, to about 17 per mm. in Aeschrion, Cadellia (2374), Guilfoylia,
Mannia (2374), and Samadera, and up to 29 per mm. in Suriana (2374);
homogeneous (Kribs's Types I, II, and III) in Aeschrion, Ailanthus,
Amaroria (938), Cadellia (938), Castela, Desbordesia (938), Eurycoma (938),
Hannoa, Holacantha, Irvingia, Klainedoxa, Odyendea, Picrasma, Quassia,
Simaba, Simaruba, and Soulamea markedly heterogeneous (Kribs's Type II A),
;

with 4 or more marginal rows of square or upright cells, in Brucea, Picramnia,

and Samadera and, according to Heimsch (938), in Alvaradoa; Heimsch (938)
lists the following as having moderately heterogeneous rays (Kribs's Type
II B): Aeschrion p.p., Brucea, Harrisonia, Holacantha, Irvingia, Kirkia,
Mannia, Picrodendron, and Simaba cells in most species wider vertically than
;

transversely as seen in tangential sections. Occasional crystals present in the

ordinary cells in some species, but not numerous, and in chambered cells in
Samadera cells containing dark gum in Irvingia and Kirkia, and gum
;

reported by Webber (2374) as sometimes present in Alvaradoa, Cadellia,

Picramnia, and Suriana. Cell walls thin to very thin in some species, parti-
cularly in Brucea, Odyendea, and Simaruba and with small, but numerous,
intercellular spaces in Brucea, Castela, Hannoa, Odyendea, Samadera, and
Simaruba. Pits between ray cells small and very numerous in Picrasma.
Storied in some species of Aeschrion, Castela, Hannoa (2374), Odyendea
(2374), Picrasma, Quassia, Simatuba, and Soulamea (2374), and occasionally
with a tendency to stories in some specimens of Kirkia. Fibres with simple
or bordered pits, more numerous on the radial than on the tangential walls,
except in Irvingia and Kirkia; borders, when present, usually small and
inconspicuous but moderately distinct in Ailanthus and Simaruba] Webber
(2374) distinguishes the following woods as having fibre-tracheids, Alvaradoa,
Picramnia, and all the genera of the sub-family Simaruboideae other than
Castela and Holacantha. Commonly septate in Alvaradoa and Kirkia, the
septate fibres of the former grouped in zones or patches occasionally septate
;

in Ailanthus malabarica DC. (1334), Guilfoylia, Picramnia, Soulamea fraxini-

folia Brongn. et Gris., S. muellertRrongn. et Gris. (2374), and Suriana (2374);
lumina occasionally filled with gum in Suriana and Cadellia (2374); walls
thick in Castela, Guilfoylia, Holacantha, Irvingia, and Klainedoxa. Mean
324 SIMARUBACEAE
length 0-9-1 -7 mm. Vasicentric tracheids reported by Webber (2374) in
Alvaradoa, Castela, and Holacantha; those of Alvaradoa with spiral thicken-
ing (938). Intercellular canals. Vertical intercellular canals occur normally
near the protoxylem of many species of this family (see p. 320). Vertical
canals have also been observed in the secondary wood of many of these species,
but appear to be traumatic, though Record (1843) classifies those occurring
in Simaruba as normal. Webber (2374) reports such canals in Ailanthus
altissima, A. philippinensis Mern, Castela nichokonii, Eurycoma longifolia Jack,
Picraena paloamarga (Speg.) Soeg., Simaruba amara Aubl., and Soulamea
amara Lam., and Heimsch (938) in Brucea macrophylla Oliv. The canals vary
considerably in size. Gummosis of unusually broad multiserate rays has been
reported in Odyendea ovalis A. Peter (2168) and in Ailanthus altissima and
Klainedoxa gabonensis Pierre (2374).

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
There are very few anatomical characters common to the whole of the
Simarubaceae. This lack of homogeneity, which occurs also in the external
morphological characters, seems to indicate that the family is unriatural, but
consists of a number of groups which are themselves relatively uniform. It
was pointed out by Jadin (1138) that, on anatomical grounds, the Irvingeae
are clearly differentiated from the Simarubeae. He also suggested, although
on somewhat inadequate anatomical grounds, that Suriana should form
the basis of a separate family, the Surianaceae, having affinities with the
Geraniaceae. It is interesting to note that Brunellia, which was included in
the system of Bentham and Hooker amongst the Simarubaceae, differs from
most other members of the family which have been investigated anatomically
in the possession of a mixture of simple elliptical perforations and scalariform
perforation plates to the vessels. Brunellia has, since then, on external mor-
phological evidence, been transferred to a separate family placed by Engler
and Hutchinson in their respective systems near the Cunoniaceae, where
similar vessel perforations also occur. Reference to the description of the
Cneoraceae will show that its members exhibit quite notable differences from
the Simarubaceae, so that its separation into a distinct family seems to be fully
justified. Koeberlinia, which was included in the Simarubaceae in the Bentham
and Hooker system, was described under Capparidaceae by Pax and Hoffman
(1678). On anatomical grounds it does not agree very well with either family,
and there seem to be good reasons for treating it separately as the Koeber-
liniaceae.

(ii)
FROM WOOD STRUCTURE
The systematic anatomy of the woods of this family has been investigated
by Webber (2374), who has drawn the following conclusions. Each of the
sub-families Kirkioideae, Irvingioideae, and Alvaradoideae represents a
distinct,homogeneous group, whereas the Surianoideae shows some diver-
gence and the Simaruboideae exhibits somewhat wide variation. The struc-
ture of the Surianoideae supports Solereder's abolition of the monotypic
family Surianaceae, his close grouping of Suriana and Cadellia, and his with-
drawal of Guilfoylia from Cadellia. Within the Simaruboideae, the structure
 SIMARUBACEAE 325
of Holacantha and Castela differs markedly from that of all other genera.
However, the difference does not support Jadin's erection of the monotypic
family Holacanthaceae, since Castela resembles Holacantha. The genus
Picrodendron^ formerly included in the Irvingioideae, bears considerable
resemblance to the members of this sub-family in wood structure.
The wood anatomy of the sub-families Kirkioideae, Irvingioideae, Picram-
nioideae, and Alvaradoideae might support the ranking of these groups as
distinct families or as components of other families if this were suggested on
other morphological grounds.
Heimsch (938) in general supports Webber's conclusions. Record and
Hess (1886) place Picrodendron apart in the Picrodendraceae and Suriana in
the Surianaceae, and comment that the classification of the family would be
simplified still further by excluding Alvaradoa and Picramnia.

ECONOMIC USES
Quassia chips Aeschrion exceka (Sw.) Kuntze (syn. Picraena excelsa Lindl.)
possess tonic and anthelminthic properties. This material is also used in the
preparation of insecticides. The West African Dika Nut consists of the
kernels of Irvingia gabonensis Baill. The powdered leaflets of Ailanthus
altissimaSwingle (syn. A. glandulosa Desf.) have sometimes been used as an
adulterant in belladonna, mint, and other leaves which occur in commerce.
Ailanthus leaves may often be detected by the presence of the characteristic
glands (see 'Leaf'), but other diagnostic characters include the striated cuticle;
the large unicellular trichomes; the clustered crystals which accompany the
veins; the ranunculaceous stomata; the single layer of palisade cells; the
straight anticlinal walls of the epidermal cells. None of the genera furnishes
timber of general commercial importance, though some, e.g. Ailanthus and
Simaruba, produce timber that is used locally, chiefly for packing-cases.
According to Record and Hess (1886) limited amounts of Marupa (believed
to be Simaruba amara Aubl.) have been exported from Brazil for interior
trim; the wood is used locally for house sheathing and boxes and is said to be
resistant to insect attack on account of its bitterness.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aeschrion,* Ailanthus,* Alvaradoa, Amaroria, Brucea, Cadellia, Castela,
Desbordesia, Eurycoma, Guilfoylia, Hannoa, Harrisonia, Hebonga, Hola-
cantha, Hyptiandra, Irvingella, Irvingia, Kirkia, Klainedoxa, Mannia,
Odyendea, Perrieria, Picraena,* Picramnia, Picrasma,* Picrella, Picroden-
dron, Picrolemma, Quassia,* Rigiostachys, Samadera, Simaba, Simaruba,
Simarubopsis, Soulamea, Spathelea, Suriana.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Aeschrion, Ailanthus, Alvaradoa, (Amaroria), Brucea, (Cadellia), Castela,
(Desbordesia), Dictyoloma, (Eurycoma), Guilfoylia, Hannoa, (Harrisonia),
Holacantha, Irvingia, Kirkia, Klainedoxa, (Mannia), Odyendea, Picramnia,
Picrasma, (Picrodendron), Quassia, Samadera, Simaba, Simaruba, (Sou-
lamea), (Suriana).
326 SIMARUBACEAE
LITERATURE
(i) On General Anatomy
Boas 208, Cronquist 503, 504, Davis 551, Engler 642, Jadin 1138, Mailer 1572, Pax
and Hoffmann 1678, Petaj 1700, Schneider 2043, Spiekerkoetter 2168.

(ii) On Wood Structure

de Bastos 147, Benoist 170, den Berger 182, B.H.For.D. 274, Chalk 364, Cooper 461,
Greguss 2522, Heimsch 938, Howard 1088, Jaccard 1130, Janssomus 1154, Jentsch 1174,
Kanehira 1206, 1209, 1214, Kribs 1283, Lecomte 1334, O'Donnell 1629, Pearson and
Brown 1679, Pfeiffer, J. Ph. 1713, Record 1780, 1781, 1783, 1787, 1801, 1809, 1843,
1851, Record and Hess 1886, Record and Mell 1894, Riera 1937, Spiekerkoetter 2168,
Stone 2202, 2206, 2207, Tang 2231, Webber 2373, 2374, 2377, 2378, Williams 2430,
Yamabayashi 2478.

)C
81. BRUNELLIACEAE
(Fie. 77 on p. 334)

SUMMARY
(i) GENERAL
A family of tropical American trees belonging to the single genus Brunellia.
The young stems are typically 2- or 3 -angled at first, and contain a large
proportion of pith. The most characteristic anatomical characters include the
thick-walled, curved, unicellular hairs, which often form a felt on the young
twigs, and the very striking appearance of the mesophyll of the leaf where, in
transverse sections, the palisade and spongy tissues are confined to definite
areas between the transcurrent columns of fibres which extend from the
vascular bundles to the upper epidermis.

(ii) WOOD
Vessels small, with numerous multiples, perforation plates simple and
scalariform, intervascular pitting scalariforin and intermediate, pits to ray
cells similar. Parenchyma absent. Rays up to 6 cells wide or exclusively

uniseriate, heterogeneous. Fibres septate and with simple pits.

LEAF
Dorsiventral.
Hairs, especially on the lower surface, very thick-walled,
unicellular, curved. Cells of the lower epidermis densely papillose. Stomata
not easily observed, but apparently confined to portions of the lower epidermis
above the spongy tissue. Hypoderm, composed of i or 2 layers of thick-
walled pitted cells, present beneath the upper epidermis. This tissue is less
well developed in B. comocladiifolia H. et B. than in B. tomentosa H. et B.
Mesophyll composed of about 3 layers of very tall, narrow cells and a
relatively smaller region of spongy tissue. Vascular bundles of the veins
embedded in the spongy mesophyll, but surrounded by sclerenchymatous
sheaths, and vertically transcurrent by narrow columns of sclerenchymatous
elements extending to the upper epidermis. Midrib with a closed but
dorsally flattened vascular bundle. Petiole, in transverse sections, also
exhibiting a closed vascular strand (dorsally concave in B. tomentosa but more
flattened and interrupted in B. comocladiifolia (Fig. 77 j)), surrounding a
parenchymatous pith, and supported externally by a fairly broad, sinuous
ring of thick-walled fibres. A few subsidiary vascular strands also occur in
 BR UNELLIA CEAE 327
a latero-superior position. Large solitary and clustered crystals sporadically
distributed in the palisade tissue, and clustered ones in the parenchymatous
tissues of the petiole.

Axis
YOUNG STEM
Cortex containing fairly numerous clustered crystals and stone cells.
Pericycle with a broad, continuous, somewhat sinuous ring of sclerenchyma,
consisting mostly of thick- walled fibres. Phloem containing rather infrequent
groups of fibres and fairly numerous clustered crystals, the latter smaller
than those in the cortex. Xylem forming a closed cylinder traversed by
uniseriate rays. Vessels in radial groups of 12 or occasionally more (groups
rather shorter in B. comocladiifolia H. et B. than in B. tomentosa H. et B.,)
seldom exceeding 60 in radial diameter perforations and pitting as described
//, ;

below for the mature secondary xylem. Pith large, very heterogeneous, com-
posed of cells with thin but pitted walls. Crystals, see 'Cortex* and Thloem'.

WOOD
Vessels very to moderately small, mean tangential diameter often less than
50 /A; solitary and with numerous multiples of z to several cells; 5-20 per

sq. mm. Perforation plates simple and scalariform. Intervascular pitting

scalariform or transitional and, according to Tippo (2261), rarely opposite;
pits to ray cells similar, mostly elongated and in scalariform arrangement
(1886). Tyloses present. Parenchyma absent. Rays according to Tippo,
1-6 cells wide; often exclusively uniseriate and more than 12 per mm. often
;

more than i mm. high; heterogeneous (Kribs's Types I and III); with very
few procumbent cells when exclusively uniseriate. Fibres with pits usually
described as simple or indistinctly bordered, e.g. by Record and Hess (1886),
but the cells classified as fibre-tracheids by Tippo (2261); septate and with
thin walls.

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The scalariform perforation plates of the vessels serve to differentiate
Brunellia from the Simarubaceae, where the genus was included in the
Bentham and Hooker system. It is also of interest that the Cunoniaceae, near
which the family Brunelliaceae is placed in the respective systems of Engler
and Hutchinson, likewise possesses scalariform perforation plates.

(ii) FROM WOOD STRUCTURE

Tippo (2261) makes the following comments on the position of the family.
'The Brunelliaceae are higher than the Hamamelidaceae in the following ways ;

presence of fibre-tracheids, larger vessel diameter, and presence of some vessel

elements with simple perforations. The Brunelliaceae are also higher than
the Hydrangeaceae on the basis of these three characters. However, the latter
have less oblique vessel end walls, some alternate intervascular pitting, and a
higher type of ray in some species.*
328 BRUNELLIACEAE
GEJNUS DESCRIBED
(i) FOR GENERAL ANATOMY
The above description is based on specirhens of Brunellia comocladiifolia
H. et B.* and JS. tomentosa H. et B.* in the Kew herbarium.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Brunellia.

LITERATURE
(i) On General Anatomy
Engler 637.
(ii) On Wood Structure
Bausch 154, Record 1851, Record and Hess 1886, Tippo 2261.

^82.
CNEORACEAE
(FiG. 76 on p. 330)

SUMMARY
(i) GENERAL
Small shrubs, confined to the Mediterranean region and the Canary Islands.
The following description of the leaf and stem, except where stated to the
contrary, refers to Cneorum tricoccum Linn., grown at Kew.

(ii) WOOD
Vessels mostly in clusters, with radial chains or flame-like clusters in some
species, perforations simple, intervascular pitting alternate and minute, pits
to parenchyma similar, members extremely short. Parenchyma pre-
dominantly paratracheal, scanty to aliform, and terminal, sometimes con-
sisting almost entirely of fusiform cells and storied. Rays up to 2 or 3 cells
wide, with few uniseriates, low and homogeneous in mature material. Fibres
with simple pits, very short.

LEAF
Dorsiventral.Hairs frequently 2-armed and T- or Y-shaped. Multi-
cellular external glands also present. Epidermis on both surfaces moderately
cutinized, composed of cells with slightly sinuous anticlinal walls. Stomata
occur sporadically on the upper and abundantly on the lower surface, ranun-
culaceous, each surrounded by 5 or 6 ordinary epidermal cells. Mesophyll
consisting of a single layer of palisade cells and a broader, very lacunar,
spongy region. Vascular bundles of the smaller veins embedded in the
mesophyll, each surrounded by a sheath of amyliferous cells. Petiole sup-
plied by an almost completely closed but dorsally flattened vascular strand
accompanied by minute, latero-superior, accessory bundles. Secretory cells,
with oily or resinous contents, present in the mesophyll, but rather incon-
spicuous unless differentiated by staining.
 CNEORACEAE 329

Axis
STEM
Cortex mainly parenchymatous, apart from the secretory cells (see below).
Pericycle containing small, inconspicuous, widely spaced strands of fibres
in C. tricoccum Linn, outer part in C. pulverulentum Vent, demarcated by an
;

almost continuous ring of sclerenchymatous elements accompanied, according

to Engler (640), by cells containing large solitary crystals. Xylem and
phloem forming continuous cylinders traversed by narrow rays. Vessels
unevenly distributed, solitary or in irregular clusters, up to about 18 /x in
radial diameter, perforations simple. Pith consisting of loose parenchyma.
Secretory cells, with oily or resinous contents, present in the primary
cortex, especially in the sub-epidermal region. Crystals, see Tericycle'.

WOOD (Fig. 76 C-D)

Vessels moderately small (50-100 \L mean tangential diameter) or slightly
smaller; commonly in multiplesand chains in Cneorum pulverulentum Vent.
(938), mostly in small multiples and irregular clusters in C. trimerum (Urb.)
Chodat, and in flarne-like clusters in C. tricoccum Linn. (938); about 15 per
sq. mm. in C. trimerum', ring-porous in C. tricoccum (938) and sometimes
tending to be so in C. trimerum with spiral thickening in C. tricoccum (938).
;

Perforations simple. Intervascular pitting alternate and minute; pits to

parenchyma typically similar to the intervascular pitting, but, according to
Record and Hess (1886), often unilaterally compound. Mean member length
about 0-17 mm. Parenchyma predominantly paratracheal; scanty in twigs
of C. pulverulentum and C. tricoccum (938), but moderately abundant in
mature material of C. trimerum (Fig. 76 D), aliform and occasionally linking
adjacent vessel groups, with some scattered cells and terminal bands, i, some-
times 2, cells wide. Consisting in this species almost entirely of fusiform
cells, with only occasional strands of 2 cells, and irregularly storied. Rays in
mature material of C. trimerum up to 2, occasionally 3, cells wide, with few
and very low uniseriates that are often only 2 cells high; multiseriate rays also
very low (less than 10 cells and 250 ju, high) about 7 per mm. homogeneous
; ;

(Kribs's Type I) tending to be arranged in echelon. The rays of immature

;

twigs, according to Heimsch, higher and heterogeneous. Fibres with numer-

ous simple pits on the radial walls. Walls thick. Mean length about 0-7 mm.
Vascular tr ache ids, according to Heimsch (938), 'probably occur' in C.
tricoccum.

TAXONOMIC NOTES
Cneorum was included in the Simarubaceae in the Bentham and Hooker
system. Heimsch (938) concludes that, on the basis of wood anatomy, the
Cneoraceae are closer to the Rutaceae than to the Zygophyllaceae.

GENUS DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Cneorum.*
* Kew
Represented in the slide collection.
 6

K
FIG. 76. KQEBERLINIACEAE, A-B; CNEORACEAE, C-D; OCHNACEAE, E-L
A, Canotia holacantha Torr. B, C. holacantha Torr. C, Cneorum trimerum (Urb.) Chodat. D, C.
trimerum (Urb.) Chodat. E, Bkutemantkus grandiflorus Spruce. F, B. grandiftonts Spruce. G, Lopkira
alata Banks. H, L. alata Banks, I, Ochna arborea Burch. J, O. arborea Burch. K, Ouratea amplectam
(Stapf) Engl. L, Brackenridgea hookeri (Planch.) A. Gray.
 CNEORACEAE 331

LITERATURE
(i) On General Anatomy
Engler 640.
(ii) On Wood Structure
Heimsch 938, Record 1783, 1843, 1851, Record and Hess 1886.

< 83. KOEBERLINIACEAE

(FiG. 76 on p. 330; FIG. 77 on p. 334)

SUMMARY
(i) GENERAL
This family is represented by the small shrub Koeberlinia spinosa Zucc.
which forms dense thickets in Texas and Arizona. The tips of the small twigs
are differentiated as spines. Minute, deciduous leaves occur. The family also
includes the genus Canotia.

(ii) WOOD
Vessels very to extremely small,solitary and with spiral thickening in the
late woodring-porous, intervascular pitting very small and alternate, pits to
;

parenchyma similar members moderately short. Parenchyma apotracheal,

;

as scattered cells or short tangential lines. Rays up to 2-7 cells wide, low or
high, almost homogeneous, with intercellular canals in Koeberlinia. Fibres
with distinctly bordered pits, sometimes with spiral thickening, very short.

Axis
YOUNG STEM (The following description applies only to Koeberlinia spinosa
Zucc.)
Hairs fairly frequent, consisting of short, conical, thick-walled trichomes.
Stomata situated in deep pits. Outer part of the primary cortex differen-
tiated as palisade tissue ; inner part parenchymatous and somewhat lacunar.
Cork arising in the pericycle, consisting of cells with strongly thickened
outer tangential walls. Pericycle demarcated by large, conspicuous, thick-
walled strands of fibres opposite the vascular bundles, connected to form a
ring by narrower groups of pitted, lignified cells with comparatively wide
lumina. Phloem containing isolated or grouped secretory (resin) canals,
tangentially compressed in transverse sections of herbarium specimens. Vas-
cular bundles separated by relatively broad medullary rays, whose distal
ends are slightly enlarged. Vessels not very numerous, unevenly distributed,
solitary in irregular clusters, or, less frequently in radial rows, seldom
and
exceeding 15 /z in radial diameter; perforations simple. Wood fibres with
thick walls, and slit-shaped, inconspicuously bordered pits. Pith composed
of slightly lignified parenchymatous cells, many containing solitary crystals
which are mostly cubical. A few longitudinal columns of vertically elongated
with relatively thick walls and granular contents also occur in the pith.
cells

Secretory canals, see Thloem*. Secretory cells and crystals, see Tith*.
The somewhat similar stem of Canotia holacantha Torr. which has narrower
rays than Koeberlinia is illustrated in Fig. 77 o.
 33* KOEBERLINIACEAE
WOOD (Fig, 76 A-B)
Vessels of the late wood very small (25-50 ^ mean tangential diameter) to
extremely small (less than 25 //,); solitary; very numerous in Canotia (200 per
sq. mm.); often ring-porous; spiral thickening present in the late wood
vessels of stemwood of Canotia and Koeberlinia but absent from the pore-zone
vessels; present in all vessels of branchwood of Koeberlinia (1806). Inter-
vascular pitting alternate, minute; pits to ray and wood parenchyma cells
similar to the intervascular pitting. Dark deposits common in Koeberlinia

(1806). Mean member length 0-3 mm. Parenchyma apotracheal, as

scattered cells or short tangential lines. Strands usually of 4 cells. Rays up
to 7 cells wide and high in Koeberlinia (1886), up to z cells wide, low and
about 14 per mm. in Canotia; uniseriates numerous in Canotia, often only
i or 2 cells
high, composed almost entirely of procumbent cells; almost
homogeneous (Kribs's Type I); with numerous solitary crystals in Canotia.
Record (1806) notes the presence in Koeberlinia of 'numerous radial inter-
cellular canals, varying in size from a pin-hole to a lenticular cavity one-
sixteenth inch high, observed in larger stems but not in small ones. They
resemble those found in certain Apocynaceae and Euphorbiaceae/ Fibres
with distinctly bordered pits on both radial and tangential walls; walls thick;
spiral thickening sometimes present but not very distinct. Mean length about
0-5 mm. A
tendency to ripple marks has been noted (1886) in one specimen
of Koeberlinia.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
Koeberlinia differs from the Simarubaceae, amongst which it was included
in the system of Bentham and Hooker, in having pericyclic cork, and widely

separated vascular bundles. It resembles many of the Simarubaceae, on the

other hand, in possessing secretory canals, but whereas those of the Sima-
rubaceae commonly occur in the pith, those of Koeberlinia are present in the
phloem. These facts seem to justify the exclusion of Koeberlinia from the
Simarubaceae.

GENUS DESCRIBED
(i)
FOR GENERAL ANATOMY
Koeberlinia.* The description is based mainly on an examination of
material from the Kew herbarium.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Canotia, (Koeberlinia).

LITERATURE
On Wood Structure
Record 1806, 1851, 1864-6, Record and Hess 1886.
 (333)

Vfr
84. OCHNACEAE
(Fie. 76 on p. 330; FIG. 77 on p. 334)

SUMMARY
(i) GENERAL ANATOMY
A
tropical family of trees, shrubs, or very rarely herbs. Hairs are in-
frequent, but unicellular, uniseriate, or multicellular when present. The
leaf is usually dorsiventral. Stomata in the leaf are usually but not in-
variably confined to the lower surface, and are sometimes rubiaceous. Cells
with strongly thickened inner tangential and radial walls, the lumen of each
cell being completely filled with a large clustered, or, more rarely, a solitary

crystal, occur in the parenchymatous tissues, especially in Brackenridgea,

Elvaria, Ochna, and Ouratea. These elements, termed 'cristarque cells'
(Fig. 77 F), are most frequent in the outer or inner part of the cortex of the
petiole and young stem, as well as in the region of the lateral veins of the leaf.
Those in the young stem and petiole are often arranged in a continuous or
interrupted layer just below the epidermis. Variously orientated medullary
bundles have been recorded in the petiole of Btastemanthus, Cespedesia^
Godoya, Hilairella Planchonella, and Poecilandra; and similar medullary
>

bundles of various types in the pith of the stem of Cespedesia, Godoya, and
Planchonella. Cortical bundles are nearJy always present in the stem. The
cork is superficial in origin, sometimes arising in the epidermis itself, and is
composed of thin-walled cells or of cells with thickened tangential walls. The
pericycle of the stem nearly always contains isolated groups of fibres.
The xylem and phloem of the young stem are usually in the form of con-
tinuous cylinders traversed by narrow rays.

(ii) WOOD
Vessels sometimes almost exclusively solitary, occasionally with some
radial pattern, perforations typically simple, intervascular pitting alternate,

very small to minute, pits to ray cells similar; with vestured pits in the
Exalbuminosae members of medium length. Parenchyma typically para-
;

tracheal and often scanty in the Albuminosae of Gilg, apotracheal (abundant

diffuse) in the Exalbuminosae, though with a tendency towards abaxial
aliform types; in broad bands in Lophira. Rays up to 2-8 cells wide, often
high; markedly heterogeneous, except in Lophira, Fibres with simple to
distinctly bordered pits of medium length to moderately long.
;

LEAF
Generally dorsiventral, but occasionally centric in certain species of Ouratea.
Hairs infrequent; unicellular, uniseriate, or multicellular when present.
Glandular shaggy hairs recorded on the stipules of Godoya and on the leaf
teeth of Lavradia glandulosa St. Hil. Cuticle on the upper epidermis stated
by Beauvisage (163) to be thick in Strasburgeria. Epidermis composed of
cells with straight anticlinal walls in Blastemanthus and Luxemburgia with y

sinuous walls in Ochna\ inner walls often mucilaginous and sometimes

penetrating into the mesophyll in Brackenridgea, Elvasia, Hilairella, Luxem-
burgia, Ochna, and Ouratea; sometimes papillose. Epidermal cells specially
large and serving for water storage in Sauvagesia racemosa St. Hil. Stomata
 o
MELIACEAE, A and D; RUTACEAE, B,
FIG. 77. E, and G-H; OXALIDACEAE, C and K;
OCHNACEAE, F and N; SIMARUBACEAE,'l and L-M; BRUNELLIACEAE, J;
KOEBERLINIACEAE, O
A, Khaya senegalensis A. Juss. Petiole X 12. B, Ruta graveolens Linn. Petiole X 15. C, Averrhoa
carambola Linn. Petiole Xis. D, Turraea obtusifolia Hochst, Petiole X i<;. E, Barosma betulina
Bartl. et Wendl. Petiole X 15. F, Gomphia sp. Cristarque cells Xa83. G, Choisya ternata H. B. et
K, Petiole Xis. H, Citrus aurantium Linn. Young stem Xi5.' I, Aeschrion exceha (Sw.) Kuntze.
Petiole X ii. J, Brunellia comocladiifolia Humb. et Bonpl. Petiole X 9. K, Averrhoa carambola Linn.
Young stem Xi5. L, Ailanthus altissima Swingle. Petiole Xii. M, A, altissima Swingle. Young
stem X8. N, Gomphia sp. Young stem Xis. 0, Canotia holacantha Torr. Stem Xi.
c.c Layer of cristarque cells. c*e, Cutinized epidermis. $. Secretory canals. s,c. Secretory cavity.
t

t,p. Spongy pith, t,w. Zone of small cells with thin walls.
 OCHNACEAE 335
often but not invariably confined to the lower surface, sometimes rubiaceous,
but subsidiary cells reported to be absent from Lophira and Strasburgeria.
Stomata arranged in crowded groups between the network of veins in Godoya.
Hypoderm present beneath the upper epidermis in Strasburgeria (consisting
of mucilaginous cells) and Lophira. 'Spicular cells' (sclerenchymatous idio-
blasts) form a continuous layer below the upper epidermis in Blastemanthus,
Cespedesia,Ehasia,Hilairella,Luxemburgia,Poecilandra, Trichovaselia, Vaselia.
Similar cells sometimes accompany the vascular bundles of the veins and have
branches extending into the mesophyll. Mesophyll including a single layer
of palisade tissue in Brackenridgea, Elvasia, Ochna, and Ouratea, and large
mucilage cells in Euthemis and Strasburgeria. Vascular bundles of the lateral
veins accompanied by abundant sclerenchyma, the smaller ones often ver-
tically transcurrent. Separate bundles enter the base of, but generally unite
within, the petiole to form a closed or slightly dissected ring, although the
vascular strand sometimes remains open in Elvasia and Ouratea or crescent-
shaped as in Ochna atropurpurea DC. Variously orientated medullary
bundles recorded in the petiole of Blastemanthus, Cespedesia, Elvasia, Godoya,
Hilairella, Planchonella, Poecilandra, Trichovaselia, Vaselia. Petiole of
Wallacea containing a ring of bundles surrounding 2 medullary strands, one
of the latter normally and the other inversely orientated, together with centric
cortical bundles consisting of a central mass of xylem surrounded by phloem
fibres. Petiole of Strasburgia with 5 vascular strands at the base, but the
median central and 2 adjoining lateral ones appear as 2 rings in transverse
sections through the distal end, the 2 remaining laterals being closed or some-
times U-shaped according to Beauvisage (163). Isolated bundles present in
the petiole of Lophira. 'Cristarque cells' frequent below the epidermis of the
petiole in Brackenridgea, Elvasia, Gomphia, Ochna, and Ouratea, sometimes
occurring in the endodermal region or outer cortex as well. Crystals mostly
clustered but frequently solitary, generally accompanying the vascular bundles
of the veins, but also in Sauvagesia scattered in the mesophyll; sometimes
situated in so-called 'cristarque cells' with U-shaped thickenings to the walls.
The 'cristarque cells' accompany the vascular bundles in Brackenridgea,
1

Elvasia, Ochna, and Ouratea. (See also under Tetiole ). Crystals said to be
absent from Lavradia spp.

Axis
YOUNG STEM (Fig. 77 N)
Cork superficial in origin, sometimes arising in the epidermis itself in
Ochna and Ouratea. Cortex containing 'cristarque cells' (Fig. 77 F) (see also
'Summary* and 'Leaf') in Brackenridgea, Elvasia, Gomphia, Ochna, Ouratea,
most frequently situated below the epidermis, but also occurring in the region
of the endodermis as well as in the central tissues of the cortex. The mode of
differentiation and distribution of the 'cristarque cells' is stated by Solereder
to be of specific diagnostic value. Cortical vascular bundles, consisting of
the bases of the lateral vascular strands to the leaves which branch off from
the conducting system of the stem in the region below the nodes, occur
throughout the family, but their number and size are variable. Twenty-four
or more cortical bundles are present in Lophira, whilst they are also well
developed and numerous in Lavradia and Schuurmansia, but less frequent in
 Leitgebiaand Sauvagesia. Bases of the internodes sometimes devoid of cortical
bundles. Cortex also including stone cells as well as abundant clustered
and/or solitary crystals in most of the genera including Lophira. Pericycle
nearly always containing strands of fibres, but sometimes with a continuous
or almost continuous ring of sclerenchyma in Ouratea owing to the cells
between the fibre bundles becoming sclerosed. Pericycle in Lophira and

Strasburgeria at first containing isolated strands of fibres, but these unite

subsequently to form a continuous ring. Xylem and phloem usually con-
stituting closed cylinders. Secondary phloem sometimes containing fibres;
stratified into hard and soft portions in Godoya, the structure as seen in trans-
verse section in this genus recalling that of Titia. Vessels mostly scattered in
Ochna and Ouratea', radially arranged in Luxemburgia; perforations usually
simple, andcircular or elliptical; but in some genera, including Lavradia and
Sauvagesia, accompanied by scalariform plates with a few bars, or with a
larger number of bars in Luxemburgia. Scalariform perforation plates also
recorded by Beauvisage (163) in Strasburgeria. Pith often becoming lignified
at an early stage; sometimes containing medullary bundles, the latter con-

sisting of vessels and fibrous cells in Godoya and Planchonella, but of phloem
and fibrous cells in Cespedesia. Medullary bundles assume a more normal
structure in the axis of the inflorescence. Bundles of fibres occur in the pith
of Lophira, where some are divided into chambers, each containing a solitary
crystal. Medullary bundles stated to be absent from the axis in Blastemanthus,
Hilairella, Luxemburgia, and Poecilandra, although occurring in the petiole of
some of these genera (see 'Leaf ). Crystals, see 'Cristarque cells*, 'Cortex*,
and Tith*. Mucilage sacs or passages present in the cortex and pith of
Euthemis, Sauvagesia, Schuurmansia, and Strasburgeria. Irregularly dis-
tributed secretory cells, containing granular or refractive, probably tannini-
ferous material which is stained readily by safranin, occur in the cortex,
phloem, ray cells, and pith of Gomphia sp. and Ochna atropurpurea DC. and
probably in other unexamined genera and species as well

WOOD (Fig. 76 E-L)

Vessels mostly small (mean tangential diameter less than 100 /x), some-
times very small (25-50 p), medium-sized in Blastemanthus (1886), Cespedesia,
and Wallacea and moderately large in Lophira] almost exclusively solitary in
Brackenridgea, Elvasia p.p., Ochna, and Ouratea, with multiples of 2-3 cells
moderately common in the other genera, with a loose oblique pattern in
Lophira\ according to Solereder, with a radial arrangement in Luxemburgia;
very variable in number, ranging from about 2 per sq. mm. in Cespedesia and
Lophira to 80-150 per sq. mm. in Ochna arborea Burch. (360), mostly between
20 and 50 per sq. mm. Perforations exclusively simple except in Lavradia,
Luxemburgia, and Sauvagesia, for which Solereder records some scalariform
plates. Intervascular pitting alternate, very small to minute; pits to ray
cells
similar. Vestured pits reported by Bailey (78) in 5 genera of the Exalbumi-
nosae. Tyloses observed only in a single specimen of Blastemanthus grandi-
fiorus Spruce; solid deposits present in Blastemanthus, Lophira, and Ochna.
Mean length o-5~o*9 mm. Parenchyma (a) paratracheal, in Blastemanthus
(Fig. 76 E), Cespedesia, Tyleria, and Wallacea, typically scanty and confined
to a few cells round the vessels, but more abundant and irregularly vasicentric
 OCHNACEAE 337
in Cespedesia macrophylla Benth, ; (b) predominantly apotracheal in Bracken-
ridgea, Elvasia, Ochna 76 i), and Ouratea, typically diffuse and in short
(Fig.
uniseriate lines (sometimes wider in Elvasid) that often tend to be associated
with the abaxial sides of the vessels, e.g. in Brackenridgea hookeri A. Gray
(Fig. 76 L), Elvasia essequiboensis Engl., and some species of Ouratea, e.g. O.
amplectans Hutch, et Dalz. and that formerly known as Gomphia oblongifolia
Ridl. x and (c) as regular, broad, continuous bands 3-4 cells wide in Lophira
;

(Fig. 76 H). Distinct terminal bands observed only in Blastemanthus and

Tyleria. Crystals in chambered cells present in small numbers in Cespedesia
and Elvasia, abundant in Lophira', dark-coloured gum abundant in all the
species examined. Strands commonly of 8 or more cells. Rays multiseriate,
2-3 cells wide in Blastemanthus, Elvasia p.p., Lophira, and Wallacea, 4-8 cells
wide in the remainder commonly i mm. or more in height except in Lophira
; ;

uniseriates typically numerous and composed entirely of high upright cells,

but with both square and upright cells in Blastemanthus grandiflorus Spruce ;

uniseriates often rather few in Lophira and composed entirely of procumbent

cells; 10-20 per mm., most numerous in Blastemanthus and fewest in Lophira;
homogeneous (Kribs's Type I) in Lophira but markedly heterogeneous
(Kribs's Type I) in the other genera, commonly with 10 or more rows of
upright or square marginal cells sheath cells present in Brackenridgea, Ochna,
;

and Ouratea. Single crystals occasionally present in the ordinary cells; large
thick- walled idioblasts, each containing a single crystal present in Ochna and
Ouratea (except Ouratea oblongifolia)', similar, but smaller, cells occur in
Elvasia; cells containing abundant, dark-coloured gum. The multiseriate
rays, except in Blastemanthus and Lophira, appear to be derived from the
splitting of very high primary rays; very high rays (up to about 8 mm.) occur
sporadically and the rays tend to be of 2 distinct sizes. Fibres with simple
pits in Tyleria and Wallacea, with small to moderately distinct bordered pits
in the other genera pits usually rather more numerous on the radial than on
;

the tangential walls; pits numerous in Elvasia, Ochna, and Ouratea; some
fibres septate in Tyleria. Walls very thick, the secondary wall often showing
Mean length 1-2-2-0
distinct zones. mm. Vasicentric tracheids present in
small numbers in Lophira.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
The only outstanding characteristic for the whole family is the possession
of cortical bundles. in the genera whose inclusion in the
These occur not only
family has been long established but also in Lophira which was treated in the
Bentham and Hooker system as belonging to the Dipterocarpaceae and in
Lavradia, Leitgebia, Sauvagesia, and Schuurmansia which the same authors
included in the Violaceae. The presence of 'cristarque cells* in Bracken-
ridgea, Elvasia, Gomphia, Ochna, and Ouratea is an important feature which
confirms the existence of close affinities between these genera, but serves to
differentiate them from other genera in the family in which these special cells
have apparently not been recorded. The presence of mucilage cells and
1
This species is now considered to be an Ouratea, but the specific epithet has already been
used for Ouratea oblongifolia Rusby and no new combination appears to have been published.
338^ OCHNACEAE
passages in Euthemis, Sauvagesia, Schuurmansia, and Strasburgeria serves to
distinguish these genera rather clearly from the others.

(ii)
FROM WOOD STRUCTURE
The genus Lophira differs from all the others, particularly in its parenchyma
and its much more highly specialized rays. Such affinities as it has with the
family would appear to be with the Exalbuminosae, possibly through Elvasia y

rather than with the Albuminosae. Vestal (2329) suggests that there may be
a connexion between the Ochnaceae and the Dipterocarpaceae through
Lophira.
Gilg's two sub-families Exalbuminosae and Albuminosae are clearly distin-
guishable in their wood structure by the nature of the parenchyma and the
presence of vestured pits in the Exalbuminosae, Vestal also notes a greater
elongation of the upright cells in the rays as characteristic of the Albuminosae,
but this does not appear to be entirely dependable.
wood of Testulea gabonensis Pellegr. given by Normand
Illustrations of the

(1612) show a general resemblance to the Ochnaceae in the cross-section, but

a much more highly specialized ray type, comparable with that of Lophira.

ECONOMIC USES
The
only timber of commercial importance is Ekki, the product of Lophira
alata var. procera Burtt Davy. This is a very hard, durable wood, used chiefly
for sleepers and marine piling. The wood of Ochna arborea Burch. is used in
South Africa for the handles of tools. Meni oil is obtained from the seeds of
Lophira alata Banks.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Blastemanthus, Brackenridgea, Cespedesia, Elvasia, Euthemis, Godoya,
Gomphia,* Hilairella, Lavradia, Leitgebia, Lophira, Luxemburgia, Ochna,*
Ouratea, Planchonella, Poecilandra, Sauvagesia, Schuurmansia, Strasburgeria,
Trichovaselia, Vaselia, Wallacea.
* in the Kew slide collection.
Represented

(ii) FOR WOOD STRUCTURE

Blastemanthus, Brackenridgea, Cespedesia, Elvasia, (Lavradia), Lophira,
(Luxemburgia), Ochna, Ouratea, (Sauvagesia), Tyleria, Wallacea.

LITERATURE
(i) On General Anatomy
Beauvisage 163, Gilg 770, Privault 1759.

(ii) On Wood Structure

Bailey 78, Bausch 154, Besson 186, Chalk 360, 364, Cooper 461, Foxworthy 705,
Hopkinson 1083, Jentsch 1173, Lecomte 1335, M6niaud 1491, Normand 1612, Record
1843, 1851, Record and Hess 1886, Record and Mell 1894, Riera 1937, Stone 2206, Thieme
2247, Vestal 2329, Williams 2430.
 (339)

85. TETRAMERISTACEAE
(FiG. 79 on p. 346)

SUMMARY
(i) GENERAL
The family is represented by the sole genus Tetramerista which includes
a few arboreal species in the Malayan region. The most interesting anatomical
character is the occurrence of raphides.

(ii) WOOD
Vessels large and mostly in multiples of 2-4 cells, perforations simple,
intervascular pitting alternate and minute, pits to ray cells similar, members
very long. Parenchyma apotracheal, diffuse. Rays up to 4-5 cells wide, very
high, markedly heterogeneous; with raphides in enlarged cells. Fibres with
simple pits and very thin walls, very long.

LEAF
Dorsiventral. Hairs. None observed. Epidermis on both surfaces com-
posed of pentagonal and hexagonal, moderately thick- walled cells, covered
externally with a striated cuticle. Stomata confined to the lower surface;
each surrounded by 4, or sometimes more, ordinary epidermal cells (ranuncu-
laceous), 2 of which are parallel to the pore in some instances (rubiaceous).
A single layer of hypoderm with pitted cell walls occurs beneath the upper
epidermis. Mesophyll composed of i or 2 rows of palisade cells and a much
broader region of spongy tissue. Vascular bundles of the smaller veins
embedded in the mesophyll, partly surrounded by thick-walled fibres. Mid-
rib and leaf base supplied by a solitary, annular vascular strand which is
flattened or slightly concave towards the adaxial surface, and supported at the
periphery by a continuous ring of moderately thick-walled fibres. A few
sclerenchymatous idioblasts occur in the midrib and also in the central part
of the mesophyll. Crystals. Large cells containing raphides or crystal-sand
present in the midrib and in the palisade layer of the mesophyll.

Axis
YOUNG STEM
Cork arising in the inner part of the cortex. Cortex containing scattered
stone cells, either isolated or in groups. Pericycle not well defined in the
material available. Phloem and xylem in the form of continuous cylinders
traversed by narrow rays. Scattered stone cells similar to those in the cortex,
also present in the phloem. Vessels mostly in radial rows, somewhat angular,
seldom exceeding 60 p, in radial diameter; perforations mostly simple, very
oblique, but others with scalariform plates with about 8 thin bars. Wood
fibres with narrowly bordered pits with elongated, slit-shaped apertures.
Pith composed of moderately thick-walled, pitted cells. Crystals. Raphides
occur in special cells in the pith and phloem, similar cells filled with crystal-
sand also being present in the phloem.
340 TETRAMERISTACEAE
WOOD (Fig. 79 H-I)
Vessels large (mean tangential diameter more than 200 /u) solitary and in
;

numerous multiples of 2-4 cells; about 2-3 per mm. Perforations simple,
moderately oblique. Intervascular pitting alternate, minute; pits to ray and
wood parenchyma cells similar. Mean length about 1-5 mm. Parenchyma
apotracheal, typically as scattered cells, but with a tendency to continuous
bands in some specimens. Strands usually of 8 cells. Rays up to 4-5 cells
wide and often several millimetres high; uniseriates moderately numerous
and composed of square and upright cells, but largely replaced in some
specimens by the marginal rows of the multiseriate rays; about 10 rays per
mm. markedly heterogeneous (Kribs's Type I), with up to 10 to very many
;

marginal rows of square to upright cells, these cells often biseriate near the
procumbent cells. Enlarged procumbent cells containing raphides present in
the multiseriate rays. Fibres with simple pits that occur mainly in the radial
walls. Walls very thick. Mean length about 2-5 mm.

TAXONOMIC NOTES
The taxonomic position of Tetramerista is not well established. It differs
from the Ochnaceae in lacking cortical bundles in the young stem, and from
the Theaceae in possessing raphides. On the advice of Dr. J, Hutchinson it
is here treated as a separate family.
The wood anatomy generally does not support the inclusion of Tetramerista
in either the Ochnaceae or the Marcgraviaceae, though the occurrence of
raphides in the rays of both Tetramerista and Marcgravia may indicate some
affinity with the last genus. The wood of Tetramerista shows a greater general
resemblance to that found in the Theaceae, but the genus stands out as
exceptional if included in this family. The evidence of wood anatomy sup-
ports Dr. Hutchinson's advice to treat Tetramerista as a distinct family.
Although, as already stated, Tetramerista has some features in common with
the Marcgraviaceae, it does not agree very closely with any one genus. For
example, while it has the raphides of Marcgravia and Souroubea, its vessel
groups, non-septate fibres, and diffuse parenchyma link it with Norantea
rather than Marcgravia; it differs from all three genera in having minute
intervascular pitting.
Vestal (2329) states that in minute wood anatomy Tetramerista Resembles
its

in all characters the Caryocaraceae, except in ray type and the presence of
crystals in the genus Caryocar* but in the material examined by the author
;

the resemblance is less marked, particularly as regards type of intervascular

and vessel-ray pitting; the parenchyma, too, may differ considerably and the
occurrence of raphides in the rays of Tetramerista is a further point of
difference.

ECONOMIC USES
The timber, according to Desch (574), is of local importance in Malaya and
has been commonly obtainable in the Singapore market from the adjacent
Dutch islands.
 TE TRA MERIS TA CEAE 341

GENUS DESCRIBED
(i) FOR GENERAL ANATOMY
Tetramerista.* The material examined was T. glabra Miq. from the Kew
herbarium.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Tetramerista.
LITERATURE
On Wood Structure
Desch 574, Hess 959, Vestal 2329.

86. BURSERACEAE
(FiG. 78 on p. 342; FIG. 79 on p. 346)

SUMMARY
(i) GENERAL
A tropical family of resinous trees and shrubs. The hairs are of various
types and include simple and stellate clothing trichomes, as well as glandular
hairs which may be either capitate or snail-shaped. The epidermis, parti-
cularly on the upper surface of the leaf, often contains a proportion of
mucilage cells. Similar cells also occur, although more rarely, in the meso-
phyll or in the parenchymatous tissues of the stem. Stomata sometimes occur
on both surfaces of the leaf but are more frequently confined to and always
more numerous on the lower side; they are ranunculaceous. Hypoderm, on
the upper side of the leaf, is infrequent and limited to a few genera. In the
mesophyll the palisade tissue most frequently consists of a single layer, but
exceptions occur. The petiole is usually cylindrical or dorsally flattened,
sometimes winged, and, in transverse sections, nearly always exhibits a com-
plete circle of vascular bundles, whilst medullary bundles are enclosed within
the circle in Canariellum, Canarium, Dacryodes, Pachyhbus, Santiria, and
Trattinickia. Cork generally arises in the sub-epidermis of the young stem,
whilst the cortex sometimes contains sclerenchymatous idioblasts. The
pericycle is characterized either by arcs or a composite continuous ring of
sclerenchyma. The xylem and phloem constitute closed cylinders traversed
by medullary rays 1-2 cells wide. The phloem often includes a proportion of
fibres. The pith of the stem may be homogeneous or heterogeneous, usually
lignified. Inversely orientated medullary bundles occur
in the stem of all

species of Canarium. Secretory canals generally present in the phloem

of both leaf and stern, and occur in the pith as well in a few species. For
mucilage cells see 'Epidermis*. Solitary and clustered crystals are frequent,
e.g. in the cortex.
Whilst the Burseraceae as a whole are fairly well defined anatomically, the
component genera cannot easily be distinguished. The genus Protium for
instance is anatomically homogeneous, but in Commiphora the range of
structure amongst different species is almost as wide as in the whole of the
family. A
considerable range of variation occurs also in Bursera.
342 BURSERACEAE
(ii) WOOD
Vessels moderately small to medium-sized; perforations exclusively simple,
intervascular pitting alternate, with large hexagonal borders, pits to ray cells
and parenchyma large, simple and unilaterally compound; members of
medium length. Parenchyma paratracheal; varying from scanty to vasi-
centric and occasionally with some diffuse. Rays up to 2-4 cells wide with
few uniseriates; heterogeneous; enlarged cells containing single crystals
present in some species in echelon or storied locally in some genera. Fibres
;

septate with small simple pits of medium length to moderately long. Inter-
; ;

cellular canals present in the rays of many species.

LEAF
Generally dorsiventral; occasionally isobilateral. Hairs include the follow-
ing diverse kinds, (i) Simple, sometimes either hooked or 2-armed. (ii) Stellate

FIG. 78. BURSERACEAE

A-B, Elevated stomata of Santiria mollis Engl. C, Branched trichome of Santiria mollis. D-G,
External glands: D, Boswellia papyrifera (Delile) A. Rich.; E, Protium spruceanum (Benth.) Engl.;
F, Canarium hispidum El. G, Crepidospermum rhoifolium, Tr. et Planch.
; By Solereder.

and tufted, e.g. in species of Canarium and Santiria (Fig. 78 c). (iii) Glan-
dular, either capitate or snail-shaped. Glandular hairs with stalks of 1-2 cells
and 2- to 4-celled heads recorded in Boswellia papyrifera (Delile) A. Rich.
(Fig. 78D), similar hairs with rather longer stalks in Canarium acutum EngL
and Garuga pinnata Roxb., thread-like hairs broadened at the apex in Protium
spruceanum (Benth.) Engl. (Fig. 78 E). Snail-like glands reported by Solereder
in species of Canarium (Fig. 78 F), Crepidospermum (Fig. 78 G), Pachylobus,
Santiria. These various kinds occur either alone or together and their distribu-
tion is of specific diagnostic value. Lower surface of the lamina provided with
open grooves filled with small hairs in Trattinickia rhoifoliaWilld. Epidermis
somewhat variable in structure, frequently including a varying proportion of
mucilage cells. Mucilaginous cells recorded in the upper epidermis of Bursera
angustata Griseb., B. aptera Ramir, B. bipinnata (Schlecht) EngL (whole
 BURSERACEAE 343

epidermis), J5. multijuga Engl. (scattered), Canariwn boivini Engl., Commi-

phora spp, Pachylobus spp., Protium spp. Mucilage cells present on the lower
side in Bursera aptera and rarely in Pachylobus absent from Tetragastris. Lower
;

epidermis papillose in Canariellum, Commiphora spp. (papillae described as

external idioblasts), and Trattinickia sp. containing groups of silicified cells in
;

Protium. Stomata present on both surfaces or limited to, and always more
numerous on, the lower side ranunculaceous. Stomata recorded on both
;

surfaces in certain species of Commiphora and Tetragastris confined to the

;

lower surface in other species of Commiphora situated at the tips of columnar

;

elevations above the network of veins in Santiria mollis Engl. (Fig. 78 A-B).
Hypoderm recorded below the upper epidermis in Aucoumea (cells elongated
and thickened), Canariellum (z or 3 layers), certain species of Commiphora,
Dacryodes, and Triomma. Mesophyll generally dorsiventral, but palisade
tissue recorded beneath both surfaces in a few species, notably in Commiphora,
Palisade tissue consisting of i layer in Canarium (sometimes double in the
section Monodelpha Engl.), Commiphora, Crepidospermum (pro parte),
Dacryodes, Garuga, Pachylobus (usually), Protium (very rarely 2 layers),
Tetragastris, Trattinickia, Triomma\ consisting of more than i layer in
Aucoumea, Boswellia, Canariellum, Crepidospermum (pro parte), Pachylobus
(sometimes) and Santiria. Some of the palisade cells mucilaginous in
Canarium, Protium, Santiria, and Trattinickia, those of Protium occasionally
contain a crystal as well. Crystalliferous cells also occur in the palisade layer
of Scutinanthe. Mesophyll exhibiting a considerable range of structure in
different species of Trattinickia containing mucilage cells in certain species
;

of Commiphora. Spongy mesophyll well developed, in the particularly thick

lamina of Canariellum. Vascular bundles of the small veins containing
secretory canals in the pith in Canarium, and in the phloem of the larger
veins in most genera. Bundles vertically transcurrent in Dacryodes and
Protium. Petiole, in transverse sections, nearly always exhibiting a circle of
vascular bundles, e.g. in Canariellum, Canarium, Pachylobus, Santiria, Tratti-
nickia, the outer ring enclosing medullary bundles, often inversely
orientated, in Canariellum, Canarium, Dacryodes, Pachylobus (excluding
Sorindeia juglandifolia Oliv. var. dahomensis A. Chev. (syn. P. dahomensis
Engl.), and Santiria. Petiolar bundles normally in a circle, but occasionally
in arcs in Commiphora. Petiole of 'Scutinanthe boerlagii Hochr/ containing a
ring of xylem and phloem surrounding a central, normally orientated mass of
xylem and phloem. Petiole of Bursera very variable in transverse sections
through the distal end, with a ring of xylem and phloem supported by a closed
ring of sclerenchyma in B. tonkinensis Guill. with an arc of 4 bundles and
;

2 lateral masses of collenchyma in B. galeottiana Engl. with an open arc

;

of bundles surrounded by a collenchymatous sheath in B. aptera. A layer of

cork has been recorded on one side of the petiole in Tetragastris. 'Cortex' of
the petiole containing numerous sclerenchymatous idioblasts in Canariellum
and a smaller number in Dacryodes, and very numerous crystals in Commi-
phora. Petiolules provided with basal swellings, believed to have a motor
function, in Aucoumea, Canarium, and Pachylobus. Crystals, see 'Mesophyll*.
Secretory canals, see Veins'. Mucilage cells, see 'Epidermis' and
'Mesophyll'.
344 BURSERACEAE
Axis
YOUNG STEM
Cork originating in the sub-epidermis except in Santiria where it is deeper.
Layers of cork, peeling off like paper, in certain species of Boswellia and
Commiphora. Primary cortex containing sclerenchymatous idioblasts (pre-
sumably identical with the stone cells mentioned by Solereder) in Bursera
(scattered or in small groups), Canariellum (abundant), Canarium (sometimes
in groups of 6-10 ; absent from C. balansae EngL), Crepidospermum, Dacryodes,
Garuga, Protium (often), Santiria (small, sometimes in groups), Trattinickia,
Triomma. Cortex traversed longitudinally by secretory canals in Tetra-
gastris*, including mucilaginous cells in species of Hedwigia, Garuga, and
Protium, and sclerenchymatous fibres in Pachylobus. Pericycle with a com-
posite and continuous ring of sclerenchyma, the latter consisting of arcs of
fibres with stone cells between them, in Aucoumea, Bursera (most species),
Canarium, Commiphora (certain species), Garuga, Santiria, Tetragastris,
Triomma; with separate arcs of sclerenchyma in Commiphora (very small in
some species; limited to groups of fibres in the region of the largest secretory
canals in C. fraxinifolia Bak. and C. marchandi Engl.), Crepidospermum
(almost continuous), Dacryodes, Pachylobus, Protium. Pericyclic sclerenchyma
stated by Sabnis (1977) to be absent from Commiphora mukul EngL Phloem
and xylem constituting closed cylinders. Secondary phloem sometimes
provided with fibres, notably in Aucoumea, Bursera (rare), Canariellum,
Garuga (forming a continuous ring), Santiria macrocarpa King but not in other
species of Santiria, and Trattinickia. Fibres stated by Guillaumin (839) to be
absent from the phloem in Canarium. Xylem including small vessels with simple
perforations, the vessels sometimes tending to be in radial groups parenchyma
;

which is chiefly paratracheal, but tangential bands of metatracheal parenchyma

have also been recorded in certain species of Commiphora; uniseriate or
biseriate rays which are often crowded. Pith usually homogeneous in Bos-
wellia, Canariellum, Canarium, Dacryodes, Protium, Trattinickia, Triomma;
heterogeneous in Aucoumea, Garuga, Pachylobus, Santiria (outer part scleri-
fied, inner part thin-walled), Tetragastris. Pith of Canarium secundum A. W.
Benn differentiated into a central, non-sclerified region ; a completely sclerified
region; an outer zone similar to the central one and containing a circle of
medullary bundles. Secretory canals recorded in the pith of Boswellia,
Canarium balansae Engl., Commiphora (certain species), Sorindeia juglandi-
folia Oliv. var. dahomensis A. Chev. (syn. Pachylobus dahomensis Engl.) (at
the periphery; not occurring in other species of Pachylobus). Medullary
bundles present throughout Canarium, but not in the closely related genus
Pachylobus. The number and arrangement of the medullary bundles varies,
even within a single branch in Canarium, and is therefore of no specific
diagnostic value. Secretory canals (see also 'Cortex' and Tith') almost
invariably present in the phloem, and therefore of diagnostic value for the
family, but stated to be absent from the phloem in all organs of 'Scutinanthe
boerlagii Hochr.'.
Certain species of Commiphora are characterized by spiny or non-spiny
shoots of limited growth. Non-spiny short shoots of C. marchandi Engl.
and C. orbicularis Engl. provided with a star-shaped pith and exceedingly
large medullary rays.
 BURSERACEAE 345

WOOD (Fig. 79 A~G)

Vessels moderately small (50-100 /u,
mean diameter) in Balsamodendron,
Boswellia, Bursera, Elaphrium, Pachylobus p.p., Protium, and Tetragastris,
medium-sized (100-200 ^ mean diameter) in the other genera, largest in
Garuga\ solitary and in multiples of 2 or 3, occasionally 4 or 5, cells and
commonly with some irregular clusters with a tendency to an oblique radial
;

pattern in some species of Aucoumea, Canarium, Commiphora, Pachylobus,

Protium, Santiria, Tetragastris, and Trigonoclamys; usually varying in number
between 4 and 15 per sq. mm., occasionally more numerous, e.g. in Protium
with 20-40 per sq. mm,, fewest (about 5 per sq. mm.) in some species of
Aucoumea, Bursera, Canarium, Garuga, Santiria, and Trattinickia. Perfora-
tions exclusively simple; intervascular
pitting with hexagonal
alternate,
borders, large, except in some species of Dacryodes, Pachylobium, Protium,
Trigonoclamys, and Triomma, coalescent apertures present in Aucoumea and
some species of Canarium pits to ray cells and parenchyma large and simple,
;

commonly unilaterally compound. Thin-walled tyloses abundant in some

species of Bursera, Canarium, Elaphrium, Tetragastris, and Trattinickia, and
occasionally containing starch grains, crystals, or gum (1154, 2378). Mean
member length usually between 0-3 and 0*6 mm.; Webber (2378) notes
extremes of o-i mm. in Crepidospermum and 0-9 mm. in Canarium. Paren-
chyma paratracheal, as sheaths round the vessels, usually rather scanty
(Fig. 79 A) when vasicentric, usually limited to a single row, but biseriate in
;

some species of Canarium and Tetragastris', very sparse in some species of

Bursera, Canarium, Elaphrium, Hemisantiria, and Pachylobus diffuse paren- ;

chyma in addition has been reported (592, 1207, 1208, 2378) in species of
Boswellia, Canarium, Dacryodes, and Protium', broken bands have been
reported (592, 2168, 2378) in some species of Commiphora, Protium, and
Santiria. Sometimes containing dark gum crystals not observed. Sometimes
;

with uniseriate terminal bands in Dacryodes and Protium, and, according to

Spiekerkoetter (2168), in broad tangential bands in Commiphora subcrenata
A. Peter, C. mildbraedii EngL, and C, heterozygia A. Peter. Solereder notes
that according to J. Moller there are large thin-walled oil cells in the wood of
Bursera aloexylon EngL Silica present in Santiria oblongifolia Bl. (794).
Strands usually of 4 cells, but sometimes of up to 8 cells. Some of the cells
reported (2378) to be septate. Rays exclusively uniseriate in some species of
Crepidospermum, Dacryodes, Hemisantiria, Protium, Santiria, Tetragastris
(938), and Trigonoclamys; 2-3 cells wide in most of the other species and up
to 4 cells wide in some species of Boswellia, Bursera, and Garuga; Heimsch
(938) notes that the rays may be up to 5 or 6 cells wide in a few species; less
i mm.
than high; uniseriate rays few in species with multiseriate rays and
composed of both upright and procumbent cells; mostly 4-6 rays per mm. in
woods with multiseriate rays, more numerous (up to 12 mm.) in woods with
uniseriate rays only; heterogeneous (Kribs's II B and III), with 1-3
Type
rows of square or upright marginal cells, up 4 rows in some species of
to
Pachylobus', the marginal cells commonly containing single crystals, such cells
sometimes wider tangentially than the non-crystalliferous cells and con-
spicuous in tangential sections, as in Balsamodendron, Bursera p.p., Canarium
p.p., Commiphora, Dacryodes (2378), Garuga, Protium, and Tetragastris \
346 BURSERACEAE
procumbent and upright cells commonly containing dark gum. Silica reported
(574, 794) in some species of Canarium, Dacryodes, Protium, and Santiria.
Intercellular spaces moderately distinct in some species. With distinct echelon
arrangement and sometimes storied locally in Canarium p.p., Pachylobus p.p.,

FIG. 79. BURSERACEAE, A-G; TETRAMERISTICACEAE, H-I

A, Aucoumea klaineana Pierre. B, A, klaineana Pierre. C, Santiria griffithii Engl. D, Pachylobus
rostrata (Bl.) HJ.L. E, Canarium schweinfurthii Engl. F, Protium heptaphyllum March. G, Boswellia
sertata Roxb. H, Tetramerista glabra Miq. I, T. glabra Miq.
i.e. Intercellular canal, r. Cell containing raphides.

most species of Santiria, Trattinickia, and Trigonochlamys. Fibres often thin-

walled, but with moderately thick walls in Boswellia, Bursera p.p., Canarium
p.p., Dacryodes, Pachylobus, Protium, Santiria p.p., Tetragastris, and Triomma\
sometimes with a mucilaginous layer and sometimes containing gum septate ;

in all the material examined, but Spiekerkoetter (2168) states that septa are

absent from Commiphora holziana Engl. and Heimsch (938) notes their
absence from two species of Canarium, whose identity is suspect (see under
Taxonomic Notes); pits simple, small, slit-like to almost round, and more
 BURSERACEAE 347
numerous on the radial than on the tangential walls; fibres often in regular
radial rows; Janssonius (1154) refers to occasional large crystals in the fibres
of Garuga pinnata Roxb., but these do not appear to be present ift all material
of this species. Webber (2378) notes the occasional occurrence of starch
grains, crystals, and brown, gummy masses. Silica present in Santiria spp.
(794). Mean length 0-8-1-4 mm. Intercellular canals in the secondary
rays a constant feature of some species, but of sporadic occurrence in others
(Fig. 79 G); small or large; Webber (2378) notes unusually large canals in
Protium puncticulatum Macbr. ;
canals observed or reported in at least some
species of Boswellia, Bursera, Canarium, Canariellum (2378), Commiphora,
Dacryodes (185), Elaphrium, Garuga, Protium^ Tetragastris (2378), and
Triomma. Desch (574) notes the absence of radial canals from Scutinanthe
brunnea Thw., but there appears to be some doubt about the correct identity
of this wood. The epithelial cells usually distinctly smaller than the other
cells; according to Heimsch (938) the canals in Commiphora zimmermannii
Engl. are lined with sclerotic cells. Webber (2378) notes abundant tylosoids
in Bursera microphytta Gray. Traumatic vertical canals have been reported
(2378) in Canarium, Protium, and Santiria. Webber (2378) notes the occur-
rence of gum-filled pith flecks in various species of Bursera, Canarium,
Crepidospermum, Dacryodes, Protium, and Tetragastris, 'suggesting that
traumatic vertical canals may be present under certain conditions*. Besson
(186) shows rather low ash and silica percentages for woods of this family.

ROOT
Examined only in Aucoumea and Canarium spp. Cortex containing a
continuous or interrupted zone of sclerenchyma. Numerous layers of dis-
continuous sclerenchyma occur in Aucoumea. Phloem containing frequent
secretory canals of large diameter, and a few fibres. Xylem provided with
vessels of large diameter filled with tyloses.

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The Burseraceae constitute an anatomically homogeneous group. Guillau-
min (839) considers that they have affinities with the Rutaceae, Anacardiaceae,
Simarubaceae, Cneoraceae, Meliaceae, Coriariaceae, Sapindaceae, and Hippo-
castanaceae, but of these, the most marked affinities are with the Rutaceae,
Anacardiaceae, Meliaceae, and Simarubaceae. Guillaumin (838) has also
drawn attention to the fact that the anatomical characters of Pachylobus
dahomensis Engl. differ somewhat from those of other species of Pachylobus.
For this reason he suggests that this species may have been wrongly included
in the Burseraceae, and that it shows closer affinities with the Anacardiaceae
and in particular with Sorindeia. In another paper Guillaumin (840) has
shown that, on anatomical grounds, Canarium sumatranum Boerlage et
Koorders is correctly included in Canarium. While anatomical characters
such as the occurrence of resin canals in the phloem of both families have
been used as evidence of the close relationship of the Burseraceae and
Anacardiaceae, Engler (643) considers that the constant floral differences
between the two groups are too great to justify the view that there is any very
348 BURSERACEAE
close affinity between them. Hutchinson treats the families in separate orders,
which are, nevertheless, regarded as being phylogenetically related.

(ii) FROM WOOD STRUCTURE

Both Webber (2378) and Heimsch (938) consider that the Rutaceae,
Simarubaceae, Meliaceae, Sapindaceae, Burseraceae, and Anacardiaceae form
4
a natural group. Heimsch states that The Burseraceae and Anacardiaceae are
highly similar in xylem structure and together are distinct from the others',
basing this opinion mainly on the occurrence of radial intercellular canals,
less advanced ray types, the lack of banded parenchyma other than terminal,
the type of ray- vessel pitting, and the common occurrence of septate fibres in
both families. Webber, on the other hand, concludes that 'the wood structure
of the Burseraceae rather than presenting evidence for grouping this family
with the Anacardiaceae instead of with the Simarubaceae, Rutaceae and
Meliaceae, is more suggestive of the probable common ancestry of all five
families*. She considers the Burseraceae and Anacardiaceae to be the least

specialized of this group and suggests that the traumatic canals of the
Simarubaceae and Rutaceae may indicate their origin from plants such as
the Burseraceae and Anacardiaceae that have normal intercellular canals in
their rays.
Janssonius (1154, vol. v, p. 464) notes a close similarity between the woods
of this family and those of a group of the Euphorbiaceae (see Thyllanthoideae',
Group B, p. 1221) that includes the genera Acalypha, Antidesma, Bischofia,
Bridelid) and Glochidion. No radial canals, such as are characteristic of the
Burseraceae, however, occur in this group.
Webber (2378) has considered the evidence of wood anatomy on the internal
specialization of the family. She has found that there is no evidence from the
wood (a) that the Asiatic species of Protium are more specialized than the
American species of this family, (b) that species having many leaflets are more
specialized than those with few, or (c) that the woods of Canarium and
Commiphora are more specialized than those of Protium. From the basic
similarity between the woods of all the genera she concludes that, in this family,
specialization in the structure of the fruits, flowers, and leaves has proceeded
at a more rapid rate than in the woods.
The wood of Canarium sumatranum does not differ significantly from that
of other species of Canarium. Heimsch (938) notes that the woods of C.
bengalense Roxb. and C. commune Linn, differ not only from other species of
Canarium but from the family as a whole in such features as the type of
parenchyma and the absence of septate fibres and suggests that they may be
wrongly placed in this family.

ECONOMIC USES
Frankincense is a gum resin obtained from the bark of Boswellia carteri
Birdw. Myrrh is a similar substance from various species of Commiphora.
A gum resin, also used as incense, is obtained from Protium heptaphyllum
(Aubl.) March. The wood of various species of Bursera is perfumed. Black
Dammar is the product of Canarium strictum Roxb.
The family includes one very important timber tree, Aucoumea klaineana
 BURSERACEAE 349
Pierre,which furnishes the Gaboon Mahogany or Okoume of commerce. The
timber is very
extensively used for veneers and plywood.
The timbers of the other genera are also light and easy to work and some
of them are used locally for packing-cases, cheap furniture and planking.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Aucoumea, Boswellia, Bursera, Canariellum, Canarium, Commiphora,
Crepidospermum, Dacryodes, Garuga, Hedwigia, Pachylobus, Protium,
Santiria, Scutinanthe, Tetragastris, Trattinickia, Triomma.

(ii) FOR WOOD STRUCTURE

Aucoumea, Balsamodendron, Boswellia, Bursera, (Canariellum), Canarium,
Commiphora, (Crepidospermum), Dacryodes, Elaphrium, Garuga, Hemi-
santiria, Pachylobus, Protium, Santiria, (Scutinanthe), Tetragastris, Tratti-
nickia, Trigonochlamys, and Triomma.

LITERATURE
(i) On
General Anatomy
Engler 643, Guillaumin 838, 839, 840, Sabnis J977, Solereder 2163, Spiekerkoetter
2168.

(ii) On Wood Structure

Beekman 167, den Berger 179, 182, Besson 186, Br. Hond. F.D. 274, Chevalier et al.
312, Collardet 448, Cooper 461, Dadswell 525, Desch 574, Dixon 592, Gonggrijp 794,
Heimsch 938, Howard 1088, Janssonius 1154, Jentsch 1172, Kanehira 1207, 1208, Kribs
1283, Lecomte 1332, Martm-Lavigne 1450, M6niaud 1491, Pearson and Brown i679,Pfeiffer
J.Ph. 1713, Record 1780, 1783, 1787, 1801, 1843, 1851, Record and Hess 1886, Record
and Mell 1894, Riera 1937, Spiekerkoetter 2168, Stone 2202, 2207, Torres 2269, Webber
2377, 2378, Williams 2430.

87. MELIACEAE
(FiG. 77 on p. 334; FIG. 80 on p. 350; FIG. 81 on p. 354)

SUMMARY
(i) GENERAL
A
family from tropical and warm temperate regions consisting almost
exclusively of trees or shrubs. The hairs are of various kinds, (i) Unicellular
or uniseriate. (ii) Two-armed, (iii) Stellate, (iv) Peltate, (v) Glandular hairs
of various types. The leaf is always dorsiventral, and the lamina is particu-
larly characterized by the presence of variously shaped secretory cells con-
taining resin, which are usually situated at the boundary between the palisade
and spongy mesophyll. Secretory cells are sometimes present in the cortex
and pith of the axis as well, while secretory cavities occur in the same
positions in a few 'genera. Stomata in the leaf have been recorded only on
the lower surface they are ranunculaceous. Hypoderm, which is confined
;

to the upper side of the leaf, has been observed only in Carapa sp. and in
Entandrophragma. The mesophyll of a few genera includes 'spicular cells'
and crystal idioblasts. The midrib contains variously orientated, isolated,
collateral bundles in Khaya, Melia, Trichilia and Turraea, but a closed ring
y
 350 ME LI ACE AE
of xylem is The petiole of species with simple
present in Entandrophragma.
an arc of bundles or a single arc-shaped
leaves, in transverse sections, exhibits
vascular strand: in pinnate leaves there is a closed vascular ring. Medullary
bundles have been recorded in the petiole in a few genera. In the axis the
cork originates in the sub-epidermis; the cortex contains stone cells of
various types (sometimes present in the pith as well); the pericycle is
characterized by separate crescentic strands of fibres, which are, however, very
close together in some species. The xylem and phloem in young stems

FIG. 80. MELIACEAE, A; RUTACEAE, B; DICHAPETALACEAE, C

A, Lower epidermis of the leaf of Cedrela toona Roxb. with a glandular hair. B Lower epidermis
of the leaf of Flindersia maculosa (Lindl.) Benth. with a peltate and a glandular hair. C, Lower epidermis
of the leaf of Tapura amazonica Poepp, et Endl. By Solereder,

constitute a closed cylinder traversed by narrow rays. The secondary phloem

contains fibres, either scattered, in small groups, or in bands. Crystals, both
solitary and clustered, are common in the tissues of all of the organs.

(ii) WOOD
Vessels moderately small to medium-sized; radial multiples of 2 or 3 cells
common; perforations exclusively simple; intervascular pitting typically
minute, but occasionally larger; pits to ray cells similar; members of medium
length. Parenchyma most characteristically paratracheal (vasicentric, aliform,
or confluent) and often with terminal in addition diffuse parenchyma some-
;

times present and often consisting of crystalliferous cells; broad, moderately

regular and apparently apotracheal bands characteristic of some species;
sometimes storied. Rays usually exclusively uniseriate or 2-4 cells wide, but
considerably wider (up to 9cells) in some species uniseriate rays usually rare
;

in woods with multiseriate rays; mostly heterogeneous, with 1-2 marginal

rows, but homogeneous in some genera; often distinctly storied or in echelon.
Fibres with simple or narrowly bordered pits, septate in a majority of species ;

of medium length to moderately short. Intercellular canals of the traumatic

vertical type occur sporadically.

LEAF
Dorsiventral. Hairs include the following kinds, (i) Simple, unicellular in
Aitonia, Chisocheton, Dysoxylum, Ptaeroxylon, TrichiUa, Turraea. (ii) Uni-
seriate in Cedrela and Melia. (iii) Two-armed hairs in certain species of
Amoora, Dysoxylum, Epicharis, and occasionally in other genera as well.
 ME LI ACE AE 351

(iv) Stellate hairs in Aglaia, Melia, Pterorhachis, Trichilia. (v) Peltate scales,
sunk in small pits in the strongly cuticularized leaf surface, in Aglaia, Amoora.
(vi) Glandular hairs, always multicellular but variously shaped, in Cabralea,
Carapa, Cedrela (Fig. 80 A), Entandrophragma, Melia, Ptaeroxylon, Turraea.
Cells of the epidermis mucilaginous in Ghukrasia sp. and in the Cedreleae;
papillose in certain species of Ekebergia, Heynea, Walsura. Stomata confined
to the lower surface ranunculaceous in certain species of Aitonia, Carapa,
;

Cedrela, Khaya, Melia, Ptaeroxylon, Swietenia, Trichilia, Turraea. Hypoderm

below the upper epidermis recorded only in species of Aglaia, Carapa,
Entandrophragma, Sandoricum. Mesophyll. Palisade tissue i-layered in
species of Melia and Turraea more than i -layered in Carapa moluccensis
;

Lam., Entandrophragma sp., Khaya sp., Trichilia (up to 4-layered). Scleren-

chymatous idioblasts present in the mesophyll in certain species of Dysoxylum
and in Khaya senegalensis A. Juss. Vascular bundles of the midrib exhibiting
a considerable range of structure in different genera and species; with 2
bundles in most species of Turraea, but united to form a single strand in
i species; with 2 superimposed bundles in Khaya senegalensis and Melia

volkensii Giirke, with an arc-shaped strand with incurved ends in Trichilia;

with a triangular strand with incurved ends towards the adaxial side in
Entandrophragma casimirianum De Willd. et Dur. (syn. E. candotteanum Willd. et
Dur.). Petiole with 3 bundles entering the base, the bundles becoming united
towards the distal end to form a single crescent-shaped strand with incurved
ends in most species of Turraea (Fig. 77 D). Medullary bundles occur in
certain species of Cedrela, Soymida, Trichilia, Vavaea, and Walsura; and
have been described by Spiekerkoetter (2168) in Turraea. According to
Solereder the petioles of species with simple leaves, especially in Quivisia,
Turraea, and Walsura, exhibit, in transverse sections, an arc of bundles, and
those with pinnate leaves a ring of bundles. This may in general be true but
the matter needs further investigation. A cylindrical vascular strand was noted
in Khaya senegalensis A. Juss. (Fig. 77 A). Secretory cells of various sizes and
shapes occur throughout the family. They are sometimes branched, and most
commonly to be found at the boundary between the palisade and spongy
mesophyll, although confined to the region of the bundles in Megaphyllaea,
and recorded in the centre of the midrib and in the petiole of Entandrophragma
casimirianum. Crystals, both solitary and clustered, common; their size,
nature, and distribution believed to be of value for specific diagnosis. Skutch
(2127) has recently investigated a compound leaf of Guarea which exhibits
annual increments of growth.

Axis
YOUNG STEM
Stem surface papillose in Melia volkensii Giirke. Cork invariably arising in
the sub-epidermis in all of the species examined; thickening of the cork cells
very variable in different genera and species, e.g. considerably thickened in
Trichilia and Turraea, weakly suberized in Khaya senegalensis A. Juss. Intume-
scences which arise from the phelloderm and consist of thin-walled phelloid
cells filled with fatty material occur on the stem surface in Khaya ivorensis A,
Chev. and K. senegalensis A. Juss. They have been described by Ossowski
(1643). Cortex containing stone cells. The latter solitary in certain species of
352 MELIACEAE
Amoora, Entandrophragma, Hearnia, Khaya, and Vavaea; sometimes grouped
in Cabralea, Chisocheton, Dasycoleum, Megaphyllaea, and Synoum\ branched
in Aglaia sp. ; arranged in a ring in certain species of Amoora and Sandoricum.
Slightly sclerosed parenchyma sometimes present in Guarea and Trichilia.
Resinous secretory cells occur in the cortex of Entandrophragma casimirianwn
De Willd. et Dur. (syn. E. candolleanum Willd. et Dur.), Khaya senegalensis
A. Juss,, and probably other species. Pericycle nearly always containing
isolated strands of fibres, but a composite and continuous ring recorded in
Cabralea sp. Phloem and xylexn constitute closed cylinders traversed by
narrow rays. Secondary phloem containing fibres, either scattered, in small
groups, or in bands or rings, e.g. in isolated groups in some species of Turraea,
but forming tangential bands in other members of this genus; with concentric
rings of fibres in Khaya senegalensis A. Juss. and Melia volkensii Giirke. Vessels
with simple perforations. Pith very variable in size homogeneous or hetero-
;

geneous, variations in this respect being exhibited by different members of

a single genus. Perimedullary region clearly differentiated from the remainder
of the pith in some species, but not in others. Spiekerkoetter (2168) suggests
that this character may be of specific diagnostic value. Resinous secretory
cells present in the pith of Entandrophragma casimirianum and secretory cells
in Melia azedarach Linn. Groups of stone cells or prosenchymatous elements
recorded in the pith of certain species of Cabralea, Chisocheton, Hearnia,
Megaphyllaea, and Sandoricum. Secretory cells, see 'Cortex' and Tith'.

FRUIT STALK
Fruit stalks stated by Solereder to be polystelic in Swietenia mahogani Linn.

BARK
According to Wenzel (2412) the tanniniferous bark of Carapa moluccensis
Lam. exhibits the following characters. Bark covered externally by a layer of
cork 275-400 broad cork composed of radially arranged cells of variable
fj, ;

size and form and commonly containing cluster crystals. Rays extending to the
inner boundary of the cork layer, mostly 2 cells wide but sometimes up to 4
and very rarely 5 cells wide; up to 2, but mostly i mm. tall. Ray cells twice
as long in radial as in tangential diameter. Cells of the phloem parenchyma
with thicker walls than those of the rays; vertically elongated, containing
rhomboidal crystals of calcium oxalate. Phloem parenchyma traversed by
tangential bands of sclerenchyma; bounded externally by a tissue of dis-
organized sieve tubes (Keratenchym). Calcium oxalate crystals sometimes
of unusual form, and capable of being mistaken for stone cells. For the
histology of Cocillana and substitute (Guarea) barks see 'Economic Uses'.

WOOD (Fig. 81)

Vessels usually moderately small to medium-sized (50-200 /x, mean tan-
gential diameter), very small (less than 50 ^) in Ptaeroxylon, large (more than
200 IJL) in at least some specimens of Amoora, Carapa, Dysoxylum, Entandro-
phragma, Khaya, and Lovoa, multiples of 2 or 3 cells common, with some
multiples of 4-6 cells, in Cabralea, Heynea, Owenia, Ptaeroxylon, Trichilia,
Turraea, and Walsura p.p. clusters moderately common in Heynea, Melia,
;

Synoum, Turraea, and Walsura p.p. with a tendency to oblique arrangement

;
 ME LI ACE AE 353
locally in Heynea, Turraea, and Vavaea and in distinct radial rows in Cipadessa
(938), Ptaeroxylon, and Pterorhachis (938) ; varying in number from fewer than
5 per sq. mm. in the species with the larger vessels to about 20 per mm. more
;

numerous in Ptaeroxylon', ring-porous or semi-ring-porous in some species of

Cedrela and Melia spiral thickening present
; in Melia and sometimes in the
smaller vessels of Cedrela. Perforations exclusively simple. Intervascular
pitting alternate ; minute except in Amoora, Aphanomixis, Cedrela, Ekebergia,
Melia, Pseudocarapa, and Ptaeroxylon, and sometimes with coalescent aper-
tures in these genera; pits to ray cells similar to the intervascular pitting,
Heimsch (938) states that this pitting *tends to be oblong or gash-like* in
Cedrela and Melia, but this could not be confirmed in the material examined.
Tyloses absent, deposits of gum common. Mean member length o-3~O'7 mm.
Parenchyma very variable in type and amount, terminal, diffuse, vasicentric,
aliform, confluent, and broad bands being all represented in different genera,
and even the specie^ of single genera are often distinguished by widely different
forms. Paratracheal parenchyma is always present, except in Ptaeroxylon,
and is typically predominant; even in species with regular and apparently
metatracheal bands the parenchyma in many cases becomes aliform or con-
fluent where it is locally less abundant. Terminal bands, often conspicuous
and 4 or more cells wide, present in Azadirachta, Carapa, Cedrela, Chuk-
rasia, Dysoxylum, Ekebergia, Entandrophragma, Epicharis, Heynea, Khaya
(sporadic), Lansium (938), Owenia, Pseudocedrela, Ptaeroxylon, Soymida,
Swietenia, Trichilia, Turraea, Wahura (938), and Xylocarpus', Kribs (1285)
reports terminal parenchyma also in Cipadessa and Sandoricum; diffuse
parenchyma, sometimes containing crystals, and in some genera consisting
almost entirely of scattered files of crystalliferous cells, present in Azadirachta,
Carapa p.p., Cedrela (Fig. 81 N), Chukrasia, Guarea (rare), Khaya p.p.,
Lovoa, Melia p.p., Owenia, Pseudocarapa, Pseudocedrela, Ptaeroxylon, Sando-
ricum, Soymida, Swietenia p.p., Synoum, and Xylocarpus, and reported by
Heimsch (938) in Vavaea and Walsura', parenchyma, other than terminal and
diffuse as described above (a) absent from Ptaeroxylon, (V) scanty to vasi-
centric in Chukrasia, Cipadessa, Ekebergia, Epicharis, Khaya, Melia, Owenia,
Pseudocedrela, Swietenia (Fig. 81 j), Turraea, Turraeanthus, and Xylocarpus',
vasicentric to predominantly aliform in Azadirachta, Carapa, Cedrela, Lovoa
(Fig. 81 i), Pseudocarapa, and Vavaea, (c) predominantly aliform
*
or confluent,
or in moderately regular bands in Aglaia, Guarea (Fig. 81 E), Moschoxylum* ,
and Trichilia, and (d) consistently in bands in Cabralea, Lansium, Reinwardtio-
dendron, Synoum, and Walsura (Fig. 81 A); single cells scattered among the
fibres (diffuse) in some species of Cedrela, Pseudocarapa, and Pseudocedrela.
Crystals present in chambered cells in Azadirachta, Cabralea, Chisocheton,
Chukrasia, Cipadessa, Dysoxylum, Ekebergia, Entandrophragma, Epicharis,
*
Guarea, Melia, Moschoxylum' Owenia, Ptaeroxylon, Reinwardtiodendron,
,

Sandoricum, Trichilia, Turraeanthus, Vavaea, and Xylocarpus, and in the

ordinary cells in some other species silica pre&entinAphanomixisgrandifolia Bl.
;

and Chisocheton spp. (794). Sometimes containing gum. According to Welch

(2396) the parenchyma of Dysoxylum fraseranum Benth. contains a volatile
oleo-resin that may cause 'sweating* and spoil a polished surface, particularly
in timber not properly seasoned. Usually storied in woods with storied rays
and sometimes where the rays are not storied, e.g. in Carapa. Strands
4594 Aa
 'B

FIG. 81. MELIACEAE

A, Walsura villosa Wall, B, Amoora maingayi Hiern. C, Ptaerdxylon obliquum Radlk. D, P. obliquum
Radlk. E, Guarea cedrata (A. Chev.) Pellegr. F,Chukra$ia tabularisjuss. G, Guarea cedrata (A. ChevJ
Pellegr. H, Lovoa hlaineana Pierre. I, L.^klaineana Pierre. J, Swetenia macrophylla King. K, S*
,

macrophylla King. L, Khaya ivorensis A. Chev. M, Cedrela odorata Linn. N, C. odorata Linn.
i.e. Intercellular canal.
 MELIACEAE 355

commonly with strands of 4-6 cells in some species

typically of 8 cells, but
of Ekebergia, Entandrophragma, Heynea, Melia, Trichilia, and Turraeanthus;
strands of up to 12 cells common in some specimens of Dysoxylum. Rays
exclusively uniseriate or nearly so in Aglaia p.p., Amoora p.p., Aphanomixis,
Cabralea p.p., Cipadessa, Dysoxylum p.p., Elutheria (1886), Guarea p.p.,
Lansium, Pseudocarapa, Ptaeroxylon, Pterorhachis (938), Synoum, Trichilia
p.p., and Turraea p.p. (938); the remainder mostly 2-3 cells wide, but up to
4-6 cells wide in some species of Carapa, Entandrophragma, Khaya, Lovoa,
Owenia, Turraeanthus, Walsura, and Xylocarpus and up to 7-9 cells wide in
some species of Khaya and in Melia commonly more than i mm. in height
;

in some species of Amoora, Carapa, Dysoxylum, and Khaya uniseriate rays ;

usually rare in species with multiseriate rays and commonly only i or 2 cells
high; varying from all square or upright cells to all procumbent cells; uni-
seriates more numerous in Khaya than in Entandrophragma or Swietenia',
Panshin (1650) describes the rays of Khaya and Soymida as being of 2 sizes;
mostly between 4 and 14 per mm., up to 20 per mm. in Cipadessa and
Turraea] rays, where exclusively uniseriate, varying from heterogeneous
(Kribs's Type III) to homogeneous (Kribs's Type III), often even within
i genus; typically homogeneous in species of Aphanomixis, Guarea, Ptero-
rhachis, and Trichilia (938). Multiseriate rays commonly heterogeneous
(Kribs's Type II B), with i or 2 marginal rows of square or upright cells, but
with 4 or more rows in Sandoricum and Vavaea\ species with rays 2-3 cells
wide commonly with several uniseriate marginal rows, but with only the
extreme i or 2 rows composed of square or upright cells; homogeneous
(Kribs's Types I and II) in Ekebergia p.p., Entandrophragma p.p., Guarea,
Lovoa p.p., Melia, Ptaeroxylon, Reinwardtiodendron, Turraeanthus, and Wai-
sura, and nearly homogeneous in some species of Aglaia, Azadirachta, Cedrela,
'

Ghisocheton, Dysoxylum, Moschoxylum? Soymida, Toona, and Trichilia and,

',

according to Heimsch (938), in occasional specimens of Swietenia and

Turraea', crystals common in the marginal cells of Carapa, Cedrela, Entan-
drophragma, Khaya, Owenia, Pseudocedrela, Soymida (1285), Swietenia, and
Xylocarpus Janssonius (1154) refers to occasional druses in Cedrela; cells
\

often containing gum; silica present in Aphanomixis grandifolia Bl. and

Chisocheton spp. (794). Storied in most specimens of Pseudocedrela, Ptaero-
xylon, Swietenia, and Xylocarpus and in some species of Carapa, e.g. C.
moluccenns Lam,, and Entandrophragma, e.g. E. cylindricum Sprague, and
occasionally, according to Record (1851), in Cedrela, Chukrasia, and Khaya;
in echelon in some other genera. Fibres with pits usually confined almost
entirely to the radial walls, simple or with very narrow borders.
1
Septate in
most of the genera but non-septate in Cedrela, Chukrasia, Ekebergia, Heynea,
Lovoa, Melia, Odontandra (938), Owenia, Pterorhachis (938), Quivisia (1285),
Sandoricum, Soymida, Trichilia, Turraea, Turraeanthus, and Walsura] some-

1
There appears to be some difference of opinion as to the nature of the pitting in the fibres
in this family. Kribs (1285) uses the presence of simple pits as one of the characters by which
he distinguishes the genera of his proposed sub-families Swietenioideae and Lovoinoideae,
i.e. Carapa, Cedrela, Chukrasia, Entandrophragma, Khaya, Lovoa, Pseudocedrela, and
Soymida. Janssonius (i 154) describes the pits as simple in Carapa and Walsura and bordered
in Aglaia, Amoora, Cedrela, Chisocheton, Dysoxylum, and Melia. Panshin (1650) describes
the pits of Chukrasia as bordered and those of Khaya and Entandrophragma as bordered in
some species. Many other discrepancies in the literature might be cited.
356 MELIACEAE
times with a few septate fibres in Cedrela and Chukrasia (1285). Sometimes
containing gum. Walls usually thin to moderately thick, but thick in Aglaia,
Lansium, Owenia, Pseudocarapa, and Soymida. Mean length 0*6-1-9 mm '

Intercellular canals of the vertical, traumatic type have been reported in

Carapa, Cedrela, Dysoxylum, Entandrophragma, Khaya, Lovoa, Melia, San-
doricum, and Swietenia. Groom (826) states that in Khaya, Lovoa, and
Smetenta small canals are usually lysigenous, but those in well-developed
bands are schizo-lysigenous or schizogenous and that the contents give
reactions typical of wound gum. Growth ring development has been studied
by Coster (481) in Azadirachta, Melia, Swietenia, and Toona, by Chowdhury
(415) in Cedrela, and by Hummel (1109) in Entandrophragma and Khaya.
Besson (186) shows a relatively high percentage of silica for Entandrophragma
candollei Harms.
ROOT
The erect, peg-like pneumatophores of Amoora cuculata Roxb. and
Carapa moluccensis Lam. have been investigated by Groom (826) and Liebau
(1368). They arise from horizontal roots as horn-like wings produced by the
activity of the cambium, and continue to grow in height and thickness by
means of a cambium. Forked pneumatophores sometimes arise by the fusion
of two situated side by side. Lenticels are always present. The xylem of
the pneumatophores consists of vessels, fibres, and parenchyma arranged in
periclinal arches, but the structure becomes more complex with increase in
age. The scanty vessels are partly blocked by callus-like plugs in Amoora.
Groundwork of the wood composed of thin-walled fibres and parenchyma, or
of parenchyma alone.
Gallager (736) has recorded a few facts about the roots of twenty-two
species of Meliaceae, without including any features of great taxonomic
interest beyond the occurrence of secretory cells in all of the species examined
except in Cedrela, but even here their absence was not definitely affirmed.

TAXONOMIC NOTES
(i)
FROM GENERAL ANATOMY
The family is anatomically
similar to and undoubtedly has affinities with
the Rutaceae and Burseraceae. The boundaries between these families are,
in fact, by no means well defined, and taxonomists have not always agreed in
which of them some of the genera should be placed. Chloroxylon, for example,
has been variously included in the Rutaceae and Meliaceae. The anatomical
features are not conclusively more in favour of one of these views than the
other. Aitonia and Ptaeroxylon also have a somewhat indefinite status since
the Sapindaceae by some authors and in the
they have been placed in
Meliaceae by others. Flindersia is yet another genus of uncertain position,
which in this book has been described under Rutaceae,

(ii) FROM WOOD STRUCTURE

Kribs (1285) suggests that Carapa, Cedrela, and Xylocarpus should be
placed in the sub-family Swietenioideae and that Lovoa should be set aside by
itself in a sub-family Lovoinoideae. Panshin (1650) also comments on the
different structure of Lovoa. Kribs considers that the Swietenioideae (as thus
 M ELI ACE A E 357
re-defined by himself) is the only
sub-family in which the genera form a
distinct homogeneous group in respect to anatomical and morphological
characters and suggests that it should be raised to the rank of a family.
Kribs notes that of the four genera with simple leaves, Reinwardtiodendron>
Turraea, Vavaea, and Quivisia, the wood anatomy of the three latter is closely
similar and is relatively more primitive than that of the other genera, but that
the wood of Reinwardtiodendron closely resembles that of Lansium.
Commenting on the confusion existing over the identity of certain genera
of the Melioideae, he lists characters of the wood by which Aphanomixis can
be separated from Amoora, Azadirachta integrifolia Merr. from Melia, and
Carapa from Xylocarpus.
Kribs considers that, on the basis of the anatomy of their woods, the genera
Chloroxylon, Flindersia, and Ptaeroxylon should be placed in the Rutaceae.
Panshin finds that the wood anatomy of Entandrophragma spp. bears out
the separation of the four sub-genera established by Harms.
The woods of Cedrela and Toona are indistinguishable. Janssonius (1154)
draws attention to the marked difference between the woods of Cedrela and
Melia and those of the other genera and suggests that these two genera should
be placed in a separate family. Heimsch (938) considers that the Meliaceae,
Rutaceae, Simarubaceae, Sapindaceae, Burseraceae, and Anacardiaceae con-
stitute a more or less natural group of plants.

ECONOMIC USES
This family is possibly of greater importance as a source of hardwood
timber than any other. It includes several woods that are well known all over
the world, such as Mahogany (Swietenia), African Mahogany (Khaya),
Sapele Mahogany (Entandrophragma), Spanish Cedar (Cedrela), and African
Walnut (Lovoa), and several others that, though less important, are well
known to commerce outside the countries where they are grown, e.g. Bosse
(Trichilia), Rose Mahogany (Dysoxylum), various Cedars (Cedrela and Toona),
and Crabwood or Andiroba (Carapa). Many others are of importance locally.
The best-known timbers are typically red in colour, lustrous, and easy to
work; they are often highly figured and some, e.g. the True and African
Mahoganies, are unusually free from distortion under changing conditions of
moisture. Such characters make these woods eminently suitable for cabinet
work. Some
of the woods make excellent constructional timber, e.g. the
Entandrophragmas, and are often used locally for such purposes. The bark of
Carapa moluccemis Larn. yields tannin, and that of Guarea rusbyi Rusby,
known to pharmacognoscists as Cocillana, possesses medicinal properties
resembling those of Ipecacuanha. The microscopical characters of Cocillana
and related barks have been described by Ballard (115). The tanniniferous
bark of Carapa moluccemis is described above on p. 352.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Aglaia, Aitonia, Amoora, Cabralea, Carapa, Cedrela, Chisocheton, Chloro-
xylon, Chukrasia, Dasycoleum, Dysoxylum, Ekebergia, Entandrophragma,
Epicharis, Guarea, Hearnia, Khaya,* Megaphyllaea, Melia,* Ptaeroxylon,
358 MELIACEAE
Pterorhachis, Quivisia, Sandoricum, Soymida, Swietenia,* Synoum, Trichilia,*
Turraea,* Vavaea, Walsura.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Aglaia, Amoora, Aphanomixis, Azadirachta, Cabralea, Carapa, Cedrela,
Chisocheton, Chukrasia, (Cipadessa), Dysoxylum, Ekebergia, (Elutheria),
Entandrophragma, Epicharis, Guarea, Heynea, Khaya, Lansium, Lovoa,
Melia, 'Moschoxylum', Owenia, Pseudocarapa, Pseudocedrela, Ptaeroxylon,
(Pterorhachis), (Quivisia), Reinwardtiodendron, Sandoricum, Soymida,
Swietenia, Synoum, Trichilia, Turraea, Turraeanthus, Vavaea, Walsura, and
Xylocarpus.
LITERATURE
(i) On General Anatomy
Ballard 115, Gallager 736, Groom 826, Harms 904, Kienholz 1236, Ledoux 1336, *337
1338, Liebau 1368, Ossowski 1643, Skutch 2127, Spiekerkoetter 2168, Wenzel 2412.

(ii) On Wood Structure

AubrSville 49, 50, 51, 53, 54, Baker 104, Becking 164, Beekman 167, Benoist 170,
den Berger 179, 182, Besson 186, Boulger 244, B.H. For. D. 274, Burgerstein 310, 312,
Busch 322, Chalk et al. 360, 364, Chattaway 368, Chowdhury 411, 415, Collardet 449,
450, Cooper and Record 461, Coster 481, Dadswell 525, 530, Dixon 592, Duchesne 612 A,
Eggeling 624, Esmans 662, Fanshawe 2519, Fiebrig-Gertz 685 >r poxworthy 705, Garratt
744, Gleason 788 A, Gonggrijp 794, Groom 826, Hale 870, 'Harrar 906, He'din 931,
Heimsch 938, Howard 1088, Hummel 1109, Janssonius 1154, Jayawardana 1159, Jolly
1x88, Jones 1191, Kanehira 1206, 1209, Keen 1229, Koehler 1261, 1262, 1263, Kribs
1283, 1285, Lecomte 1333, I334 Ledoux 1337, 1338, Louis 1393, Martin- Levigne I45i
Mell 1481$, Meniaud 1491, Meyer 1506, Normand 1608, Panshin 1649, 1650, Pearson and
Brown 1679, Pereira 1687, Pfeiffer J. Ph. 1713, Piccioli 1716, Record 1776, 1780, 1781,
1782, 1783, 1784, 1787, x8oi, 1809, 1818, 1825, 1839, 1843, I ^S I > 1080, Record and
Hess 1886, 1891, Record and Mell 1894, Rendle 1918, Riera 1937, Scott 2075, 2076,
Sim 2099, Spiekerkoetter 2168, Staner 2183, Stone 2202, 2203, 2206, 2207, Sudworth
2219, Swain 2224, Tang 2231, Torres 2269, Watt 2370, Welch 2396, Williams 2428,
2430, Yamabayashi 2478.

88. DICHAPETALACEAE
(Fic. 80 on p. 350; FIG. 82 on p. 360)

(Description of the leaf and young stem based mainly on those given
respectively by Solereder and Engler and Krause (645).)

SUMMARY
(i) GENERAL
A
tropical family of small trees and shrubs. Particularly noteworthy
features include: the presence of unicellular hairs with conical or wart-
shaped papillae on the surface; rubiaceous stomata confined to the lower
surface of the leaf; the frequent occurrence of mucilaginous cells in the
epidermis, hypodermis, and sometimes in the ground tissue of the petiole and
branch.

(ii) WOOD
Vessels small, perforations simple or simple and scalariform, intervascular
pitting usually alternate and minute, pits to ray cells similar, members of
 DICHAPETALACEAE 359
medium length to moderately long. Parenchyma predominantly para-
tracheal, vasicentric to aliform, accompanied by varying amounts of diffuse
parenchyma. Rays up to 5-6, occasionally 8-10, cells wide, markedly hetero-
geneous and often with sheath cells. Fibres with numerous, very small,
bordered pits; of medium length.

LEAF
Usually dorsiventral. Simple, unicellular hairs with conical or wart-like
papillae recorded in species of Dichapetalum and Tapura (Fig. 80 c).
Epidermis consisting of i or several layers of cells. Hypoderm also recorded
in species of Dichapetalum, Stephanopodium, Tapura. Stomata confined to
the lower surface; rubiaceous in species of Dichapetalum and Tapura
(Fig. 80 c). Mesophyll often partly composed of short palisade cells; spongy
tissue absent from Dichapetalum cymosum (Hook.) EngL Vascular bundles of
the smaller veins accompanied by sclerenchyma in species of Dichapetalum
and Tapura^ branches of the sclerenchymatous elements sometimes extending
into the mesophyll. Mucilage cells recorded in the epidermis, hypodermis,
and sometimes in the ground tissue of the petiole, especially in certain species
of Stephanopodium and Tapura. Secretory cells with brown contents also
reported to occur in the mesophyll.

Axis
YOUNG STEM
Cork arising in the sub~epidermis. Phelloderm frequently sclerotic.
Primary cortex often containing stone cells; mucilaginous cells recorded in
the same region and in the ground tissue generally in species of Stephano-
podium and Tapura. Pericycle including isolated groups of fibres. Secondary
phloem containing sclerenchyma in certain species of Stephanopodium.
Xylem including narrow vessels usually with simple perforations but occa-
sionally (Tapura guianensis Aubl.) with scalariform perforation plates; fibres
with bordered pits; rays of z distinct sizes, the smaller 1-4 and the larger up
to 10 cells wide.

WOOD (Fig. 82 A-D)

Vessels usually moderately small (50-100 /z mean tangential diameter) but
sometimes larger; solitary and ir_ multiples of 2 or 3 cells and sometimes,
according to Heimsch (938), in clusters and pore chains in Gonypetalum and
Tapura\ about 40 per sq. mm. in the material examined. Perforation plates
simple in Dichapetalum, simple and scalariform, with up to 10 or more bars,
in most species of Tapura (simple in the other species). Intervascular pitting
typically alternate and minute and pits to ray cells similar, but Heimsch notes
scalariform and transitional intervascular pitting in Tapura obovata Britton
et P. Wils. and Record and Hess (1886) refer to occasional unilaterally com-

pound pitting to ray cells. Tyloses reported by Heimsch in one species of

Dichapetalum. Mean length 0-6-0-9 mm. Parenchyma typically pre-
dominantly paratracheal, about the vessels and as short, irregular wings, but
always with some diffuse parenchyma in addition. Heimsch (938) notes more
abundant apotracheal parenchyma (scattered cells to narrow bands) in Tapura
cubensis (Poepp. et End!,) Griseb. A specimen of Tapura guianensis Aubl is
 FIG. 82. DICHAPETALACEAE, A-D; OLACAGEAE, E-N
A, Chailletia gelonimdes Hook. f. B, Chailletia gelonioides Hook. f. C, Dtchapetalum sp. D, Dichape-
talum sp. E, Strombosia javanica Bl. F, Schoepfia schreberi Gmel. G, Ximenia americana Linn. H,
Liriosma gracilis A. C. Smith. I, Aptandra zenkeri Engl. J, Ongokea klaineana Pierre. K, Strombona
javanica Bl. L, Ochanostachys amentacea Mast. M, Olax subscorpioides Oliv. N, Liriosma gracilis
A. C. Smith.
 DICHAPETALACEAE 361
described by Williams as having numerous fine metatracheal bands, but
differs in this respect and some others from material of this species examined

by the author. Strands commonly of up to 8 cells. Rays usually up to 5 or 6

cells wide, sometimes up to 8-10 cells wide in Dichapetalum (938); more than
irnm. high in some species; uniseriates numerous and composed entirely of
square and upright cells; about n
rays per mm.; heterogeneous (Kribs's
Types I and II A), with 2 or 3 to several marginal rows of square and upright
cells; sheath cells usually present. Solitary crystals often present in both
upright and procumbent cells. Fibres with moderately to very numerous
small pits on both radial and tangential walls, the pits with small borders;
Heimsch (938) describes these cells as fibre-tracheids and tracheids. Walls
moderately to very thick, occasionally gelatinous. Mean length about 1:5 mm.
The wood of a single specimen of Chailletia gelonoides Hook. f. differs in
many respects from the rest of the family, particularly in having numerous,
bi-seriate apotracheal bands of parenchyma, shorter vessel members (0*35 /x),
the smaller rays and parenchyma storied, and numerous fusiform parenchyma
cells.

TAXONOMIC NOTES
The affinities of the Dichapetalaceae are not well established, and the few
anatomical facts which have been recorded about the leaf and young stem are
insufficient to be of much assistance in determining its taxonomic position.
On the basis of wood structure, Tippo (2261) places the family at about the
same level of anatomical specialization as or slightly higher than the Cuno-
niaceae and Brunelliaceae. The material examined by the author, however,
appears to be at a distinctly higher level than many of the genera in the
Cunoniaceae.
Heimsch (938) suggests that the linking of this family with the Mal-
pighiaceae, Vochysiaceae, Tremandraceae, Polygalaceae, and Trigoniaceae,
as in the systems of Hallier and Engler and Prantl, is preferable to linking it
with either the Euphorbiaceae or Rosaceae as suggested by Wcttstein and
Hutchinson respectively.
ECONOMIC USES
The poisonous seeds of Dichapetalum toxicarium Baill. are used in West
Africa for destroying rats.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
(Chailletia), Dichapetalum, Stephanopodium, Tapura.

(ii) FOR WOOD STRUCTURE

Chailletia, Dichapetalum, (Gonypetalum), Tapura.

LITERATURE
(i) On General Anatomy
Engler and Krause 645.

(ii) On Wood Structure

Heimsch 938, Record 1810, 1843, 1851, Record and Hess 1886, Tippo 2261, Tupper
2295, Williams 2430.
 (362)

89. OLACACEAE
(As understood by Sleumer (2129))
(Fie. 82 on p. 360)

SUMMARY
(i) GENERAL
A
tropical and sub-tropical family of trees, shrubs, and climbers. Anatomical
characters which are common to the whole family appear to be lacking, except
in so far as solitary and/or clustered crystals are generally present. The more
interesting features include the following, (i) The presence of partially or
wholly silicified cells, solitary or in groups, which occur in the mesophyll of
certain genera (see list under 'Leaf* below), (ii) Schizogenous secretory
cavities with resinous contents in the leaf and in the cortex of the axis in the
Couleae (but not in other genera), (iii) Laticiferous tubes, which are usually
branched and non -septate, in the primary cortex, phloem, pith, and sometimes
in the leaf as well in the Couleae and Heisteria. The laticiferous tubes are,
however, not accompanied by resinous cavities in Heisteria, although, as
mentioned above, these occur in the Couleae. (iv) Spicular fibres (sometimes
described as idioblasts) are to be found in the mesophyll of a number of
genera.

(ii) WOOD
Vessels typically small, occasionally medium-sized exclusively solitary or
;

with numerous radial multiples with a tendency to a radial or oblique pattern.

;

Perforations exclusively scalariform or simple, with perforations to ray cells

in many genera, intervascular pitting scalariform, opposite or alternate, the
latter often very small; always with some large simple pits to ray and paren-
chyma cells; members very long in the genera with scalariform perforation
plates, others of medium length to moderately short. Parenchyma typically
apotracheal, diffuse to uniseriate, metatracheal bands in Schoepfia, exclusively
;

of paratracheal, storied fusiform cells. Rays 1-4 cells wide and usually
1*5-4 mm
high. The genera fall into two groups, (i) with 1-4 marginal rows
*

of upright cells and a continuous multiseriate part composed of almost square

cells that arerounded in tangential section and with intercellular spaces, and
(2) with 10 or more marginal rows of upright cells, the middle part of the rays
composed of alternating layers of upright and truly procumbent cells, the
latter angular in tangential section and without intercellular spaces. Fibres
with few or numerous distinctly bordered pits, rarely septate, moderately to
very long in woods with scalariform perforation plates, of medium length in
the others. Vasicentric tracheids present in Olax.

LEAF
Usually dorsiventral, but centric structure recorded in Olax stricta R. Br.
and Ximenia coriacea Engl. Hairs mostly simple, but dendritic types present
in the Couleae. Lower epidermis papillose in Liriosma and many species of
Olax. Stomata confined to the lower surface in Minquartia and Ochano-
stachySy present on both surfaces in certain species of Olax and in Ximenia;
rubiaceous stomata recorded in Coula and in one species of Olax, but not in
 OLACACEAE 363
other members of the last genus. Hypoderm below the upper epidermis
recorded in Cathedra and in certain species of Schoepfia. Partly or wholly
silicified cells, solitary or in groups, occur in the mesophyll of Cathedra,

Olax, Schoepfia, and Ximenia.

Spicular cells (also described as sclerenchymatous fibres or idioblasts)
occur in the mesophyll of Eganthus, Endusa, Heisteria, Minquartia, Ochano-
stachys, and Scorodocarpus. Terminal tracheids of the veins strongly
developed in Phkbocalymna, Schoepfia, and Ximenia. Three bundles enter
the base of the leaf in Coula, Minquartia, Ochanostachys, Olax, Schoepfia, and
Strombosia, but transverse sections through the distal end of the petiole
exhibit a solitary vascular strand in certain species of Heisteria, Ochanostachys,
Scorodocarpus and Strombosia, or an arc of separate bundles in species of
,

Liriosma, Olax, and Ximenia. Schizogenous secretory cavities (see also

'Axis') with resinous contents (becoming blue when treated with eau de
javelle) in the mesophyll of Coula, Eganthus, Endusa, Minquartia, and

Ochanostachys, i.e. in all genera of the Couleae. Mesophyll cells sometimes

containing drops of resin in Cathedra, Schoepfia, and Ximenia. Laticiferous
tubes (see also 'Axis'), stated to be branched and non-septate, accompany the
veins of Heisteria, Minquartia, and Ochanostachys. Similar elements lie free
in the mesophyll of Heisteria, being recognizable with a lens as a network of
transparent lines in the leaf. Similar but articulated tubes accompany the
veins and also lie freely in the mesophyll of Endusa.

Axis
YOUNG STEM
Cork arising in the sub-epidermis, mostly composed of thin-walled cells,
but intermixed with some having strongly thickened tangential walls in Coula,
Minquartia, Ochanostachys. Pericycle containing a ring of sclerenchyma,
usually composite and continuous, in Heisteria, Minquartia, Ochanostachys,
certain species of Scorodocarpus, and in Strombosia; with bands or isolated
strands of fibres in Liriosma, Olax, Schoepfia, and Ximenia. Secondary
phloem partly sclerenchymatous in Liriosma. Vessels with simple and/or
scalariform perforations. Schizogenous secretory cavities with resinous
contents, similar to those described above under 'Leaf, occur in the cortex
of Coula, Minquartia, and Ochanostachys. Branched, non-septate laticiferous
tubes, present in the primary cortex, phloem, and pith of Coula, Eganthus,
Heisteria, Minquartia, and Ochanostachys.

WOOD (Fig. 82 E-N)

Vessels typically small (less than 100 ^ mean tangential diameter), medium-
sized (100-200 IJL) in at least some species of Anacolosa, Chaunochiton, Coula,
Ongokea, and Scorodocarpus, very small (25-50 p) in Aptandra, Heisteria,
Liriosma, Ptychopetalum, and Ximenia p.p.; almost exclusively solitary in
Agonandra (1886), Heisteria, Liriosma, Ongokea, Phlebocafymna, and some-
times in Ximenia, radial groups and multiples abundant in the other genera
except Liriosma, and many species with a radial or oblique pattern, due to the
radial multiples or to the grouping of solitary vessels and multiples, the pattern

very distinct in Olax (Fig. 82 M) and rather vague in Anacolosa, Aptan-

dra, Coula, Endusa, Minquartia, Ochanostachys p.p., Ptychopetalum, and
364 OLACACEAE
Strombosia; sometimes with a tendency to an oblique tangential pattern in
Schoepfia; very variable in number in different genera, fewest (less than 5 per
sq. mm.) in Aptandra^ Chaunochiton^ and Ongokea, most numerous (40-60 per
sq. mm.) in Heisteria and Ximenia; diffuse-porous. Perforation plates scalari-
form in Coula, Endusa, Heisteria, Minquartia, Ochanostachys, Scorodocarpus,
Strombosia, and Strombosiopsis, usually with moderately large perforations and
few (6-1 o) bars, but with more than 20 bars in Heisteria. Chalk and Chatta-
way (358) have drawn attention to the common occurrence in this family of
perforations between vessels and ray cells, whereby a vessel may pass through
a ray and emerge on the other side, particularly in Ptychopetalum anceps Oliv.
and Strombosia javanica Blume. It has been observed that, in addition to the
type of perforated ray cells referred to in the above publication, the ray and
parenchyma cells of Chaunochiton, Ongokea, and Schoepfia, where they adjoin
vessels, often have what appear to be small round perforations in their walls;
there is, however, no evidence that the vessels pass right through these rays,
a fact that may possibly be explained by the different character of the rays
in these genera. Solereder records a similar feature in Cathedra rubricaulis
Miers. In Ongokea the contents of a ray cell can sometimes be observed
protruding into the vessel through such a 'perforation', having the appearance
of a minute tylosis. Intervascular pitting scalariform or opposite in Heisteria,
Strombosia, and Strombosiopsis, opposite in Coula and Scorodocarpus, alternate
in the remainder; pits small in Anacolosa, Olax, Ongokea, Phlebocalymna,
Ptychopetalum, Schoepfia, and Ximenia', some of the pits to rays or parenchyma
cells large, often simple, more rarely unilaterally compound, sometimes

numerous, but rather rare in Ongokea and Schoepfia. Tyloses present in some

species of Minquartia, Ochanostachys, Scorodocarpus, Strombosia, Strombo-

siopstSy and Ximenia, sometimes sclerosed in Strombosiopsis tetrandra Engl. ;

sometimes with gum-like deposits. Mean member length about 1*3-1*5 mm.
in the genera with scalariform perforation plates, e.g. Coula, Heisteria,
Ochanostachys, and Strombosia; about 0-3-0-5 in the genera with simple
perforations, e.g. Aptandra, Olax, and Ongokea. Parenchyma typically
apotracheal, varying from scattered cells in Strombosia (Fig. 82 E) and Ximenia
to numerous, irregular, uniseriate bands forming a reticulate pattern with the
bands more regular and biseriate in Olax (Fig. 82 M); with a few
rays, the
round the vessels (paratracheal) in addition to diffuse parenchyma in
cells

Ongokea (Fig. 82)) and, according to Janssonius (1154), in Anacolosa.

Chambered crystalliferous cells common in Anacolosa, Aptandra, Chaunochi-
ton, Coula, Endusa, Liriosma, Minquartia, Ochanostachys, Olax, Ongokea, and
Ptychopetalum', gum-like deposits present in a few species. Strands usually
of 8 or more cells and sometimes up to 16, e.g. in Strombosia, but commonly
of only 4 cells in Aptandra, Chaunochiton, Liriosma, Olax, Ongokea, and
Ximenia. In Schoepfia the parenchyma is aliform and confluent, consists
almost entirely of fusiform cells and is storied* Rays. The genera, other than
Schoepfia, can be divided into two groups according to their ray characters,
(i) Aptandra, Chaunochiton, Liriosma (Fig. 82 N), Olax, Ongokea, and
Phlebocalymna: rays usually 2-3 cells wide, but up to 9 cells in Phlebocalymna
and almost exclusively uniseriate in Aptandra (Fig. 82 i); 8-14 per mm. cells
;

large, rounded in tangential section, and with intercellular spaces; hetero-

geneous (Kribs's Type II A), with 1-4 marginal rows of uniseriate upright
 OLACACEAE 365
cells, the multiseriate portions forming the bulk of the ray and composed of
cells that are almost square in radial section and not very distinct from the
upright cells in tangential section; uniseriates moderately numerous, com-
posed of cells similar to those of the multiseriate rays; seldom more than
0*6 mm.
in height in Ongokea, up to 2-3 mm. in the other genera; crystals
abundant in chambered cells in Ongokea, not observed in the other genera.
(2) Anacolosa, Coula, Endusa, Minquartia, Ochanostachys (Fig. 82 L), Scoro-
docarpus, Strombosia (Fig. 82 K), Strombosiopsis, and Ximenia: rays 2-4 cells
wide, uniseriate in Anacolosa, Ptychopetalum, and Strombosia javanica BL;
12-20 per mm.; cells angular in tangential section and without intercellular
spaces; markedly heterogeneous (Kribs's Type I), except in Ximenia
(Type with 10 or many more marginal rows of uniseriate upright cells,
II B),
which are very high axially and narrow tangentially and radially, and are very
distinct on tangential sections from the procumbent cells; the latter truly

procumbent and occurring commonly in small groups alternating with rows

of upright cells; the distribution and shape of the upright cells are sometimes
very suggestive of tile cells. Uniseriates numerous, composed of square to
upright cells. Crystals in chambered or ordinary cells present in most of the
genera, abundant in Endusa, and gum-like deposits not uncommon seldom ;

more than 0-6 mm. in height in Ximenia, up to 2-5 mm, in the other genera.
The rays of Heisteria and Ptychopetalum (uniseriate) are intermediate in
character, the inner cells of the rays being comparable to the almost square
multiseriate cells of group (i), but with many marginal rows of upright cells
comparable to those of group (2). In Schoepfia (Fig. 82 F) the rays are up to
4 cells wide, short (seldom more than 15 cells and 250 p high), with numerous
rays only 1-3 cells high, 8~io per mm., homogeneous (Kribs's Type I).
Fibres with and numerous bordered pits in Heisteria, Liriosma,
distinct

Ongokea, Phlebocalymna, and Ximenia, the other genera with rather few,
simple pits mainly limited to the walls in contact with ray or parenchyma
cells, with funnel-shaped canals and slit-like inner and rounded outer aper-
tures. With occasional septa in Strombosia. Walls usually very thick and
often showing distinct zones. Vascular tracheids present in Olax. Mean
length about i -9-2-5 mm. in woods with scalariform perforation plates, e.g.
Coula, Heisteria, Ochanostachys, and Strombosia', 1-0-1-8 mm. in woods with
simple perforations, e.g. Aptandra, Olax, and Ongokea.

ROOT
Particulars concerning the anatomy of Liriosma root have been published
by Youngken (2493).

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The Olacaceae, in combination with the Icacinaceae, constituted the
Olacineae as understood in the system of Bentham and Hooker. The Olacaceae
are here treated as a distinct family as understood by Sleumer (2129). The
anatomical evidence at present available is not sufficiently comprehensive to
give any clear indication of how closely related the Olacaceae and Icacinaceae
may be, but this subject is discussed more fully below under Icacinaceae.
366 OLACACEAE
(ii) FROM WOOD STRUCTURE
The sub-family Schoepfioideae(with only one genus, Schoepfia) differs
very considerably from the rest of the family, e.g. paratracheal parenchyma
composed of fusiform cells, and short homogeneous rays, such differences
being more marked and more numerous than the differences between many
families. It has, however, in common with some of the other genera, the
curious 'perforations' between the vessels and ray or wood parenchyma cells.
It is more highly specialized than the other genera of the family and some of
the differences may be due to this.
The family, evenif Schoepfia is excluded, is
by no means uniform in struc-
ture, but though separate groups of genera can be distinguished by single
characters, such as radial grouping of vessels, solitary vessels, fibre-tracheids,
&c., these groups overlap and have more points of resemblance to than differ-
ences from the remaining genera. The family includes genera at very different
levels of specialization and specialization may have taken place very unevenly
in different elements, e.g. advanced vessel pitting may be accompanied by a
primitive type of ray.
'Ray Type i* includes all the genera examined of Sleumer's Olacoideae,
i.e.Aptandra, Liriosma, Olax, and Ongokea, except for Ptychopetalum, which
has uniseriate rays of intermediate type, but this group also includes Chauno-
chiton of the Discalacoideae. *Ray Type 2', on the other hand, includes all the
genera examined of the Discalacoideae, i.e. Anacolosa, Coula, Endusa, Min-
quartia, Ochanostachys, Scorodocarpus, Strombosia, Strombosiopsis, and
Ximenia, except for Heisteria, which is intermediate in character, and
Chaunochiton. A similar difference in type of ray occurs in the Octoknemaceae,
between Octoknema borealis Hutch, et J. M. Dalz. and Okoubaka aubrevillei
D. Normand (p. 378).
Pellegrin et

ECONOMIC USES
The Gaboon nut (Coula edulis Baill.) are edible after
kernels of the Coula or
cooking, and the fruits and sometimes the kernels of the Wild Olive (Ximenia
americana L.) are also eaten, although their value has been much disputed.
The woods are of little commercial importance owing to small dimensions or
quantities, though some are of good quality. Black Man wood, Minquartia sp.,
ishighly valued in Central America for its durability and has been used for
railway sleepers, posts, and poles. The wood of Ximenia americana Linn, is
sometimes used as a substitute for sandalwood (Metcalfe, 1497).

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Cathedra, Coula, Eganthus, Endusa, Heisteria, Liriosma, Minquartia,
Ochanostachys, Olax, Phlebocalymna, Schoepfia, Scorodocarpus, Strom-
bosia7 Ximenia.

(ii)
FOR WOOD STRUCTURE
(Agonandra), Anacolosa, Aptandra, Chaunochiton, Coula, Endusa, Heis-
Liriosma, Minquartia, Ochanostachys, Olax, Ongokea, Phlebocalymna,
teria,

Ptychopetalum, Schoepfia, Scorodocarpus, Strombosia, Strombosiopsis, and

Ximenia.
 OLACACEAE 367
LITERATURE
(i) On General Anatomy
Sleumer 2129, Youngken 2493.

(ii) On Wood Structure

Baker 104, Benoist 170, den Berger 179, 182, 183, Besson 186, Burgerstein 312, Chalk
and Chattaway 358, Cooper 461, Foxworthy 705, Howard 1088, Janssonius izj>4> Jentsch
1178, Metcalfe 1497* Normand 1619, Pfeiffer, J. Ph. 1713, Record 1843, 1851, 1853,
Record and Hess 1886, Record and Mell 1894, Williams 2426.

90. ICACINACEAE
(FiG. 84 on p. 368; FIG. 85 on p. 370; FIG. 86 on p. 372; FIG. 88 on p. 382)

SUMMARY
(i) GENERAL
A
tropical family of woody plants including erect trees and shrubs as well
as climbers. The general anatomy of the family is still imperfectly known, in
spite of the fact that it was subdivided into tribes by Engler (629, 630), many
years ago on an anatomical basis. The characters employed included the
presence or absence of scalariform perforation plates to the vessels, as well
as of interxylary phloem, together with variations in the equal or unequal
development of xylem around the periphery of the stem. Recent work of
Bailey and Howard (81) has shown this classification to be unreliable (see also
under 'Taxonomic and Phylogenetic Notes' on p. 375). There have also been
considerable differences of opinion concerning the taxonomy and nomen-
clature of the family, both of which facts make it difficult to interpret such
anatomical facts as have been recorded in the literature. The phylogenetic
conclusions drawn from the anatomical work by Bailey and Howard have
aroused much interest in recent years. Anomalous structure occurs in
certain genera.

(ii) WOOD
Vessels showing an unusual range from unspecialized to specialized forms;
the former typically numerous, small, solitary, with exclusively scalariform
perforation plates, scalariform or opposite pitting and very long members,
the latter small to medium-sized, often in radial multiples, and irregular
groups, and occasionally with a radial or tangential pattern, with simple
perforations, alternate pitting and members of medium length. Pits to ray
cells or parenchyma usually similar to the intervascular pitting, occasionally

larger. Parenchyma apotracheal in the genera with exclusively scalariform

perforation plates, mostly diffuse, but sometimes in definite bands; in the
more advanced woods varying from banded apotracheal with a little vasicentric
to predominantly paratracheal usually vasicentric where sparse and aliform
where more abundant. Rays very commonly of 2 distinct sizes, the larger
rays from 3 to ic or more cells wide, heterogeneous and often with sheath
cells; the large rays exhibiting varying stages of dissection into smaller units
and 'aggregate' in some species. Fibres typically with bordered pits, often
large and conspicuous, with simple pits in 2 genera; varying from very long
in the unspecialized to medium length in the more highly specialized genera.
368 ICACINACEAE
Included phloem of various types present in the tribes of climbing plants;
with included phloem in the Sarcostigmateae.

LEAF
Hairs simple, unicellular of varied length not uncommon in Apodytes,

FIG. 83. OPILIACEAE

A-B, A group and a pair of cystolithsfrom the mesophyll of Cansjera timorensis Decne. After Van
Tieghem.

FIG. 84. OPILIACEAE, A; ICACINACEAE, B-F

Hairs of: A, Cansjera parvifolia Kurz; B-t),
sjera parvtfoha natea excelsa Bl.; E-F, Hydathodes of Gonocaryum
B-D, Platea
pynforme SchefF. A-D by Solereder, E-F after Haberlandt.

Discophora, Liriosma, Mappia, Poraqueiba. Spherical unicellular

Emmotum,
hairs andmulticellular, peltate hairs also recorded in Platea excelsa BL
84 B-D). Tufted hairs in Phytocrene bracteata
Wall. Unicellular bladder-
(Fig.
like hairs recorded in Cardiopteris rumphii H. Baill. Cells of the epidermis
sometimes mucilaginous in Apodytes and Mappia. Lower epidermis papillose
inApodytes dimidiata E. Stomata ranunculaceous to cruciferous in a
Mey.
in Apodytes
few species of Alsodeiopsis, Lasianthera, Stemonurus, rubiaceous
dimidiata E. Mey. Hydathodes recorded in Gonocaryum pynforme Scheff.
(Fig. 84 E-F) and small ovoid bodies at the ends of the veins serving for water
 ICACINACEAE 369
secretion in Pyrenacantha malvaefolia Eng., P. scandens Harv., and Tremato-
sperma cordatum Urb. Mesophyll containing sclerenchymatous fibres in
Desmostachys renschii O. Hoffm. and Discophora guianensis Miers. Scleren-
chymatous fibres in the veins of Emmotum provided with numerous,
oblique,
slit-shaped, spirally arranged pits giving the appearance of spiral thickening.
Petiole of Apodytes dimidiata E. Mey. (Fig. 88 i) exhibiting an arc-shaped
vascular strand in transverse sections through the distal end. Articulated
laticiferous tubes recorded in the veins and petiole of Cardiopteris lobata
Baill. Solitary crystals present in certain species of Icacina, Mappia,
Poraqueiba, clustered crystals in species of Apodytes, Citronella mucronata
(Ruiz, et Pav.) D. Don, Desmostachys, Discophora, Emmotum, Poraqueiba ;
crystal-sand in Apodytes andamanica Kurz.

Axis
YOUNG STEM (Fig. 88
L-M)
Outer part of the cortex of the young stem containing well- developed stone
cells in Phytocrene macrophylla BL Cork arising in the epidermis in species of
Apodytes, lodes, Lasianthera, Pennantia, Phytocrene, Trematosperma. Cork
cells sometimes with strongly thickened, inner tangential walls in Apodytes,
Lasianthera, Pennantia. Pericycle recorded by Solereder as having a com-
posite and continuous ring of sclerenchyma in Emmotum fagifolium Desv.,
Gonocaryum gracile Miq., Leptaulus daphnoides Benth., Pennantia endlicheri
Russ., Platea excelsa BL, Poraqueiba guianensis Aubl., Stemonurus javanicus
Bl., Urandra apicaulis Thur., containing isolated bundles of fibres in Cardio-
pteris, or a loose ring of fibres in Pyrenacantha volubilis Hook. Young stem
with a circle of unevenly distributed vascular bundles around the pith in
Phytocrene macrophylla Bl. Inversely orientated medullary bundles stated
to occur in lodes tomentilla Miq. Howard (1096, 1097), writing of the xylem
of Codiocarpus, Discophora, Gastrolepis, Irvingbaileya, Lasianthera, Medu-
santhera as represented in twigs taken from herbarium sheets, states that all
of these genera belonging to the Icacinoideae possess at least a proportion of
vessel elements with scalariform perforation plates, or transitions between
these and simple perforations. Further particulars concerning the individual
genera are given as follows:
(i) Discophora.
Vessels aggregited or clustered and provided with scalari-
form perforation plates, simple perforations, and transitional forms. Inter-
vascular and vessel-parenchyma pits large, scattered, circular, alternate.
Ground tissue of the wood consisting of fibre-tracheids with pits 3-5 p in
diameter. Wood parenchyma abundant, diffuse to semi-banded or scanty
paratracheal. Rays heterogeneous, i to several cells wide; uniseriates tall,
composed of high cells.
(ii) Gastrolepis. Vessels in loose clusters and provided with scalariform
perforation plates as well as simple, and transitional perforations. Inter-
vascular and vessel-parenchyma pits showing transitions from scalariform to
bordered types. Ground tissue of the wood consisting of dense fibre-tracheids.
Wood parenchyma tending to be scanty paratracheal with vestiges of diffuse
elements.
(iii) Irvingbaileya. Vessels more or less aggregated and provided with
4594 Bb
370 ICACINACEAE
scalariform perforation plates as well as simple, and transitional perforations.
Intervascular and vessel-parenchyma pits tending to be scalariform-opposite.
Ground tissue of wood consisting of tracheids with wide lumina. Wood
diffuse with little or no
parenchyma scanty tendency to form bands.
(iv) Lasianthera. Vessels scattered or isolated. Vessel-parenchyma pits
scalariform with enlarged apertures. Ground tissue of wood consisting of

FIG. 85. ICACINACEAE

Phytocrene macrophylla Bl. A, Transverse section through the axis (magnification about 2:1).
B, Transverse section through a piece of a xylem wedge. After Robinson,

tracheids with inconspicuously bordered pits. Wood parenchyma scanty,

diffuse, or abaxial-paratracheal.
(v) Medusanthera. Vessels arranged in irregular multiples; with a smaller
proportion of scalariform perforation plates than in the related genera. Inter-
vascular and vessel-parenchyma pits small; ground tissue of wood consisting
of fibre-tracheids with minutely bordered pits, 2-0-2-5 p, in diameter. Rays
similar to those of Discophora.

Bailey and Howard (81) describe a study of the nodal anatomy in which it
was shown that amongst the Icacinoideae some of the genera possess trilacunar
(primitive) nodes and others unilacunar (advanced) nodes. Even within the
Icacineae there are genera in each of these categories, whilst in the lodeae,
Sarcostigmateae, and Phytocreneae, which are chiefly climbers, only uni-
lacunar nodes were observed. Members of the last three tribes are climbing
or twining in habit, whereas the trilacunar Icacineae are erect. These tribes
are regarded as phylogenetically more advanced than the usually erect
trilacunar Icacineae, whilst the unilacunar Icacineae are taken to represent a
transitional stage between the trilacunar Icacineae and the other tribes.
 ICACINACEAE 371
These differences in phylogentic specialization are correlated with others
described below under 'Wood', whilst further reference to the subject is
made under Thylogenetic and Taxonomic Notes'.
Mucilage spaces recorded in the soft tissues of the stem of Phytocrene
macrophylla Bl. The secretion in these spaces, when present, stated to be
strongly fluorescent. Mucilage canals said to occur in Trematosperma, and
lysigenous canals, arising secondarily on the inside of the vascular bundle's of
the axis, in Cardiopteris.

WOOD (Fig. 86 A-I)

Vessels small (less than 100 p, mean tangential diameter), just medium-
sized in a few genera, very small (less than 50 fi) in Alsodeiopsis, Apodytes,
Leptaulus, Medusanthera, Ottoschulzia, and Pennantia Exclusively solitary or
nearly so in genera with exclusively scalariform perforation plates (except
Pittosporopsis and Plated) and also in some species of Cantleya, Gonocaryum,
Lasianthera, Leptaulus, and Stemonurus, in which the perforation plates are
both scalariform and simple (82). Radial multiples common and sometimes
of 4 or more cells in Alsodeiopsis, Discophora, Mappia (82), Nothapodytes (82),
and Platea p.p., and with occasional irregular clusters in some of these woods;
with a tangential pattern in Desmostachys (Fig, 86 G) and Ottoschulzia.
Varying in number from about 6 to 80 per sq. mm., fewest (6-10 per sq. mm.)
in Alsodeiopsis, Cantleya, Medusanthera, and Platea, most numerous (50 or
more per sq, mm.) in Leptaulus and Pennantia. Spiral thickening reported by
Record (1801) in Citronella (Villaresia) mucronata D. Don. According to
Bailey and Howard (82) the perforation plates are (a) scalariform only in
Anisomallon, Apodytes, Calatola, Cassinopsis, Citronella, Dendrobangia,
Emmotum, Oecopetalum, Ottoschulzia, Pennantia, Pittosporopsis, Platea, and
Poraqueiba, (b) scalariform and simple in Cantleya, Discophora, Gastrolepis,
1

Gonocaryum, Grisollea, Leptaulus, Medusanthera, Stemonurus, and Urandra,

and (c) simple only in Alsodeiopsis, Chlamydocarya, Desmostachys, Hosiea,
Humirianthera, Icacina, Lavigeria, Leretia, Mappia, Mappianthus, Merrillio-
dendron, Miquelia, Natsiatum, Nothapodytes, Phytocrene, Pleurisanthes, Poly-
cephalium, Polyporandra, Pyrenacantha, Rhaphiostylis, Rhyticaryum, and
Sarcostigma\ scalariform perforation plates with thin and numerous bars
(20-50) in Anisomallon, Apodytes, Calatola, Citronella, Pennantia, and Platea\
the perforations commonly subdivided into reticulate patterns in Calatola and
Pennantia. Intervascular pitting usually transitional between scalariform and
opposite or opposite in woods in which scalariform perforation plates occur,
and alternate where the perforations are simple, the pits very small in
Alsodeiopsis, Apodytes, Medusanthera, and Pennantia, with striations due to
coalescent apertures in Alsodeiopsis and Desmostachys', pits to ray and paren-
chyma cells usually similar to the intervascular pit-pairs, but enlarged and
almost simple in Discophora and Gonocaryum, Tyloses sometimes present in
Cantleya and Emmotum, thick- walled in the former. Bailey and Howard (83)
give the following mean member lengths: Icacineae (a) Trilacunar, with
scalariform perforation plates, 1*47 mm., (b) Trilacunar, with simple and
scalariform perforation plates, 1-05 mm., (c) Unilacunar, 0-59 mm. For the

1
Williams (2430) describes both simple and scalariform plates in Poraqueiba sericea Tul.
 H
FIG. 86. ICACINACEAE, A-I; OCTOKNEMATACEAE, J, K
A, Alsodeiopsis staudtii Engl. B, A. staudtii Engl. C, Pennantia cunninghamii Miers. D, Apodytes
dimidiata E, Mey. E, v4. dimidiata E. Mcy, F, Leptaulus daphnoides Benth. G, Desmostachys vogeKi
Stapf. H, Cantleya corniculata (Becc.) Howard. I, C. corniculata (Becc.) Howard, J, Octoknema
borealis Hutch, et Dalz. K, O. borealu Hutch, et Dalz.
 ICACINACEAE 373
lodeae, Sarcostigmateae, and Phytocreneae, 0-47 mm. Parenchyma varying
considerably in type and amount and often unstable. Bailey and Howard (83)
have shown that the type of parenchyma is closely related to the degree of
specialization of the vessels. Woods with exclusively scalariform perforation
plates typically with abundant apotracheal parenchyma in numerous short
lines (Fig. 86 E), sometimes with a tendency to definite bands in Ottoschulzia
and Poraqueiba, and with some vasicentric parenchyma in Citronella and
Emmotum', of the woods with both simple and scalariform perforation plates,
Discophora and Stemonurus have banded apotracheal parenchyma only, the
others have at least some paratracheal parenchyma, as in Gonocaryum with
diffuse and vasicentric, or predominantly paratracheal parenchyma (Fig. 86 if),
e.g. aliform in Cantleya p.p. and Stemonurus or with a few cells round the
vessels in Cantleya p.p. and Gastrolepis (83), and with short rows attached to
the vessels on the abaxial surfaces in Leptaulus (Fig, 86 F) woods with
;

exclusively simple perforations commonly with some or predominantly para-

tracheal parenchyma with both apotracheal bands and paratracheal (vasicentric)
;

parenchyma in Alsodeiopsis (Fig. 86 A), Mappia, Medusanthera, Merrillio-

dendron, and Nothapodytes (83); according to Bailey and Howard (83), in
Desmostachys (Fig. 86 G) and the following genera with a scrambling or
climbing habit of growth Humiriantfiera, Icacina, Lavigeria, Leretia, Pleuri-
santhes, Rhaphiostylis, and Rhyticaryum the banded apotracheal parenchyma
is conspicuously reduced, giving varied transitions between banded apotracheal
and paratracheal types of distribution and forms having a predominantly
paratracheal or vasicentric distribution. These authors note similar unstable
types in small-vesselled parts of the stem of certain representatives of the
Phytocreneae and lodeae, whereas in the large-vesselled anomalous wood of
these plants the parenchyma tends to be more dominantly paratracheal they ;

also note that in Sarcostigma, and at times in certain of the Phytocreneae, the
strands of wood parenchyma tend to be replaced by curious septate parenchy-
matous elements. Strands usually of 8 cells, sometimes more in the un-
specialized genera. Crystals not observed. Rays very variable in width and
height, varying from up to 3 or 4 cells wide in Anisomallon, Apodytes, Cantleya,
Discophora, Leptaulus, and Platea excelsa Bl. up to 10 or more cells wide in
Alsodeiopsis schumannii Engl., Citronella, Emmotum, Medusanthera, Otto-
schulzia, Pennantia, Poraqueiba, and Stemonurus^ varying in height from about
i mm. in
Leptaulus to 4 mm. or more; commonly of 2 distinct sizes, except
where the multiseriate rays are narrow, as in Apodytes (Fig. 86 D), Calatola,
Cantleya, and Discophora, or where uniseriate rays are very scarce, as in
Tylecarpus\ the large primary multiseriate rays commonly dissected into
smaller units (Fig. 86 B) and producing typical Aggregate* rays in some
species of Alsodeiopsis and Poraqueiba (84); the uniseriate rays typically
numerous, often very high and usually composed of very high upright cells,
exceptionally high in Desmostachys vogelii Stapf. (84), least numerous in some
species of Cantleya, Citronella, Discophora, Gastrolepis, Gonocaryum, and
Medusanthera] varying in number from about 4 to 16 rays per mm., fewest in
some species of Cantleya, Citronella, and Medusanthera', very distinctly
heterogeneous (Kribs's Type I in the unspecialized genera, Type II A and B
in the others), with several marginal rows of markedly upright cells, rows
seldom exceeding 4 in Cantleya, Citronella, and Medusanthera and frequently
374 ICACINACEAE
with 10 or more rows in Anisomallon, Calatola, Ottoschulzia, Platea> and
Stemonurus; commonly with sheath cells. One or more crystals occur in the
ordinary ray cells of several species and, less frequently, small quantities of
dark-coloured deposits. According to Bailey and Howard (84) in the highly
specialized climbing plants of the lodeae, Sarcostigmateae, and Phytocreneae
there is a conspicuous tendency for the elimination of multiseriate rays from
the first-formed secondary xylem multiseriate rays occur in some species of
;

the lodeae, either extending outwards from gaps in the primary body or
arising abruptly at some distance from the stele; the first-formed secondary
xylem of most species of lodes and Polyporandra has multiseriate rays with or
without varying numbers of biseriate or aggregate rays, a type of structure
that predominates through relatively thick stems in the Sarcostigmateae; in
the lodeae much-modified multiseriate rays with very thin-walled cells are
retained in the later-formed anomalous wood. In the Phytocreneae multi-
seriate rays are usually eliminated from young stems except for pairs of wide

rays, which flank the inwardly projecting strands of phloem. Fibres, accord-
ing to Bailey and Howard (83), with large conspicuous bordered pits in the
genera with exclusively scalariform perforation plates and also in some species
of Cantleya in Discophora, Gastrolepis, Gonocaryum, Grisollea, Lasianthera,
;

Leptaulus, and Medusanthera the pits are bordered, but the borders are smaller
and the chambers flatter and less obvious in section in the aborescent genera
;

with exclusively simple perforations there is a conspicuous reduction in the

size of the bordered pits (less marked in Merrilliodendron) and the pits are

simple in Mappia and Nothapodytes. Walls usually thick, sometimes with

extremely small lumina, but thin-walled in Platea. Bailey and Howard (83)
give the following mean lengths: Icacineae (a) Trilacunar, with scalariform
perforation plates, 2*49 mm., (b) Trilacunar with simple and scalariform
perforation plates, 2-58 mm., (c) Unilacunar, 1-60 mm.; for the lodeae,
Sarcostigmateae, and Phytocreneae, 0-87 mm. Bailey and Howard (83) note
that the sectional area of the fibres is much smaller in the members of the
Icacinaceae with exclusively simple perforations. In the unilacunar Icacina-
ceae, having a scrambling or scandent habit of growth, as in the scandent
lodeae Sarcostigmateae and Phytocreneae, Bailey and Howard state that
'there are reversals in the specialization of the imperforate tracheary elements,
which culminate in the formation of curious short tracheids. These tracheary
cells are characterized by having relatively conspicuous lumina and numerous
bordered pits. They are most typically developed in the large-vesselled
anomalous wood of Lavigeria, Pleurisanthes and of various representatives of
the lodeae and Phytocreneae. Intermediate or transitional stages of their
phylogenetic differentiation occur in Hunlirianthera, Icacina, Leretia, Mappi-
anthus, Rhaphiostylis, and Sarcostigma, as well as in the denser, smaller-
vesselled parts of the stems of certain lodeae and Phytocreneae.' Included
(interxyiary) phloem. Bailey and Howard (81) refer to the occurrence of
interxylary phloem in the climbing shrubs of the tribe Sarcostigmateae, and
to various forms of anomalous structure in young stems in the lodeae and

Phytocreneae. For further particulars see next paragraph*

ANOMALOUS STRUCTURE
Tree trunks flanged in Citronella moorei (F. Muell.) Howard, according to
 ICACINACEAE 375
Francis (708). The xylem and secondary phloem of Phytocrene (Fig. 85 A-B)
are differentiated as alternating wedges, the xylem ring consisting of well-
developed teeth of wood with phloem patches between them. The normal
phloem strands are also well developed but with weak areas of xylem between
them. This abnormal structure is absent from young stems. Zones of growth,
each exhibiting the abnormal type of structure described above, arise from
successive cambia in certain species of Chlamydocarya, Phytocrene^ and Sarco-
stigma. Each zone of growth either completely encircles the branch or is
confined to certain segments, according to Pfeiffer (1712), Solereder, and
Timmermans (2261). Alternating wedges of xylem and phloem also recorded
in Pyrenacantha. Interxylary phloem present in Chlamydocarya and
Sarcostigma. Instances where unequal increases in the thickness of the wood
sometimes become obliterated in old material have been recorded in lodes and
Natsiatum. For other information concerning interxylary phloem see 'Wood*.

TAXONOMIC AND PHYLOGENETIC NOTES

Many of the genera of the Icacinaceae were included in the system of
Bentham and Hooker in the tribes Icacineae and Phytocreneae of the Olacineae.
In the system of Engler and Prantl the Icacinaceae are treated as a separate
family under Sapindales. The Icacinaceae are also recognized as a separate
family by Hutchinson, who, like Wettstein, places them in the Celastrales.
When considering the relationship between the Icacinaceae and Olaceae, it is
interesting to note the occurrence of articulated laticiferous tubes in Cardio-
pteris similar to those which are to be found in the tribe Couleae of the
Olacaceae. Then again the presence of sclerenchymatous fibres in the meso-
phyll of Desmostachys and Discophora is reminiscent of those in Eganthus,
Endusa, Heisteria, Minquartia Ochanostachys, and Scorodocarpus.
y

It will be seen from the various references to the work of Bailey and Howard

given above under the description of the axis that these authors have used
the anatomical method to investigate the phylogenetic relationships within
the family. Starting with the assumption, discussed on p. xxxix of this book,
that the trilacunar node is more primitive than the unilacunar type, Bailey
and Howard first showed that trilacunar nodes occur only in the Icacineae,
whereas in certain other genera of the Icacineae, as well as in the lodeae,
Sarcostigmateae, and Phytocreneae, only unilacunar nodes were observed.
These last three tribes, consisting mostly of twining or climbing plants, are
therefore regarded as more specialized than the trilacunar Icacineae. The
unilacunar Icacineae are taken to be intermediate between these two
categories.
The same authors also examined the vessel members, wood fibres, wood
parenchyma, and types of ray which occur in the family from the phylogenetic
standpoint. In general there was found to be good agreement concerning the
lines of specialization within the family no matter which of these characters
is used as an indicator, but the evidence provided by the vessel elements and

wood fibres seems more definite than that derived from the parenchyma or
rays. For instance, when using the characters of the vessel members, the
authors show how the Icacinoideae may be divided into three major categories :

I. Those in which the vessel perforation plates are exclusively scalariform.

II. Those in which a mixture of simple and scalariform perforation plates

376 ICACINACEAE
occur. III. Those in which the perforations are exclusively simple. Genera
with unilacunar (advanced) nodal structure were all found to be in group III
with exclusively simple (advanced) perforations. Conversely, trilacunar nodes
and scalariform perforation plates were found to occur together. Bailey and
Howard suggest that these facts are of taxonomic as well as of phylogenetic
significance. They wisely state, however, that Tarallel developments and
convergent evolution are of such common occurrence in foliar and cauline, as
well as in floral structures that evidence from all parts of the plant must be
harmonized and integrated in attempting to determine actual genetic affinities
within an order, family, sub-family or tribe'. After describing and discussing
the evidence from the wood fibres, rays, and parenchyma as well as that from
the vessels, the same authors conclude that, although the trends of specializa-
tion in all of these organs tend to run parallel with one another, 'the rates of
evolutionary modifications in one category of these tissue cells not infrequently
are retarded or accelerated in relation to the changes that are occurring in the
other tissue cells'. Bailey and Howard's researches have clearly demonstrated
some most interesting correlations.

ECONOMIC USES
A substitute for Mate Tea (Ilex paraguariensis St. Hil.) is obtained from
Citronella gongonha (Mart.) Howard (syn. Villaresia gongonha Miers), the
substitute having a strong resemblance to the genuine article. The
anatomical structure has been described by Lendner (1357). Starch and
oil are obtained from the fruit and seed of Poraqueiba in the Para region of
Brazil. Humirianthera is cultivated for the sake of its fleshy tubers and root-
stocks, but the starchy material must be washed before use as it is otherwise
toxic. A blue dye is obtained from the bark, leaves, and fruits of Calatola.
The woods of this family are of little commercial importance. That of
Apodytes dimidiata E. Mey. is used in South Africa for cart and wagon felloes
and Record and Hess (1886) note species of Dendrobangia and Ottoschulzia
as being used locally in tropical America and the West Indies.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Alsodeiopsis, Apodytes, Cardiopteris, Chlamydocarya, Citronella, Codio-
carpus, Desmostachys, Discophora, Emmotum, Gastrolepis, Gonocaryum,
Icacina, lodes, Irvingbaileya, Lasianthera, Leptaulus, Liriosma, Mappia,
Medusanthera, Natsiatum, Pennantia, Phytocrene, Platea, Polyporandra,
Poraqueiba, Pyrenacantha, Sarcostigma, Stemonurus, Trematosperma,
Urandra.

(ii) FOR WOOD STRUCTURE

Alsodeiopsis, Anisomallon, Apodytes, Calatola, (Cantleya), (Cassinopsis),
(Chlamydocarya), Citronella, (Dendrobangia), Desmostachys, Discophora,
Emmotum, (Gastrolepis), Gonocaryum, (Grisollea), (Hosiea), (Humiri-
anthera), (Icacina), (Lasianthera), (Lavigeria), Leptaulus, (Leretia), (Mappia),
(Mappianthus), Medusanthera, (Merrilliodendron), (Miquelia), (Natsiatum),
(Nothapodytes), (Oecopetalum), Ottoschulzia, Pennantia, (Phytocrene),
(Pittosporopsis), Platea, (Pleurisanthes), (Polycephalium), (Polyporandra),
 ICACINACEAE 377
Poraqueiba, (Pyrenacantha), (Rhaphiostylis), (Rhyticaryum), (Sarcostigma),
Stemonurus.
LITERATURE
(i) On General Anatomy
Bailey and Howard 81, Engler 629, 630, Francis 708, Heinzelman 2523, Howard 1091,
1092, 1093, 1094, 1095, 1096, 1097, 1098, Lendner 1357, Pfeiffer 1712, Scala 2018,
Timmermans 2261.
(ii) On Wood Structure
Bailey and Howard 81, 82, 83, 84, Benoist 170, den Berger 182, Burgerstein 310, Chalk
et al. 360, Cockrell 434, Cooper 461, Cozzo 494, Dadswell 525, Dadswell and Record 533,
Desch 574, Francis 706, Howard 1088, Janssonius f\54, Kanehira 1206, Lecomte 1334,
Metcalfe 1497, Normand 1619, Pfeiffer, H. 1712, Record 1843, 1851, 1853, Record and Hess
1886, Record and Mell 1894, Scott 2075, Tupper 2295, Williams 2426, 2430, 2431.

91. OCTOKNEMACEAE
(Fic. 86 on p. 372)

SUMMARY
This West African family has, until recently, been treated as comprising
the trees and shrubs belonging to the single genus Octoknema. Some
authorities now regard Okoubaka aubrevillei Pellegrin et D. Normand (syn.
Octoknema okoubaka Aubr. et Pellegrin) as a distinct genus, a view which
appears to be supported by differences in the structure of the wood. The
anatomical features of the leaf and young stem of Octoknema which are given
below are those recorded by Solereder and by Mildbraed (1534). The wood
of the 2 genera exhibits the following structure. Octoknema. Vessels in
numerous multiples, perforation plates scalariform, members very long.
Parenchyma absent. Rays up to 3 cells wide, markedly heterogeneous.
Fibres with simple pits, sometimes septate, moderately long. Okoubaka.
Vessels solitary, perforations simple, members moderately to very short.
Parenchyma apotracheal, diffuse. Rays up to 4 or 5 cells wide, homo-
geneous. Fibres with simple pits, but with bordered pits round the vessels,
moderately long.
LEAF
Hairs tufted, e.g. in Octoknema klainena Pierre, and stellate, particularly in
O. affinis Pierre. Epidermis composed of small, irregularly polyhedral cells.
Stomata confined to the lower surface ranunculaceous. Palisade tissue not
;

but mesophyll more compact on the adaxial than on

clearly differentiated,
the abaxial side. Vascular bundles of the veins accompanied by strongly
thickened fibres and crystals in the endodermis.

Axis
YOUNG STEM
Cork arising in the sub-epidermis, the component cells thickened on the
outer tangential and radial walls. Primary cortex containing stone cells with
thickened, lignified walls, isolated or in small groups. Cortex also containing
slightly thickened cells enclosing solitary crystals. Pericycle with a com-
posite and continuous ring of sclerenchyma. Phloem including numerous
378 OCTOKNEMACEAE
small bundles of fibres surrounded by crystalliferous cells. Xylem with
evenly distributed vessels of equal size walls with bordered pits perforation
; ;

plates frequently scalariform. Rays 1-2 cells wide; component cells often
containing solitary crystals. Pith including groups of stone cells. Crystals,
see Cortex', 'Phloem', and 'Rays'.

WOOD
Octoknema borealis Hutch, et J. M. Dalz. (Fig. 86 and K)
j

Vessels moderately small (mean tang, solitary and in

diameter 50-100 /it);
numerous multiples of 2 or 3 cells; about 35 per sq. mm. Perforation plates
scalariform, usually with fewer than 20 bars. Intervascular pitting opposite
or intermediate between opposite and alternate, large; pits to ray cells
commonly large, horizontally elongated and simple. Mean member length
about -2-i -6 mm. Parenchyma absent. Rays up to 3 cells wide ; commonly
i

fused vertically to give rays 2 mm. or more high; uniseriates numerous,

composed entirely of upright cells (apart from chambered crystalliferous
cells);about 16 rays per mm.; markedly heterogeneous (Kribs's Type I),
commonly with 10 or more marginal rows of square to upright cells.
Chambered crystals numerous in the upright cells. Fibres with simple pits
which have long apertures and trumpet-shaped canals and are almost entirely
limited to the radial walls; sometimes septate. Walls moderately thin to thick.
Mean length about 2-2 mm.
Okoubaka aubrevillei Pellegrin et D. Normand (Octoknema okoubaka Aubr.
et Pellegr.)

Based on the description by D. Normand (1617). Vessels medium-sized

(100-200 /*); exclusively solitary; about 4 per sq. mm. Perforations simple.
Intervascular pitting alternate and small ( ?). Mean length about 0-25 mm.
Parenchyma apotracheal, as numerous scattered cells and short uniseriate
lines. Strands commonly of 2 cells. Tending to be storied. Rays up to

4-5 cells wide and about 1-5 mm. high; uniseriates rare or absent; about
4 rays per mm. homogeneous (Kribs's Type II), composed of rather large
;

procumbent cells. Crystals sometimes present in the ordinary cells. Fibres

with simple pits. Walls very thick. Mean length about 2-0 mm. Fibre-
tracheids occur about the vessels, with numerous bordered pits and thin walls.

TAXONOMIC NOTES
Although there has been much uncertainty concerning the taxonomic
position of Octoknema, it is treated as a unigeneric family both by Mildbraed
(1534) and Hutchinson (1113). The absence of any outstanding anatomical
features makes it difficult to decide whether it has affinities with the Olacaceae.
If it is related to this family the type of hair would suggest affinities with the
Couleae.
Normand (1617) has drawn attention to the marked differences between
the woods of Octoknema and Okoubaka and the separation of the genus
Okoubaka appears to be well justified. Similar differences occur between
genera recognized as belonging to the Olacaceae, and most of the features
(with the exception of the homogeneous rays in Okoubaka and the absence of
parenchyma from Octoknema) in the two genera which we are now considering
 OCTOKNEMACEAE 379
can be found in one or other of the genera which are generally accepted as
members of the Olacaceae.
The wood structure of Octoknema is unspecialized, whereas that of
Okoubaka is moderately highly specialized. Normand suggests that, while
Octoknema has many wood characters in common with the Olacaceae, the
structure of Okoubaka suggests affinity with the Santalaceae.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Octoknema.

(ii)
FOR WOOD STRUCTURE
Octoknema, (Okoubaka).
LITERATURE
(i) On General Anatomy
Hutchinson 1113, Mildbraed 1534.
(ii) On Wood Structure
Cooper 461, Normand 1617, 1619, Record 1843, 1851.

92. OPILIACEAE
(Fie. 83 on p. 368; FIG. 84 on p. 368; FIG. 87 on p. 380; FIG. 89 on p. 388)

SUMMARY
A small family of trees, shrubs, or woody climbers from tropical regions,
especially in Asia and Africa. Characteristic features include the occurrence
of cystoliths in leaf and stem and branched, lignified cells in the mesophyll.
The wood exhibits the following characters. Vessels exclusively solitary or
with some multiples, perforations simple, intervascular pitting rare, alternate
and small, pits to parenchyma similar, members moderately short. Paren-
chyma diffuse, strands usually of 2 cells. Rays up to 2-5 cells wide, with
very few uniseriates, homogeneous; large cystoliths present in most of the
genera. Fibres with simple or bordered pits, of medium length.

LEAF
Usually dorsiventral, but centric in Opilia amentacea Roxb. Multicellular
hairs, branched like a stag's horn, recorded in certain species of Cansjera
(Fig. 84 A). Stomata present on both surfaces in Cansjera parvifolia Kurz.,
rubiaceous in Champereia and Opilia. Veins connected by a branched system
cells, in Agonandra, Cansjera, Lepionurus, and
of spirally thickened, lignified
Opilia. Secretory elements. Mucilage cells situated in the spongy meso-
phyll of Agonandra sp. and Opilia sp. Secretory cells with finely granular
contents, readily stained with iodine, recorded in the lowest layer of the
mesophyll of Opilia amentacea. Crystals absent. Cystoliths, arranged in
groups of 2 to several in special cells, occur in the mesophyll of Agonandra,
Cansjera (Fig. 83 A-B), Lepionurus, and Opilia.

Axis
YOUNG STEM
Cork arising in the epidermis in certain species of Champereia and Opilia.
380 OPILIACEAE
Pericycle with a composite and continuous ring of sclerenchyma in
Agonandra, Cansjera, Champereia, Lepionurus, and Opilia. Xylem including
vessels which are 33 in diameter in Champereia, exhibit spiral striation in
//,

Opilia amentacea Roxb., and have simple perforations in Champereia and

Opilia. Rays 2-3 cells wide in Champereia', wood fibres provided
with
bordered pits and xylem parenchyma scanty in the same genus. Cystoliths
in the parenchyma
(Fig. 87 A-D), similar to those in the leaf, generally present
of the axis, especially in the rays, of all of the genera.

FIG. 87. OPILIACEAE

A, Double cystoliths from the phloem of the axis in Champereia
griffithiana Planch. (Griffith, n. 4388, Herb. Monac.), calcified. B,
The same after decalcification with acetic acid. C, Secondary meta-
morphosis of the double cystoliths of Champereia sp, (Cuming, 1 129,
Herb. Monac.). D Group of cells with silicified cystolith-like bodies
t

from the leaf-tissue of Champereia griffithiana. By Solereder.

WOOD (Fig. 89 A-C)

p, mean tangential diameter) to very
Vessels moderately small (50-100
small (25-50 /i); exclusively solitary in Agonandra and Champereia, with some
multiples in Opilia\ about 25 per sq. mm. Perforations simple. Intervascular
pitting very rare, alternate and small pits to ray and wood parenchyma small
;

and round, sometimes striated. Mean member

length 0-2-0-35 (Cham- mm -

pereia). Parenchyma apotracheal, as numerous scattered cells that some-

times tend to form short uniseriate lines. Strands usually of 2 cells, sometimes
of 4 cells in Champereia, and with some fusiform cells in Agonandra. Rays up
to 5 cells wide in Champereia manillana Merr., but seldom more than 2 cells
wide in Agonandra', uniseriates very few, composed of procumbent cells;
about 7 rays per mm. homogeneous (Kribs's Type I). Large cells containing
;

cystoliths of calcium carbonate (Fig. 89 B) observed or reported (1206,

1799, an d *8i8) in Cansjera, Champereia, Lepionurus, Melientha, Opilia, and
Rhopafapilia, but absent from Agonandra. Fibres* Pits simple in Cham-
pereia but with small, moderately distinct borders in Agonandra, and distinctly
bordered in Opilia. Walls thick. Mean length i '0-1-4 mm *

TAXONOMIC NOTES
The genera Agonandra, Cansjera, Lepionurus, and Opilia constituted the
tribe Opilieae of the Olacineae in the system of Bentham and Hooker. The
Opiliaceae are treated as a distinct family by Sleumer (2130), and as a family
 OPILIACEAE 381
in the Olacales in Hutchinson's (1113) classification. The occurrence of a
branched system of lignified cells in the leaf, and the development of cystoliths
throughout four of these genera are characters which indicate that they
all
constitute a homogeneous group, and at the same time serve to differentiate
them from the Olacaceae and Icacinaceae in which the same diagnostic
characters have not been recorded. Champereia was included in the Santa-
laceae in the Bentham and Hooker system, but placed in the Opiliaceae in the
classifications of both Engler and Hutchinson. On anatomical grounds it
seems to belong to the Opiliaceae rather than the Santalaceae, the presence
of the cystoliths which are such a characteristic feature of the Opiliaceae
being especially interesting in this connexion. The family also appears to be
unique in having cystoliths in the secondary xylem.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Agonandra, Cansjera, Champereia, Lepionurus, Opilia.

(ii) FOR WOOD STRUCTURE

Agonandra, (Cansjera), Champereia, (Lepionurus), (Melientha), Opilia,
(Rhopalopilia).
LITERATURE
(i) On General Anatomy
Hutchinson 1113, Sleumer 2130.

(ii) On Wood Structure

Kanehira 1206, 1209, Record 1799, 1818, 1851, 1853.

93. AQUIFOLIACEAE
(Fia. 88 on p. 382; FIG. 89 on p. 388)

SUMMARY
(i) GENERAL
A
widely distributed family of trees and shrubs, most species being ever-
green. Practically all of the anatomical observations which have been made
refer to the single genus Ilex. The description which follows may be taken to
refer to this genus only, except where stated to the contrary. The family is
anatomically homogeneous, except in so far as there are minor differences,
especially in leaf structure, between the deciduous and evergreen species.
The epidermis of the leaf is frequently many-layered and the component
cells mucilaginous. Hypoderm has also been recorded on the upper side of
the leaf of certain species, but it seems probable that true hypoderm has not
always been clearly distinguished from an epidermis of more than one layer.
Cork warts on the lower surface of the leaf are characteristic of some species,
and the leaf margin is frequently strengthened mechanically by cuticlarized
or sclerenchymatous tissues. The hairs are usually infrequent but, where
present, generally consist of small unicellular trichomes, although
multi-
cellular, thin-walled trichomes have been recorded in certain deciduous
species. Stomata ranunculaceous; confined to the lower surface. In the
 FIG. 88. GOUPIACEAE, A; CELASTRACEAE, B, D, and G; HIPPOCRATEACEAE, C;
AQUIFOLIACEAE, E~F, H, and J-K; ICACINACEAE, I and L-M
A, GoMpifl #/a6ra Aubl. Petiole X22. B, Kurrimia pulckerrima Wall. Petiole X 15, C, Salacia
roxburghii Wall. Petiole X 15. D, Euonymus radicans Sieb. Stomata on lower surface x 250. E, Ilex
aquifolium Linn. T.S, lamina; adaxial part, x 167. F, /. paraguariensis St. Hil, Stomata on lower sur-
face X 167. G, Cflt/ra e^ttfo's Forsk. Petiole x 15. H, Ilex paraguariensis St. Hil. T.S. lamina X 167.
I, Apodytes dimidiata E. Mey. Petiole X 15. J, Ilex paraguariensis St. Hil. Petiole x 15. K, /. aqui-
folium Linn, Petiole xis. L, Apodytes dimidiata E. Mey. Stem Xis. M, Miquetta sp. Stem Xis.
c. Cortex with abundant tanniniferous cells, f.b. Fat bodies, s.c. Secretory cavities.
 AQUIFOLIACEAE 383

xylem of the young stem the wood fibres are provided with bordered pits,
and the vessels with scalariform perforation plates.

(ii)
WOOD
Vessels small, often in marked radial groups or lines, spiral thickening
usually present in Ilex and Byronia, perforation plates exclusively scalariform
and mostly with many bars, intervascular pitting usually opposite; members
moderately to very long. Parenchyma moderately abundant, diffuse. Rays
of 2 distinct sizes, the larger 5-15 cells wide and 2-5 mm. high, heterogeneous.
Fibres with distinctly bordered pits; spirally thickened in Ilex] of medium
length to moderately long.

LEAF
Dorsiventral. Cuticle very thin in Nemopanthus and in certain species of
Ilex, e.g. /. laevigata (Pursh.) Gray and /. verticillata (L.) Gray, but very
thick in most species of Ilex and consisting of 2 distinct layers in some. Hairs
rare, but where present usually consisting of unicellular trichomes with the
lumina almost or completely obliterated. Long, multicellular, thin-walled
trichomes recorded in /. laevigata and /. verticillata. Cork warts, resembling
lenticels and consisting of hemispherical groups of cells with suberized walls

radiating from the centre of the hemisphere, abundant on the lower surface
of certain species of Ilex. Margins of the leaf strengthened mechanically in
various ways in different species (i) by cuticular thickenings (ii) by scleren-
:
;

chymatous tissue containing chlorophyll (/. insignis Hook, f.); (iii) by a sheath
of sclerenchyma (/. aquifolium Linn.). Epidermis consisting of 1-3 layers of
cells in different species of Ilex\ the component cells partly or wholly muci-

laginous in a considerable number of species, sometimes penetrating deeply

into the mesophyll. Lower epidermis with considerable thickening and pitting
of the inner walls in /. affinis Gardn. Hypoderm, not to be confused with
a many-layered epidermis, occurs in certain species, notably J. aquifolium
(Fig. 88 E). Stomata (Fig. 88 F) confined to the lower surface; sometimes
surrounded by a rampart of cuticle ranunculaceous. (Solereder's statement
;

that some semblance of subsidiary cells occurs in /. paraguariewis St. HiL

has not been confirmed by observation.) Mesophyll including 2 or more
layers of palisade tissue in species with coriaceous leaves, but only i -layered
in deciduous species such as /. laevigata and /. verticillata. Spongy mesophyll
somewhat variable in different species; intercellular spaces few in /. decidua
Walt, but well developed in /. insignis Hook. f. and /, latifolia Thunb.
Vascular bundles of the veins accompanied by a varying amount of scleren-
chyma in different species of Ilex. Phloem of the midrib stated to form a
continuous band round the xylem in the evergreen species /. opaca Ait. and
7. glabra (L.) Gray, but according to Holm (1053) the bundle is crescent-

shaped in /. laevigata and /. verticillata. Midrib very prominent on the upper

side in /. glabra, but most prominent on the lower side in a majority of the
species. Petiole, in transverse sections, exhibiting an arc-shaped vascular
strand, relatively open in some species, e.g. Ilex aquifolium (Fig. 85 K), but
with strongly incurved ends in others, e.g. in Ilex paraguariensis (Fig. 85 j) ;

often accompanied by 2 very small accessory strands. Vascular strand accom-

panied externally by sclerenchyma varying in amount in different species, and
384 AQUIFOLIACEAE
almost completely absent from /. glabra. Fat bodies common in the meso-
phyll. Crystals exclusively clustered, sometimes occurring in roundish
idioblasts. Epidermis stated to contain sphaerocrystalline masses, possibly
consisting of hesperidin, in varieties of /. paltorioides Reiss. and a few other
species.

Axis
YOUNG STEM
Cuticle very thick, smooth in some species (e.g. /. aquifolium Linn.), but
granular in others (e.g. /. glabra (L.) Gray). Cork arising in the epidermis or
hypodermis; cork cells with unusually thick walls. Epidermis tending to
persist even after the formation of cork. Primary cortex parenchymatous
when young, but outer part tending to become slightly collenchymatous in
/. aqutfolium and other species. Stone cells have been recorded in the cortex,
but none were observed in any of the species of Ilex available for examination.
Pericycle containing isolated strands of fibres in young stems, but becoming
united by stone cells to form a composite and continuous ring in species of
Byronia and Ilex. Xylem and phloem commonly traversed by broad rays.
Vessels with scalariform perforation plates. Solitary and clustered crystals
occur in the cortex.

WOOD (Fig. 89 D-E)

Vessels mostly very small (less than 50 p mean tangential diameter) and
sometimes extremely small, but rather larger in some species of Ilex, particu-
larly in 7. cymosa Bl. (75 /LC) radial multiples of 4 or more common in Ilex
;

(except /. cymosa Bl.) and Byronia and often imparting a marked radial pattern
to these woods, sometimes in clusters, and with a tendency to local tangential
arrangement in some species of Ilex difficult to count satisfactorily owing to
;

the groups, fewest (about 7 per mm.) in /. cymosa Bl. Kanehira (1206) gives
;

the upper limit for Ilex as 120 per mm.; more numerous at the beginning
of the ring in some species of Ilex and ring-porous in Nemopanthus (533),
typically with well-marked spiral thickening in temperate species of Ilex
and Byronia, but Record (1894) lists 3 species of /fee without spiral thickening,
to which may be added /. cymosa Bl. Perforation plates exclusively scalari-
form, with more than 20 bars except in Nemopanthus. Intervascular pitting
opposite, sometimes scalariform in Ilex (Record, 1818, 1853) anc* sometimes
nearly alternate in /. aquifolium L. pits to ray and parenchyma cells similar
;

to the intervascular pitting, but sometimes elongated or unilaterally com-

pound in Ilex. Deposits and tyloses not observed but calcium oxalate re-
corded in callus tissue of/, opaca Ait. (Record, 1818). Mean member length
0-9-1 3 mm. Parenchyma moderately abundant, apotracheal, diffuse, as
scattered cells or short uniseriate lines of a few cells. Chambered cells
containing crystals moderately common in /. cymosa Bl. Strands commonly
of 8 cells. Rays of 2 distinct sizes, uniseriate or wide and multiseriate, the
larger rays mostly between 4 and 10 cells wide, more than 10 cells wide in a
few species of Ilex> e.g. /. cymosa Bl., /. hanceana Max., and /. mitts (L.)
Radlk., and in Phelline\ maximum height 2-5 mm. ; uniseriate rays, composed
of high upright cells, usually abundant; Loesener, according to Solereder,
states that the rays are exclusively uniseriate in /. vertidllata\ about 7-15 rays
 AQUIFOLIACEAE 385
per mm. heterogeneous (Kribs's Type I), the larger rays with between 4 and
;

10 marginal rows of upright cells in most species, usually with fewer than
4 rows in /. hanceana Max., /. mitts (L.) Radlk., and Phelline and with more
than 10 rows in 7. aquifolium L. and Byronia with sheath cells, except in
;

Nemopanthus', crystals in chambered cells observed in /. cymosa Bl. and

recorded by Kanehira (1206) for several Formosan species of Ilex. Fibres
with distinctly bordered pits (borders rather small in /. cymosa BL), more
numerous on the radial than on the tangential walls with moderately thin to
;

moderately thick walls spiral thickening characteristic of temperate species

;

of Ilex; absent from /. cymosa Bl. and 3 species listed by Record (1894).
Mean length 1-3-2-1 mm. Growth ring formation in Ilex has been described
by Coster (481).
ROOT
The root structure of Ilex has been described by Weber (2380).

ECONOMIC USES
Mate\ Yerba Mate, or Paraguay Tea is the product of various cultivated
forms and varieties of Ilex paraguariensis St. Hil. Other species of Ilex, as
well as unrelated plants such as species of Citronella, Rapanea, Rudgea (see
'Rubiaceae'), and Symplocos, have been used as substitutes according to
Lendner (1357) and Scala (2024). Yaupon or Yapon tea is the product of
Ilex cassine Walt.
The leaf of Ilex paraguariensis (Fig. 85 H and j) exhibits the following
structure. Cells of the upper epidermis with thick, straight anticlinal walls.
Cuticle thick and often striated, but the amount of striation somewhat variable,
Stomata confined to the lower surface, surrounded by about 3-5 ordinary
epidermal cells with sinuous anticlinal walls. Palisade tissue very variable
in different samples, usually consisting of about 3 layers of cells, but the
individual cells almost isodiametric in some samples and very elongated in
others. Fat bodies sometimes present in this tissue. Spongy mesophyll with
abundant intercellular spaces. Large cluster crystals fairly common in the
mesophyll as well as in the parenchyma of the midrib. Midrib with a single
vascular strand, consisting of a cylinder of xylem enclosing a lignified pith;
xylem almost or completely surrounded by phloem, the whole conducting
system being enclosed in a ring of sclerenchyma. The main vascular strand
of the petiole is similar to that of the midrib, but the xylem has the form of
an arc with very much incurved margins in some specimens. Small accessory
strands also occur in the petiole.
The European and American Hollies (Ilex aquifolium L. and /. opaca Ait.)
furnish white woods that are widely used for marquetry and inlay work and
are sometimes dyed to imitate ebony.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Byronia, Ilex,* Nemopanthus.
* Kew slide
Represented in the collection.

(ii)
FOR WOOD STRUCTURE
(Byronia), Ilex, (Nemopanthus), (Phelline).
4594 CC
386 AQUIFOLIACEAE
LITERATURE
(i) On General Anatomy
Holm 1053, Lendner 1357, Rivett 1943, Scala 2024, Weber 2380.
(ii) On Wood Structure
den Berger 179, 182, Brown, F. B. H. 280, Brown, H. P. 288, 289, Chalk and Rendle
365, Coster 481, Cozzo 494, Dadsweil and Record 533, Desch 574, Descole 576, Giordano
786, Greguss 2522, Howard 1088, Jones 1191, Kanehira 1206, 1209, Kribs 1283, Lecomte
1334, Record 1783, 1800, 1803, 1818, 1843, 1851, 1853, Record and Hess 1886, Record
and Mell 1894, Tang 2231, Tupper 2295, Williams 2430, Yamabayashi 2478.

94. CYRILLACEAE
(FiG. 89 on p. 388)

SUMMARY
A small family of shrubs from the
U.S.A., Brazil, and intervening regions.
Itsanatomy has been investigated by Solereder and Beauvisage (163). The
wood exhibits the following characters. Vessels very small and very numer-
ous; perforation plates exclusively scalariform with numerous fine, closely
spaced bars; intervascular pitting scalariform to opposite, pits to ray cells and
parenchyma similar to the opposite intervascular pits. Members of medium
length to moderately long. Parenchyma diffuse, or diffuse and scanty para-
tracheal. Rays mostly 4-5 cells wide, with few uniseriate rays and these

commonly only i or 2 cells high; heterogeneous. Fibres with numerous

distinctly bordered pits; of medium length.

LEAF
High, irregularly arranged ridges of cuticle on the lower side in Cliftonia
nitida Gaertn. f. Cells of the epidermis with straight anticlinal, sometimes
mucilaginous walls. Stomata in Cliftonia nitida and Cyrilla racemiflora Linn,
confined to the lower surface; surrounded by fairly numerous ordinary
epidermal cells. Mesophyll consisting wholly of palisade tissue in Cliftonia
nitida and Cyrilla racemiflora according to Solereder, but 2 layers of palisade
tissue present in Cyrilla according to Beauvisage (163). Vascular bundles
of the veins accompanied by sclerenchyma. Petiole of Cyrilla stated by
Beauvisage (163) to be complex in structure; exhibiting, in transverse sections,
a vascular strand having the form of a complete ring, depressed on the adaxial
side and accompanied by an accessory collateral strand, the whole vascular
system being surrounded by a continuous sheath of sclerenchyma. Both
solitary and clustered crystals occur; especially large clustered ones recorded
by Beauvisage (163) in Cyrilla.

Axis
YOUNG STEM
Cork very precocious in development, arising in the innermost part of the
primary cortex; accompanied by phelloderm. Primary cortex containing
numerous sclerosed parenchymatous cells with wide lumina; but soon
becoming detached owing to the early formation of the deep-seated cork.
Pericycle devoid of fibres. Parenchyma in the outer part of the phloem
becoming sclerosed after cork has been formed according to Solereder, but
 CYRILLACEAE 387
isolated and infrequent fibres stated by Beauvisage (163) to occur in the
phloem, Xylem including numerous fairly uniformly distributed vessels,
having scalariform perforation plates with 20-30 bars. Wood fibres with
bordered pits. Pith sclerosed.

WOOD (Fig. 89 F-G)

Vessels very small (25-50
/JL
mean tangential diameter), mostly solitary,
except for the apparent tangential pairs produced by overlapping ends, but
with a tendency to true tangential groups in the late wood of Cliftonia;
extremely numerous, especially in Cliftonia (about 140 per mm.). Occasionally
with a tendency to be semi-ring-porous (Heimsch 938). Perforation plates
exclusively scalariform, with up to 30-50 fine, closely spaced bars. Inter-
vascular pitting scalariform to opposite, small; pits to ray parenchyma cells
similar to the opposite intervascular pits pits to ray cells usually limited to
;

the upright cells; walls unusually thin. Mean member length about 0*8 mm.
(100). Parenchyma moderately abundant in Cyrilla (Fig. 89 F), diffuse, as
isolated cells scattered among the fibres; less abundant in Cliftonia and mostly
as occasional cells touching the vessels. Sometimes containing dark deposits.
Strands of 4-8 cells. Rays usually up to 4, occasionally 5 or 6 cells wide, and
up wide in a few species (1886); less than i mm. high; uniseriates
to 8 cells
i or 2 cells high and
few, often only composed almost entirely of square to
upright cells; about 6 rays per mm. heterogeneous (Kribs's Type II A, some-
;

times almost II B), usually with only a single marginal row of distinctly
upright dark deposits common. Heimsch (938) notes crystals in a
cells;
single species. Fibres with numerous distinctly bordered pits on both radial
and tangential walls, with thin to thick walls in different parts of the growth
ring; mean length about 1*0 mm. (100).

TAXONOMIC NOTES
The wood structure is of an unspecialized type.

GENERA DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Cliftonia and Cyrilla.
LITERATURE
(i) On General Anatomy
Beauvisage 163.
(ii) On Wood Structure
Heimsch 938, Howard 1088, Record 1783, 1843, z ^5 z f l8 53 Record and Hess 1886.

95. CELASTRACEAE
(Fic. 88 on p. 382; FIG. 90 on p. 392)

SUMMARY
(i) GENERAL
A
widely distributed family consisting mostly of small trees or shrubs but
a few species are climbers. Outstanding anatomical characters for the whole
family appear to be lacking. Whilst the xylem in young stems of most of
 M
FIG. 89. OPILIACEAE, A-C; AQUIFOLIACEAE, D-E; CYRILLACEAE, F-G.
GOUPIACEAE, H and L; HIPPOCRATEACEAE> I-J and M
A, Agonandra brasiliensis Benth. et Hook. B, Champereia manillana Merrill. C, Agonandra brasi-
liensis Benth. et Hook. D, Ilex mitis Radlk. E, 7. aquifolium Linn. F, Cyrilla racemiflora Linn.
G, Cliftonia ligustrina Spreng. H, Goupia glabra Aubl. I, Salacia megistophylla Standley. J, S.
reticulata Wight. K, Htppocratea malpighiaefolia Rudge. L, Goupia glabra Aubl. M, Hippocratea
malpighiacfolia Rudge.
c. Cell containing cystolith.
 CELASTRACEAE 389
the species which have been investigated is characterized by small vessels
which are frequently solitary and possess simple perforations, the genera
Kurrimia and Perrottetia are distinguishable from the other Celastraceae
examined in possessing scalariform perforation plates in the vessels. Accord-
ing to Solereder scalariform perforation plates also occur in Elaeodendron and
Glossopetalon, but their presence was not confirmed in Elaeodendron glaucum
Pers. by direct observation. Characteristic secretory sacs or canals filled
with granular material, which is stated to resemble rubber, occur in the
phloem of the axis as well as in the vascular bundles of the leaf veins in certain
genera. Hairs are infrequent, but when present are unicellular or uniseriate,
the former sometimes more like papillae. The epidermis of the leaf some-
times consists of several layers, or hypoderm may be present. The meso-
phyll contains sclerenchymatous idioblasts in certain genera, whilst fat bodies
occur in some of the cells in the same tissue.

(ii) WOOD
Vessels typically small, numerous and solitary, though multiples of 2-3
cells are characteristic of some
species; with flame-like arrangement in
Otherodendron ; sometimes ring-porous or semi-ring-porous, and occasionally
with spiral thickening; perforation plates usually simple but scalariform in a
few species; intervascular pitting and pits to ray and parenchyma alternate,
very small in most genera. Members of medium length to moderately long.
Parenchyma very variable in type and amount; commonly very sparse or
absent, diffuse or in broad multiseriate bands in some genera, vasicentric or
transitional between banded and aliform in a few others. Rays in some genera
exclusively uniseriate, homogeneous, and with conspicuous intercellular
spaces; in other genera, 2-8 (mostly 3-4) cells wide and markedly hetero-
geneous. Fibres. The ground tissue typically composed of fibre-tracheids
with numerous, distinctly bordered pits, but occasionally of septate fibres
with simple pits septate fibres often occur in multiseriate bands, comparable
;

in distribution to parenchyma bands; very occasionally with spiral thickening;

of medium length to moderately long.

LEAF
Generally dorsiventral, but isobilateral in certain species, for example, of
Gymnosporia, Maytenus, and Mortonia. Hairs infrequent; short unicellular
trichomes recorded in species of Euonymus, Fraunhofera, Mystroxylon\
papillose hairs of i or 2 cells in Tripterygium\ papilla-like trichomes accom-
panied by long hairs with thin transverse walls in Fraunhofera and Wimmeria\
unicellular hairs, sometimes 2-armed, in Myginda. Cuticle striated in certain
species of Microtropis, Polycardia, and Zinowiewia. (It is not quite clear what
is meant when Solereder describes the cuticle as being pitted in species of

Catha and Polycardia. Examination of a few species indicates that the cuticle
is
capable of becoming separated into 2 layers, also that there are small areas
rather suggestive of pits, in which the cuticle is thinner. Whether these thin
places in the cuticle represent the pits mentioned by Solereder is uncertain.)
Epidermis palisade-like in certain species of Cassine, Gymnosporia, Kokoona,
Maurocenia, Mortonia, Polycardia^ Pterocelastrus and Putterlickia\ consisting
>
390 CELASTRACEAE
wholly or locally of 2-3 layers in species of Catha, Elaeodendron, Gymnosporia,
and Plenckia. Cells of the upper epidermis occupy nearly half the width of
the lamina in Tripterygium forrestii Loes. Hypoderm frequently present,
either on the upper side alone or towards both surfaces, in species of Cassine,
Celastrus, Denhamia, Elaeodendron, Euonymus, Gyminda, Gymnosporia,
Maurocenia, Maytenus, Myginda, Mystroxylon, Rhacoma, and Schaefferia. In
the literature true hypoderm has probably not always been clearly distin-
guished from an epidermis of more than i layer. Epidermis containing scanty
or plentiful crystal cells (crystal idioblasts) in species of Catha, Denhamia,
Elaeodendron, Euonymus, Gyminda, Kurrimia, Lophopetalum, Maytenus,
Microtropis, Myginda, Pleurostylia, Siphonodon, Wimmeria (see also 'Crystals'
below). Stomata (Fig. 88 D) usually confined to the lower surface, but present
on the upper side as well in species of Gymnosporia, Maytenus, and Mortonia ;

usually cruciferous to ranunculaceous but rubiaceous in Kurrimia', guard cells

surrounded by a rosette of smaller epidermal cells in Mortonia. Guard cells
stated by Rehfous (1904) to become divided, the daughter cells being situated
below them, in species of Catha, Celastrus, and Euonymus. MesophylL
Palisade cells transversely divided in species of Mortonia and Pachystima;
homogeneous in species of Gymnosporia, Myginda, and Zinowiewia. Scleren-
chymatous idioblasts present in the mesophyll of species of Maurocenia,

Maytenus, Microtropis, Pterocelastrus, and Schaefferia. Vascular bundles of

the veins generally accompanied by sclerenchyma smaller veins vertically
;

transcurrent in species of Cassine, Plenckia, Tripterygium, and Wimmeria.

Terminal tracheids enlarged in Myginda and Schaefferia. Midrib with a
simple vascular strand in species of Catha and Euonymus with 2 central
;

bundles in Glyptopetalum and Microtropis. Vascular strand, in transverse

sections through the distal end of the petiole, appearing arc-shaped in most
of the plants investigated, which include species of Calha (Fig. 88 G), Celastrus
(ends of the strand very much incurved in some species), Euonymus (nearly
closed insome species), Glyptopetalum, Microtropis, Perrottetia, Tripterygium',
vascular strand closed in members of certain other genera including Cassine,
Elaeodendron, Lophopetalum, and Maytenus. Petiole of Kurrimia pulcherrima
Wall. (Fig. 88 B) with an almost closed but interrupted cylinder of xylem and
phloem surrounding 2 well-defined medullary bundles. Medullary bundles
in the petiole also recorded by Loesner (1389) in Lophopetalum. Accessory
bundles present outside the main petiolar strand in species of Celastrus,
Euonymus, Perrottetia, and Tripterygium. Pcricyclic region of the petiole
containing abundant sclerenchyma in species of Cassine, Catha, Celastrus,
Kurrimia, Maytenus sclerenchyma not well developed in, or almost absent
;

from, species of Celastrus, Elaeodendron, Euonymus, Glyptopetalum, Micro-

tropis, Perrottetia, and Tripterygium. 'Cortex' of petiole containing irregularly
shaped sclerotic cells in Maytenus sp. Long secretory sacs with rubber-like
contents (see also 'Axis') recorded in the pericycle of the veins in certain
species of Celastrus', similar sacs also recorded in the position normally
occupied by phloem fibres, and sometimes in the mesophyll as well in numer-
ous species of Wimmeria. Secretory canals with yellow contents stated to
accompany the veins and also to be situated at the leaf margins of Mortonia

greggii Gray; similar canals but devoid of contents recorded in the leaf
margins of certain species of Pachystima. Tanniniferous cells recorded in
 CELASTRACEAE 391
the mesophyll of species of Cassine, Euonymus, Gymnosporia, Maytenus,
Microtropis, Myginda, Pachystima, Siphonodon, Wimmeria, and Zinowieutia.
Similar cells observed in the 'cortex* of the petiole of species of Cassine,
Catha, and Elaeodendron. Crystals solitary or clustered, sometimes accom-
panied by crystal-sand cluster crystals often abundant in the cortical region
;

of the petiole. Solereder's more complete statements concerning their

occurrence and distribution need reinvestigation. Crystals stated to occur in
continuous layers of special cells in certain species of Pleurostylia and Ptero-
celastrus', sometimes present in septate palisade cells in Elaeodendron and
Mortonia or in crystal idioblasts in the mesophyll of Cassine, Catha, Celastrus,
and Euonymus. See also 'Epidermis* and 'Hypoderm'.

Axis
YOUNG STEM
Cork usually arising in the sub-epidermis, but originating in the epidermis
itself in Euonymus and Microtropis; formed in the outer part of the primary
cortex in Elaeodendron and Lauridia, and in the inner part of the cortex in
Myginda and Tripterygium. Cork cells often tabular and thin-walled, but
thickened tangential walls occur in species of Celastrus, Elaeodendron,
Kurrimia, Maytenus, Microtropis, and Tripterygium (very thick- walled, pitted
cells). Corky outgrowths, resembling but apparently distinct from lenticels,
stated to arise from the cortical parenchyma of Euonymus verrucosa Scop.
Cortex frequently containing tanniniferous cells, notably in species of Cassine,
Catha, Celastrus, Elaeodendron, Gymnosporia, Microtropis. Stone cells present
in the cortex of species of Cassine, Elaeodendron, Maytenus, Microtropis, and
bundles of fibres in the same region in Euonymus. Outer layers of the cortex
consisting of palisade cells in Euonymus americana Linn, but not in E. atropur-
purea Jacq. Pericycle often containing strands of fibres, notably in species
of Catha, Elaeodendron, Euonymus, and Perrottetia; with a composite and
continuous ring of sclerenchyma in species of Cassine, Celastrus, Fraunhofera,
Glyptopetalum, Kurrimia, Lophopetalum, Maurocenia, Maytenus, Microtropis,
and Tripterygium pericyclic fibres apparently absent from certain species of
;

Celastrus and Euonymus, but stated to occur in the form of a continuous ring
in other members of the last genus. Phloem and xylern usually constituting
closed cylinders traversed by narrow rays, but stated by Colas (446) to be in
the form of bundles separated by conspicuous medullary rays in Maytenus sp.
Phloem containing large stone cells in old material of Euonymus europaeus
Linn, and in Kurrimia pulcherrima Wall. Fibres stated by Beau visage (163)
to occur in the phloem of species of Catha. Xylem including vessels which
are commonly of uniform, small diameter, notably in species of Cassine, Catha
(radial diameter seldom exceeding 33 /x in first-year stems of Catha edulis
Forsk.), Celastrus, Elaeodendron, Euonymus (radial diameter seldom exceeding
50 p, in first-year sterns of Euonymus europaeus), Maytenus] mostly solitary or
in pairs and up to about 85 p in radial diameter in Tripterygium forrestii Loes. ;
in radial rows of Kurrimia pulcherrima Wall, and tending to be similar in
Microtropis discolor Wall. Vessels of the first year's wood in certain species
of Celastrus are very small and infrequent compared with those developed
during subsequent seasons. Perforations believed to be exclusively simple in
392 CELASTRACEAE
species of Cassine, Celastrus, Elaeodendron, Euonymus, Microtropis, and
observed in Kurrimia and
Tripterygium, but scalariform perforation plates
Perrottetia. Pith generally homogeneous, but frequently heterogeneous in

FIG. 90. CELASTRACEAE

E. europaeus Linn. D, Lopho-
A, Celastrus peduncularis Sind. B, Euonymus europaeus Linn. C,
Eckl. ct ^eyh.
petalum fimbriatum Wight. E, Myginda eucymosa Loss et Pitt. F, Elaeodendroncapense
G, ,E. dEpns Eckl. et Zeyh. H, Kurrimia paniculata Wall. I, /C. paniculata Wall. J, Pterocelastrus
tricuspidatus Walp.

and
Gymnosporia, Lophopetalum, Microtropis, Perrottetia, Polycardia,
celastrus; sometimes containing stone cells in Kokoona
and Maytenus; tannini-
ferous cells in species of Cassine, Catha, Celastrus, Elaeodendron, Microtropis,
and cells sometimes occur in the phloem
Tripterygium. Elongated secretory
 CELASTRACEAE 393
of species of Catha, Celastrus, Elaeodendron (very elongated, comparable with
canals), and Microtropis. See also 'Cortex* above. Secretory cells observed
in the phloem of Celastrus rugosus Rehd. et Wil. and Euonymus europaeus Linn.
The secretory cells were examined particularly in Euonymus europaeus,
where they are apparently absent from the very young stem but occur in the
secondary phloem of older material, especially when this tissue clearly con-
stitutes part of the bark. They appear to have rare transverse septa and to be
filled with granular material, stained yellow by chlor-zinc-iodide.

WOOD (Fig. 90)

Vessels typically very small (less than 50 /x mean tangential diameter),
extremely small in some genera, e.g. Gymnosporia, Maurocenia, and Plenckia,
moderately small (100-200 p) in Caryospermum, Kurrimia, Lophopetalum, and
Perrottetia exclusively solitary, or nearly so, in Cassine, Celastrus, Elaeoden-
;

dron, Lophopetalum littoralis Ridl. and L. maingayiRidL, Maurocenia, Mortonia,

Myginda, Plenckia, Pleurostylia, Perrottetia, Pterocelastrus, and Schaefferia,
radial flames reported in Otherodendron (Kanehira 1209), radial multiples
of 2 or 3 cells common in the remainder, sometimes up to 4 cells in some
species of Lophopetalum and Maytenus; typically numerous; fewer than 5 per
sq. mm. in Perrottetia, 5-20 per sq. mm. in Kookoona, Kurrimia, Lopho-
petalum, and Siphonodon, 20-40 per sq. mm. in Caryospermum, Cassine,
Maytenus p.p., Myginda, and Pterocelastrus, 40 to over 200 in the remaining
genera; ring-porous or semi-ring-porous in Celastrus (Record 1853), Gymno-
sporia buxifolia (L.) Szysz., Maurocenia, Mortonia, Pachystima (Record 1853),
Plenckia, and Rhacoma', spiral thickening sometimes present in Austroplenckia
(1886), Celastrus, Euonymus, Maytenus p.p., Mortonia (1886), Pachystima
(1886), and Plenckia\ Solereder records spiral thickening also in Tripterygium
and Record (1851) in Lophopetalum, though the latter could not be confirmed
in the material examined. Perforation plates scalariform in Cassine p.p.,
Elaeodendron, Kurrimia, and Perrottetia with fewer than 10 bars, except in
1

Perrottetia (1209). Solereder quotes Loesner as describing the perforations

of Cassine as exclusively simple, which though applicable to C. capensis Linn.,
is not true of C. crocea (Thunb.) O. Kze. in which the perforation plates are

exclusively scalariform. Intervascular pitting alternate, typically very small

except in Caryospermum, Euonymus, Kurrimia, Mortonia, and Perrottetia, pits
to parenchyma and rays cells similar to the intervascular pitting except in
Kurrimia, in which they are commonly unilaterally compound or large and
elongated. Tibriform vessel members* reported (1886) in Euonymus and
Pachystima. Mean member length o-6-i-o mm. Parenchyma typically
apotracheal or absent; diffuse in Denhamia, Maytenus p.p., Myginda (Fig. 90 E),
Rhacoma, Schaefferia, 2 and Wimmeria, in regular multiseriate bands, frequently
up to 4 or more cells wide, in Celastrus p.p. (Fig. 90 A), Elaeodendron p.p.,
Gymnosporia p.p., Kookoona, Lophopetalum (bands 1-2 cells wide), Maytenus
p.p., Pleurostylis, Pterocelastrus, and Torralbasia; with similar bands, but com-
posed of septate fibres instead of parenchyma, in several genera (see below
under 'Fibres'); parenchyma absent from or very sparse in Austroplenckia
1
With simple perforation in a single species of Perrottetia from New Guinea.
2
Record and Garratt (1884) describe the parenchyma in Schaefferia frutescens Jacquin as
'apparently absent'.
394 CELASTRACEAE
(1886), Caryospermum (1154), Celastrus p.p. (1886), Elaeodendron p.p.,
Euonymus, Maytenus p.p., Mortorda, Pachystima (1886), and Siphonodon',
vasicentric in Perrottetia\ transitional between aliform and narrow bands in
Kurrimia (Fig. 90 i). Seldom containing crystals. Strands usually of up to
4 or 8 cells.

Rays r predominantly uniseriate in Austrtplenckie (1886),

exclusively
Eutnymus, H&rtogia, Kookoona, Lophopetalum; some rays up to 2 cells wide
in Maurocenia, Microtropis (1886), Mortonia, Myginda, Pachystima (1853),
PerrMetin, Rhacoma, Torralbasia, and Wimmeri&\ usually 3-6 cells wide in
the *ther genera, but sometimes more in species of Celastrus and occasionally
nly 2 cells wide in Plenckia\ Janssonius (i 154) classes the rays as of 2 distinct
sizes some species of Caryospermum, Elaeodendron, Microtropis, and
in
Siphonodon', commonly about i mm. in height and slightly exceeding this in
some species; species with multiseriate rays also have numerous uniseriates
composed wholly of upright cells; uniseriate rays only i or 2 cells high
common in Celastrus p.p., Elaeodendron p.p., Gymnosporia, Maurocenia, and
Pterocelastrus p.p.; uniseriate rays rare in Celastrus p.p., Kurrimia, and
Maytenus p.p.; more than 12 rays per mm. in most species and up to 20 or
more per mm. in Kookoona, Plenckia, and Pterocelastrus fewer than 12 per
;

mm. in some species of Kurrimia, Maytenus, and Siphonodon] homogeneous

(Kribs's Type III), and with rounded cells (tang, section) and conspicuous
intercellular spaces in the woods with exclusively uniseriate rays, except in
Hartogia (heterogeneous Type III); similar, but less marked, intercellular
spaces present also in Maurocenia, Maytenus p.p., and Pleurostylia; distinctly
heterogeneous (Kribs's Types I and II A) in the genera with multiseriate rays,
with 4 or more marginal rows of upright cells except in Denhamia, Maytenus
p.p., Mortonia, Pterocelastrus p.p., and Wimmeria, and commonly with 10 or
more rows in some species of Cassine, Catha, Elaeodendron, Gymnosporia,
Rhacoma, and Siphonodon groups of multiseriate procumbent cells sometimes
;

alternating with, and not much wider than, the uniseriate upright cells, e.g. in
Catha, Elaeodendron p.p., and Pleurostylia. Single crystals occur in the
ordinary cells of most species and large crystals in idioblasts occur in Siphono-
don celastrineus Griff. dark deposits present in some species. Fibres of most
;

species with numerous, distinctly bordered pits on all walls, the borders often
very large, but rather smaller in Gymnosporia p.p.; borders absent from
or very indistinct in Perrottetia and Siphonodon', with occasional septa in
Euonymus; septate fibres with simple, slit-like pits, form the ground tissue in
Caryospermum (1154) and Siphonodon and multiseriate bands of septate
fibres occur the fibre-tracheids in Cassine, Catha, Celastrus p.p.,
among
Elaeodendron Fraunhofera (2158),
p.p., Gymnosporia p.p., Hartogia,
Maurocenia, Maytenus p.p., and Plenckia; these bands of septate fibres are
exactly comparable in distribution with the multiseriate bands of parenchyma
occurring in other species (cf. A and G in Fig. 90), and in a single genus the
bands may be composed of septate fibres in one species and of parenchyma in
another; with spiral thickening in Euonymus, except E.javanicus Blume, and
very occasionally in Pachystima (Record 1853), walls varying from thin,
e.g. most species of Lophopetalum, to extremely thick, e.g. Kurrimia and
Pterocelastrus', mean length o-8-i'8 mm. Vasicentric tracheids present in
Microtropis bivalvis Wall. (1154).
 CELASTRACEAE 395
ROOT
Structure described by Colas (446) for Maytenus sp. as follows: Cork
consisting of 12-15 layers of sub-hexagonal cells. Phelloderm well developed,
containing sclerotic idioblasts. Phloem, in transverse sections, appearing as
elongated conical groups of rounded cells and sieve tubes separated by
triseriate rays. Hexagonal fibres also present towards the centre. Xylem

consisting of rounded or sub-hexagonal vessels, irregularly distributed in

lignified parenchyma. Vessels frequently containing tyloses. Ground tissue
of older roots stratified into zones of lignified parenchyma, alternating with
hexagonal fibres. Tannin abundant in the tissues. The root structure in
species of Cassine, Ccl&strus, Elae&dendr&n, Euonymus, and Maytenus has been
described by Weber (2380).

ANOMALOUS STRUCTURE
Excentric growth in thickness recorded in the stem and root of certain
species of Maytenus.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
It was pointed out by Dunn (617) that Kurrimia, Perrottetia, and Triptery-
gium differ considerably from one another and also from the remainder of the
Celastraceae. is reflected to some extent in the anatomical
This distinction
structure. Dunn
drew attention to the lack of taxonomic agreement
also
between Dipentodon and Goupia, both of which were included amongst the
Celastraceae in the Bentham and Hooker system, and the remainder of the
family. In discussing the taxonomic positions to which these genera should
be assigned, Dr. T. A. Sprague has expressed the view (not published) that
there is no reason for excluding Tripterygium from the Celastraceae so far as
external morphology is concerned. He points out that it is the type genus of
Loesner's (1389) sub-family Tripterygioideae. Very little is known at present
concerning the anatomy of the other genera included in this sub-family. In
Dr. Sprague's opinion Perrottetia and Kurrimia are not sufficiently distinct
for them to be excluded from the Celastraceae. Goupia, on the other hand, is
more definitely aberrant, and has been described in this book under the
distinct family Goupiaceae, while Dipentodon has been dealt with under
Flacourtiaceae. For further views concerning Goupia see p. 398.

(ii) FROM WOOD STRUCTURE

The wood anatomy seems tobear little relation to the sub-families and the
whole family appears to be rather heterogeneous. Broad bands of either
parenchyma or septate fibres are a characteristic feature of Gassine, Catha,
Celastrus, Elaeodendron Gymnosporia, Hartogia, Kookoona, Lophopetalum,
y

Maurocenia, Maytenus, Plenckia, Pleurostylia, Pterocelastrus, and Torralbasia

a list that is composed mainly of Eucelastreae and Eucassineae. Diffuse
parenchyma is present in Denhamia, Maytenus, Myginda, Rhacoma, Schaef-
feria and Wimmeria i.e. mostly from the same groups but with Wimmeria
t

(Tripterygioideae) in addition; parenchyma absent from Euonymus and

Mortonia. This leaves Caryospermum, Kurrimia, Perrottetia^ and Siphonodon
as odd genera with paratracheal parenchyma and without fibre tracheids.
396 CELASTRACEAE

ECONOMIC USES
The root bark of Euonymus americana Linn, and E. atropurpurea Jacq.,
which was at one time reputed to possess medicinal properties, has been
described anatomically by Holm (1035). Members of the same genus have
also aroused interest in Russia as a possible source of rubber. An oil with
medicinal properties is obtained from the seeds of Celastrus paniculata Willd.
R. H. et
According to Colas (446) the medicinal bark of Maytenus chuchuasha
R. Colas from the Amazon region exhibits the following anatomical characters.
Cork consistingof 7-8 rows of cells with thin, sinuous, suberized walls, and
the phelloderm of 10-12 rows of cells. Region below the phelloderm including
numerous sclerotic cells. Phloem arranged in tangentially elongated strands,
separated by rays 1-3 cells wide,
and differentiated into the following con-
centric zones: (i) The innermost consisting of polygonal cells arranged in
radial rows, (ii) The
second somewhat collenchymatous, consisting of rounded
elements in radial rows, with sieve tubes present amongst them, the whole
zone being stratified by strands of hexagonal fibres with small lumina. (iii) The
third zone similar in structure to the innermost one, but cells more collenchy-
matous. (iv) The fourth zone composed of radially elongated bands of phloem
radial bands
tissue, consisting of collenchymatous elements accompanied by
of regularly arranged fibres, separated by large ovoid cells. Pyramidal
crystals and tanniniferous cells present, the latter chiefly in the region
between the cork and the cones of phloem elements,
Arabian or African tea, which consists of the leaves and twigs of Catha
edulis Forsk., exhibits the following leaf structure. Cuticle on both surfaces
with somewhat sinuous anti-
very thick. Epidermal cells from both surfaces
clinal walls. Stomata confined to the lower surface, surrounded by 3-5

ordinary epidermal Palisade tissue consisting of 1-3 layers in different

cells.

specimens. Spongy mesophyll often occupying

more than half the width of
the lamina. Tanniniferous cells, with yellowish-brown contents, scattered
throughout the lamina, but particularly numerous in the cortical region and
in the
phloem of the midrib. Cluster crystals present with varying frequency
veins
mesophyll and epidermis. Vascular bundles
of the smaller accompanied
strand of the
by thick-walled fibres, especially on the lower side. Vascular
midrib, in transverse sections, appearing arc-shaped with
incurved ends, and
a broad band of thick-walled fibres on the abaxial side.
accompanied by
Smaller subsidiary strands with similar structure sometimes present in the
midrib as well. Petiole with abundant tanniniferous cells in the cortex, and
vascular strand; pericyclic sclerenchyma
supplied by a crescent-shaped
poorly developed (Fig. 88 G).
Few of the species of this family are large enough to produce timber, but
two species of Lophopetalum furnish soft, even-textured, and easily worked
timbers that are used in India, and some species of Maytenus are used locally
in South America. Some of the smaller trees and shrubs produce dense woods
that are used locally as boxwood substitutes and the wood of Euonymus spp.
is used for 'orange' manicure sticks and for small articles of turnery.
 CELASTRACEAE 397
GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Cassine,* Catha,* Celastrus,* Denhamia, Elaeodendron,* Euonymus,*
Fraunhofera, Glossopetalon, Glyptopetalum, Gyminda, Gymnosporia,
Kokoona, Kurrimia,* Lauridia, Lophopetalum, Maurocenia, Maytenus,
Microtropis,* Mortonia, Myginda, Mystroxylon, Pachystima, Perrottetia,*
Plenckia, Pleurostylia, Polycardia, Pterocelastrus, Putterlickia, Rhacoma,
Schaefferia, Siphonodon, Tripterygium,* Wimmeria, Zinowiewia.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Austroplenckia, Caryospermum, Cassine, Catha, Celastrus, Denhamia,
Elaeodendron, Euonymus, Gymnosporia, Hartogia, Kookoona, Lophope-
talum, Maurocenia, Maytenus, (Microtropis), Mortonia, Myginda, (Othero-
dendron), (Pachystima), Perrottetia, Plenckia, Pleurostylia, Pterocelastrus,
Rhacoma, Schaefferia, Siphonodon, Torralbasia, (Tripterygium), Wimmeria.
LITERATURE
(i) On
General Anatomy
Beauvisage 163, Colas 446, Dunn 617, Holm 1035, Loesner 1389, Rehfous 2904,
Sabnis 1977, Weber, W. 2380.

(ii) On Wood Structure

Benoist 170, den Berger 182, Besson 186- Burgerstein 312, Chalk and Chattaway 358,
Desch 574, Greguss 2522, Janssonius 2147, ZI 54 Jolly 1188, Kanehira 1206, 1209,
Martin -Levigne 1450, McNair, G. T. 1472, M6niaud 1492, Pearson and Brown 1679,
Pfeiffer, H. 1712, Pfeiffer, J. Ph. 1713, Record 1783, 1800, 1843, 1851, 1853, 1884,
Record and Hess 1886, Record and Mell 1894, Stone 2202, 2207, Tang 2231, Yamabayashi
2478.

96. GOUPIACEAE
(Fie. 88 on p. 382; FIG. 89 on p. 388)

SUMMARY
Small to very large trees belonging to the single genus Goupia which occurs
in the Amazon region, Colombia, &c. An interesting anatomical character is
the complex vascular structure of the petiole. The wood exhibits the follow-
ing features. Vessels medium-sized and solitary, perforation plates scalar!-
form, pits to parenchyma small, often with coalescent apertures and uni-
laterally compound, members very long. Parenchyma apotracheal, scattered
among the fibres, and paratracheal, vasicentric to aliform. Rays 2-5 cells
wide; markedly heterogeneous, relatively small multiseriate groups of pro-
cumbent cells often separated by many uniseriate upright cells. Fibres with
bordered pits, moderately to very long. Vasicentric tracheids present.

LEAF
Hairs sparse, consisting of short, unicellular trichomes with
Dorsiventral.
relatively thick walls. Epidermis on the upper surface consisting of several
layers of cells separated by thin walls; cells of the outer epidermal layer on
both surfaces polygonal, with almost straight but slightly thickened and pitted
anticlinal walls. Stomata confined to the lower surface, each surrounded by
about 4 epidermal cells, not very clearly differentiated from those of the
398 GOUPIACEAE
remainder of the epidermis (ranunculaceous). Mesophyll composed of about
2 layers of palisade tissue, and a broader spongy region; containing branched
sclerenchymatous elements (idioblasts). Vascular bundles of the smaller
veins embedded in the mesophyll larger veins with cylindrical xylem accom-
;

panied by small phloem groups, the whole being surrounded by a broad ring
of fibres. Main vascular strand of the petiole (Fig. 88 A) appearing, in trans-
verse sections through the distal end, as an abaxial arc of xylem and phloem,
accompanied by an arc of closely placed bundles on the adaxial side, the whole
strand resembling an almost closed circle surrounding about 4 medullary
bundles and accompanied externally by cortical strands. Cortical region of
the petiole containing tanniniferous cells. Cluster crystals observed.

Axis
YOUNG STEM
Cork superficial in material examined at Kew. Cortex narrow. Pericycle
containing a composite and continuous ring of sclerenchyma. Xylem with
numerous, evenly distributed, mostly solitary vessels, up to about 100 ft in
radial diameter, with scalariform perforation plates. Pith small, quadrangular,
somewhat heterogeneous, including cells with lignified pitted walls and others
with granular contents. Cluster crystals observed.

WOOD (Fig. 89 H-L)

Vessels medium-sized (100-200 p, mean tangential diameter) or slightly
larger; exclusively solitary; about 10 per sq. mm. Perforation plates scalari-
form with 3-10 thick bars. Intervascular pitting very rare, pits to ray and
wood parenchyma small and often with coalescent apertures, sometimes
compound, several pits subtending a narrow, elongated pit in the
unilaterally
parenchyma wall, particularly in the wood parenchyma cells, such pits
horizontal, vertical, or oblique. Solid deposits moderately common. Mean
member length about i -7 mm. Parenchyma scattered among the fibres and
associated with the vessels, the latter varying from a few cells on the abaxial
sides to complete sheaths, and often forming distinct, narrow wings (Fig. 89 H).
Strands composed of 8 or more cells. Rays 2-5 cells wide and commonly
more than i mm, high; uniseriates only moderately numerous, composed of
high upright cells; about 14 rays per mm.; markedly heterogeneous (Kribs's
Type I)>and consisting of z or 3 multiseriate portions of procumbent cells
separated by several rows of uniseriate upright cells (Fig. 89 L) the biseriate
;

parts scarcely any wider tangentially than the uniseriate parts with occasional
;

sheath cells in the larger rays; sometimes with occasional solitary crystals in
the upright cells. Fibres with numerous distinctly bordered pits on both
radial and tangential walls, the borders larger than those of the intervascular
pitting; walls very thick; mean length about 2*2 mm. Vasicentric tracheids
present.
TAXONOMIC NOTES
Goupia was included in the Celastraceae in the Bentham and Hooker
system, but was given the status of a sub-family Goupioideae within the
Celastraceae by Loesner (1389). In Dr. T. A. Sprague's opinion the genus
should be restored to the status of a family as recommended by Miers (15 16 A).
Dr. Sprague has pointed out (unpublished) that the venation of the leaves, the
 GOUPIACEAE 399
stipules, the aestivation of the petals, the anthers, and the gynoecium are
highly characteristic and taken together are sufficient to exclude Goupia from
the Celastraceae. For these reasons, together with the petiolar structure,
which is unlike that of the Celastraceae (with the possible exception of
Kurrimia and Lophopetalum), Goupia has been treated as a separate family in
this book. The wood is of an unspecialized type.

ECONOMIC USES
Goupia glabra Aubl. in British Guiana produces a strong, durable timber
suitable for heavy construction.

GENUS DESCRIBED
The above
description of the general anatomy is based on an examination
of Goupia glabra AubL* in the Kew herbarium.
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Locsncr 1389, Miers 1516 A.

(ii) On Wood Structure

Janssonius 1154, M6niaud 1491, Record 1853, Record and Hess 1886, Record and Mell

97. HIPPOCRATEACEAE
(Fie. 88 on p. 382 ;
FIG. 89 on p. 388 ; FIG. 91 on p. 402)

SUMMARY
(i) GENERAL
A
tropical family of small trees or shrubs, sometimes scandent. The
anatomical characters are very similar to those of the Celastraceae. One of
the most characteristic features is the occurrence of laticiferous canals
which are confined to the axis of some species, but occur in the leaf as well in
others. Some of the mesophyll cells also contain a material resembling rubber.
Although laticiferous elements are so characteristic of the family they are not
always easily seen, especially if the contents have been dissolved when pre-
paring and mounting the sections. Sclerosed cells of various kinds are
common in the leaf. The petiole structure exhibits a considerable range of
variation in different species (see 'Leaf'). Anomalous thickening has been
recorded in Salacia.

(ii) WOOD
Vessels medium-sized to large, usually solitary, perforations simple, pits
to parenchyma usually small; members of medium length. Parenchyma
paratracheal, rather sparse. Rays exclusively uniseriate or of 2 distinct sizes,
the larger up to 30 cells wide and very high. Fibres of 2 types, those of the
ground tissue with bordered pits and thick walls; scattered amongst these are
thin-walled septate fibres with various parenchyma-like distributions,
moderately short. Included phloem of the foraminate, concentric or
interrupted type present in several species.
400 HIPPOCRA TEACEAE
LEAF
Generally dorsiventral rarely centric in Hippocratea and Salacia. Hairs
;

stellate, with uniseriate rays in Hippocratea velutina Afzel.; imiseriate, but

with closely placed septa at the base in Hiotricha Loes. unicellular, papillose
.
;

in H. aspera Lam. Cork warts present on the lower surface in certain species
of Salacia. Epidermis containing crystal cells (idioblasts) in most species of
Hippocratea and Salacia. Outer walls of the epidermal cells stated to be
pitted in Salacia oblonga Wall. Stomata usually confined to the lower surface,
but exceptions recorded in a few species of Campylostemon and Hippocratea ;

mostly ranunculaceous, but instances recorded in all 3 genera of the guard

cells being surrounded by i or 2 (Salacia dulcis Benth.) circles of 4 cells each,
2 cells in each ring being parallel to the pore. Hypoderm of i or 2 layers
common below the upper surface of the leaf in Campylostemon^ Hippocratea,
and Salacia. Some of the mesophyll cells contain deposits of a rubber-like
substance. Mesophyll including various types of sclerosed cells, such as the
'spicular cells*, connected with the sclerenchyma of the veins in species of
Salacia. Vascular system of the petiole very variable, exhibiting the follow-
ing types of structure in transverse sections, (i) A
horseshoe-shaped group of
bundles in some species of Hippocratea and Salacia. (ii) A ring of bundles
surrounding an inversely orientated plate of xylem and phloem in S. micrantha
Peyr. (iii) An arc-shaped strand towards the abaxial side, but having an
adaxial plate of xylem and phloem partly surrounded by the arc in S. rox-
burghii Wall. (Fig. 88 c). (iv) With a flattened ring of bundles accompanied
on the upper side by 3 or 4 inversely orientated bundles in S. crassifolia Peyr.
Tanniniferous cells occur in the mesophyll of species of Hippocratea and
Salacia. Large secretory cavities present in the cortical tissue of the petiole
and in the margins of the lamina of S. roxburghii. Laticiferous canals (see
*
also Axis') have been recorded in species of Hippocratea and Salacia as well
as in one of Campylostemon where they accompany the phloem of the veins
and usually extend into the mesophyll. Mesophyll also containing mucilage
cells in Hippocratea velutina Afz. Solitary and clustered crystals common ;

see also under 'Epidermis'.

Axis
YOUNG STEM
Cork arising in the second to the fourth layer of the cortex. Cork cells
with fairlywide lumina; often thin walled or with strong thickening of the
inner or outer tangential walls. Primary cortex frequently including branched
stone cells, especially in the inner portion. Pericycle with isolated strands
of fibres in some species, or a composite and continuous ring of sclerenchyma
in others. Phloem
containing fibres and branched sclerenchymatous cells.
Xylem usually including vessels with simple perforations; scalariform plates
also recorded in i species of Salacia. Pith sometimes containing stone cells.
Laticiferous canals (see also under 'Leaf') recorded in species of Hippo-
cratea and Salacia and in one of Campylostemon^ generally situated in the
phloem, or, more rarely, in the primary cortex. Phloem also containing tannin
sacs in Hippocratea and Salacia. Solitary and clustered crystals common,
solitaryones sometimes very abundant. Solitary crystals in the cortex and
pith much larger than those in the phloem in Salacia roxburghii Wall.
 HIPPOCRATEACEAE 401
BARK
Bark in many members of the family stated by Loesner (1390) to contain a
reddish-yellow pigment, soluble in alcohol.
WOOD (Fig. 89 i, j, and M, Fig. 91 c)
Vessels medium-sized (100-200 /x mean tangential diameter) in Cheilo-
clinium and Salacia, large (more than 200 p) in Hippocratea in the material
examined, but reported by Record (1853) to ke minute in some species;
exclusively solitary or almost so, except in Pristimera (1886); very variable in
number even in different areas of the same stem, average about 10-16 per
sq. mm. Perforations simple; intervascular pitting rare, pits to parenchyma
small, often with striations due to coalescent, slit-like apertures, sometimes
unilaterally compound or horizontally elongated in Hippocratea. Tyloses not
observed; gum present in some species of Salacia. Mean member length
0-5-0-7 mm. Parenchyma exclusively paratracheal, as a few cells round the
vessels; strands of 4-12 cells. Rays in Cheilodinium p.p. and Salacia ex-
clusively uniseriate, about 20 per mm. and composed entirely of square or
upright cells (Kribs's Heterogeneous Type III); of 2 distinct sizes in Cuervea
and Hippocratea, the larger up to 30 cells wide and of indeterminate height,
the smaller rays uniseriate; about 6-10 rays per mm., heterogeneous (Kribs's
Type II A). Crystals and gum present in both genera. Record and Hess (i 886)
note the occurrence of patches of thin-walled cells in some genera. Fibres of
the ground tissue with conspicuous bordered pits which are usually larger
than the intervascular pitting and equally numerous on both radial and
tangential walls; interspersed among these fibres, with a parenchyma-like
distribution, are thin-walled septate fibres with simple or inconspicuously
bordered pits; as scattered cells or short uniseriate lines comparable with
diffuse parenchyma in Cuervea and Hippocratea, aliform to confluent in
Cheilodinium and Salacia; these cells may be abundant particularly in the
first-formed rings of secondary wood (362). Mean length 0-6-0*9 I*1mm
Cheilodinium and Salacia the fibre-tracheids round the vessels have wider
lumina and very numerous pits and grade into vasicentric tracheids. Vasi-
centric tracheids usually present. Included (interxylary) phloem of the
concentric type (c. I. circumvallatum) occurs in Cheilodinium gleasonianum
A. C. Sm. (1886) and Salacia (Fig. 91 c), with successive layers of xylem and
phloem separated by broad bands of conjunctive parenchyma containing
circular or irregular cells ; the xylem and phloem layers are
groups of stone
only rarely interrupted by interfascicular rays. Record and Hess (1886)
report anomalous structure of the 'foraminate' type (c. L foraminulatum) in
Cheilodinium cognatum (Miers) A. C. Sm. and Prionostemma aspera (Lam.)
Miers. Anomalous structure of the 'interrupted* type (c. L interruptum)
reported (1853) m Hippocratea^ 'the breaking of the cambial ring into several
arcs results in the formation of unequal amounts of xylem and phloem, and
the stem becomes more or less deeply grooved'; with a tendency to such
formation also in Hemiangium (1886).

ANOMALOUS STRUCTURE
Successive rings of interxylary phloem recorded in Salacia (Fig. 91 c). For
further details see preceding paragraph.
 MENISPERMACEAE, A; POL YGALACEAE, B; HIPPOCRATEACEAE,
FIG. 91. C;
MALPIGHIACEAE, D; APOCYNACEAE> E; CONVOLVULACEAE, F
Anomalous stems. A, Anomospermum grandifolium Eichl. B, Securidaca lanceolata St. Hil. Xylem
white; phloem and cortex shaded; nat. size. C, Salacia serrata Camb. Xi^. Dr Tetrapteris sp. showing
split xylem (Schenck Holzs. 500 Rio) nat. size. E, Condylocarpus sp. Cortex and interxylary phloem
shaded; pith and perimedullary xylem indicated by circles. F, Parana volubilis Burm. nat. size.
E, After Fritz Miffler. Remainder after H. Schenck.
 HIPPOCRA TEA CEAE 403
TAXONOMIC NOTES
The
general similarity of the anatomical characters to those of a majority of
the Celastraceae indicates that the two families are closely related to one
another. It is noteworthy that the unusual parenchyma-like distribution of
septate fibres in the xylem of the Hippocrateaceae has a parallel in the
Celastraceae. This close relationship seems to be supported also by the
evidence of external morphology. Smith and Bailey (2141), in discussing
the genus Brassiantha, have pointed out that the division between the Hippo-
crateaceae and Celastraceae is artificial.

ECONOMIC USES
According to Colas (446) one or more species of Salacia with medicinal
properties occur in the Amazon region. No anatomical details are given by
Colas.
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Campylostemon, Hippocratea, Salacia.*
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Cheiloclinium, Cuervea, (Hemiangium), Hippocratea, (Pristimera), Salacia.

LITERATURE
(i) On General Anatomy
Colas 446, Loesner 1390, Smith, A. C. and Bailey 2141.

(ii) On Wood Structure

Chalk and Chattaway 362, Record 1851, 1853, Record and Hess 1886, Williams 2430.

98. STACKHOUSIACEAE
SUMMARY
A
small family of herbs with woody rhizomes comprised in the two genera
Macgregoria and Stackhousia which occur chiefly in Australia, New Zealand,
&c. Tanniniferous cells occur in the parenchymatous tissues of both leaf
and stem. Deposits of a rubber-like substance have been reported, but
according to Solereder these are probably fat bodies. It is interesting to note
that crystals of calcium oxalate have not been observed in the family.

LEAF
Dorsiventral or centric. Hairs unicellular. Stomata in Stackhousia
spathulata Sieb. present on both surfaces ranunculaceous. Mesophyll con-
;

taining tanniniferous cells in Stackhousia brunonis (Endl.) Benth. Bodies,

variously interpreted as consisting of fat or rubber, also occur in the leaf.
Vascular bundles of the veins, not accompanied by sclerenchyma.

Axis
YOUNG STEM
Hypoderm usually well defined. Outer part of the cortex containing
abundantly pitted fibres in positions corresponding to ribs on the surface of
404 S TA CKHO USIA CEA E
the stem in all species of Stackhousia except S. pulvinaris F. v. M. as well as
in Macgregoria. Endodermis frequently composed of large, conspicuous
cells. Pericycle usually containing isolated bundles of fibres, but scleren-

chyma is lacking in S. maideni Pampan. and S. pulvinaris. Xylem containing

vessels with simple perforations, a small amount of parenchyma, fibres with
bordered pits; rays stated to be absent (cf. woody species of Veronica). Pith
composed of large parenchymatous cells. Tanniniferous cells common in
the cortex,

RHIZOME
Secretory cells, having wide lumina filled with dark-brown contents,
either solitary or in longitudinal rows of a few together, recorded in the phloem.

TAXONOMIC NOTES
The
Stackhousiaceae are generally regarded as having affinities with the
Celastraceae, If the presence of rubber-like deposits in the Stackhousiaceae
could be more definitely established this would provide additional evidence
of the affinities of the family since rubber-like substances occur also in the
Celastraceae.

GENERA DESCRIBED
Macgregoria, Stackhousia,
The above description is largely based on the one previously given by
Solereder.

99. RHAMNACEAE
(Fie. 92 on p. 408; FIG. 93 on p. 412)

SUMMARY
(i) GENERAL
This family occurs in both tropical and temperate regions. It consists
mainly of trees and shrubs, but includes a few typical xerophytic shrubs
such as Colletia armata Miers and Discaria toumatou Raoul. with flattened,
spiny, assimilatory stems and reduced leaves. The family exhibits but few
distinctive diagnostic characters, but the following points are worthy of note.
The usually mucilaginous epidermis of the leaf is papillose in numerous
genera. The stomata are generally ranunculaceous, but occasionally
rubiaceous. The mesophyll is usually dorsiventral, but rarely centric. Both
solitary and clustered crystals occur, whilst acicular ones are characteristic
of a few genera the crystals sometimes appear as transparent dots in the leaf.
;

Mucilage cells are common in the leaf and in the primary cortex of the axis,
whilst mucilage cavities are sometimes present as well in the latter. Other
secretory cells with tanniniferous contents are also frequent. The petiole, in
transverse sections, usually exhibits a solitary arc-shaped vascular strand, or,
more rarely, an arc of separate bundles.

(ii) WOOD
Vessels mostly small, but large in a few species, commonly in multiples
 RHAMNACEAE 405
and with a tendency to a radial arrangement, with pronounced 'flames' in
some species, ring-porous and with spiral thickening of the late wood vessels
in several genera, perforation plates simple, intervascular pitting alternate,
with minute to large borders; members of medium length to moderately
short. Parenchyma predominantly paratracheal in most species, vasicentric,
aliform or confluent, predominantly apotracheal or intermediate in a few
species. Rays mostly 2-5 cells wide but considerably wider in some species
and exclusively uniseriate in others, varying from markedly heterogeneous to
homogeneous with few uniseriate rays, sometimes composed entirely of square
to upright cells. Fibres with simple pits; of medium length to moderately
short. Vascular tracheids present in species with vessels in radial 'flames'.

LEAF
Generally dorsiventral, but sometimes centric or sub-centric. Rolled leaves
with a furrow on either side of the median vein occur in Microrhamnus
ericoides Gray. Furrows formed by the projecting network of veins also occur
in species of Ceanothus belonging to the sub-genus Cerastes, Hairs consisting
mostly of simple, unicellular or uniseriate trichomes of varying length.
Stellate hairs recorded in the Australian genera Cryptandra, Pomaderris,

Spyridium, Trymalium. Leaf teeth sometimes glandular in Ceanothus, Noltea,

Rhamnus. Epidermis mucilaginous or containing mucilaginous cells in
species of Alphitonia, Berchemia, Ceanothus, Colletia, Colubrina, Condalia,
Cormonema, Crumenaria, Cryptandra, Dallachya, Discaria, Emmenosperma,
Gouania, Hovenia, Lamellisepalum, Lasiodiscus, Maesopsis, Paliurus, Poma-
derris, Reissekia, Reynosia, Rhamnella, Rhamnidium, Rhamnus, Sageretia,
Scutia, Trevoa, Trymalium, Ventilago, Zizyphus. A double epidermis recorded
in a few species of Ceanothus and Rhamnus. Epidermis, especially on the lower
surface of the leaf, provided with papillae in species of Berchemia, Cryptandra
(upper side), Discaria, Helinus, Karwinskia, Marlothia. Stomata confined
to the lower surface in species of Ceanothus, Colletia, Gouania, Reissekia,
Rhamnus, Ventilago, Zizyphus, but recorded on both sides in Rhamnus
lycioides Linn.; usually ranunculaceous, but rubiaceous in a few species of
9

Colletia, Rhamnus, and Zizyphus ,

cruciferous in Cryptandra sp. Small
crowded stomata recorded in species of Reynosia and Sarcomphalus, and
especially large ones in Condalia, Discaria, Emmenosperma, and Rhamnus.
Hypoderm below the upper epidermis recorded in a few species of Ceanothus,
Microrhamnus, Rhamnus, Sarcomphalus. Mesophyll. Palisade tissue, con-
taining isolated enlarged cells filled with mucilage, present in species of
Condalia and tanniniferous cells in species of Condalia, Maesopsis, Pomaderris,
Rhamnella, and Scutia, the cells being more or less sclerosed in certain of these
genera; mucilage and tannin occur together in Discaria and Talguenea.
Palisade tissue consisting of a single layer composed of broad tanniniferous
cells inCryptandra oborata Sieb. Mesophyll consisting wholly of palisade
parenchyma in species of Ceanothus with furrowed leaves (see above). Spongy
mesophyll composed of elongated, interwoven cells in Reynosia and Sarcom-
phalus. Idioblasts filled with brown contents sometimes present in Colletia,
Scutia,and species of Zizyphus (Sabnis 1977). Small lateral veins vertically
transcurrent in species of Alphitonia, Ceanothus, Colubrina, Cormonema,
4o6 RHAMNACEAE
Crumenaria, Helinus, Hovenia, Karwinskia, Pomadems, Reynosia, Rhamni-
dium, Sageretia, Sarcomphalus, Zizyphus", veins embedded in the mesophyll
in the CoIIetieae, Gouanieae, and Ventilagineae (of Bentham and Hooker), as
well as in Condalia, Emmenosperma, Krugiodendron, Lasiodiscus, Rhamnus,
Scutia, Trymalium, and species of Zizyphus larger lateral veins accompanied
\

by a sclerenchymatous sheath in the Ventilagineae and also in species

of

Berchemia, Emmenosperma, Gouania, Lamellisepalum, Lasiodiscus, Phylica,

Reynosia, Sageretia, Sarcomphalus, Scutia,
and Zizyphus. Smaller veins pro-
vided with a sheath of cells in Condalia,
large, tanniniferous, parenchymatous
Krugiodendron, Microrhamnus, Rhamnus (especially the section Cermsporia),
Petiole, in transverse sections through the distal end, exhibiting
Zizyphus.
a single open or somewhat U-shaped arc of xylem and phloem in examined
species of Berchemia, Ceanothus, Discaria, Frangula (Fig. 93 B), Gouania,
the form
Noltea, Paliurus, Pomadems, Zizyphus. Three separate bundles in
of an open arc occur in Discaria serratifolia (Vent.) Benth. and Paliurus spina-
christi Mill. (Fig. 93 F), whilst the continuous, open, arc-shaped strand of
Gouania domingensis Linn, is accompanied by two very small, latero-superior
bundles. Large clustered crystals recorded in the mesophyll of species of
Colubrina, Condalia, Cryptandra, Hownia, Krugiodendron, Lariodiscus, Poma-
derris, Rhamnus, Trevoa, Trymalium, and Zizyphus. Solitary crystals
recorded
in enlarged palisade cells in species of Karwinskia, Reynosia, Rhamnidium,
Rhamnus, Zizyphus. Acicular crystals (styloids) abundant in Gouania.
Mucilage receptacles recorded in the parenchyma of the larger leaf veins,
and in the petiole in Alphitonia, Berchemia, Ceanothus (pro parte), Colubrina,
Condalia (pro parte), Cormonema, Dallachya, Emmenosperma, Gouania,
Hovenia, Karwinskia, Lasiodiscus, Maesopsis, Paliurus, Rhamnella, Rhamni-
dium, Rhamnus (pro parte), Ventilago (pro parte), Zizyphus.
various
Cooper (463) and Chodat (398) have described the leaf structure of
detail. The distribution of crystals
species of Ceanothus and Rhamnus in some
and secretory cells, the proportion of palisade tissue in the mesophyll, together
with features of the epidermis and stomata were found to provide characters
of specific diagnostic value.

Axis
YOUNG STEM (Fig. 93 A, c, and E)
Cork arising in the epidermis or sub-epidermis in at least certain species of
Ceanothus, Discaria, Pomadems, Rhamnus, Zizyphus. Pericycle containing
isolated strands of fibres in species of Alphitonia, Berchemia, Ceanothus,
Colletia, Colubrina, Condalia, Discaria, Emmenosperma, Gouania, Hovenia,
Noltea, Paliurus, Phylica, Pomaderris, Rhamnus, Soutia, Ventilago, Zizyphus ;
a composite and continuous ring of sclerenchyma recorded in other species of
in Gouania domingensis Linn.
Zizyphus. Stone cells observed in the pericycle
bands of fibres often sheathed by septate fibres
Secondary phloem including
which contain solitary crystals.Xylem in all investigated species in the form
of a continuous cylinder traversed by narrow rays; including vessels with
in
simple perforations, except for occasional scalariform perforation plates
Phylica and, according to Solereder, in Zizyphus. Wood fibres with simple
filled with mucilage or tanniniferous materials,
pits. Secretory cells, usually
occur in all unlignified tissues of the stem, but their distribution varies
 RHAMNACEAE 407

Elongated secretory cavities also present in the

slightly in different species.
cortex and pith of Frangula alnus Mill. (syn. Rhamnus frangula Linn.,
Fig. 93 E), R. californica Eschsch., and R. purshiana DC., but they tend to
disappear in old material. Solitary and clustered crystals common. Solitary
forms observed in the cortex in certain species of Berchemia, Phylica, Zizyphus,
in the phloem of species of Pomaderris and Rhamnus, in the pith in species of
Ceanothus, Colletia, Discaria, Paliurus, Pomaderris. Clustered crystals ob-
served in the cortex in species of Ceanothus, Colletia, Discaria, Pomaderris,
Rhamnus, Zizyphus, in the phloem in species of Ceanothus, Discaria, Poma-
derris, Rhamnus, Zizyphus, and in the pith in species of Ceanothus, Discaria,
Pomaderris, Rhamnus, The occurrence and distribution of crystals are of
specific rather than generic diagnostic value.
The assimilatory stems of Colletia armata Miers and Discaria toumatou
Raoul possess a very thick cuticle, numerous stomata, a well-defined hypo-
derm, several layers of palisade chlorenchyma in the outer part of the cortex,
whilst large solitary crystals occur in the medullary rays and in the peri-
medullary region.

BARK .

The anatomy of the bark of Ceanothus and Rhamnus has been described for
species with medicinal properties. Further particulars are given below under
'Economic Uses'.

WOOD (Fig. 92)

Vessels mostly small(less than 100 \i mean tangential diameter), very small
in Auerodendron, Berchemia, Ceanothus p. p., Colletia, Discaria, Krugiodendron,
Lasiodiscus, Reynosia, and Rhamnus, medium-sized in Ampelozizyphus,
Colubrina p.p., Hovenia, and Maesopsis, and large in Ventilago-, never
exclusively solitary, multiples of 4 or more cells moderately common in
Krugiodendron, Lasiodiscus, Pomaderris, Rhamnus p.p., and Ventilago, and in
pronounced radial or oblique flames (Fig. 92 A and H) in Ceanothus p.p.,
Condalia, Microrhamnus, and Rhamnus p.p., and similar or almost ulmiform
(Fig. 92 K) in Colletia and Discaria, with a tendency to radial or oblique
arrangement in Auerodendron, Ceanothus incanus T. et G., and Rhamnella;
very numerous in woods with radial flames, varying from 6 to 60 per mm. in
the woods with small vessels and from 2 to 7 per mm. in those with medium-
*
sized or large vessels, most numerous (apart from woods with radial flames')
in Krugiodendron and Lasiodiscus (more than 30 per mm.) and fewest (2-4
per mm.) in Hovenia, Maesopsis, and Ventilago; ring-porous or semi -ring-
porous in Ceanothus p.p., Colletia, Condelia p.p., Discaria (Record 1856),
Hovenia, Rhamnella, Rhamnus, and Zizyphus p.p. (Record 1851); with spiral
thickening in Adolia (Record 1783), Ceanothus, Colletia, Colubrina (1868),
Condalia, Discaria p.p., Doerpfeldia (1868), Microrhamnus, Rhammis, Sarcom-
phalus (1868), Scutia, and Zizyphus jujuba Mill. (2158). Perforation plates
exclusively simple, except that Solereder records some scalariform plates
with few bars in Phylica and Zizyphus calophytta Wall. Intervascular pitting
alternate, with coalescent apertures in many species, sometimes very marked,
as in Hovenia and Paliurus', pits small to minute in Auerodendron, Krugio-
dendron, Lasiodiscus^ Paliurus, Pomaderris, Reynosia^ Rhamnella, Sageretia,
 Fio. 92. RHAMNACEAE
A, Rhamnus cathartica Linn. B, Maesopsis eminii Engl. C, M. imtmt Engl. D, Doerpfeldia cubensis
Urb. E, Colubrina panamensis Standl. F, Zizyphus jujuba Lam.
.
"
G t Z. yii/tida Lam. H, Ceanothus
incanut T. et G, I, C. thyrsiflorus Each. J, Co/fof"a AVco/or Hook. f. K, C. rfwo/or Hook. f.
 RHAMNACEAE 409
and Scutia, moderately large to large in the other genera pits to parenchyma
;

and ray cells similar to the intervascular pitting, occasionally unilaterally

compound. Tyloses not observed but recorded by Kanehira (1206) for
Paliurus samosissimus Poir.; solid deposits present in some species and
abundant in Reynosia (Record 1856). Mean member length o-3-O'7 mm.
Parenchyma very sparse to moderately abundant, predominantly para-
tracheal in most species; most commonly as a few cells round the vessels;
aliform and sometimes locally confluent in Auerodendron, Colubrina p.p.,
Discaria, Maesopsis (Fig. 92 c), Reynosia, Scutia, and Zizyphus p.p. (Fig. 92 G),
confluent in Doerpfeldia (Fig. 92 D) and Zizyphus p.p.; associated with the
vessels where these are in 'flames' but very sparse in Ceanothus, Microrhamnus,
and some species of Rhamnus; with some diffuse parenchyma in addition to
paratracheal in Auerodendron and Doerpfeldia and also, according to Kanehira
(1206), in Paliurus ramosissimus Poir. and two species of Rhamnus\ in wavy
bands, according to Williams (2430), in Gouania lupuloides (L.) Urban.;
predominantly diffuse in Sarcomphalus and some species of Zizyphus, e.g.
Z. angolito Standl. with uni- to biseriate terminal bands in Ampelozizyphus,
;

Hovenia, Lasiodiscus, Paliurus, Sageretia, Sarcomphalus, Scutia, and Zizyphus

p.p. ; in Zizyphus the
parenchyma is very variable being, for example, pre-
dominantly apotracheal in numerous uniseriate bands in Z. angolito Standl.,
Z. cyclocardia Blake, Z. guatemalensis Hemsl., Z. hauaensis H. B. et K., and
Z. sonorensis S. Wats. (1886), in narrow irregular to broad confluent bands
in Z. melastomoides, Z. mistol Griseb., and Z. spina-ckristiWilld. p.p., aliform
in other species, often accompanied by irregularly scattered single strands.

Crystals in chambered cells observed in Auerodendron, Berchemia, Maesopsis,

and Rhamnella and also reported by Record (1856) in some species of Colletia,
Colubrina, Condalia, Doerpfeldia, Karwinskia, Rhamnidium, Sageretia, and
Sarcomphalus. Storied in Ventilago. Very thin-walled in most of the species.
Strands typically of 4 cells, but occasionally up to 6 or 8 cells, e.g. in Ampelo-
zizyphus, Paliurus, Rhamnella, Rhamnidium, and Zizyphus. Chambered
crystals sometimes present. Rays most commonly up to 2-5 cells wide,
exclusively uniseriate in Paliurus and some species of Zizyphus, e.g. Z.
jujuba Mill, and Z. mucronata Willd. up to 6-8 cells wide in Colletia,
;

Discaria (Record 1856), and Ventilago and up to 30 cells wide in Ampelo-

zizyphus\ of 2 distinct sizes in Ampelozizyphus and with relatively few inter-
mediates between uniseriate and 4 or more cells wide in Ventilago', less than
i mm.
high except in Scutia (up to i -5 mm.) and Ampelozizyphus, in which
very high primary rays occur in various stages of dissection into smaller units;
uniseriate rays scarce or i or 2 cells high in Auerodendron,
commonly only
Colletia, Condalia, Maesopsis, Rhamnella, Rhamnidium, and Rhamnus p.p.;
about 10-20 rays per mm. in woods with rays up to 3 cells wide, 4-10 per mm.
in woods with larger rays, more numerous in Sageretia and Scutia> fewest in
Maesopsis', usually heterogeneous (Kribs's Types II A and B); with 4 or more
marginal rows of distinctly upright cells in Berchemia, Colubrina, Cormonema,
Emmenosperma, Hovenia, Karwinskia (Record 1856), Lasiodiscus, Pomaderris,
Rhamnidium, Sageretia, and Zizyphus p.p. homogeneous (Kribs's Types I-II)
;

or with a single marginal row of square cells not much higher axially than the
procumbent cells in Maesopsis, Rhamnus, and Ventilago, and, according to
Record and Hess (1886), in some specimens of Doerpfeldia and Sarcomphalus\
4 io RHAMNACEAE
the other genera intermediate; the shrubby members often with their pro-
cumbent cells replaced by square cells or with uniseriate margins of square
cells that are not distinguishable from the procumbent cells in tangential

sections, e.g. Ceanothus incanus T. et G. and Microrhamnus\ wood with wholly

uniseriate rays homogeneous (Kribs's Type III), heterogeneous (Kribs's
Type III) or composed entirely of square and upright cells; sheath cells
present in Rhamnella\ cells very commonly containing gum-like deposits and
single crystals, the latter very numerous in Doerpfeldia, Krugiodendron,
Scutia, and Zizyphus mucronata Willd. ; with a tendency to arrangement in
echelon in Krugiodendron and Maesopsis. Fibres with simple pits, more
numerous on the radial than the tangential walls, usually very sparse, but
more numerous in some of the woods with thin- walled fibres; often septate,
according to Kanehira (1206) in Rhamnus formosana Mats. Walls thick to
very thick in Auerodendron, Berchemia, Ceanothus p.p., Colletia, Condalia>
Discaria (1856), Doerpfeldia, Emmenosperma, Krugiodendron, Lasiodiscus,
Paliurus, Reynosia, Sageretia, Scutia, and Zizyphus p.p., thin-walled in
Alphitonia, Ampelozizyphus^ Ceanothus p.p., Colubrina, Cormonema Hovenia, y

and Maesopsis very commonly with a gelatinous inner layer. Mean length
\

0-7-1 -7 mm. Vascular tracheids present in woods with vessels in radial

'flames' ;
often spirally thickened.

ROOT
Bottomley (241) has described nodules on the roots of Ceanothus americanus
Linn, which contain nitrogen-fixing bacteria. Root nodules are rare in
specimens of C. americanus cultivated in Great Britain but common in
American specimens. The nodules are stated to increase in size by means of
outgrowths of endogenous origin, and, when mature, are differentiated into
four distinct zones: (i) An apical meristematic zone, (ii) zone where theA
cells become infected with bacteria, (iii)
A zone containing enlarged, radially
elongated cells filled with bacteria, (iv) A basal zone almost free from
bacteria.
Holm (1064) has recorded the occurrence of concentric bands of mechanical
tissue surrounding the stele in Rhamnus purshiana DC., and Wirth (2446)
that in certain cells of the root of Ceanothus americanus there is a brownish
substance whose colour is intensified by hydrochloric acid.

TAXONOMIC NOTES
Record and Hess (1886) have drawn attention to the occurrence of two
distinct groups among the species of Zizyphus; one is characterized by diffuse
apotracheal parenchyma, as in Z. angolito Standl. and Z. sonorensis S. Wats.,
the other by paratracheal parenchyma, as in Z.jujuba Mill, Z. mistol Gris.,
and Z. spina-christi Willd.
The species of Rhamnus also fall into two distinct groups; one group,
corresponding to the sub-genus Eurhamnus, having a marked oblique or
dendritic vessel pattern, the other, corresponding to the sub-genus Frangula,
having vessels that are semi-ring-porous, but otherwise without pattern.
Record and Hess consider that the species furnishing the Red or Pink Ivory
of northern Natal, Rhamnus zeyheri Sond., is out of place in this genus.
 RHAMNACEAE 4"
ECONOMIC USES
The timbers of this family are of little importance, Maesopsts berchemoides
A, Chev. in East Africa, however, is a very large tree furnishing a soft timber
with some possibilities. Some species of Zizyphus are, or have been, used
locally, e.g. the Jamaican Cogwood
Z. chloroxylon (L.) Oliv. and Z. jujuba
Mill, in India. Record and Hess (1886) note that Colubrina, KarwiwUa, and
Rhamnidium are the source of durable woods that are sometimes used locally,
and Sarcomphalus laurinus Gris., though a small tree, is reputed to produce
one of the best timbers in Jamaica. The charcoal of Frangula alnus Mill,
was in great demand during the recent war for making powder for fuses.
The wood of Krugiodendron ferreum (Vahl.) Urban, with a specific gravity
of up to 1-42 air dry, is reputed to be the densest known (1805).
The barks of various species of Rhamnus and Ceanothus are used in medicine
because they possess laxative properties, whilst the fruits of Rhamnus cathartica
Linn, are also used as a laxative in veterinary practice. The medicinal barks
have been described anatomically and their chemical constituents discussed
by Gathercoal (751), Greenish and Wallis (812), Hasler (917),
Holm
(1064), Maeder (1413), Taylor (2238), and Wirth (2446).
The bark of Rhamnus purshiana DC., the source of cascara sagrada, exhibits
the following characters when sufficiently mature. Exposed outer surface
of 10-12 rows of thin-
purplish in colour when scraped. Cork consisting
walled cells, sometimes filled with a brown deposit. Middle portion of the
bark mainly parenchymatous, but containing elongated mucilage sacs (which
become compressed and difficult to recognize in old material), stone cells, and
rosette crystals. The inner portion of the bark, which arises only when the
stems are sufficiently mature, consists of longitudinally elongated cells, and
includes bundles of fibres sheathed by cells containing prismatic crystals.
of the inner bark
Large groups of stone cells also arise in the outer part
between the somewhat sinuous, medullary rays which are 2-4 cells wide. The
parenchymatous cells of the bark contain a yellow substance which is coloured
violetby sodium hydroxide solution.
The bark of Frangula alnus Mill. (syn. Rhamnus frangula Linn.) is very
similar to that of R. purshiana from which it may generally be distinguished
by the absence of stone cells.
Hasler (917) experienced considerable difficulty in finding reliable charac-
ters for distinguishing between the barks from closely related species of
Rhamnus, especially when variations in structure corresponding to different
consideration. It was demonstrated, more-
stages of maturity were taken into
over, that the frequency of crystals varies according to
the amount of calcium
in the soil.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Berchemia,* Ceanothus,* Colletia,* Colubrina, Condalia,
Alphitonia,
Cormonema, Crumenaria, Cryptandra, Dallachya, Discaria,* Emmenosper-
ma, Frangula,* Gouania,* Helinus, Hovenia, Karwinskia, Krugiodendron,
Lamellisepalum, Lasiodiscus, Maesopsis, Marlothia, Microrhamnus,
Noltea,* Paliurus,* Phylica,* Pomaderris,* Reissekia, Reynosia, Rhamnella,
FIG. 93. RHAMNACEAE, A-C and E-F; AMPELIDACEAE, H-I; ACERACEAE, and D G
c
A, Discaria toumatou Raoul. mX2i. B,Frangula alnus Mill.- (Rhafttnus frangula Linn.). Petiole
X 31. C, Pomaderris apetala Labuj. Stem X 13. D, Acer campestre Linn. Petiole X 32. E, Frangula
alnus Mill,(Rhamnus frangula Linn.). Stem X 17. F, Paliurus spina-christi Mill. Petiole X48.
G, Acer campestre Linn. Stem X 13. H, Leea angulata Korth. Petiole x8. I, Vitis vinifera Linn.
Stem Xis.
The small. circles in B and E represent secretory (mucilage) cavities.
 RHAMNACEAE 413
Rhamnidium, Rhamnus,* Sageretia, Sarcomphalus, Scutia, Spyridium,
Talguenea, Trevoa, Trymalium, Ventilago, Zizyphus.*
* Slides in Kew collection.

(ii) FOR WOOD STRUCTURE

(Adolia), Alphitonia, Ampelozizyphus, Auerodendron, Berchemia, Ceano-
thus, Colletia, Colubrina, Condalia, Cormonena, Discaria, Doerpfeldia,
Emmenosperma, (Gouania), Hovenia, (Karwinskia), Krugiodendron, Lasio-
Microrhamnus, Paliurus, (Phylica), Pomaderris, Reynosia,
discus, Maesopsis,
Rhamnella, Rhamnidium, Rhamnus, Sageretia, Sarcomphalus, Scutia, Venti-
lago, Zizyphus.

LITERATURE
(i) On General Anatomy
Bottomley 241, Chodat 398, Cooper 463, Gathercoal 751, Greenish and Wallis 812,
Hasler 917, Heppeler 954, Holm 1064, Lamorlette 1316, Maeder 1413, Sabnis 1977,
Taylor 2238, Watkins 2367, Weberbauer 238 i, Wirth 2446.

(ii) On Wood Structure

Baker 104, Besson 186, Brown, F. B. H. 280, Brown, H. P. 289, Burgerstein 310, 312,
Chowdhury 411, Cooper 461, Greguss 2522, Howard 1088, Janssonius 1154, Kanehira
1206, 1209, Messeri 1493, Normand 1613, Pearson and Brown 1679, Record 1783, 1805,
1843, 1851, 1856, Record and Hess 1886, Record and Mell 1894, Tang 2231, Williams
2430, Yamabayashi 2478.

100. AMPELIDACEAE
(VITACEAE)
(Fie. 93 on p. 412; FIG. 94 on p. 414; FIG. 95 on p. 416)

SUMMARY
(i) GENERAL
A tropical and sub-tropical family which includes many typical lianes,
although some members, especially in the genus Leea, are erect and arboreal.
The lianes are frequently much swollen at the nodes, and bear tendrils whose
structure varies considerably in response to the nature of the substratum to
which they become attached. For instance, they become coiled around stems
and other cylindrical supports, but adhere to flat surfaces by means of cushion-
like endings. Some species such as Cissus currori Hook. f. have a tuberous
stem base serving mainly for the storage of water and from which erect branches
arise. The morphology of the plants is often difficult to interpret. Some of
the vines, especially those from the tropics, are provided with an extensive
system of aerial roots. In correlation with the liane habit the young stem often
exhibits a characteristic appearance in transverse section owing to the vessels
being particularly wide in diameter, whilst the xylem is split up into lamellae
by the broad primary and secondary medullary rays. This structure recalls
that of the Aristolochiaceae. The family is also characterized by the presence
of raphides, which are contained in sacs which sometimes appear under the
lens as transparent dots or fine striae. Other forms of crystals include acicular,
solitary and clustered types. The raphide sacs usually contain mucilage as
well as the crystals, and in other instances mucilage cells occur from which
4*4 AMPELIDACEAE
raphides are absent. Mucilage cells or large crystals sometimes appear as
transparent dots in the same way as the raphide sacs.
The hairs are of the following types: simple unicellular; uniseriate;
unicellular 2-armed; short stalked, glandular. Pearl glands and peltate scales
also occur. The stomata are ranunculaceous. The cork arises superficially
in some species, but in the cortex, pericycle, or even in the phloem in others.
The pericycle contains isolated bundles of fibres or a composite and con-
tinuous ring of sclerenchyma. Anomalous structure has been recorded in
Cissus scdriosa Bl. (Tetrastigma scariosum Planch.) and related species.

FIG. 94. AMPELIDACEAE

A, Pearl-gland of the vine. B-C, Leea aequata L.: B, Section through the leaf and through a gland;
C, Surface-view of the gland. A, after Penzig; B-C by Solereder.

(ii) WOOD
Vessels large in the climbers, small in Leea, radial multiples sometimes
moderately common;
perforation plates simple; intervascular pitting usually
scalariform, sometimes opposite or alternate, pits to parenchyma simple and
oblong, or bordered and round, members moderately short to very long.
Parenchyma paratracheal, usually rather sparse, sometimes moderately
abundant; forming the ground tissue in some species of Cissus sometimes ;

containing raphides. Rays broad and very high, up to 20 cells wide in some
species, sometimes of 2 distinct sizes but more commonly with few or no
uniseriate rays; cells varying from all procumbent to all square or upright;
raphides present in many species. Fibres septate, very short to very long.

LEAF
Usually dorsiventral, with i or z layers of palisade tissue in Vitis\ sometimes
tending to be centric. Hairs of the following types, (i) Simple, uniseriate
trichomes, with blunt or pointed ends, the component cells sometimes
distinctly articulated,(ii) Simple, unicellular, sometimes very long
and pro-
ducing a felt resembling a spider's web. (iii) Unicellular, 2-armed, with or
without stalks, notably in certain species of Cissus. (iv) Stalked glandular
trichomes rare. Deciduous pearl glands common in Cissus, Leea (Fig. 94 B-C),
and VitiS) but varying in frequency within a species, sometimes in relation to
the vigour of the plant or the humidity of the atmosphere. These glands are
 AMPELIDACEAE 415

usually spherical structures each with a short stalk, the interior consisting of
large, polygonal cells, surrounded by an epidermis perforated by a stoma, the
latter usually but not invariably situated opposite the stalk of the gland. Cells
of the glands rich in protein, oil, and sugar. Somewhat similar glands, com-
posed of comparatively small, isodiametric cells and each surrounded by a palisade
epidermis, and opening to the exterior by a solitary apical stoma, described
in detail by Solereder in Leea aequata Linn. Walter (2354) has also investi-
gated the structure and physiology of pearl glands in this family. Leaf teeth
secreting mucilage in the bud of Vitis vinifera Linn. Stomata ranunculaceous
in the few species examined. Stomatal distribution very imperfectly known.
Petiole not examined in many species, but, in transverse sections, exhibiting
a ring of isolated bundles, accompanied by two, or less frequently more,
cortical bundles on either side of the groove in the adaxial surface of the

petiole in certain species of Leea (Fig. 93 H) and Vitis. Secretory cells with
amorphous contents (probably mucilaginous and tanniniferous) widely
distributed in the parenchymatous tissues, notable in the petiole of Leea
angulata Korth. and Vitis vinifera Linn, and probably in other genera and
species. Large mucilage cells, also appearing as dots, recorded in species of
Cissus, ParthenocissuSy and Vitis. Raphide sacs (see also 'Summary'), of
variable lengths and often containing mucilage, occur throughout the family,
the individual raphides stated to be pointed at one end but bidentate at the
other in Cissus and Vitis. Solitary and clustered crystals also common;
druses in Leea especially large and sometimes appearing as dots.
Dune forms of Psedera quinquefolia (L.) Greene and Vitis vulpina Linn,
have been recorded by Starr (2188) as having taller palisade cells, and
epidermal cells which are larger in surface area but shorter than those in
mesophytic specimens of the same species. In Vitis vulpina hairs were
observed by the same author above the veins on both surfaces in dune
specimens, but were confined to the same position on the upper surface in
mesophytic forms.

Axis
YOUNG STEM (Fig. 93 i)
Often very characteristic, as seen in transverse section, owing to the parti-
cularly wide diameter of the vessels and the appearance of the lamellae of
xylem and phloem separated by the broad, primary, and secondary medullary
rays. Cork in Parthenocissus qmnquefolia Planch, arising immediately below
the epidermis. In the same species successive annual growths, produced from
the same phellogen, remain separated from one another by a layer of sclerotic
cork. Cork stated to arise in a similar manner in Cissus^ Leea, and certain
species of Vitis; originating in the pericycle and consisting mainly of thin-
walled cells, with solitary sclerotic cells amongst them in Vitis vinifera Linn.
(Fig- 93 i) and other closely related species of Vitis. Primary cortex said to
contain cells with perforations in Cissus antarctica Vent. Pericycle contain-
ing isolated strands of fibres in Leea angulata Korth. as well as in Vitis vinifera
and closely related species, but exhibiting transitions between this arrange-
ment and a composite continuous ring of sclerenchyma in other species.
Pericyclic fibres with wide lumina and thin walls in Vitis vinifera and closely
related species, but short-lived owing to the development of deep-seated
4 x6 AMPELIDACEAE
cork. Secondary phloem stratified into sclerosed and soft portions in the
Euvitis section of Vitis, but with the fibres arranged in more or less radial
rows beside the medullary rays and exhibiting a scattered arrangement in the

FIG. 95. AMPELIDACEAE

A, Parthenoctssus vitacea Hitchc. B, Vitis arizonica Engelm. C, Leea guineense Don. D, Vitis
labrusca Linn. E, Leea philippinensis Merr. F, Cissus sicyoides Linn.

inner part of each phloem strand in the Muscadinia section of the same genus.
Phloem and xylem forming crowded but individually distinct vascular
bundles separated by broad primary medullary rays in Vitis vinifera (Fig. 93 i)
and other species. Vessels wide in climbers; perforations simple.
Pith com-
posed of cells with walls of variable thickness in different species; homo-
geneous or heterogeneous. Cluster and solitary crystals and raphides
 AMPELIDACEAE 417
common in the parenchymatous tissues. Acicular crystals, smaller than typical
raphides and not embedded in mucilage, also recorded in the cortex and
phloem of various members of the family, notably in species of Ampelopsis
and Cissus. Secretory cells with amorphous contents (probably mucilaginous
and tanniniferous) widely distributed in the parenchymatous tissues of the
stem in Leea angulata Korth, Vitis vinifera, and probably in other genera and
species as well.

WOOD (Fig. 95)

Vessels small in Leea (less than 100 p mean tangential diameter), large
(more than 200 ^) in the climbers Cissus, Parthenocissus, Psedera, Tetrastigma,
and Vitis largest vessels up to 350 // in tangential diameter; of 2 types in
',

Parthenocissus and some species of Vitis, the larger solitary and often over
300 p. in diameter, the smaller seldom exceeding 100 p, in diameter and either
commonly in radial multiples (Vitis) or grouped round the larger vessels
(Parthenocissus) radial multiples common also in Psedera; very variable in
;

number even in the same genus, from 2 to 22 per mm. ; ring-porous in some
species of Vitis (1851). Perforations simple. Intervascular pitting usually
scalariform, locally opposite in Psedera and alternate in Cissus sicyoides Linn.,
Parthenocissus, and Tetrastigma; pits to parenchyma round and bordered in
woods with alternate intervascular pitting, large, oblong, and often simple
in the others, usually with the long axes horizontal. A
few tyloses present in
most species of Cissus, Parthenocissus , and Vitis, sometimes locally abundant.
Mean member length very variable, e.g. Parthenocissus 0-3 mm., Vitis 0-7 mm.,
and Leea i -7 mm. Parenchyma paratracheal, varying from a few cells round
the vessels (Fig. 95 c and D) to a well-defined vasicentric sheath, but never
abundant; according to Kanehira (1206) sparse and scattered among the
fibres in Leea sambucina Willd. In certain soft-stemmed species of Cissus,

e.g. C. sicyoides L., according to Solereder (2158) 'the groundwork of the wood
consists of unlignified parenchymatous tissue with thin walls, in which the
vessels are embedded in groups, each group surrounded by a sheath consisting
of a little lignified wood-parenchyma and wood-prosenchyma' (Fig. 95 F).
A similar type of structure occurs in Parthenocissus. Hess (959) reports the
occurrence of raphides in this tissue in Cissus and there may also be chambered
solitary crystals on the margins of the sheaths where they join the fibres.
Strands usually of 4 cells, sometimes up to 8 cells. Storied in Parthenocissus
and Tetrastigma. Rays broad (more than 100 ^) and usually very high;
commonly of indeterminate height, but exhibiting various stages of dissection
into smaller units in some species; the larger rays 4-6 cells wide in Cissus and
Leea p.p., up to 15-20 cells wide in some species of Psedera and Vitis; of
2 distinct sizes (uniseriate or broad) in Leea, Vitis arizonica Engelm., and V.
californica Benth. (8); with a fairly complete gradation of sizes in Cissus
sicyoides L. and Parthenocissus', normally without uniseriate rays in most
species of Vitis (8) and with few or none in Cissus, some species of Leea,
Psedera, and Tetrastigma; uniseriates varying from high upright cells in Leea
p.p. to procumbent in Parthenocissus; Adkinsoh (8) notes that, as a result of
injury, uniseriate rays may occur in woods from which they are normally
absent. Between 2 and 6 rays per mm. in woods with large rays only, up to
12 per mm. in woods with uniseriate rays; cells of the multiseriate rays all
4594 Ee
418 AMPELIDACEAE
upright or square in Leea (Fig. 95 E) and Sundaica, almost all procumbent in
Parthenocissus, Psedera^ and Vitis p.p., in other species a mixture of square to
procumbent cells without any distinct marginal rows; cells very variable in
tangential width in different species, relatively small where the cells are all
procumbent (Fig, 95 A and B), e.g. Psedera, and large where true procumbent
cells are rare (Fig. 95 E); sheath cells present in some species; cells typically
filled with dark contents and containing occasional single crystals in some

species. Raphides occur in the rays of Leea, Tetrastigma^ and Vitis. The
small rays storied in Parthenocissus. Fibres septate and frequently containing
dark deposits; pits simple and usually more numerous on the radial than on
the tangential walls, with small borders in Leea tetramera B. L, Burtt. With
moderately thin walls in most species. In Cissus sicyoides and Parthenocissus
the fibres do not form the ground tissue but occur in narrow bands or patches
separated by more abundant thin-walled parenchyma (Fig. 95 F); they are
non-septate, very thick-walled, much elongated (5 times the length of the
vessel members or more) and very long (nearly 3 mm. in Cissus). Storied in
Tetrastigma lauterbachianum Gilg. Mean length very variable, e.g. Partheno-
cissus 0-6 mm., Vitis 1-4 mm., and Leea 2-4 mm. Vasicentric tracheids
reported by Record and Hess (1886); with scalariform thickening and some-
times with delicate spiral thickening. Intercellular canals of the vertical
traumatic type observed in Cissus sicyoides L.

ROOT
The particulars recordedby Turner (2300) about the aerial roots of Vitis
rotundifoliaMichx. include the following. Epidermis short-lived; apparently
devoid of stomata. Primary cortex 15-25 cells wide; outer part collenchy-
matous, but provided with well-developed intercellular spaces towards the
interior. Raphide sacs present in the cortex, but crystals becoming dissolved

during development of the root, and the cells containing them disorganized.
Endodermis consisting of cells with thickened walls and serving mainly for
storage; casparian thickenings only rarely observed. Primary vascular
system hexarch to octarch. Secondary thickening proceeds normally.
Vessels in the secondary xylem up to 200 /* in diameter; tyloses fairly
common. Pith composed of cells with slightly thickened walls; still persisting
in y-year-old roots.

ANOMALOUS STRUCTURE
Anomalous secondary thickening has been described in Cissus scariosa Bl.
(Tetrastigma scariosum Planch.) by Schenck whose description was followed
by Solereder (2158) and is repeated here. The strap-shaped stem grows in
thickness in the normal way until it is I cm. thick. At this stage the individual
segments of xylem and phloem are separated by broad parenchymatous rays.
In each segment of xylem and phloem on the narrow side of the stem a strip
of cambial tissue arises in the phloem parenchyma, immediately on the inside
of the first-formed strand of phloem. The newly formed strip of cambium
then gives rise to fresh xylem and phloem. Certain other species of Tetrastigma
are said to exhibit a similar anomaly.
 AMPELIDACEAE 419
PHYLOGENETIC AND TAXONOMIC NOTES
Adkinson making a comparative anatomical study of a selection of
(8), after
the Vitaceae, concluded that the erect members of the family such as Leea
retain more primitive characters than the climbers. In the primitive erect
types the broad medullary rays are accompanied by linear rays which have
been lost in most of the climbing species, although still retained in Vitis
californica Benth. In some other members of the Vitoideae vestiges of linear
rays persist in the seedling and 'conservative* parts of the plant. Adkinson
9
also concluded that 'Ampelopsis and Cissus, where the xylem is more reduced
and dissected than in Vitis, are the more primitive genera. The vine is
regarded as an approach to the herbaceous (advanced) type of stem.
Hess (959) suggests that the wood of Leea bears some relation to Dilleniaceae.

ECONOMIC USES
The most important economic plant in the family is Vitis vinifera Linn, the
grape vine, from the fruits ofwhich wines are prepared, while those of certain
varieties, when dried, are most familiar as raisins, currants, &c. The Virginia
Creeper of gardens is Parthenocissus tricuspidata Planch.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Ampelopsis, Cissus, Leea,* Parthenocissus, Psedera, Tetrastigma, Vitis.*
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Cissus, Leea, Parthenocissus, Psedera, Tetrastigma, Vitis.

LITERATURE
(i) On General Anatomy
Adkinson 8, Bowman 250, Esau 2518, Gilg 767, La Riviere 1320, Rehder 1903, Starr
2188, Turner 2300, Walter 2354.
(ii) On Wood Structure
Adkinson 8, Dadsweli and Record 533, Desch 574. Greguss 2522, Greiss 817, Hess 059,
Howard 1088, Janssonius 1154, Kanehira 1206, Pfeiffer, H. 17x2, Record x8x8, 1843,
1851, Record and Hess 1886.

. SAPINDACEAE
(Fio. 96 on p. 424; FIG. 97 on p. 426; FIG. 100 on p. 438)

SUMMARY
(i) GENERAL
Trees, shrubs, and lianes mainly from tropical but also from temperate
regions. The lianes frequently exhibit anatomical peculiarities in the stems,
and also possess tendrils. Thorns, which represent reduced axillary branches,
occur in Stocksia. Of the erect members of the family some are branchless
with a terminal collection of leaves and thus resemble palms or tree ferns.
Species of Pseudima, Pseudopteris, Talisia, Toulicia, Tripterodendran possess
this unusual habit form. The hairs may be either (i) simple, unicellular with
420 SAPINDACEAE
a tendency to be 2-armed; (ii) tufted; (iii) small peltate scales. Small glan-
dular hairs are also frequent, particularly in young leaves. The colour of the
leaf in herbarium specimens is somewhat distinctive in certain genera (see
'Leaf'). The epidermis of the leaf often includes mucilaginous cells which
appear as transparent dots. Small translucent areas are also caused by the
presence of secretory cells. The lower surface of the leaf is often papillose.
The stomata, usually confined to the lower surface, are rammculaceous
except in Conchopetalum and Harpullia. Hypoderm is often present. The
mesophyll is usually dorsiventral, but wholly or partly centric in certain
genera; it often includes sclerenchymatous fibres. Crystals are mostly
solitary or clustered, whilst crystal-sand has been recorded in Exothea and
styloids in Diatenopteryx. The cork in the young stem is superficial in origin
in most instances. The pericycle contains a composite and continuous ring
of sclerenchyma except in Valenzuelia and Xanthoceras. Several types of
anomalous thickening have been described in a number of genera. Saponin
is commonly present in the tissues.

(ii) WOOD
Vessels typically small and numerous, with many multiples of 2-3 cells,
but without any definite pattern; perforations simple, intervascular pitting
alternate and small to minute; pits to parenchyma similar. Very occasionally
ring-porous or with spiral thickening; members of medium length to very
short. Parenchyma most commonly paratracheal only and sparse, sometimes
vasicentric, with isolated crystalliferous strands or diffuse parenchyma
scattered among the fibres, or in confluent bands terminal parenchyma some-
;

times prominent. Rays typically exclusively uniseriate or nearly so, low,

numerous and homogeneous; 2-3 cells wide in a few genera and occasionally
heterogeneous. Fibres with simple pits and typically septate, though septa
may be few or lacking where the fibres are very thick walled; of medium
length to very short.

LEAF
Usually dorsiventral; wholly or partly centric in certain species of Cardio-
spermum, Diplopeltis, Dodonaea, Elattostachys, Erythrophysa, Heterodendron,
Koelreuteria Lecaniodiscus, Paullinia> Stocksia, Xanthoceras. Hairs, (i) Uni-
>

cellular, or tending to be uniseriate, often with swollen, sometimes striate

bases sunk below the epidermis in species of Cardiospermum (pointed),
Chytranthtis, Matayba, Pancovia> Xerospermum. (ii) Unicellular but tending
to be 2-armed in Alectryon, Matayba, Nephelium, Pometia, Stadmannia.
(iii) Multicellular,
tufted or stellate hairs, sometimes accompanied by simple
ones in Arfeuillea, Cossignia, Euphoria, Harpullia, Nephelium, Xanthoceras.
(iv) Glandular hairs, each with a short uniseriate stalk and an oval head
of
a few cells, very common, possibly present throughout the family on young
leaves, (v) Small multicellular peltate scales recorded in species of Arytera,

Cupaniopsis, Dictyoneura, Dodonaea, Filidum, Ganophyllum, Lecamodiscus,

Lepiderema, Lychnodiscus, Smelophyllum, Stadmannia. Glandular hairs some-
times specially large in Crossonephelis, Melanodiscus, and Talisia\ complex in
structure and occasionally containing solitary and clustered crystals in the
head in Llagunoa and Loxodiscus. Glandular hairs not observed in all or
 SAPINDACEAE 421
certain species of Castanospora, Chytranthus, Conchopetalum, Cupania,
Dodonaea, Eriandrostachys, Gleniea, Harpullia, Macphersonia, Matayba,
Melicocca, Molinaea, Otophora, Pancovia, Phialodiscus, Porocystis, the
Schleichereae, Storthocalyx, Tina, Toechima, Toulicia, Trigonachras, Tristira,
Tristiropsis, Valenzuelia, Xanthoceras. Dried leaf lead-grey or yellowish-
green in external appearance in the Aphanieae, Cupanieae, and Lepisantheae
owing to special contents of the epidermis; lower side chocolate-brown owing
to tanniniferous contents of the mesophyll in Guioa, Nephelium, and Otophora ;

blackish-brown in Elattostachys, blackish-green in Harpullia; young dried

leaves reddish in Nephelium, Otophora, Pometia, Schleichera, Talma. Leaves
and branches sometimes covered with a resinous secretion from the external
glands in species Dodonaea, Filicium, Ganophyllum, and Llagunoa.
of
Epidermis composed of cells with sinuous anticlinal walls and pits in the
bays of the undulations in all or certain species of Alectryon, Aporrhiza,
Arytera, Atalaya, Blighia, Chytranthus, Cupaniopsis, Doratoxylon, Eriocoelum,
Euphoria, Exothea, Guioa Haplocoelum, Harpullia, Laccodiscus, Lecaniodiscus^
y

Lepiderema, Lepidopetalum, Lychnodtscus, Matayba, Melanodiscm, Mischo-

carpus, Pancovia, Phialodiscus, Sarcopteryx, Sarcotoechia, Storthocalyx,
Synima, Toechima, Trigonachras, Xerospermum. Some of the epidermal cells
sclerosed in Matayba purgans (Poepp. et Engl.) Radlk. Epidermal cells parti-
cularly tall in Deinbollia, Otophora^ and Sarcopteryx] provided with delicate,
vertical, secondary walls in species of Alectryon, Arytera, Cupaniopsis,
Euphoria, Heterodendron, Macphersonia, Tina, Xerospermum; with secondary
walls parallel to the leaf surface in Cupaniopsis and Gongrodiscus. Mucilaginous
cells frequent in the epidermis, sometimes appearing as transparent dots;

occurring throughout some of the genera, but limited to certain, often closely
related, species in others. Papillae often present on the lower surface, some-
times forming a network owing to their being united to one another by
cuticular ridges. (For further details of mucilaginous epidermal cells and
papillae see Solereder.) Stomata reported to be unusually small in Cupania,
Nephelium, Sapindus, Talma, and other related genera, but much larger in
Aphonia and Otophora, usually confined to the lower surface, but present on
both sides in certain species of Dodonaea, Lepiderema, Pappea, Paullinia,
Serjania; known to be rubiaceous in species of Conchopetalum and Harpullia,
elsewhere ranunculaceous. Stomata particularly numerous but small in
Arytera, Dilodendron, Elattostachys, Guioa, Lepidopetalum, Mischocarpus,
Molinaea, the Nephelieae, Paranephelium, Porocystis, Sapindus, Sarcotoechia,
Schleichera, Talisia, Toechima, Toulicia, Tristira; much larger but less numer-
ous in Aphonia and Otophora. Parenchymatous hypoderm present in certain
species of Alectryon, Arytera, Atalaya, Conchopetalum, Cupania, Harpullia,
Matayba, Molinaea, Storthocalyx, Talisia] hypodermal cells fibrous in
Euphoria gardneria Thw.; filled with a soft, homogeneous or lamellated
substance of unknown chemical nature in Cossinia. Mesophyll containing
sclerenchymatous fibres or sclerosed cells in certain species of Cupaniopsis,
Haplocoelum, Harpullia, Matayba, Paullinia, Serjania, Xerospermum', similar
cells also recorded by Le Renard (1363) in the rachis of Arfeuillea, &c.
Intercellular spaces in the spongy mesophyll particularly large in Eriandro-
stachys, Ganophyttum, Harpullia, Otonephelium, Otophora, Paranephelium,
Plagioscyphus, Pseudonephelium; almost or completely absent from species of
422 SAPINDACEAE
Alectryon, Arytera, Atalaya, Averrhoidium> Cupaniopsis, Elattostachys, Meli-
cocca, Pappea, Podonephelium, Talisia, Tina, Toulida. Palisade cells trans-
versely septate in species of Alectryon, Eriandrostachys, Macphersonia,
Magonia, Nephelium, Otonephelium, Pometia, Xanihoceras\ hour-glass shaped
in Lepidopetalum; top shaped in Harpullia\ scarcely longer than broad in
Rhysotoechia and Toechima; provided with small transverse folds in Concho-
petalum. Veins vertically transcurrent in Alectryon, Athyana, Bridgesia,
Cardiospermum (larger veins only), Diatenopteryx, Euphoria, Hornea, Litchi,
Mischocarpus, Nephelium, Pappea, Paranephelium, Plagioscyphus, Porocystis,
Stadmannia, Synima, Thouinia, Thouinidium, Toulicia, Xerospermum; sur-
rounded by a ring of sclerenchyma in Tristiropsis. Smaller veins often
embedded in the mesophyll. The presence or absence of sclerenchyma around
the veins is said to be of value in the identification of species. Petiole
exhibiting, in transverse sections through the distal end, a continuous,
approximately circular strand or a circular group of separate strands. In the
material available for examination a continuous vascular strand was observed
in species of Koelreuteria (Fig. 100 B), Nephelium, and Xanthoceras(Fig. 1001),
and separate bundles in species of Dodonaea, Paullinia, Xanthoceras. Addi-
tional vascular strands sometimes present in the cortical region, e.g. in
Paullinia cupana Kunth (Fig. 100 c); medullary bundles also observed, e.g.
in Nephelium longana Lam. and Paullinia cupana. Four isolated bundles
recorded in the relatively herbaceous species Cardiospermum halicacabum
Linn. Main vascular strand of the petiole surrounded by a circle of scleren-
chymatous fibres in all of the species examined except Dodonaea viscosa (L.)
Jacq. Le Renard (1363) observed 3 separate vascular strands in the base of
the leaf of various species of Arfeuillea, Cossignia, Harpullia, &c., but trans-
verse sections at a higher level in the rachis revealed a triangular, and, near
the distal end of the rachis, a circular vascular strand formed by the fusion of
the 3 basal bundles. Crystals mostly solitary or clustered; crystal-sand
recorded in Exothea. Crystals particularly frequent in the epidermis in species
of Chytranthus, Conchopetalum, Filicium, Ganophyllum, Pancovia, Paullinia^
Pometia, Xerospermum. Clustered and/or solitary crystals generally present
in the soft tissues of the petiole none observed in Koelreuteria paniculata
;

Laxm. Secretory cells (see also Epidermis', Toung Stem', and Tetiole'),
either tubular and sometimes forming uniseriate rows of variable lengths, or
rounded and irregular in shape, commonly, but not universally, present in the
leaf,sometimes appearing as transparent dots. The contents, believed to be
a substance resembling saponin, are stated to resemble latex in living- speci-
mens, but appear to be clear or turbid and vary in colour from yellowish-
brown to brownish-black in herbarium material.Contents of the secretory
cells inErythrophysa and Stocksia give a tannin reaction; stated to consist of
chlorophyll in Paranephelium. Further details quoted by Solereder.

Axis
YOUNG STEM (Fig. 96 AHF and 100 D and F)
Surface with ribs (collenchymatous) in some species, e.g. Cardiospermum
halicacabum Linn. Cork nearly always arising in the sub-epidermis or outer
part of the primary cortex; originating within the pericyclic sclerenchyma only
in Distichostemon and Dodonaea. The phellogen in these last 2 genera produces
 SAPINDACEAE 433

sclerenchyma as well as cork, a second sclerenchymatous ring being formed in

the phelloderm. By repetition of this process the branch eventually becomes
covered by a 2- to 4-fold layer of cork and sclerenchyma. Primary cortex
collenchymatous, especially at the angles of the stem, particularly in species
climbing by tendrils ; containing groups of stone cells in certain species of
Atalaya, Melicocca, Sapindus, and Talisia or a ring of stone cells in Toulicia
y

guianensis Aubl. Le Renard (1363) observed collenchyma to be absent from

the cortex in species of Arfeuillea, Cossignia madgascariensis Bail., and Har-
pultia pendula F. v. M., whereas it occurred in all other species of Harpullia
examined as well as in Koelreuteria paniculata Laxm. Pericycle generally
containing a composite ring of sclerenchyma when sufficiently mature, but,
when younger, provided with an interrupted or almost continuous circle of
fibres in certain species. The occurrence of stone cells has not been demon-
strated in a few instances, notably in species of Cupania, Dodonaea, Valen-
zuelia, and Xanthoceras, but this may have been because the material was not
sufficiently mature. Primary phloem containing fibres with infrequent trans-
verse partitions in species of Cardiospermum and Serjania; phloem fibres
and/or sclerosed cells also observed by Le Renard (1363) in Arfeuillea,
Cossignia, Harpullia, and Koelreuteria. Secondary phloem in other genera
containing few or no fibres, but stone cells sometimes present in this position,
Xylem usually in the form of a continuous cylinder, traversed by rays 1-2
cells wide. Vessels commonly isolated, but sometimes in groups perforations
;

simple. Concentric zones of xylem parenchyma observed by Le Renard

(1363) in young stems of Cossignia madagascariensis Bail., but not in species
of Arfeuillea, Harpullia, &c. Pith sometimes including stone cells; consisting
almost wholly of these elements in species of Molinaea; composed of thin-
walled cells in Cardiospermum halicacabum Linn. Le Renard (1363) observed
(i) groups of rounded
sclerosed cells with pitted walls in the pith of Arfeuillea
and Harpullia thanatophora Bl. ; (ii) vertical rows of elongated sclerosed cells
in Harpullia cupanioides Roxb.; (iii) irregularly ramifying strands of more
rounded cells in other genera and species. Rudimentary vascular bundles, or
bundles consisting wholly of fibres, sometimes met with in the pith of Guioa,
Mischocarpus, Toulicia, and Tristiropsis. Secretory cells, similar to but less
easily visible than those of the leaf, present in the primary cortex and secondary
phloem as well as in the pith; varying in size and distribution in different
genera and species; sometimes arranged in longitudinal series. Both solitary
and/or clustered crystals common in the parenchymatous tissues; details
concerning their occurrence and distribution probably of value for the
identification of species. Besides crystals elsewhere in the ground tissue,
Le Renard (1363) observed them in a layer of endodermal cells in species of
Arfeuillea, Cossignia, Harpullia, and Koelreuteria.

ANOMALOUS STRUCTURE
Several types of anomalous structure have been recorded in lianes belonging
to this family, (i) A compound xylem-mass (Fig. 96 A) consisting of a
central ring of bundles, surrounded by but separated from, several peripheral
rings of bundles by a small amount of cortical parenchyma. The bundles in
the central and peripheral rings are each provided with a separate pith,
and each grows in thickness by its own cambium. The central and peripheral
424 SAPINDACEAE
bundles are interconnected at the nodes. This type of structure, which
produces ribs on the stems, is confined to 91 out of 172 species of Serjania
and 1 6 out of 122 of Paullinia\ it is unknown in other families except possibly
in the Leguminosae. (ii) The divided xylem-mass (Fig. 96 B) is similar to
the structure just described, but the central vascular ring is absent, the main
pith being surrounded by 5-7 rings of xylem and phloem, which remain open

Fro. 96. SAPINDACEAE

A, Compound xylem-mass of Serjania fuscifolia Radlk, B, Divided xylem-mass of Serjania corrugata
Radlk. C-E, Corded xylem-mass of Thinouia ventricosa Radlk. and other species at three successive
stages representing different ages. F-G, Cleft xylem-mass of Urvillea laevis Radlk. at two stages in the
growth. -After Radlkofer.

on the inside for a considerable time, so that their individual piths are
connected with the main one at the centre. This type is known only in

Serjania corrugata Radlk. and closely related species, (in) The corded xylem
mass (Fig. 96 C-E). In this type growth proceeds normally for 5 or 6 years,
after which accessory cambia arise in the cortical parenchyma externally to
the original vascular ring and produce subsidiary rings of xylem and phloem
each enclosing a separate pith. The accessory vascular rings are thus con-
nected with one another but not with the original vascular ring. This type is
stated to occur in all species of Thouinia, in a few of Paullinia, and sometimes
as a secondary complication in species exhibiting compound or divided xylem
masses as described above, (iv) The cleft xylem mass (Fig. 96 F-G). Here
 SAPINDACEAE 425
also the structure is at first normal, but superficially grooved, owing to certain
portions of the cortex being rather strongly depressed. The axis subsequently
becomes split into 3 or more portions corresponding to the grooves, owing to
the appearance of cambial tissue at these points. The separate portions of the
stem then grow in thickness through the activity of a cambium, which sur-
rounds each of them on all sides. Lastly a lobed xylem-mass originates by
increased development of the wood at 5 or more places at the circumference
of the stem; 5 or more ribs are formed, which project externally, and are
separated from one another by narrow grooves. This structure occurs in
some species of Serjania and Urvillea.
The exposed surface of the sapwood beneath the bark of the following trees
exhibits a 'corduroy' appearance owing to the presence of longitudinal corruga-
tions: Alectryon connatus Radlk., Arytera lautereriana Radlk., Diploglottis
cunninghami Benth., Guioa semiglauca Radlk., Sarcopteryx stipitata Radlk.
According to Francis (708) transverse sections of young stems of Arytera
lautereriana exhibit 5 indentations alternating with an equal number of ribs
on the surface of the woody cylinder, where they correspond to the outline of
the pith. Smaller indentations occur in the region of the node where they are
related to the median and lateral leaf traces. Transverse sections of a stem
9 cm. in diameter showed some of the depressions to correspond to small
areas of cells containing a brown tanniniferous substance, associated with
thick-walled pitted parenchymatous elements not found in the normal wood.
These areas were thought to be caused by injury, possibly by insects. The
prominent indentations of mature trees were found to be connected with pairs
of large aggregate rays terminating at the surface of the sapwood where the
indentations occur. The indentations in Sarcopteryx stipitata Radlk, are
likewise associated with similar pairs of large aggregate rays.

WOOD (Fig. 97)

Vessels typically small (mean tangential diameter less than 100 ^t), very
small (less than 50 /x) in Cupania p.p., Diplokeleba, Diatenopteryx, Dora-
toxylon, Exothea, Glossolepis Hippobromus, Pappea, Stadmannia, Thouinia,
y

and Thouinidium, medium-sized (100-200 p!) in Blighia, Castanospora^

Chytranthus, Cubitta, Cupaniopsis, Lecaniodiscus, Lepisanthes, Paranephelium,
Paullinia, Pometia, Sapindus, Schleichera, Schmidelia, Talisia, Tristira, Tristi-
and Ungnadia typically with numerous multiples of 2 or 3, the multiples
ropsis, ;

tending to be 8- rather than 0-shaped, with multiples of 4 or more moderately

common in Dodonaea, Hippobromus^ Laccodiscus, Nephelium and Placodiscus,
y

with fewer multiples and most of the vessels solitary in Blighia, Cubilia>
Cupaniopsis Deinbollia, Eriocoelum, and Pometia, with a tendency to a loose
y

oblique pattern (Fig. 97 H) in some specimens of Athyana, Blighia, Cupania

(1886), Hypelate (1886), and Lecaniodiscus; with a tendency to form longer
multiples locally in many species, probably in response to injury; Heimsch
(938) notes clusters in species of Dodonaea, Koelreuteria^ Pappea, Podo-
nephelium, Sapindus, and Tina; of 2 distinct sizes in Paullinia, the larger
solitary, the smaller in long radial multiples; varying iii number from about
2 to 50 per mm., mostly 2-5 and not more than 10 per mm. in woods with
medium-sized vessels, 10-50 per mm. in woods with very small vessels,
sometimes relatively few and widely spaced in woods with moderately small
426 SAPINDACEAE
vessels, e.g. Cupania, Erioglossum, Glossolepis, Placodiscus, and Stadmannia\
ring-porous or semi-ring-porous in some species of Hypelate (1864), Koel-
reuteria, and Sapindus\ spiral thickening not observed but reported (1206) in
species of Allophylus, Blighia, Cupaniopsis, Exotheca, Guioa, Koelreuteria,

FIG. 97. SAPINDACEAE

A, Placodiscus pseudostipularis Radlk. B, Sapindus mukorosi Gaertn. C, Ellatostachys nervosa (F.
Muell.) Radlk. D, Blighia sapida Kon. E, Cubilia rumphii Bl. F, Schleichera trijuga Willd. G,
Hemigyrosa deficiens Bedd. H, Blighia sapida Kon. I, Pometia pinnata Forst.

Sapindus, Thouinia, and Thouinidium. Perforations simple. Intervascular

pitting typically alternate and very small to minute, but rather larger (up to
moderate sized) in Allophyllus, Dembollia, Dodonaea, Eriocoelum, Harpullia y
Hippobromus, Lecaniodiscus, Placodiscus, Sapindus p.p., Schleichera, and
Tristira\ occasionally with distinct striations due to coalescent apertures,
e.g. in Blighia, Lecaniodiscus, Lepisanthes, Phialodiscus, and Pometia, and,
according to Solereder, in Dodonaea, Koelreuteria, Stocksia, and Valenzuelia,
and according to Kanehira (1206) in Otophora alata Blume and Xerospermum
noronhianum Blume; pits to parenchyma similar to the intervascular pitting
 SAPINDACEAE 427

except in Deinbollia and Eriocoelum p.p. in which some large oblong pits
occur. Solid deposits present in nearly all the species and often abundant,
gum-like and granular deposits and even crystals (Diplokeleba and Lepisanthes)
often occurring together in the same wood; Janssonius (1154) refers to the
presence of calcium carbonate in the vessels of certain species of Aphonia,
Lepisanthes, Mischocarpus, Sapindus, Schleichera, and Xerospermum', tyloses
rare but recorded by Janssonius in occasional vessels of Schleichera trijuga
Willd. and by Kanehira (1206) in Dodonaea, Nephelium, and Pometia. Mean
member length p*2-O'7 mm. Parenchyma in about 50 per cent, of the genera
scanty paratracheal only (apart from terminal), often very sparse and limited
to occasional cells adjoining the vessels; similar but also with crystalliferous
strands scattered among the
fibres in Arytera (1154), Cupania, Euphorianthus,
Exothea, Filicium, Koelreuteria (1206), Mischocarpus, Nephelium longana Lam.
(1206), Pappea, Ratonia (Baker 104), Schleichera (Fig. 97 F), Schmidelia, and
Stadmannia', Heimsch (930) notes 'diffuse and diffuse-in-aggregate' paren-
chyma as occurring, together with abundant vasicentric types, in Diateno-
pteryx, Elattostachys, Paullinia p.p., Serjania p.p., Stadmannia, Thouinidium,
Toulicia, and Xerospermum, and also in Pseudina, but with transitions to short
apotracheal bands; predominantly aliform, locally confluent but without
1
forming definite bands, in Dodonaea, Ganophyllum (1154), Hippobromus,
Melanodiscus, Nephelium, and Placodiscus (Fig. 97 A) similar but with
;

crystalliferous strands scattered among the fibres in Atalaya, Athyana,

Cupaniopsis, Diatenopteryx (crystals in idioblasts), Harpullia, and Vouarana;
abundant, in broad confluent bands containing numerous crystalliferous
strands, particularly on their margins, in Aphonia, Erioglossum, Hemigyrosa
(Fig. 97 G), Lepisanthes, Meliococca, Octophora (1154), Sapindus, Tristiropsis,
and Zanha\ similar but with narrower bands and without crystalliferous
strands in Diplokeleba and Napeodendron. Record and Hess (1886) note
broad bands also in Dipterodendron, Pseudina, and Talma, though not
observed by the author in the latter. Predominantly diffuse, though with
abundant vasicentric parenchyma in Toulicia guianensis Aubl. With terminal
bands in Atalaya, Cupania (938), Exothea, Filicium (938), Glossolepis, Har-
pullia,Hippobromus, Hypelate (938), Melanodiscus, Napeodendron, Nephelium,
Placodiscus, Podonephelium (938), Pometia (938), Sapindus, and Toulicia', with
terminal parenchyma only in Paranephelium\ with idioblasts containing large
single crystals in Diatenopteryx. Strands typically of 4 cells, but strands of
2 cells moderately common in Athyana and Filicium and predominant in
Diplokeleba and Hippobromus sometimes up to 6 or 8 cells in Napeodendron,
\

Nephelium p.p., Pometia, and Toulicia', some fusiform cells present in

Diplokeleba and Hippobromus. Storied in Diplokeleba. Baker (104) describes
chambered crystals in Ratonia tenax Benth. Rays of most genera exclusively
uniseriate or predominantly uniseriate with a few narrow biseriates commonly ;

biseriate in Deinbollia, Eriocoelum p.p., Exothea, Filicium, Harpullia, Melano-

discus, Meliocca, Talisia, Tristira, and Zanha, up to 3 cells wide in Aphania,
Diplokeleba, Dodonea p.p., Sapindus (though sometimes up to 5 cells wide
in S. mukorosi Gartn. and in S. rarak DC. (1154)), Schleichera p.p., and,

1
The material of Dodonaea viscosa (L.) Jacq. described by Janssonius (1154) falls into
this group, but the two specimens examined by the author are typical of the main group with
scanty vasicentric parenchyma only.
428 SAPINDACEAE
according to Heimsch (938) in somespecies oi Paullinia and Serjania. Solereder
records that comparatively broad rays are found in places in Erioglossum
rubiginosum Bl. and Serjania faveolata Radlk, and Francis (706) notes
'aggregate' rays associated with deep indentations in Sarcopteryx\ aggregate
rays also reported (1881) in some species of Matayba\ typically low and
rarely reaching 'i mm. in height except in species of Paullinia and Serjania
(938). Uniseriates often more numerous in the species with multiseriate rays
than is usual with homogeneous rays and suggestive of transition to wholly
uniseriate types; occasionally very few (Kribs's Homogeneous Type II),
e.g. m some species of Exotheca, Sapindus, and Tristiropsis (938); composed
wholly of procumbent cells, except in the few species with heterogeneous rays.
Mostly between 8 and 14 rays per mm., fewer than 4 per mm. in Sapindus
mukorosi, 4 to 7 per mm. in Deinbollia, Eriocoelum p.p., and Exothea, 15 or
more per mm. in Blighia, Cubilia, Cupania, Diatenopteryx, Diploglottis,
Diplokeleba, Euphoria, Glossolepis, Laccodiscus, Phialodiscus, Placodiscus,
Schleichera p.p., and Stadmannia, exceptionally numerous (20 per mm. in
Diatenopteryx', typically homogeneous (Kribs's Types I and III, rarely II)
and commonly with cells small in tangential section (about 10 ^ wide tan-
gentially), cells larger and mostly square or upright in Cupaniopsis, Deinbollia,
Glossolepis, Placodiscus, and Pometia p.p., heterogeneous (Kribs's Types II B
and ill) with 1-2 marginal rows of upright cells or with occasional rows of
square or upright cells interspersed among the procumbent cells, in some
species of Allophylus, Cupaniopsis, Deinbollia, Eriocoelum, Exotheca, Glossolepis,
Harpullia, Pometia, and Talisia; heterogeneous in most species of Paullinia
and Serjania (938); crystals present in the ordinary cells of many species,
sometimes abundant, e.g. in Paranephelium, Phialodiscus, and Ratonia (Baker
104), chambered crystals present in Euphorianthus, Laccodiscus, and Stad-
mannia, and with dark gum-like deposits in many species. With echelon
arrangement in Athyana, Cupaniopsis, Diatenopteryx, Eriocoelum, Erioglossum,
Euphorianthus, Hippobromus, Lepisanthes, Meliococca, Sapindus p.p., Sar-
copteryx, and Xerospermum ; storied in Diplokeleba. Fibres with simple pits,
more numerous on, the radial than on the tangential walls. Septate in the
majority of species, the septa relatively rare or absent from some of the woods
with thick- walled fibres; septa comparatively rare in Aphania, Atalaya,
Athyana, Doratoxylon, Filicium, Lecaniodiscus, Nephelium, and Pometia\ septa
not observed in Diplokeleba, Elattostachys, Erioglossum, Glossolepis, Guioa,
Harpullia> Hemigyrosa, Hippobromus, Hypelate, Lepisanthes, Napeodendron,
Placodiscus, Schleichera, and Stadmannia\ absent also, according to Solereder,
from Cossignia 9 Diplopeltis, Distichostemon, Llagunoa, Loxodiscus, Serjania,
Urvittea, and Valenzuelia. With thick walls in Aphania, Athyana, Cupania
p.p., Diplokeleba, Dodonaea, Doratoxylon, Filicium, Hemigyrosa, Hypelate,
Lepisanthes, Napeodendron, Nephelium> Pappea, Paranephelium, Placodiscus,
and Stadmannia. Solereder refers to the absence of really wide lumina, cf.
Tilta, and the consequent dense character of the woods of this family, stating,
however, that relatively wide lumina occur in Koelreuteria paniculata Laxm.,
Porocystis toulicioides Radlk., and Toulicia guianensis Aubl. to this list may be
;

added Jagera pseudorhus (A. Rich.) Radlk., Matayba ingaefolia Standl., and
Xerospermum glabratum Radlk. and, according to Janssonius (1154), the
septate fibres of Guioa diplopetala Radlk. In Allophyllus there is often a
 SAPINDACEAE 429
tendency to form irregular, parenchyma-like bands of septate fibres with
thinner walls; according to Heimsch (938), these fibres may be rounded in
cross-section and with pronounced intercellular spaces. Similar bands are
noted by Heimsch in Paullinia and Serjania* In Paranepheliwn macrophyllum
King, the fibres are sharply segregated into 2 types, those of the early wood
thin-walled, with wide lumina and numerous septa, those of the late wood
thick- walled, with narrow lumina and only occasional septa; a similar distinc-
tion, thoughless marked, occurs in some other woods with distinct growth

rings, e.g. Sapindus spp. Many of the species with little parenchyma have
vasicentric sheaths of wide septate fibres that resemble parenchyma strands.
Intercellular spaces conspicuous in some species, particularly those with
thinner walls, e.g. Jagera and Matayba. Starch or gum-like substances some-
times present and occasionally abundant, e.g. Laccodiscus. Solereder quotes
Ilohnel as stating that the fibres, but not the rays, are storied in Aphonia
senegalensis (Juss.) Radlk. Mean length 0-6-1*5 mm., more than i mm. in
Matayba (Kribs 1283), Pometia, and Sapindus. Intercellular canals
observed in the rays of one specimen 1 of Deinbollia grandiflora Hook, f., and
vertical canals of the gummosis type described by Solereder in Dilodendron
bipinnatum Radlk. and by Record and Hess (1886) in Diptokeleba. Anomalous
structure, see previous section. According to Besson (186), the wood of
Blighia sapida Kon. has a very low silica content. The development of the
growth ring in Schleichera is described by Coster (481).

TAXONOMIC NOTES
Akania, which was at one time included in this family, has been described
under Akaniaceae, see p. 436.
The wood anatomy of this family is very homogeneous and is moderately
highly specialized, particularly as regards the rays. Heimsch (938) places the
family in a group composed of the Anacardiaceae, Burseraceae, Meliaceae,
Rutaceae, Sapindaceae, and Simarubaceae and considers that there is strong
anatomical evidence for believing this to constitute a natural group of plants.
Differences exist between the families, e.g. the Anacardiaceae and Bur-
seraceae stand apart owing to the invariable occurrence of intercellular canals
in the phloem and cortex, and the occurrence of septate fibres links the
Sapindaceae with the Meliaceae rather than with the Rutaceae. Nevertheless,
Heimsch considers that these differences are not absolute and do not serve to
delimit one group from another without exceptions. He notes that the
Sapindaceae are more highly specialized with respect to rays than any of the
other families in this group.
Heimsch also considers that the wood anatomy, particularly the highly
specialized rays, supports the view that the Aceraceae and Hippocastanaceae
are close to the Sapindaceae. It should be noted, however, that these families
lack septate fibres, which are characteristic of the Sapindaceae.

ECONOMIC USES
Apart from those members of the family which yield useful timbers, the
fruits of various species are of economic importance. The saponin present in

1
G. Proctor Cooper no. 357.
430 SAPINDACEAE
the tissue enables the fruits of various species of Sapindus, known as 'Soap-
Berries', to be used for washing purposes. Other fruits such as Litchis (Litchi
chinensis Sonn.) are edible, and sometimes sold by greengrocers in Great
Britain. The Akee Apple of West Africa is obtained from Blighia sapida Koen.
The seeds of Paullinia cupana Kunth. are used in Brazil for the preparation
of Guarana Bread. The timbers, many of which are dense, tough, and fine-
textured, are of little importance. Pearson and Brown (1679), however, include
Schleichera trijuga Willd. and Filicium decipiens Thw. among the commercial
timbers of India.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Alectryon,* Allophylus, Aphania, Aphanococcus, Aporrhiza, Arfeuillea,
Arytera, Atalaya, Athyana, Averrhoidium, Blighia, Bridgesia, Cardio-
spermum, Castanospora, Chytranthus, Conchopetalum, Cossignia, Cotylo-
discus, Crossonephelis, Cupania,* Cupaniopsis, Deinbollia, Diatenopteryx,
Dictyoneura, Diploglottis, Diplopeltis, Distichostemon, Dodonaea,* Dora-
toxylon, Elattostachys, Eriandrostachys, Eriocoelum, Erioglossum, Erythro-
physa, Euphoria, Exothea, Filicium, Ganophyllum, Gleniea, Gongro-
discus, Guioa, Haplocoelum, Harpullia, Hebecoccus, Heterodendron,
Hippobromus, Hornea, Hypelate, Jagera, Koelreuteria,* Laccodiscus, Leca-
niodiscus, Lepiderema, Lepidopetalum, Lepisanthes, Litchi, Llagunoa,
Loxodiscus, Lychnodiscus, Macphersonia, Magonia, Matayba, Melanodiscus,
Melicocca, Mischocarpus, Molinaea, Nephelium,* Otonepheliurn, Otophora,
Pancovia, Pappea, Paranephelium, Paullinia,* Phialodiscus, Placodiscus,
Plagioscyphus, Podonephelium, Pometia, Porocystis, Pseudima, Pseudone-
phelium, Rhysotoechia, Sapindus,* Sarcopteryx, Sarcotoechia, Schleichera,
Scyphonychium, Serjania,* Smelophyllum, Stadmannia, Stocksia, Stortho-
calyx, Synima, Talisia, Thinouia, Thouinia, Thouinidium, Tina, Toechima,
Toulicia, Trigonachras, Tripterodendron, Tristira, Tristiropsis, Ungnadia,
Urvillea, Valenzuelia, Xanthoceras,* Xerospermum, Zollingeria.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Allophyllus, Aphania, Arytera, Atalaya, Athyana, Blighia, Castanospora,
Chytranthus, (Cossignia), Cubilia, Cupania, Cupaniopsis, Deinbollia, Dia-
tenopteryx, Diploglottis, Diplokeleba, (Diplopeltis), (Distichostemon),
Dodonaea, Doratoxylon, Elattostachys, Eriocoelum, Erioglossum, Euphoria,
Euphorianthus, Exothea, Filicium, (Ganophyllum), Glossolepis, Harpullia,
Hemigyrosa, Hippobromus, (Hypelate), Jagera, (Koelreuteria), Laccodiscus,
Lecaniodiscus, Lepisanthes, (Llagunoa), (Loxodiscus), Matayba, Melano-
discus, Melicocca, Mischocarpus, Nephelium, (Otophora), Pappea, Parane-
phelium, Paullinia, Phialodiscus, Placodiscus, (Podonephelium), Pometia,
(Porocystis), Pseudima, (Ratonia), Sapindus, Sarcopteryx, Schleichera,
(Serjania), Stadmannia, Talisia, Thouinia, Thouinidium, (Tina), (Toulicia),
Tristira, Tristiropsis, Ungnadia, (Urvillea), (Valenzuelia), Vouarana, Xero-
spermum, Zanha.
 SAPINDACEAE 431
LITERATURE
(i) On General Anatomy
Francis 708, Guillaumin 840, Le Renard 1363, Radlkofer 1770, Sabnis 1977.
(ii) On Wood Structure
Baker 104, Beekman 167, den Berger 179, 182, Besson 186, Brown, F. B, H. 282,
Burgerstein 310, 312, Cooper 461, Coster 481, Francis 706, Giordano 786, Greguss 2522,
Heimsch 938, Holden 987, Howard 1088, Janssonius 1154, Jolly 1188, Jones 1x91,
Kanehira 1206, 1209, Kribs 1283, Lecomte 1334, Pearson and Brown 1679, Pfeiffer, H.
1711, 1712, Pfeiffer, J. Ph. 1713, Record 1781, 1783, 1787, 1801, 1809, 1843, 1851,
1881, Record and Hess 1886, Record and Mell 1894, Stone 2202, Torres 2269, Williams
2426, 2430, Yamabayashi 2478.

102. ACERACEAE
(FiG, 93 on p. 412; FIG. 98 on p. 434)

SUMMARY
(i) GENERAL
Trees or shrubs which occur in temperate regions in the northern hemi-
sphere. Nearly all of the anatomical work on this family has been confined to
the genus Acer. The hairs are (i) unicellular or uniseriate, with a tendency
to be 2-armed in certain species; (ii) club-shaped and glandular with a uni- or
biseriate stalk; (iii) transitional between (ii) and shaggy types. The leaf
epidermis is often partly or wholly mucilaginous. Stomata, mostly confined
to the lower surface, are ranunculaceous. Secretory sacs, which are most
clearly visible in. sections treated with eau de javelle, occur in the phloem of
the leaf veins and axis, or, less frequently, in the mesophyll. In some instances
they contain typical latex, but in others are filled with highly refractive con-
tents which can be made clear by mounting dry sections in olive-oil. Idio-
blasts containing mucilage or crystals also occur in the leaf. The crystals
are mostly solitary or clustered, but sphaerites and rod-shaped types some-
times occur as well. Smaller leaf veins generally vertically transcurrent. The
petiole, in transverse sections of the few species examined, exhibits a ring
of vascular bundles, enclosing medullary strands in some species.

(ii) WOOD
Vessels moderately small, with spiral thickening, perforations simple,
intervascular pitting alternate and moderately large, pits to parenchyma
similar; members of medium length to moderately short. Parenchyma in
terminal bands composed mainly of fusiform cells, otherwise absent or with
a few cells round the vessels or, more rarely, with diffuse chambered crystalli-
ferous cells. Rays mostly 5-7 cells wide and homogeneous. Fibres with
simple pits, moderately to very short.

LEAF
Usually dorsiventral, but sometimes tending to be centric. Hairs uni-
cellular or uniseriate; 2-armed types recorded from one species of Acer]

glandular club-shaped with a uni- or biseriate stalk together with transitions

between these and shaggy forms. Cuticle occasionally striated; sometimes
coloured white by a coating of wax on the lower surface. Epidermis com-
monly mucilaginous; papillose on the lower surface in numerous species of
432 ACER ACE AE
Acer; including tanniniferous idioblasts on the lower side in A. vittosum Wall.
Stomata nearly always confined to the lower surface; sometimes rather small
in diameter, recorded as infrequent on the upper side of the leaf in the region
of the veins in Acer japonicum Thunb. ranunculaceous. Hypoderm not
;

recorded in Acer. Mesophyll generally including a single layer of palisade

cells, but up t6 4 layers recorded in A. negundo Linn. Vascular bundles of the

larger veins usually surrounded by a ring of sclerenchyma, except in species

of Acer belonging to the section Indivisa where arcs of or isolated sclerenchy-
matous elements occur. Small veins vertically transcurrent by thin- or thick-
walled tissue in all species of Acer. Petiole (Fig. 93 D), in transverse sections
through the distal end, exhibiting an adaxially flattened ring of separate
bundles in all investigated species; medullary bundles sometimes present as
well, e.g. in Acer negundo, A. opalus Mill., A. rubrum Linn, (a single medullary
strand only (Plowman 1732)). Main ring of vascular bundles in the petiole
surrounded by fibres in A. campestre Linn, and A. opalus, but not in A.
platanoides Linn. Plowman records very small medullary strands in A. sac-
charinum Linn., but observed none in A. saccharum Marsh or A. platanoides.
It is possible that the medullary strands are somewhat variable within a
species, since they were seen to be present in a variety of A. platanoides
examined at Kew. Watari (2365) has published a very detailed account of the
petiole structure in 42 species of Acer from Japan and Formosa. In all of
them the fundamental structure is similar to that described above, if allowance
be made for complications which occur at different levels in a single petiole,
especially at the apex and base. The number and size of the individual
bundles both in the main outer circle and in the medullary system vary in
different species. So far as the medullary bundles are concerned, these are
absent from some species, extend throughout the length of the petiole in
others, whilst in a third group of species they are confined to the distal end.
Variations in the petiolar structure are thus of diagnostic value in the identi-
fication of species provided that comparisons are made between sections
taken from the corresponding positions in each species. The petiole structure
of Acer has, more recently, been studied by Cortesi (477) who observed
medullary bundles in 6 out of the 25 species examined. Solitary and/or
clustered crystals fairly frequent in parenchymatous tissues. Idioblasts con-
taining small crystalline masses and a large solitary crystal, often with its long
axis at right angles to the surface of the leaf, recorded in Acer negundo and
other related species of Acer; often appearing as transparent dots in the leaf.
Similar idioblasts, but containing clustered crystals or sphaerites, present in
species of Acer belonging to the sections Indivisa and Macrantha as well as in
A. glabrum Torr. Secretory sacs, i.e. elongated secretory cells sometimes
occurring in series, stated to be always present in the phloem of the veins,
and less frequently in the mesophyll of Acer; clearly demonstrated by treating
sections with eau de javelle; containing typical latex in some instances, but
with strongly refractive contents, made clear by mounting dry sections in
olive-oil in others.

Axis
YOUNG STEM (Fig. 93 G)
Cork usually superficial in origin, generally arising in the outer part of the
 ACER ACE AE 433
cortex, but more deeply seated in species secreting wax from the branches.
Cortex containing isolated or clustered crystals in various species of Acer,
and stone cells in A. negundo Linn, and A. opalus Mill. Pericycle somewhat
variable with a composite and continuous ring of sclerenchyma in A. negundo
; ;

with an interrupted composite ring of sclerenchyma in A. pseudoplatanus

Linn. with isolated strands of fibres in various other species of Acer as well
;

as in Dipteronia, Secondary phloem containing bundles of sclerenchyma,

which are stratified in at least certain species of Acer, e.g. A. platanoides Linn,
and Dipteronia. Phloem in species of Acer and Dipteronia also containing
groups of stone cells, chambered parenchyma containing solitary crystals,

secretory sacs or cells similar to those described for the leaf. Other secretory
cells with amorphous contents of a different kind, possibly mucilaginous,
occur in the cortex, phloem, and pith in at least certain species of Acer.
Xylem soon forming a continuous cylinder, but consisting of a circle of
individually distinct bundles in transverse sections through very young stems
of Acer. Vessels usually with simple perforations. Pith usually large. Stems
of A. negundo readily giving rise to adventitious roots when in water or moist
soil (Plowman 1732). Secretory elements, and crystals, see 'Cortex' and
Thloem'.

WOOD (Fig. 98 A-B)

Vessels moderately small (50-100 [L mean tangential diameter); solitary
and in multiples of 2 or 3 cells in Acer and with numerous clusters in
Dipteronia (938); 30-60 per sq. mm.; with spiral thickening. Perforations
simple; the smallest vessels occasionally with foraminate perforation plates.
Intervascular pitting typically alternate, but, according to Heimsch (938),
with not- infrequent regions of opposite pitting; sometimes moderately large,
and often with hexagonal borders in Acer; pits to ray and wood parenchyma
similar to the intervascular pitting in Acer, but tending to be oblong in
Dipteronia. Usually empty; occasionally with deposits of gum; Solereder
9
refers to deposits of calcium carbonate in Acer rubrum L. and 'A. illyricum
(A. monspossulanum L. ?), and Record (1818) mentions similar deposits in
knots, bird's-eye, and dark streaks in various species. Mean member length
o-3~-o-5 mm. Parenchyma very sparse or absent except at the boundary of
the growth ring; usually with a few cells round the vessels, e.g. A. ginnala
Maxim, and A. pseudosieboldianum Komar, occasionally with scattered cells
containing chambered crystals, e.g. A. yiacrophyllum Pursh. and A. opalus
Mill. the terminal parenchyma varying from a few scattered cells to bands
;

i to several cells wide and

typically composed of thick-walled fusiform cells
with an occasional strand divided into 2 cells, strands of 2-4 cells moderately
common in a few species, e.g. A. thompsoni Miq. and sometimes containing
crystals; strands of paratracheal parenchyma usually of 4 cells. Rays 2-10,
mostly 5-7 cells wide, with a tendency to be of 2 distinct sizes in some species
or specimens, e.g. A. nigrum Michx., A, saccharum Marsh, and A. pseudo-
platanus L. less than i mm. in height except in a few specimens, e.g. of
;

A. pictum Thunb. and A. saccharum Marsh; with moderately numerous uni-

seriates composed of procumbent cells, and often only a few cells high; 3-12,
mostly 6-10 rays per sq. mm.; homogeneous (Kribs's Type I) in Acer
and composed of small cells; heterogeneous (Kribs's Type lie) to almost
4594 F f
434 ACER ACE AE
homogenous in Dipteronia (938). Usually with gum-like contents; crystals
recorded (938) in Acer obhngum Wall. Fibres typically with simple pits,
rare,
though narrow borders occur in a few species; pits more numerous on the

H
FIG. 98. ACERACEAE, A-B; AKANIACEAE. C-D; MELIANTHACEAE, E-F;
HIPPOCASTANACEAE, G-H
A, Acer saccharum Marsh. B, A. pseudoplatanus Linn. C, Akania hillii Hook. f. D, A. hillii Hook. f.
E, Bersama paullinioides Baker. F, B. leiostegia Stapf* G, Aesculus hippocastanum Linn. H,, A. hippo-
castanwn Linn.

radial than on the tangential walls; often with conspicuous intercellular spaces.
Mean length 0-6-0-9 mm. Holden (987) has pointed out that the fibres have
noticeably thicker walls in the neighbourhood of the vessels and H. P. Brown
(1679) refers to elements transitional between fibre-tracheids and terminal
parenchyma in the outer part of the ring of some species. Heimsch (938)
states that bands or areas of starch-storing fibres are characteristic of Acery
but points out that they may be rendered obscure by common section-cutting
 ACER ACE'AE 435

techniques. Growth ring formation of Acer in Java is described by Coster

(481) and in the U.S.A. by Hanson and Brenke (888) and Lodewick (1386);
the seasonal starch content and cambial activity of the stem and roots in
England has been investigated by Cockerham (439).
BARK
The
bark of various species of Acer has been investigated by Plowman
(1732). Its thickness varies in different species, as also the width and degree
of undulation of the medullary rays. Sclerotic elements are always present;
they form definite zones in some species but are more irregularly distributed
in others. Acer platanoides Linn, contains fewer lignified elements than the
other species examined. Crystalliferous and tanniniferous cells occur in
different proportions according to the species.

ROOT
Plowman (1732) has recorded the following information about the root of
A. negundo Linn. Bark of young roots thin, containing very few sclerotic,
crystalliferous, and tanniniferous cells, but including large sap-storage cells
and canals. Rays numerous, straight, 1-2 cells wide. Vessels large and
numerous often in radial clusters of 5. Ground tissue of the wood consisting
of groups of 2 distinct sorts of tracheids with thin and thick walls respectively.
Tannin abundant in old roots.
TAXONOMIC NOTES
Acer and Dipteronia, which are now generally regarded as members of the
Aceraceae, were included in the Sapindaceae in the Bentham and Hooker
system. The absence of septate fibres distinguishes these genera from the
Sapindaceae, though the wood anatomy in general suggests a close relation-
ship. Plowman (1732) has drawn attention to the morphological distinctions
between Acer negundo Linn, and the other species of Acer, and, after consider-
ing these and the geological history of A. negundo, concludes that the species
should be given generic rank with the name Negundo aceroides Moench.
Heimsch (938), however, points out that differences in the xylem are too
slight to support this change. The species is still generally known as A.
negundo.
ECONOMIC USES
The bark of mostspecies of Acrr contains sugar, but in only a few of them,
such as A. saccharinwn Wang., does it occur in sufficient quantities to make
its extraction possible on a commercial scale. The sap containing the sugar is

obtained by tapping the trunks. The genus Acer furnishes some very impor-
tant timbers, e.g. the Hard or Rock Maple (A. saccharum Marsh), Soft Maple
(A. rubrum and A. saccharinuni), and the Sycamore (A. pseudoplatamis L.).
Not only is the normal straight-grained timber in demand, but the wood is
often figured and highly prized for particular purposes, e.g. the curly grain
or 'fiddle-back* figure for violins and 'bird's-eye' Maple for veneers.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acer,* Dipteronia,
* Kew
Represented in the slide collection.
436 ACER ACE AE
(ii) FOR WOOD STRUCTURE
Acer, (Dipteronia).
LITERATURE
(i) On General Anatomy
Cortesi 477, Hanson and Brenke 888, Plowman 1732, Sinnott 2108, Watari 2365.

(ii) On Wood Structure

Bienfait 197, Brown, H. P. 283, 288, Chalk 365, Cocker-
Beekman 167, den Berger 179,
ham 439, Coster 481, Desch 574, Greguss 2522, Hale 870, Hanson and Brenke 888,
Heimsch 938, Hess 961, Holden 987, Holmes 1079, Howard 1088, Janssonius 1154,
Jones 1191, Kanehira 1204, 1206, Lodewick 1386, Moiseeva 1545, Nicoloff 1593, Pearson
and Brown 1679, Record 1778, 1783, 1818, 1843, ^S 1 ? Record and Hess 1886, Stone
2203, Tang 2231, Yamabayashi 2478.

103. AKANIACEAE
(FiG. 98 on p. 434; FIG. 99 below)

SUMMARY
A family of small trees from eastern Australia belonging to the single genus
Akania. The following general account is based on material of Akania hillii
Hook, grown at Kew, and the wood anatomy on a single specimen of the same

B
FIG. 99. AKANIACEAE
A, Akania hillii Hook. f. Petiole x 8. B, A. hillii Hook. f. Portion of stem X 18.

species from Australia. The wood exhibits the following characters. Vessels
with simple or rare scalariform perforation plates, alternate to opposite inter-
vascular pitting and elongated pits to ray cells members of medium length. ;

Parenchyma paratracheal, scanty. Rays very wide and high, heterogeneous,

uniseriates absent. Fibres with bordered pits, septate, of medium length,

LEAF
Dorsiventral. Hairs very infrequent. Upper epidermis consisting of
cellswith slightly curved anticlinal walls; cells of the lower epidermis with
more sinuous anticlinal walls. Stomata confined to special depressions in the
lower surface of the leaf, where they occur amongst very characteristic, closely
congested, slender papillae with crowned apices. Mesophyll including 2
 AKANIACEAE 437
layers of rather short palisade cells, but about two-thirds of its thickness is
made up of very lacunar, spongy tissue. Vascular bundles of the smaller
veins embedded in the mesophyll, each surrounded by a sheath of fibres.
Vascular system in the midrib of the leaflets consisting of dorsal and ventral
arcs of bundles, strongly supported by fibres both externally and internally,
the centre of the strand being occupied by pitted cells with fairly wide lumiiia.
Petiole (Fig. 99 A), in transverse sections towards the apex, exhibiting a
dorsally flattened circle of individually distinct but closely packed collateral
bundles, each supported by slightly less thickened fibres on the inside of each
xylem group. Centre of the petiole occupied by an extensive parenchymatous
tissue. Cortical region narrow. Scattered secretory cells with amorphous
contents present in all unlignified tissues; more frequent in the palisade than
in the spongy portion of the mesophyll. Large solitary crystals present in
some of the cells immediately external to the sclerenchymatous sheaths around
the vascular bundles of the veins.

Axis
YOUNG STEM (Fig. 99 B)
Cork arising in the sub-epidermis. Cortex narrow, including scattered
secretory cells with amorphous contents. Pericycle marked by a broad,
almost continuous ring of thick-walled fibres, the fibrous ring being inter-
rupted by small groups of secretory cells opposite the medullary rays. Xylem
and phloem forming individually distinct vascular bundles separated by
fairly broad medullary rays. Phloem consisting of conspicuous groups of
thin-walled tissue devoid of sclerenchymatous elements. Vessels solitary and
variously clustered; up to about 60 p, in radial diameter; perforations mostly
simple, but a few scalariform perforation plates with numerous bars also
observed ; sometimes with a few ty loses. Pith very broad, consisting mostly of
pitted parenchymatous cells, but also including vertical columns of secretory
cells with amorphous contents. Coarse cluster crystals fairly frequent in the

parenchymatous tissues; only a few solitary crystals observed. Secretory

elements, see 'Cortex, Tericycle', and Tith*.

WOOD (Fig. 98 C-D)

Vessels moderately small (50-100 p mean tangential diameter), solitary
and in radial, tangential, or oblique multiples and clusters; about 20 per sq.
mm. Perforation plates simple and moderately oblique, or very rarely scalari-
form. Intervascular pitting moderately small, alternate or opposite (533),
pits to ray and wood parenchyma cells commonly large, horizontally elongated
and simple; mean member length about 0*75 mm. Parenchyma para-
tracheal, limited to a few cells round the vessels, and terminal, in bands i or
2 cells wide. Strands mostly of 6-8 high cells. Rays up to 16 cells and over
400 fi wide; up to 6 mm. high; with very few or no uniseriates; about 2*5 per
mm.; heterogeneous, commonly with 2-6 marginal rows of upright cells,
which are often uniseriate only at the extreme tips; sheath cells present;
solitary crystals present in chambered and ordinary cells. Fibres with
numerous, small bordered pits, on both radial and tangential walls; commonly
septate and with moderately thick walls; mean length about 1*5 mm.
 FIG. 100. HIPPQCASTANACEAE, A and E; SAPINDACEAE, B-D, F, and I;
STAPHYLEACEAE, G and J; MEL1ANTH ACEAE, H and K
A, Aesculu* hippocastanum Linn. Petiole x6. B, Koelreuteria paniculata Laxm. Petiole xn.
C, Pauttinia cupana H. B. et K. Petiole X 17. D, Koelreuteria paniculata Laxm. Stem X ig. E,
Aesculus hippocastanum Linn. Stctn X 7. F, Pauttinia cupana H. B, et K. Stem X 10. G, Staphylea
hohcarpa Hemsl. Petiole X ig. H, Metianthus major Linn. Petiole X 9. I, Xanthoceras sorbifoha Bgc.
Petiole X 19. J, Staphylea holocarpa Hemsl. Stem X 19. K, MeUanthus major Linn. Young stem X 9.
K
Small concentric circles in represent bundles with central phloem.
 AKANIACEAE 439
TAXONOMIC NOTES
The taxonomic position of Akania has been much discussed. A. hUlii
Hook., originally described somewhat imperfectly by J. D. Hooker, was first
of all assigned to the Sapindaceae. F. Mueller and other botanists sub-
sequently treated it as a member Stapf (2185) made
of the Staphyleaceae.
Akania the basis of a new family the Akaniaceae, and fully discussed his
reasons for so doing. Harms (905) likewise treats the Akaniaceae as a distinct
family. The very wide rays and the absence of uniseriates render the secondary
xylem of Akania quite distinct from any of the woods of the Sapindaceae.
For further details the articles by Stapf and Harms should be consulted*

GENUS DESCRIBED
Akania.*
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Harms 905, Stapf 2185.
(ii) On Wood Structure
Dadswell and Record 533, Heimsch 938, Record 1843, 1851.

104. HIPPOCASTANACEAE
(FiG. ioo on p. 438)

SUMMARY
(i) GENERAL
A small family of trees and shrubs consisting of the genera Aesculus from
north temperate regions and Billia from tropical America. In a general way
the anatomical characters resemble those of the Sapindaceae, and it will
suffice to record the few characters which follow. Unicellular and uniseriate
hairs have been recorded in both genera. The cells of the epidermis of the
leaf sometimes have delicate, vertical, secondary walls in Billia> whilst,

according to Solereder, mucilaginous cells are absent from the epidermis in

both genera. In this last respect they differ from many of the Sapindaceae.
Sclerosed cells have been recorded in the mesophyll in species of Billia.
The vascular bundles of the veins are often vertically transcurrent in
Aesculus. Transverse sections through the distal end of the petiole show a
continuous cylindrical vascular strand in Aesculus californica Nutt. and A.
hippocastanum Linn. (Fig. ioo A) whilst in the last species medullary bundles
are present as well. Groups of stone cells have been recorded in the primary
cortex of the young stem in species of Billia, whilst a composite, continuous
ring of sclerenchyma in the pericycle is characteristic of all the species
which have been examined. Xylem (Fig. ioo E) in the form of a continuous
cylinder traversed by narrow rays. Vessels usually with simple perforations;
but occasional scalariform plates also recorded. Cluster and a few solitary
crystals observed.
440 HIPPOCA S TA NA CEA E
(ii) WOOD
Vessels small and with numerous multiples, usually numerous, perfora-
tions simple, intervascular pitting alternate, pits to parenchyma usually
similar, sometimes larger or unilaterally compound, solid deposits and tyloses
often present; members of medium length. Parenchyma (a) terminal,
(b)scanty paratracheal, sometimes very sparse or absent; often storied. Rays
exclusively uniseriate, homogeneous or weakly heterogeneous, often storied.
Fibres with simple to distinctly bordered pits, moderately short.

Axis
WOOD (Fig. 98 G-H)
Vessels in most species of Aesculus very small (less than 50 mean tan-
//,

gential diameter) to moderately small (50-100 /x); often angular, except in

Billia (938); with numerous multiples and clusters, multiples of 4 or more
cells common in most species; more than 40 per sq. mm. and often up to

70 or more; with spiral thickening in all species of Aesculus. Perforations

typically simple, but some scalariform plates have been reported (938, 2158)
in a few species of Aesculus and in twigs of Billia. Intervascular pitting
typically alternate, but some opposite pitting commonly present (938); pits
to ray cells often limited to the marginal rows, usually similar to the inter-
vascular pitting, but sometimes horizontally elongated or unilaterally com-
pound, e.g. in A. glabra Willd. According to Bailey (78), the sieve-like
structures described by Jonsson (i 192) are not true vestured pits. Sometimes
with solid deposits or tyloses; Record (1818) reports deposits of calcium
carbonate as occasionally present in the wood of old knots. In A. punduana
Wall, and Billia (1275) the vessels are larger, not angular, less numerous
(10 per sq. mm.) and with fewer multiples, and Kramer (1275) distinguishes
Billia by the absence of tyloses. Mean member length (Aesculus) 0-4-0-5 mm.

Parenchyma in narrow, uni- to biseriate terminal bands in Aesculus, com-

posed of strands of 4 cells that tend to be storied the terminal bands wider
;

in BilUa (938); parenchyma other than terminal, absent, or very sparse, as

occasional cells touching the vessels chiefly on the tangential sides. In A.
panduana Wall, the terminal bands are up to 4 or 5 cells wide, the strands
often up to 6 cells and not storied, and the parenchyma round the vessels
forms a complete sheath (vasicentric). Heimsch (938) notes fusiform cells in
A. chimnsis Bunge. Solereder refers to more abundant parenchyma, often
including chambered crystals, in Billia. Rays exclusively uniseriate; or with
some biseriates in Billia (938), 10-15 P er nun.; usually composed entirely of
upright cells (Kribs's Homogeneous Type III), occasionally, e.g. in A. pun-
duana with single marginal rows of square or upright cells often containing
t ;

small amounts of gummy deposits. Storied in some species. Fibres with pits
more numerous on the radial than on the tangential walls, varying from
simple, e.g. in A. punduana Wall, to distinctly bordered, e.g. in A. glabra
Willd. Walls thin and sometimes with a gelatinous layer. Mean length
O'6-O'9 mm.
TAXONOMIC NOTES
It is generally accepted that the Hippocastanaceae are closely allied to the
Sapindaceae. Pax (1660) pointed out that it is a matter of opinion whether
 HIPPOCASTANACEAE 441

they should be regarded as distinct families or whether the Hippocastanaceae

should be treated as a sub-family of the Sapindaceae, Hutchinson (1113)
treats the two families as one.
Heimsch (938) considers the wood anatomy to show that this family is
closely related to the Sapindaceae. Both he and Tang (2233) treat Bret-
schneidera as a monotypic family closely related to the Sapindaceae, and both
agree that there is no indication of real relationship between this genus and
Moringa.
ECONOMIC USES
The
timbers of a few of the species of Aesculus are of some commercial
importance, e.g. the Ohio Buckeye, A.glabra Willd., and the Horse Chestnut,
A. hippocastanum Linn., and several others are used locally for purposes
requiring a fine-grained, light but fairly tough wood.

GENERA DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Aesculus,* (Billia).
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Hutchinson 1113, Pax 1660, Render 1903.

(ii) On Wood Structure

Bailey 78, Bienfait and Pfeiffer 197, Brown, H. P. and Panshin 288, 289, Chalk and
Rendle 365, Greguss 2522, Heimsch 938, Holden 987, Howard 1088, Kanehira 1209,
Kramer 1275, Nicoloff 1593, Record 1781, 1783, 1809, 1818, 1843, 1851, Record and
Hess 1886, Stone 2203, Tang 2230, 2233.

105. MELIANTHACEAE
(FiG. 100 on p. 438)

SUMMARY
(i) GENERAL
A small family of shrubs or tre^s from tropical or sub-tropical Africa. The
most noteworthy anatomical features include the occurrence of styloids
(elongated prismatic crystals) in Bersama and Melianthus and of raphides in
Greyia. Concentric medullary bundles occur in the axis of certain species of
Bersama and Melianthus,

(ii) WOOD
Vessels rather small, perforations simple, intervascular pitting alternate
and very small or scalariforin to transitional (Greyia)> pits to ray cells similar
to the intervascular pitting, members moderately to very short. Parenchyma
paratracheal, scanty, with scattered crystalliferous cells in Bersama\ storied.
Rays 3-9 cells wide, homogeneous or heterogeneous (Greyia), without uni-
seriates. Fibres with simple pits, storied, very to extremely short.
442 MELIANTHACEAE
LEAF
Dorsiventral in species of Bersama and Melianthus. Stellate hairs with 4 or
more rays recorded in Melianthus. Epidermis not mucilaginous. Stomata
confined to the lower surface ; ranunculaceous. Vascular bundles of the veins
known to be accompanied by sclerenchyma only in Bersama abyssinica Fres.
Petiole (Fig. 100 H) of Melianthus major Linn, hollow and exhibiting, in
transverse sections, a circle of vascular bundles. Small accessory bundles,
similar to the medullary bundles of the stems, present on the inside of the
main ring of bundles in the same species. Medullary bundles also recorded
in the petiole of Bersama abyssinica Fres. Styloids lying parallel to the surface
of the leaf, often situated at the boundary between the palisade and spongy
mesophyll, present in Bersama and Melianthus.

Axis
YOUNG STEM (Fig. 100 K)
Cork arising in the middle of the cortex or immediately on the outside of
the phloem; composed of cells with wide lumina and delicate walls in Greyia
and Melianthus^ but with the outer tangential walls sclerosed in Bersama.
Outer part of the primary cortex of Melianthus major Linn, composed of small
thick-walled cells and the inner part of much larger, thin-walled parenchy-
matous Pericycle some what collenchymatous in Melianthus including
cells. ;

strands of fibres, some having wide lumina and delicate transverse walls in
Bersama, or somewhat elongated sclerenchymatous elements intermediate
between parenchyma and prosenchyma in Greyia. Secondary phloem con-
sisting wholly of soft tissue. Xylem in Melianthus major (Fig. 100 K) con-
sisting of a circle of widely spaced bundles when very young, but soon forming
a continuous cylinder; vessels in the same species exhibiting well-developed
spiral thickening; perforations simple. Concentric medullary bundles,
each consisting of a central strand of soft phloem surrounded by a ring of
simple-pitted, occasionally septate, wood fibres, the strand rarely including
isolated vessels, in Bersama abyssinica Fres. and Melianthus major. Medullary
bundles originate at the nodes as branches from the normal bundle ring
extending into the pith. Styloids recorded from the primary cortex, phloem
and pith of Bersama and Melianthus raphides and/or clustered crystals occur
;

in the corresponding tissues in Greyia.

WOOD (Fig. 98 E-F)

Vessels moderately small (50-100 mean tangential diameter) in Bersama,
//,

very small in Melianthus (2158); solitary and in multiples of 2 or 3 cells; about

30 per sq. mm. spiral thickening reported by Solereder and by Heimsch
;

(938) in Melianthus comosus Vahl. Perforations simple. Intervascular pitting

alternate, very small and commonly with striations due to coalescent apertures
in Bersama, rather larger in Melianthus (938), scalariform and transitional in
Greyia; pits to ray and wood parenchyma similar to the intervascular pitting.
Tyloses present in Greyia (938). Mean member length 0-2-1-3 mm
Paren- *

chyma paratracheal, rather sparse; sometimes with uniseriate vasicentric

sheaths in Bersama and also with some scattered fusiform cells with a
crystalline substance. Dadswell and Record (533) note some diffuse paren-
chyma also in Greyia. Strands of the vasicentric parenchyma usually of
 MELIANTHACEAE 443
2 cells but sometimes with some fusiform cells. Storied. Rays 3-5 cells wide
in Bersama, to 7-9 cells in Greyia; more than i mm., and several stories
up
high; with few or no uniseriates; about 3-4 per mm. homogeneous (Kribs's
;

Type II) in Bersama, but with some sheath cells in B. paullinioides (Planch.)
Bak, heterogeneous and composed entirely of square and upright cells in
;

Greyia and sometimes containing raphides (533). Fibres with numerous

simple pits on all walls; Heimsch (938) records a few septate fibres in 2 species
of Bersama; walls moderately thick to thick; storied; mean length about
0-7 mm. (o'3 mm. in a small shoot of Greyia).

TAXONOMIC NOTES
The genera comprising the Melianthaceae were included under the
Sapindaceae in the Bentham and Hooker system. They were treated as a
distinct family by Giirke (851), who regarded them as having affinities with
the Sapindaceae. The same author draws attention to the rather notable
difference between Greyia on the one hand and Bersama and Melianthus on
the other. In this connexion it is interesting to note that Hutchinson (1113)
has raised Greyia to the status of a new family the Greyiaceae in the Cuno-
niales whilst retaining Bersama and Melianthus in the Melianthaceae under
the Sapindales. The secondary xylem of Greyia differs in some respects from
that of Bersama particularly in having scalariform intervascular and vessel-
parenchyma pitting, tyloses, heterogeneous rays, and occasional raphides.
Heimsch (938) considers the differences sufficient to justify the segregation
of the genus as a separate family, which he places next to the Melianthaceae.
He finds no support for including the family in the Cunoniales.
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Bersama, Greyia, Melianthus.*
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Bersama, Greyia, (Melianthus).

LITERATURE
(i) On General Anatomy
Giirke 851, Hutchinson 1113.

(ii) On Wood Structure

Dadswell and Record 533, Heimsch 938, Hess 959, Record 1843, 1851.

106. STAPHYLEACEAE
(Fie. ioo on p. 438; FIG. 101 on p. 444)

SUMMARY
(i) GENERAL
The known anatomical facts concerning the trees and shrubs belonging to
this family may best be considered under the organs in which they occur. The
family occurs in the Far East, Malaya, and in North and South America, &c.
 H
FIG, 101. STAPHYLEACEAE, A-E; SABIACEAE, F-H
A, Euscaphis japonica Pax, B, Turptnia carnosa Spruce. C, T.pami/era DC. D, Staphylea bumalda
DC. E, S, hohcarpa Hemsl. F, Seina paniculate Edgew. G, Meliosma panamensis Standley. H, M,
panamensis Standley,
 S TAPH YLEA CEAE 445

(ii) WOOD
Vessels small, solitary, numerous, sometimes with spiral thickening, with
oblique scalariform perforation plates, pits to parenchyma simple and round
to oblong, sometimes with spiral thickening; members moderately to
extremely long. Parenchyma typically paratracheal, limited to a few cells
on the abaxial side of the vessels; often with a few scattered cells in addition.
Rays mostly 4-7 cells wide and usually more than i mm. high, heterogeneous.
Fibres with distinctly bordered pits, very rarely septate, moderately to
extremely long. Tracheids often present.

LEAF
Dorsiventral. Glandular hairs absent. Glandular leaf teeth, composed of
parenchymatous with mucilage, situated at the ends of the vas-
cells filled
cular bundles and provided with stomata on the upper surface, in Staphylea
pinnata Linn. Epidermis sub-papillose on the lower surface in Tapiscia;
upper surface mucilaginous in Tapiscia and Turpinia. Stomata confined to
the lower surface; surrounded by 3 subsidiary cells (cruciferous type) in
Euscaphis, Staphylea, Turpinia. Veins accompanied by abundant fibres
in Staphylea and Turpinia] fibres absent from the corresponding position in
Euscaphis. Petiole (Fig. 100 G) containing a circular vascular system, com-
posed of a large arc of xylem and phloem towards the abaxial and 3 smaller
bundles towards the adaxial side in the only z species of Staphylea examined.
Clustered crystals present in Euscaphis, Staphylea, Turpinia,

Axis
YOUNG STEM (Fig. 100 j)
Cork itself in Euscaphis and Staphylea or immedi-
arising in the epidermis
ately below in Turpinia. Pericycle containing isolated strands of fibres in
all examined species of Euscaphis, Huertea,
Staphylea, Tapiscia, Turpinia.
Secondary phloem provided with strands of or isolated fibres in some species
of Euscaphis, Huertea, and Turpinia, and groups of stone cells in Staphylea
and Turpinia. Xylem in the form of a continuous cylinder traversed by
narrow rays in the 2 species of Staphylea examined, but individual bundles
somewhat distinct immediately behind the growing-point in S. cokhica Stev.
Vessels with scalariform perforation plates. Pith large, consisting of thin-
walled parenchyma in Staphylea becoming septate owing to the disorganiza-
;

tion of cells with mucilaginous membranes in Tapiscia. Secretory cells

with amorphous (probably tanniniferous) contents in the outer part of the
cortex in Staphylea. Only clustered crystals recorded in Euscaphis, Staphylea,
and Turpinia, but solitary ones stated to occur as well in Huertea.

WOOD (Fig. 1 01 A-E)

Vessels moderately small (50-100 mean tangential diameter), occasionally
//,

very small (about 40 /x), largest in Turpinia; typically exclusively solitary, or

nearly so, but with radial multiples of 4 or more cells common in some species
of Turpinia (Fig. 101 B); varying in number from about 20 per sq. mm. in
446 STAPHYLEACEAE
woods with the about 100 per sq mm. in woods with the
largest vessels to
smallest vessels. Sometimes with a tendency to ring-porousness in Staphylea;
spiral thickening sometimes present in Euscaphis and Staphylea (938). Per-
foration plates scalariform and oblique, usually with 20-30 fine bars but with
up to 50 in som$ species of Turpinia. Intervascular pitting rare owing to lack
of paired vessels, opposite or transitional between scalariform and opposite;
Heimsch (938) notes some alternate pitting in Staphylea', pits to ray and wood
parenchyma commonly simple, varying from mostly round in Euscaphis to
nearly allhorizontally elongated in Turpinia. Contents not observed. Mean
member length 1-0-2 -2 mm. Parenchyma rather sparse, typically as a few
cells along the abaxial sides of the vessels, though Holden (987) refers to it as
limited to the radial walls in Staphylea; rather more abundant and very
occasionally forming a complete sheath in Euscaphis and Turpinia pomifera DC. ,

and with some scattered cells (diffuse) in Euscaphis and Turpinia. Strands of
4-8 cells. Rays mostly 4 cells wide, up to 10 cells in Euscaphis and some

species of Staphylea (987) usually about i mm. high, but often up to 2 mm.
;

or more in Turpinia uniseriate rays numerous and composed of high upright

;

cells, the rays themselves high in Turpinia', 5-15 rays per mm., least numerous
in Euscaphis and most numerous in Turpinia heterogeneous (Kribs's
;

Types I-II A), with 4-10 rows of marginal upright cells except in Euscaphis
(3 rows), and more than 10 rows in Turpinia; with sheath cells except in
Staphylea. Crystals not observed. Perforation plates between vessels and ray
cells moderately common in Turpinia, often reticulate. Fibres with numerous

distinctly bordered pits on all walls, the borders of about the same size as those
of the intervascular pit-pairs; Heimsch (938) records septa in Huertea cubensis
Griseb. and Tapisda sinensis Oliv. Walls moderately thin to very thick.
Mean length 1-5-3 -4 nun., longest in Turpinia. Vasicentric and vascular
tracheids. Heimsch (938) refers to the occurrence, in several species, of
imperforate tracheary elements that are either tracheids or fibre-tracheids;
those of Euscaphis with spiral thickening.

TAXONOMIC NOTES
The genera included in the Staphyleaceae were treated under Sapindaceae
in the Bentham and Hooker system. They are generally regarded as related
to the Sapindaceae. The wood anatomy of the Staphyleaceae is very much
less highly specialized than that of the Sapindaceae there are many differences
;

between the woods of these two families, but most of these could conceivably
be accounted for by the differences in the level of specialization, Heimsch
(938), however, considers the wood anatomy to be consistent with the placing
of the Staphyleaceae in the Geraniales.
Commenting on Hutchinson's sub-phylum Tinnatae*, Heimsch points out
that the Staphyleaceae and Sabiaceae 'stand out among these families with
prevailingly pinnately compound leaves in that they show the most primitive
level ofxylem organization', and concludes that in spite of the pinnate-leaved
character, the family is not allied with this group.

ECONOMIC USES
Some members of the family are cultivated as ornamental shrubs.
 STAPHYLEACEAE 447
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Euscaphis, Staphylea,* Tapiscia, Turpinia.
* Kew slide
Represented in the collection.

(ii) FOR WOOD STRUCTURE

Euscaphis, (Huertea), Staphylea, (Tapiscia), Turpinia.

LITERATURE
(i) On General Anatomy
Pax 1659.

(ii) On Wood Structure

den Berger 179, 182, Chalk and Chattaway 358, Greguss 2522, Heimsch 938, Holden
987, Janssonius 1x54, Kanehira 1206, Record 1783, 1843, 1851, Record and Hess 1886,
Tang 2231, Williams 2430, Yamabayashi 2478.

107. DIDIEREACEAE
SUMMARY
Plants with a cactus-like habit which occur in Madagascar, The following
notes concerning the structure of the leaf and stem of Didierea have been
recorded by Solereder, whilst the description of the wood is based entirely on
that of Heimsch (938).

LEAF
Mesophyll homogeneous or with a slight tendency for palisade tissue to be
developed in certain species. Stomata infrequent, slightly depressed. Vas-
cular system, in transverse sections, appearing as an almost closed arc of
7-9 bundles. Clustered crystals sometimes fill the cells of the epidermis.
Secretory elements. Cells containing tannin and mucilage present in the
mesophyll.
Axis
STEM
Cortex mainly composed of parenchymatous cells with brown, tannini-
ferous contents. Inner part of the cortex including a layer of stone cells*
Phloem characterized by strands of fibres which appear circular in transverse
sections. Xylem traversed by broad medullary rays; composed mainly of
fibres with simple pits, but including vessels with simple perforations. Large
cluster crystals occur in the outer part of the cortex. Secretory elements*
Very large mucilage cavities present in the cortex.
WOOD*
Vessels solitary and in irregular clusters and groups of multiples. Perfora-
tions simple. Intervascular pitting transitional, opposite and alternate; pits
to ray cells similar. Parenchyma paratracheal, scanty, and terminal. Rays
up to 4 cells wide; more than i mm. high; uniseriates absent; heterogeneous.
Fibres with simple pits.
1
Based entirely on the description given by Heimsch (938),
448 DIDIEREACEAE
TAXONOMIC NOTES
The in the Sapindaceae in the Bentham and Hooker
family was included
system, but doubts have frequently been expressed concerning its position
there. Both Diels and Hutchinson treat the group as a separate family.
Heimsch (938), from a study of the wood anatomy of Didierea and of a twig
of Alluaudia^ concludes that the family does not belong with the Sapindaceae.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Didierea.

(ii) FOR WOOD STRUCTURE

(Alluaudia), (Didierea).

LITERATURE
On Wood Structure
Heimsch 938.

J 108. SABIACEAE
(FiG. ioi on p. 444; FIG. 1 02 on p. 450)

SUMMARY
(i)
GENERAL
A
mainly tropical and sub -tropical family of woody plants, mostly included
in the genera Sabia and Meliosma, Phoxanthus and Ophiocaryon each consist of
only i species. Meliosma is more arboreal in habit than Sabia, and the leaves
aremuch more hairy than those of the last of these genera. Le Renard (1362),
who examined the stem and leaf structure of 14 species of Sabia and of 39 of
Meliosma, found the structure to be fairly uniform in all important respects.
In the mesophyll the palisade tissue of Meliosma differs somewhat from that
of Sabia (see 'Leaf), but Sabia campanulata Wall, was found to resemble
Meliosma in this respect. Transverse sections through the distal end of the
petiole in both genera show a cylindrical vascular strand slightly interrupted
by ray tissue but whereas the strand in Meliosma is surrounded by an almost
;

continuous ring of sclerenchyma, in Sabia there is a more interrupted scleren-

chymatous ring in the corresponding position. Cluster crystals are generally
abundant in the inner cortical tissues of the petiole. The xylem of the young
stem in both genera is traversed by fairly broad medullary rays in Sabia and
by narrower ones in Meliosma.

(ii)
WOOD
Vessels mostly medium-sized, exclusively solitary or with some groups,
perforation plates simple or simple and scalariform, and sometimes reticulate ;

intervascular pitting alternate, pits to parenchyma simple and often elongated ;

members moderately to very long. Parenchyma paratracheal, very to rather
scanty, occasionally vasicentric. Rays usually up to 3-9 cells wide, sometimes
up to 1 5-20 cells high, usually heterogeneous. Fibres with simple or bordered
pits, sometimes septate, of medium length to moderately long.
 SABIACEAE 449
LEAF
Dorsiventral. Rare, unicellular hairs recorded in Sabia ; simple unicellular,
uniseriate and glandular types with relatively long uniseriate stalks and
heads of i or more cells occur in Meliosma. Stomata confined to the lower
surface; ranunculaceous in some species, but rubiaceous in others. Meso-
phyll in Meliosma composed of arm-palisade cells, said by Solereder to
resemble coral when folded, together with somewhat variable spongy tissue,
often containing stellate cells. Arm-palisade cells not recorded in Sabia apart
from S. campanulata Wall., which resembles Meliosma in respect of this
character. Vascular bundles of the veins usually surrounded by sclerenchyma
and accompanied externally by cluster crystals. Petiole of Meliosma beaniana
Rehd. et Wils. (Fig. 102 L) exhibiting, in transverse sections through the
distal end, a dorsally flattened cylindrical vascular strand, accompanied by
2 small, latero-superior bundles. Main vascular strand almost surrounded by
a ring of fibres. Cluster crystals fairly abundant in the cortical tissues of the
petiole, especially in a position immediately external to the main vascular
strand. Petiole of M. cuneifolia Franch. (Fig. 102 N) similar in structure to
that of the previous species, but crystals less numerous. The above petiolar
structure, which was observed in material grown at Kew, appears to be
common to most species of Meliosma and Sabia judging from the information
recorded by Le Renard (1362). Crystals, see 'Petiole* and 'Veins'.

Axis
YOUNG STEM (Fig. 102 M)
Cork always arising superficially. Pericycle with a composite and con-
tinuous ring of sclerenchyma in Sabia when sufficiently mature; isolated
strands of sclerenchyma generally occur in the corresponding position in
Meliosma. Pericyclic sclerenchyma forming an almost continuous ring in very
young stems of M. cuneifolia Franch., but becoming more interrupted in
slightly older material. Xylem traversed by fairly broad rays in Sabia and
by rather narrower ones in Meliosma. Portions of the rays sclerosed where
traversing the phloem in Sabia] distal portions between the phloem groups
tending to be triangular in Meliosma. Vessel perforations mostly simple, but
a proportion of scalariform plates with numerous bars also observed. Outer
part of the pith soon becoming sclerosed in Meliosma and Sabia, or wholly
lignified in Meliosma. Cluster crystals present in the cortex, phloem, and
pith of Meliosma beaniana Rehd. et Wils. and M. cuneifolia, and probably in
other species as well. Secretory cells with amorphous contents occur in the
parenchymatous tissues in the same 2 species of Meliosma, and probably in
others as well. Le Renard (1362) records them in the pith and ray cells of
most species of Meliosma. Secretory cells, arranged in longitudinal columns,
occur in the pith of, M. beaniana.

WOOD (Fig. 101 F-H)

Vessels medium-sized (mean tangential diameter 100-200 ft) in most
species; exclusively solitary in Sabia, solitary and in radial multiples of 2 or 3
cells in Meliosma and Ophiocaryon, and commonly in apparent tangential

pairs owing to the overlapping ends of members; sometimes in irregular

4594 G g
 N
FIG. 102. ANACARDIACEAE, A-F and H; CORYNOCARPACEAE, G and J;
CORIARJACEAE, I and K; SABIACEAE, L-N
A, Rhus trichocarpa Miq. Petiole X 15. B, Cotinus coggygria Scop. Petiole X 19. C, Schinus
terebinthifolius Raddi. Petiole x 16. D, Schinopsts lorentzii (Griseb.) Engl, Petiole x 32. E, Rhus
trichocarpa Miq. Stern xn. F, Pistacia chinensis Bunge. Petiole X 19. G Corynocarpus laevigata
Forst. Petiole x 12. H, Mangifera indica Linn. Petiole x 14. I, Coriaria sinica Maxim. Petiole X 12.
J, Corynocarpus laevigata Forst. Stem x8. K, Coriaria sinica Maxim. Stem X8. L, MeKosma
beaniana Rehder et Wils. Petiole x n. Wing bundles not shown. M, M. cuneifolia French. Stem
X 16, N, M. cuneifolia Franch. Petiole x 13.
f.c. Secretory canals.
 SABIACEAE 451

clusters; 2-10 per mm.; with spiral thickening in i specimen of M. rhoifolia

Max. Perforation plates simple in Sabia p.p., simple and scalariform in
Meliosma and Ophiocaryon and, according to Solereder, in Sabia p.p. multi- ;

perforate plates usually scalariform with few bars, but with up to 30 bars in
some species, occasionally reticulate ;
sometimes distinctly foraminate in M.
panamensis Standl. Intervascular pitting alternate, usually moderately large,
but small in a few species; pits to parenchyma simple; large and elongated in
some species. With
occasional tyloses in Meliosma (1154) and Ophiocaryon.
Mean member length 0-8-1-2 mm. Parenchyma paratracheal, scanty in
Meliosma and very sparse or absent from Sabia] in narrow vasicentric sheaths
and terminal bands 3-5 cells wide in Ophiocaryon; 1 adjacent vessels in
Meliosma occasionally linked by a uniseriate band (1154). Strands usually of
8 cells. Rays usually of 2 sizes, sometimes muitiseriate only; the larger rays
up to 3 or 4 cells wide in Ophiocaryon, 4-15 cells in Meliosma, and up to
20 Sabia; usually more than 2 mm. high; shorter in M. macrophylla
cells in
Merrill, and Ophiocaryon paradoxum R. Sch., rarely more than 3 cells wide
and with a gradation of sizes uniseriates numerous, except in Meliosma, and
;

composed of square and upright cells; 3-6 rays per mm. in Meliosma, more
numerous (9-14) in Ophiocaryon and Sabia heterogeneous (Kribs's Types II A
;

and II B) in Meliosma and Ophiocaryon, the cells of the large rays very variable
in size and shape in different species and commonly nearly all square and
upright; almost homogeneous and with distinctly procumbent cells in M.
rhoifolia Max. and Sabia paniculata Edgw. with sheath cells in most species
; ;

usually with a few cells containing dark deposits, crystals rare. Fibres with
simple or very small bordered pits, which are more numerous on the radial
than on the tangential walls, in Meliosma and Ophiocaryon, with distinctly
bordered pits, which are very numerous on both radial and tangential walls
and often biseriate, in Sabia; Heimsch (938) describes the fibrous elements in
Ophiocaryon as tracheids or fibre-tracheids some septate fibres present round
;

the vessels in Meliosma and, in some species, on the boundaries of the growth
rings ;
Heimsch notes some septate fibres in Sabia. Walls thin to moderately
thick. Mean length i -2-1 -9 mm. In Meliosma there are sometimes moderately
numerous tracheid-like fibres with wide lumina and lengths equivalent to
those of the vessel members.

TAXONOMIC NOTES
(i)
FROM GENERAL ANATOMY
The Sabiaceae were first named the Meliosmaceae by Endlicher, and
placed by him between the Hippocastanaceae and Sapindaceae. They were
subsequently treated as a tribe of the Sapindaceae, but the status of a family
was restored in the Bentham and Hooker system.

(ii) FROM WOOD STRUCTURE

Heimsch (938) states that in their wood structure the Sabiaceae resemble
certain of the Anacardiaceae and that, if the family really belongs to the
Terebinthales complex, it would seem to be closest to that family.
1
Williams 's description (2430) of Ophiocaryon heterophyllum Urb. differs in many respects
from material examined of O. paradoxum Rb, Schomb., particularly as regards the parenchyma,
which he records as in fine reticulate lines.
 452 SABIACEAE
Meliosma differs from Sabia in several respects. There appears to be some
confusion over Ophiocaryon> descriptions of different species showing marked
divergences.
ECONOMIC USES
Some species of Meliosma are cultivated as ornamental trees.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Meliosma,* Sabia.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Meliosma, (Ophiocaryon), Sabia.

LITERATURE
(i) On General Anatomy
Dihm 589, Le Renard 1362, Warburg 2359.
(ii) On Wood Structure

den Berger 179, 182, Dadswell and Record 533, Heimsch 938, Janssonius 1154, Kane-
hira 1206, 1209,Lecomte 1334, Record 1843, 1851, Record and Hess 1886, Tang 2231,
Tapper 2295, Yamabayashi 2478.

--/109. ANACARDIACEAE
(Fic. 1 02 on p. 450; FIG. 103 on p. 456)

SUMMARY
(i) GENERAL
A
mainly tropical family of trees and shrubs, often resinous. The resinous
substances secreted by some, but not all, of the Anacardiaceae are highly toxic
and may cause severe dermatitis. One of the most notorious poisonous
species is Rhus toxicodendron Linn., the Poison Ivy. The hairs are unicellular,
uniseriate, and glandular, the latter being very varied in shape. The stomata
need more complete examination, but in the few recorded instances they are
stated to be ranunculaceous. The outstanding anatomical feature of the axis
is the occurrence of resin-canals. These are universally present in the
primary phloem, immediately on the inside of the arcs of fibres in the peri-
cycle (see below). They are also arranged in concentric rings in the secondary
phloem of older stems. Medullary resin canals have been recorded in a con-
siderable number of genera, but similar canals in the primary cortex are much
less frequent. The cork originates superficially. The primary cortex,
besides the resin canals mentioned above, often includes stone cells. The
pericycle usually contains isolated, arc-shaped strands of fibres with the
convex side of each group directed towards the exterior, whilst a composite
and continuous ring of sclerenchyma is much less frequent. Elongated
tannin-sacs of varying abundance are common in the phloem throughout
the family. Solitary and clustered crystals have been recorded.
 ANACARDIACEAE 453

(ii) WOOD
Vessels never very small and often moderately large, occasionally with dis-
tinct oblique orulmiform pattern; perforation plates exclusively simple, except
in 2 genera; intervascular pitting alternate, typically large, though small to
minute in a few genera, with large, simple, elliptical pits to parenchyma;
occasionally ring-porous and with spiral thickening. Members of medium
length to moderately short. Parenchyma paratracheal scanty, vasicentric
;

or aliform and often rather sparse; with terminal or irregularly distributed

broad bands in some genera. Rays mostly 2-3 cells wide (excluding fusiform
rays containing canals) but larger in some species (up to 8-10 cells) exclusively
;

uniseriate in a few genera; usually heterogeneous and often with few uni-
seriates; sometimes in echelon. Fibres with simple pits, septate in about
half the genera; commonly with a gelatinous inner layer; of medium length
to very short. Intercellular canals in the rays of about two-thirds of the
genera.
LEAF
Dorsiventral in most species of Rhus of the section Gorontogeae Engl. con- ;

sistingwholly of palisade cells in R. incisa Linn. Mesophyll in a dune form

of Rhus toxicodendron Linn, described by Starr (2188) as having much taller
palisade cells than those in mesophytic specimens of the same species. Meso-
phyll in Anacardium occidentale Linn, consisting of 2 layers of palisade cells
and a broader region of spongy tissue, according to de Boer (214). Elongated
sclerenchymatous cells recorded by Goris (798) in the mesophyll of Bouea.
For particulars of the mesophyll in sumach leaves see 'Economic Uses'.
Hairs unicellular, uniseriate, or glandular; the last of these very variable in
type, recorded in Anacardium occidentale Linn., Buchanania, Dobinea, Phle-
bochiton, and Rhus. Peltate glands recorded in Campnosperma. The following
types of glandular hairs, believed possibly to be of specific diagnostic value, but
requiring further investigation, recorded in Rhus: uniseriate with a spherical
head; uni- or bicellular club-shaped glands; rosette-glands resembling
stellate hairs peltate glands, sometimes secreting sufficient resin to coat the
;

leaves with a thin layer of varnish. A covering of deciduous hairs recorded

by McNair (1474) on young leaves of Rhus diversiloba T. et G. Lower
epidermis papillose in certain species of Rhus and Swintonia. Upper
epidermis including mucilaginous cells in Buchanania. Hypoderm recorded
by Goris (798) in Gluta, Mangifera, and Melanorrhoea; and a pseudo-
hypoderm of divided epidermal cells in Protorhus by Courchet (484). Stomata
ranunculaceous in Anacardium occidentale Linn., Mangifera indica Linn.,
Rhus toxicodendron Linn., and Spondias lutea Linn. present on both surfaces
;

or confined to the lower side in species of Rhus of the section Gorontogeae;

deeply sunk in R. burkeana Sond. Midrib of Rhus toxicodendron described
by Holm (1042) as including a circular band of several collateral bundles
surrounded by a sclerenchymatous pericycle. Structure of midrib similar in
Rhus glabra Linn. Vascular bundles of the veins provided with resin canals
in the phloem. Resin canals also occur on the upper side of the xylem in
species with medullary bundles in the axis. Vascular bundles of the larger
veins vertically transcurrent by fibres in Protorhus according to Courchet
(484). Smaller veins in the same genus embedded in the mesophyll and not
454 ANACARDIACEAE
accompanied by canals. Petiole exhibiting, in transverse sections through
the distal end, an abaxial continuous arc accompanied by 1-3 adaxial groups
of vascular tissue, thus giving the whole strand the appearance of a dorsally
flattened but slightly interrupted ring in Mangifera indica, Pistacia chinensis
Bge. (Fig. 1 02 F),A terebinthus Linn., Schinus terebinthifolius Raddi. (Fig.
IO2C); similar but with the ventral arc dissected into several bundles in
Cotinus americanus Nutt. and C. coggygria Scop. (Fig. 102 B); with a con-
tinuous cylindrical strand having a somewhat sinuous outline in Schinopsis
lorentzii (Griseb.) Engl. (Fig. IO2D); with a circle of numerous vascular
bundles in Rhus diversiloba T. et G. (McNair 1474), R. toxicodendron (circle
dorsally flattened), R. trichocarpa Miq. (Fig. 102 A), and R. typhina Linn.
Petiolar vascular system with large resin canals in the phloem in all of the
above species, and presumably throughout the family. Clustered crystals
stated to appear as transparent dots in Rhus succedanea Linn., Spondias dulcis
Forst, and the genus Tapiria. Solitary crystals accompany the veins in
Mangifera indica, and clustered ones occur in the corresponding position in
Spondias lutea. Occasional crystalliferous cells at right angles to the leaf
surface recorded by Courchet (484) in Protorhus. Large rhombic and cluster
crystals recorded by Holm (1049) in the mesophyll of Rhus glabra Linn, and
cluster crystals in the palisade layer of Protorhus by Dubard and Dop (611).

Axis
YOUNG STEM (Fig. 102 E)
Cork nearly always arising in the sub-epidermis, but rather more deeply
seated in the cortex in Campnosperma, Protorhus, and Spondias. Cork cells
with walls sclerosed on one side in Mangifera and Rhus\ uniformly sclerotic
on all sides in Schinus thin- walled and spongy in Odina and Pistacia tabular,
; ;

with thick walls in Astronium spongy but including scattered sclerosed cells
;

in Anacardium. Phelloderm extensive in Astronium and Pistacia] consisting

of alternate zones of stone cells and thin-walled cells in the first of these
genera. Primary cortex containing thick-walled sclerosed cells with wide
lumina in species of Campnosperma, Melanorrhoea, Pleiogynium, Rhus ovata
Wats. (Watkins 2367), Sorindeia, and a ring of often somewhat elongated
stone cells in species of Astronium, Campnosperma, Drimycarpus, Haemato-
staphis, Holigarna, Loxostylis, Mangifera, Pseudospondias, Swintonia, Thyr-
sodium. Pericycle in most species with isolated, arc-shaped strands of fibres,
having the convex side towards the exterior; usually with a single, large resin-
canal on the inside of each group. A composite and continuous ring of
sclerenchyma present in species of Campnosperma, Dracontomelum, Drimy-
carpus, Euroschinus, Gtuta, Lithraea, Mauria, Metopium, Microstemon, Penta-
spadon, Phlebochiton, Pistacia, Schinopsis and a tendency towards a composite
;

and continuous ring in Pseudospondias and Swintonia. Phloem containing

elongated tannin-sacs, varying in abundance in different species; chambered
fibres containing solitary crystals occasionally present in the phloem in

Mangifera, Odina, and Schinus. Secondary phloem stated to be devoid of

fibres or stone cells in Rhus] containing stone cells but no fibres in Pistacia]
with fibres and stone cells in Anacardium, Mangifera, Odina, and Protorhus]
with fibres alone in Schinus. Xylem generally in the form of a continuous
cylinder, traversed by narrow rays, but occasionally somewhat interrupted by
 ANACARDIACEAE 455
the primary rays; vessel perforations always simple except in a few genera
(see 'Wood'). Mucilage cavities recorded in the cortex of Campnosperma.
Resin-canals not only occurring immediately on the inside of each group
of pericyclic fibres as mentioned above, but also present in concentric
circles in the secondary phloem; stated to anastomose to form a network in

species of Rhus, Schinus, and Spondias, and to be continuous with the canals
in the rays in Rhus viminalis Vahl. Medullary resin-canals, varying in number
in different members of a single genus or even in different internodes of a
single plant, recorded in Anacardium, Anaphrenium (pro parte), Astronium,
Buchanania, Campnosperma, Comocladia (pro parte), Cyrtocarpa, Draconto-
melum, Drimycarpus^ Euroschinus, Faguetia, Gluta, Haematostaphis, Har-
pephyllum, Holigarna, Loxopterygium Mangifera, Mauria, Melanochyla,
y

Metopium, Microstemon Odina, Pentaspadon, Phlebochiton, Pleiogynium, Pou-

y

partia, Pseudosmodingium,Pseudospondias, Rhus (in tropical species), Schinopsis,

Schinus (pro parte), Sclerocarya, Semecarpus, Solenocarpus Sorindeia, Spondias,
y

Swintonia, Tapirira, Thyr$odium Trichoscypha. Cortical resin-canals much

y

less frequent, but recorded in species of Anacardium, Dobinea, Holigarna, and

Thy rsodium. Secretory cells with tanniniferous contents frequently abun-

dant in the cortex, phloem, medullary rays, and pith. Solitary crystals (see
also under 'Phloem') usually abundant, sometimes with only one type present
in an individual species, but a mixture of kinds occurring in others.

ROOT
Each group of primary phloem containing a single resin canal; smaller
canals present in the secondary phloem. Xylem of Rhus diversiloba T. et G.
described by McNair (1474) as less firm than that of the young stem owing to
the fibres being thicker and broader. Adventitious roots have been described
by Buscalioni and Muscatello (320) in Rhus viminalis Ait. According to Holm
(1042) aerial attachment roots and subterranean nutritive roots of Rhus
toxicodendron Linn, are similar in structure, the aerial roots remaining active
for onlyone season. Other particulars concerning root structure recorded by
Weber (2380).

WOOD (Fig. 103)

Vessels mostly moderately small to medium-sized (50-200 p mean tan-
gential diameter), large (more than 200 /z) in some species of Anacardium,
Dracontomelum Gluta Mangifera, Melanochyla, Melanorrhoea Semecarpus,
y y y

and SpondiaSy smallest in Campnosperma^ Nothopegia and Schinus: with an

y

oblique pattern in Cotinus (1859) anc^ Pistada (Fig. 103 L), and also, though
less marked, in some species of Rhus ulmiform in Schinus (Fig. 103 P);
y

commonly with most of the vessels solitary but with a few radial and irregular
multiples of several small cells, radial multiples more common in some species
of Astronium Faguetia Nothopegia, Protorhus, Rhus, and Schinus y varying in
y y

number from 2 to 25 per mm., fewer than 5 per mm. in most species of
Anacardium, Antrocaryon, Bouea, Buchanania, Dracontomelon, Gluta Mangi- y

fera, Melanockyla,and Melanorrhoea, more than 20 per mm. in Lithraea,

Nothopegia, and Schinus ring-porous in some or all species of Cotinus (1851),
;

Pistacia, Protorhus, Rhus, Schinus, and 'Toxicodendron (Rhus) (1859); w

9
^
spiral thickening in Cotinus (1851), Lithraea, Microstemon (938), Pentaspadon
 FIG. 103. ANACARDIACEAE
A, Mangifera indica Linn. B, 'Campnosperma wallichii King*. C, Mangifera indiea Linn. D, Parishia
pubescent Hook. f. E, Gluta travancorica Bedd. F, Rhodosphaera rhodanmema Engl. G, Melanorrhoea
torquata King. H, Microstemon velutina Engl. I, Lannea acidissima A. Chev. J, Astronium balansae
Engl. K, Ltthraea nwlleoides (Veil.) Engl. L, Pistacia terebinthus Linn. M, Schinus praecox Speg.
N^ Rhu$ integrifolia Benth. et Hook. f. O, Odina wodier Roxb. P, Schinus latifolius Engl.
i.e. Intercellular c^naU
 ANACARDIACEAE 457

(938), Pistacia, afew species of Rhus, e.g. R. verniciflua Stokes, Schinus,

1

'

Schmaltzia (938), and Toxicodendron* (Rhus) (1859). Perforations exclusively

simple in most genera, but with a few scalariform plates in some species of
Anaphrenium (2158), Campnosperma, Heeria (938), and Micronychia (2158),
with fewer than 20 bars except in Anaphrenium. Heimsch (938) notes some
modified forms of scalariform plates in species of Comocladia, Euroschinus,
and Lithraea. Intervascular pitting alternate and typically large, rather
smaller in Buchanania and Loxostylis (2158), small in Lithraea, Protorhus,
Rhus, and Schinopsis, and very small in Faguetia and Trichoscypha\ sometimes
with striations due to coalescent apertures, e.g. in some species of Loxoptery-
gium, Melanochyla, Pentaspadon, and Rkus\ Heimsch (938) records occasional
transitional types in Campnosperma, Comocladia, Euroschinus, and Veatchia,
and some opposite pitting in Dobinea with at least some large elliptical, simple
;

pits where adjoining parenchyma in all the material examined, such pits
usually numerous and often with the long axes horizontal and producing a
scalariform appearance. Tyloses observed in most of the genera often with ;

dark contents. Heimsch (938) refers to sclerotic ty loses in one specimen of

Mangifera altissima Blanco, and Janssonius (1154) notes that the tyloses may
contain crystals. Mean member length 0-2-0-8 mm., mostly 0-4-0-5 rnm.;
longest in Dracontomelon. Parenchyma predominantly paratracheal, scanty
or vasicentric, with terminal bands in some genera often sparse in Antrocaryon,
;

Astronium, Blepharocarya (525), Buchanania, Comocladia, Dobinea (938),

Dracontomelum, Euroschinus, Faguetia, Gluta p.p., Koordersiodendron, Lannea,
Lithraea, Loxopterygium, Melanorrhoea, Microstemon, Mosquitoxylum, Odina,
Parishia, Pentaspadon, Pistacia, Pleiogynium, Poupartia, Rhus, Schinopsis p.p.,
Schinus, Sorindeia, Spondias, Swintonia griffithii Kurz., and Tapirira\ aliform
or locally confluent in Anacardium, Holigarna, Mangifera, Melanochyla>
Nothopegia, Oncocarpus, Protorhus, Schinopsis p.p., Semecarpus, Swintonia
*

(most species), Thyrsodium (1859), Toxico dendron* (Rhus) (1859), anc^

Trichoscypha\ transitional between banded and aliform in Nothopegia and
some species of Semecarpus and Swintonia and with irregularly distributed
broad bands, sometimes ending blindly or anastomosing, in Bouea macrophylla
Griff., Gluta, Mangifera p.p. (938), Melanorrhoea, and Metopium (938) absent ;

from Campnosperma and Rhodosphaera', in Rhodosphaera the vessels are

enclosed in a sheath of thin-walled septate fibres with wide lumina, and in
Loxopterygium similar vasicentric sheaths of septate fibres occur, but mixed
with parenchyma; terminal bands 1-5 cells wide present in Anacardium p.p.
(1859), Bouea, Faguetia, Gluta, Mangifera, Mauria (1859), Melanorrhoea,
Pistacia formosana Mats. (1206), Rhus p.p., Swintonia, and Trichoscypha.
Heimsch (938) notes sclerotic cells in Mangifera altissima Blanco. Cells
commonly with dark gum-like contents; crystals very rare, except in Lithraea
and Mauria (1859), in which
crystalliferous strands are scattered among the
fibres. Bargagli Petrucci refers to the presence of silica bodies in Melanorrhoea

obtusifolia Engl. Strands typically of 4 cells, occasionally up to 8 cells, e.g. in

Bouea and Dracontomelon. Rays i-io cells wide (excluding fusiform rays
containing canals), mostly 2-3 cells; exclusively uniseriate or only occasionally
biseriate in Comocladia (1859), Gluta, Mangifera p.p., Melanorrhoea, Swin-
1
Record (1859) states that spiral thickening is present at least in the smaller vessels of all
the American species.
458 ANACARDIACEAE
tonia p.p.,and Thyrsodium (1859), up to 4 or 5 cells wide in at least some
species of Antrocaryon, Astronium, Buchanania, Dracontomelum (1154),
Lannea, Lithraea, Metopium, Odina, Pentaspadon, Pistacia, Poupartia, Rhus,
Schinopsis, and Schinus, up to 8-10 cells wide in Rhus p.p. (1206) and
Spondias p.p. sometimes slightly exceeding i mm. in height in some species
;

of Dracontomelum, Odina, and Spondias; uniseriates typically composed of

both square to upright and procumbent cells, often few and usually fewer
than might be expected from the degree of heterogeneity of the multiseriate
rays; mostly between 5 and 10 rays per mm., fewer than 5 per mm. in
Euroschinus, Odina, Poupartia, Rhus p.p., and Spondias, more than 10 per mm.
in Astronium, Bouea, Gluta p.p., Mangifera p.p., Nothopegia, Schinus, and
Sorindeia p.p. heterogeneous (typically Kribs's Type II B), except in Gluta
;

and Melanorrhoea (Kribs's Homogeneous III); rather more markedly hetero-

geneous in some species of Antrocaryon and Campnosperma. Usually with
1-3 marginal rows of upright cells, with 4 or more rows in Antrocaryon,
Holigarna, Microstemon, Rhodosphaera, Rhus p.p., Semecarpus, and Tricho-
scypha in some of these latter genera with markedly heterogeneous rays the
;

biseriate parts, composed of procumbent cells, are often little wider tan-
gentially than the uniseriate upright cells; in Trichoscypha ferruginea Engl.
groups of procumbent cells alternate with upright cells; composed chiefly of
irregular upright cells in Dobinea (938). Heimsch (938) notes that, apart from
the rays containing canals, the rays are nearly all uniseriate and heterogeneous
in a few species of Comocladia, Parishia, Schmaltzia, and Sorindeia. Cells
commonly filled with a dark gum-like substance, and crystals present in many
species and sometimes abundant, either in the procumbent cells or in the
upright cells or, less commonly, in both; crystals in upright cells sometimes
large, filling rounded cells that are slightly wider tangentially than the cells
without crystals, e.g. in Astronium urundeuva EngL, Koordersiodendron pinna-
turn Merril., and Pistacia terebinthus L.; silica present in some species of
Melanorrhoea, Parishia, and Swintonia (794) intercellular spaces pronounced
;

in a few woods, e.g. Campnosperma macrophylla (Bl.) Hook, f., Gluta, and
Stvintonia; rays sometimes arranged in echelon in Antrocaryon, Campno-
sperma, Euroschinm, Parishia, Rhus, Sorindeia, and Swintonia. Fibres with
small simple pits that are scarce on the tangential walls except in Noihopegia\
septate in all or some species of Anacardium, Antrocaryon, Astronium,
Buchanania (938), Campnosperma, Comocladia, Dobinea (938), Dracontomelum,
Harpephyllum (938), Koordersiodendron, Lannea, Lithraea, Loxopterygium,
Mauria (1859), Metopium, Microstemon, Odina, Pleiogynium (938), Pou-
partia (938), Pseudospondias (938), Rhodosphaera, Rhus, Schinopsis, Schinus,
'

Sclerocarya (938), Sorindeia, Spondias, Tapirira, and Toxicodendron* (Rhus)

(1859); in Loxopterygium and Rhodosphaera septate fibres with wider lumina
than those of the ground tissue are associated with the vessels, in the latter
mixed with parenchyma (see also under 'Parenchyma'); in most species of
Buchanania similar vasicentric septate fibres occu'r mixed with parenchyma,
but the ground tissue is of non-septate fibres; according to Janssonius (1154)
occasional septate fibres occur scattered among the non-septate fibres of the
ground tissue in Buchanania florida Schau. and in Mangifera. Thin-walled
in many genera but with moderately thick to thick walls in Astronium, Bouea,
Faguetia, Koordersiodendron, Loxopterygium, Metopium, Nothopegia, Khodo-
 ANACARDIACEAE 459
sphaera, Schinopsis,and Swintonia p.p.; very commonly with a gelatinous
inner layer. Arranged in very distinct radial rows in some species of Bouea>
Dracontomelum, Melanorrhoea, Pentaspadon, and Swintonia and some other
genera, but with this feature often emphasized in the above-named genera
owing to the fibres being narrower radially than tangentially. Janssonius
(1154) notes the rare occurrence of solitary crystals in single specimens of
Buchanania florida Schau. and Spondias dulcis Forst. var. acida Engl. Mean
length 0'6-i-4 (mostly 07-1-0) mm. Vascular tracheids are reported by
Janssonius (1154) in 2 species of Spondias. Intercellular canals occur in
the rays of some or all species of Antrocaryon, Astronium, Buchanania,
Campnosperma, Euroschinus, Gluta, Harpephyllum (938), Koordersiodendron
(938), Lannea, Loxopterygium, Malosma (1859), Melanochyla (938), Mela-
norrhoea, Metopium Microstemon, Odina, Parishia, Pentaspadon,
(1859),
Pistacia, Pleiogynium, Poupartia, Pseudospondias (938), Rhodosphaera, Rhu$ y

Schinopsis, Schinus, Schmaltzia (938), Sclerocarya (938), Smodingium (938),

Spondias, Swintonia, Tapirira, 'Toxtcodendron' (Rfais) (1859), and Tricho-
*

scypha; canals very small, e.g. in Lannea barteri Engl., to large. The seasonal
development of the growth rings in Lannea and Spondias has been investi-
gated by Coster (481). Besson (186) notes a moderately high ash content and
a very high silica content in the wood of Melanorrhoea laccifera Pierre.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
Engler (628) believed the Anacardiaceae to be related to the Sapindaceae,
although he pointed out that the presence of resin canals throughout the
Anacardiaceae serves to distinguish this family from the Sapindaceae, where
resin canals do not occur. The presence of resin canals in both the Anacar-
diaceae and Burseraceae has led to the suggestion that these 2 families are
related, but Engler (643) disagrees with this view because of the constant
floral differences between the 2 groups. It is interesting to note that Copeland
and Doyel (466) wrote 'Juglandaceae are not derived from Anacardiaceae, but
a collateral relationship between these families remains a possibility*.

(ii) FROM WOOD STRUCTURE

Heimsch(938) has described separately the woods of the Mangiferae,
Spondiae, Rhoideae, Semecarpeae, and Dobineae, but has concluded that
there are no characters or combinations of characters that serve to differentiate
one group from the others. There do, however, appear to be certain trends in
these groups. As Heimsch has pointed out, more species of the Mangiferae
have homogeneous rays and bands of parenchyma than any other, the greatest
proportion of ring-porous forms with advanced stages of vessel aggregation
occur in the Rhoideae, and all the species of the Spondiae have septate fibres,
whereas these are lacking from the Semecarpeae.
The most striking characteristics of the family radial intercellular canals,
septate and paratracheal, often scanty, parenchyma are closely
fibres

paralleled in the Burseraceae. Heimsch considers that these 2 families

resemble each other more closely than either family resembles the other
groups covered in his study of Wettstein's Gruinales and Terebinthales.
1
Heimsch (937), on the other hand, has been unable to confirm this.
 460 ANACARDIACEAE
Heimsch considers the Julianiaceae to be closely related to the Anacardiaceae.
He, however, rejects Hallier's view that Juliania is a reduced derivative of
Pistacia on account of the relatively highly specialized wood structure of
Pistacia,
The evidence from wood anatomy is entirely against any relationship
between the Anacardiaceae and the Juglandaceae.
Heimsch (937) has studied the secondary xylem and pollen morphology in
the Rhus complex and his conclusions are in agreement with Barkley's taxo-
nomic study of this complex, in which the species are grouped into the genera
Actinocheita, Metopium, Malosma, Cotinus, Toxicodendron, and Rhus.

ECONOMIC USES
The resinous secretion obtained by tapping the bark of Rhus verniciflua
Stokes, when mixed with pigments, is used in the preparation of Japanese
lacquers. Sumach leaves, obtained from Rhus coriaria L, and certain other
species of Rhus and Cotinus, are used as a source of tannin. The resin known
as Mastic is obtainedfrom Pistacia lentiscus Linn., while leaves of the same
species are sometimes used as a substitute for Sumach. Japan wax, used in
the manufacture of floor polish, &c., is extracted from the seeds of Rhus
verniciflua Stokes. The very toxic nature of the resins secreted by some mem-
bers of the family has already been mentioned in the 'summary' above. Edible
fruits obtained from members of the Anacardiaceae include: the Mango

(Mangifera indica Linn.), Cashew nuts (Anacardium occidentals Linn.), and

Pistachio nuts (Pistacia vera Linn.).
Sumach leaves (Rhus coriaria Linn,) exhibit the following microscopical
characters.
Hairs of two kinds, (a) Simple, thick-walled, pointed, usually unicellular
but occasionally with a septum, often provided with an enlarged base.
(b) Glandular, with a unicellular stalk and multicellular head. Upper epider-
mis composed of polygonal cells with almost straight anticlinal walls, covered
externally with striated cuticle. Cells of the lower epidermis with thinner,
more sinuous anticlinal walls. Some of the epidermal cells radiate from around
the bases of the hairs. Stomata mostly confined to the lower surface ranun-
;

culaceous. Mesophyll dorsiventral; including a single, compact layer of

palisade tissue. Aggregations of cluster crystals, occasionally resembling
cystoliths, present in the lacunar spongy tissue. Midrib with 3 resin canals
in the phloem. Petiole in transverse sections through the distal end, exhibiting
a closed, dorsally flattened vascular strand with resin canals in the phloem.
Trieste Sumach is the product of Cotinus coggygria Scop. Most of the
microscopical characters are very similar to those of Rhus coriaria, but the
pointed thick-walled hairs are apparently absent, although glandular hairs of
the same type are common to both species. In Cotinus coggygria the epidermal
cells on both surfaces bear hemispherical papillae, those on the lower surface

being provided with finger-like processes.

The leaf of Pistacia lentiscus, which has been used to adulterate sumach,
can be recognized by the stomata confined to the lower surface, where they
are frequently surrounded by a circle of radiating epidermal cells and by the
2 or 3 layers of palisade tissue in the mesophyll. Further details have been
recorded by Hanausek (882). A kind of sumach derived from Coriaria myrti-
 ANACARDIACEAE 461

folia Linn, (family Coriariaceae) can be recognized by several anatomical

characters, one of the most obvious being the occurrence of rubiaceous
stomata on both surfaces.
The timbers of this family range from light, colourless, and perishable to
dense, highly coloured, and durable. Among the latter are the two widely
known South American timbers Quebracho, Schinopsis spp., and Zebra Wood
or Kingwood, Astronium spp. Some other genera, such as Gluta and Mela-
norrhoea, produce rather similar dense, attractively streaked timbers, e.g. the
Malayan Rengas, but these do not appear to be of more than local importance.
The sawdust from some species produces dermatitis. Several of the lighter
timbers such as those of Mangifera and Buchanania produce non-durable
general purpose timbers that are used locally and, according to Desch (574),
the timber of Campnospermum was one of the two principle timbers used
(before 1942) in the local match factories in Malaya.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Anacardium, Anaphrenium, Astronium, Botryceras, Buchanania, Campno-
sperma, Comocladia, Cotinus,* Cyrtocarpa, Dobinea, Dracontomelum,
Drimycarpus, Euroschinus, Faguetia, Gluta, Haematostaphis, Harpephyllum,
Holigarna, Lithraea, Loxopterygium, Loxostylis, Mangifera,* Mauria, Melano-
chyla, Melanorrhoea, Metopium, Micronychia, Microstemon, Nothopegia,
Odina, Pentaspadon, Phlebochiton, Pistacia,* Pleiogynium, Poupartia, Proto-
rhus, Pseudosmodingium, Pseudospondias, Rhus,* Schinopsis,* Schinus,*
Sclerocarya, Semecarpus, Smodingium, Solenocarpus, Sorindeia, Spondias,
Swintonia, Tapirira, Thyrsodium, Trichoscypha.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Anacardium (Anaphrenium), Antrocaryon, Astronium, (Blepharocarya),
Bouea, Buchanania, Campnosperma, Comocladia, Dracontomelum, Euro-
schinus, Faguetia, Gluta, Harpephyllum, (Heeria), Holigarna, Koordersio-
dendron, Lannea, Lithraea, Loxopterygium, (Loxostylis), (Malosma), Mangi-
fera, (Mauria), Melanochyla, Melanorrhoea, Metopium, (Micronychia),
Microstemon, Mosquitoxylum, Nothopegia, Odina, Oncocarpus, Parishia,
Pentaspadon, Pistacia, Pleiogynium, Poupartia, Protorhus, (Pseudospondias),
Rhodosphaera, Rhus, Schinopsis, Schinus, (Schmaltzia), Sclerocarya, Seme-
carpus, (Smodingium), Sorindeia, Spondias, Swintonia, Tapirira, (Thyr-
sodium), (Toxicodendron), Trichoscypha, (Veatchia).

LITERATURE
(i) On
General Anatomy
de Boer 214, Brocardet 276, Buscalioni and Muscatello 320, Copeland and Doyel 466,
Courchet 484, Dubard and Dop 6xi, Engler 628, 643, Goris 798, Hanausek 882, Holm
1042, 1049, Luthra and Sharma 1403, McNair 1473, I474i Starr 2188, Watkins 2367,
Weber 2380, Zemke 2505.
(ii) On Wood Structure
Baker 104, Benoist 170, den Berger 179, 182, Besson 186, Br. Hond, F. D. 274 Brown,
F, B. H. 282, Burgerstein 310, 312, Chowdhury 411, Cooper 460, Cooper and Record 461,
Coster 481, DadsweU 525, Desch 574, Fernandes 683, Foxworthy 705, Garratt 745,
462 ANACARDIACEAE
Gonggrijp 794, Greguss 2522, Greiss 817, Gupta 849, Heimsch 937, 938, Hess 962,
Howard 1088, Janssonius 1154, Jolly 1188, Kanehira 1206, 1209, Kribs 1283, Lecomte
1334, M&iiaud 1491, Messeri 1493, Pearson and Brown 1679, Pereira 1687, Pfeiffer, J. Ph.
1713, Record 1780, 1783, 1787, 1801, 1807, *8i6, 1825, I 843 1851, 1859, Record
and Hess 1886, Record and Mell 1894, Rendle 1913, Schmieg 2041, Stone 2202, 2207,
Sudworth 2218, Tang 2231, Webber 2377, Williams 2430, Yamabayashi 2478.
j^

110. CORYNOCARPACEAE
(Frc. 102 on p. 450; FIG. 104 on p. 470)

SUMMARY
The
family consists of a few species of trees up to about 40 feet high com-
prising the sole genus Corynocarpus, which occurs in Zealand and New some
of the Pacific Islands. The only species which has been described anatomically
is C. laevtgatus Forst. The wood exhibits the following characters. Vessels
mostly in clusters, perforations simple, intervascular pitting alternate and
small and pits to parenchyma similar, members moderately short. Paren-
'

chyma in broad, probably apotracheal, bands and vasicentric fusiform cells ;

common; storied. Rays all multiseriate, up to 16 cells wide and very high,
composed of mingled rows of square and procumbent cells. Fibres with
simple pits, of medium length.

LEAF
DorsiventraL Globular, multicellular hairs occur on both surfaces of
young leaves, but become disorganized and fall off when older, Cork warts
arise in association with the bases of these hairs after abscission, according to

Piggott (1719). Thread-like hairs occur on the upper surface of the stipules.
Stomata confined to the lower surface; rubiaceous. One or 2 layers of
hypoderm present beneath the upper epidermis, and a single layer beneath
the lower epidermis. Mesophyll consisting of 2 layers of almost isodiametric
palisade cells, not clearly differentiated from the spongy tissue which is
4 times as broad as the palisade. Vascular bundles of the veins embedded in
the mesophyll larger ones accompanied above and below by sclerenchyma-
;

tous elements with wide lumina. Three vascular bundles enter the base of
the leaf, but transverse sections through the distal end of the petiole (Fig.
1 02
G) exhibit an open arc of about 5 collateral bundles, with smaller, sub-
sidiary, vascular strands in the wings. Large cluster crystals abundant in the
hypoderm, especially on the upper side of the leaf; also frequent in enlarged
cells in the middle of the mesophyll.

Axis
YOUNG STEM (Fig. 102 j)
Cork arising in the outer part of the primary cortex; consisting of thin-
walled cells. Primary cortex including groups of sclerosed cells, chiefly in
the region of the lenticels. Endodermis not clearly differentiated. Pericycle
containing isolated groups of fibres with rather wide lumina, situated on the
outside of the phloem groups. Xylem and phloem, in transverse sections,
have the appearance of a ring of individually distinct bundles, with broad
lignified rays between the xylem strands. Phloem strands well developed,
 CORYNOCARPACEAE 463
devoid of resin canals and sclerenchymatous elements. Vessels tending to be
in clusters or radial groups, somewhat angular, up to about 50 /i in radial
diameter; perforations simple. Pith fairly broad, consisting of round, paren-
chymatous cells. Cluster crystals fairly numerous in the cortex, phloem, and
pith; solitary prismatic crystals occur in the ray cells.

WOOD (Fig. 104 B-E)

Vessels moderately small (50-100 p mean tangential diameter), mostly in
small multiples and irregular clusters, about 1 1 per sq. mm. with faint spiral
thickening. Perforations simple. Intervascular pitting alternate, small; pits
to ray and wood parenchyma similar to the intervascular pitting. Mean
member length about 0-3 mm. Parenchyma predominantly in broad,
probably apotracheal, bands, which are usually 5-10 cells wide and rather
short and irregular; vasicentric parenchyma also present. Strands mostly of
2 cells, but fusiform cells common. Storied. Rays up to 1 6 cells wide,
commonly more than 2 mm., and 20 stories high; uniseriates absent, about
3 rays per mm.; markedly heterogeneous, with procumbent to square or
slightlyupright cells intermingled, with sheath cells. Solitary crystals
moderately abundant in the square cells. Fibres with simple pits. Walls very
thick. Mean length about i -o mm.

TAXONOMIC NOTES
The taxonomicposition of Corynocarpus has always been somewhat
uncertain, the genus having been variously included in the Berberidaceae,
Myrsinaceae, and Anacardiaceae. It was placed in the Anacardiaceae in the
Bentham and Hooker system. In Engler's (631) opinion it belongs to the
Sapindales, in which, however, it constitutes a separate family. Hemsley
(949, 950), on the other hand, emphatically expressed the view that it has
affinities with the Anacardiaceae, from which, in his opinion, it should not be
excluded on the grounds that it is devoid of resin canals. The evidence from
wood anatomy lends no support to any relationship between the Coryno-
carpaceae and the Anacardiaceae. The wood does, however, have many points
in common with that of the Berberidaceae.

ECONOMIC USES
The Corynocarpus laevigatus Forst. are edible after culinary treat-
fruits of
ment, but not very palatable. They were at one time much prized by the
Maoris in New Zealand. The seeds contain a poisonous substance resembling
digitalin.
GENUS DESCRIBED
Corynocarpus,* (C. laevigatw Forst.).
* Kew
Represented in the slide collection,

LITERATURE
(i) On General Anatomy
Encler 631, Hemsley 949, 950, Piggott 1719.

(ii) On Wood Structure

Heimsch 938.
 (464)

111. JULIANIACEAE
SUMMARY
(i) GENERAL
A small family of resinous trees and shrubs consisting of the genera Juliania
and Orthopterygium. It is confined to tropical America. A very thorough
anatomical investigation of the family was made by Fritsch (724), using
material supplied by Hemsley (95 1) who first established the family. Fritsch's
description differs considerably from that of Jadin, which is quoted by
Solereder, and he questions whether Jadin's material was correctly named.
The most important anatomical character given by Fritsch is the constant
occurrence of resin canals in the secondary phloem of the axis.

(ii)
WOOD
Vessels small, perforations simple, intervascular pitting opposite to
alternate, pits to parenchyma often large and elongated. Parenchyma para-
tracheal, scanty. Rays up to 6 cells wide, with rather few uniseriates, almost
homogeneous. Fibres with simple pits, septate. Intercellular canals
present in the rays.
LEAF
Usually dorsiventral. Hairs unicellular or multicellular, frequently form-
ing a dense covering on the lower or both surfaces of the pinnae of the com-
pound leaves. Variously shaped glandular hairs with short stalks and more
or less club-shaped heads always present as well. Stomata generally confined
to the lower surface; ranunculaceous. Hypoderm absent. Mesophyll
usually including a single layer of elongated palisade cells beneath the upper
epidermis, but part of the spongy tissue towards the lower surface tends to
develop into palisade in Juliania adstringem Schlecht. and J. mollis Hemsl.
Spongy tissue generally occupying a smaller proportion of the mesophyll than
the palisade tissue. Veins on the lower surface of the leaf embedded in pro-
jecting collenchymatous ribs except in Orthopterygium. Vascular bundles of
the veins invariably include large resin canals in the phloem not accompanied
;

by sclerenchyma. Petiole of J. adstringens containing numerous small resin

canals in the outer cortical region. Similar canals not observed by Fritsch in
the corresponding position in Orthopterygium huaucui Hemsl. Vascular
system of the petiole in the same two species appearing, in transverse sections,
in the form of an arc-shaped group of vascular bundles near the base, but
as a circle of bundles at the distal end. Resin canals present in the phloem
of each petiolar bundle in all species, and in the pith of the petiole of J.
adstringens, Rachis similar in structure to the petiole. Crystals exclusively
clustered, mostly situated in enlarged idioblasts in the mesophyll; much
*
smaller crystals also present in the phloem. Secretory canals, see Veins'
and Tetiole'.
Species can be distinguished by differences in the hairs and in the nature of
the cuticle.
Axis
YOUNG STEM
Structure very uniform in all species examined by Fritsch (724). Cork
 JULIANIACEAE 465

apparently arising in the sub-epidermis; component cells usually tabular,

with rather thin walls. Many cells of the initially parenchymatous primary
cortex becoming more or less thickened and sclerosed except in J. glauca
Hems, et Rose. Phloem
invariably devoid of fibres. Xyiem including
irregularly distributed, circular vessels, 65-70 /z in diameter, mostly isolated,
but sometimes in groups of 2 or 3 ; perforations mostly simple, but occasional
scalariform plates also occur. Groundmass of the xylem composed of thick-
walled fibres provided with slit-shaped simple pits, and sometimes septate by
thin partitions. Rays 1-2 cells wide. Wood parenchyma generally absent
except from around the primary vessels. Pith well developed, composed of
rectangular, vertically elongated cells with simple pits. Abundant clustered
crystals and infrequent solitary ones occur in the cortex clusters present in
;

longitudinal rows of parenchymatous cells in the phloem; small infrequent

clusters recorded in the pith.
Details concerning the complex structure of the inflorescence as well as of
the fruit and embryo can best be obtained from Fritsch's (724) paper. Varia-
tions in the distribution of resin canals were found by Fritsch to be of some
value in the identification of species. Resin canals invariably present in the
primary cortex; at least one ring of large ones in the secondary phloem;
numerous small ones in the pith of all species except J. adstringens Schlecht.
and Qrthopterygium huaucui Hemsl.
WOOD 1

Vessels small (less than 100 // mean tangential diameter); solitary and in
multiples of 2-8 cells and occasional clusters; usually 15-25 per sq. mm.
Perforations simple. Intervascular pitting mostly alternate, sometimes
opposite; pits to ray and wood parenchyma often large and elongated.
Tyloses abundant. Parenchyma paratracheal, scanty. Rays up to 6 cells
wide; uniseriates rather few and not more than 10 cells high, composed of
square and upright cells; almost homogeneous (Kribs's Type I); crystals
present. Fibres with simple pits, septate and with thin walls. Intercellular
canals present in the rays the epithelial layers composed of very small cells.
;

TAXONOMIC NOTES
In the Bentham and Hooker system, Juliania was placed in the Anacar-
diaceae. Hemsley (951) created the family Julianiaceae from the genera
Juliania and Orthopterygium. While admitting that they resemble the
Anacardiaceae in certain respects, he considered their affinities with the
Juglandaceae and Cupuliferae to be closer. Hemsley's views were strongly
disputed on anatomical grounds by Fritsch (724), who demonstrated that
there is no clear-cut line of demarcation between the Julianiaceae and
Anacardiaceae. Fritsch also pointed out that the suggestion made by Jadin,
and quoted by Solereder, that there are affinities between thq Julianiaceae and
Simarubaceae, is probably based on an examination of wrongly named
material. Kershaw (1234), after examining the ovules of the Julianiaceae,
Juglandaceae, and Anacardiaceae, was unable to express a definite opinion
concerning the affinities of the Julianiaceae, since they were found to resemble
both of the families with which they were compared in certain respects.
1
Based entirely on the literature, particularly the description by Kramer (1275).
4594 Hh
 466 JULIANIACEAE
Wettstein (2416) treats the Julianiaceae as the first family of the Juglandales,
where they were also placed by Hutchinson (1113). Engler and Gilg (643 A)
include them in a separate order the Julianiales, which, with the Batidales,
comes between the Juglandales and Fagales. On the other hand, Heimsch
(938) and Kramer (1275), working with the secondary xylem, have confirmed
Fritsch's view that the Julianiaceae and Anacardiaceae are closely related to
one another. Copeland and Doyel (466) also conclude that the Julianiaceae
and Anacardiaceae are very closely related to one another.
It can thus be seen that there has been a cleavage of opinion between
anatomists and taxonomists concerning the affinities of the Julianiaceae. In
view of Fritsch's demonstration that the anatomical evidence is very heavily
loaded in favour of the Julianiaceae being related to the Anacardiaceae, whilst
they are anatomically unlike the Juglandaceae, the family is placed next to the
Anacardiaceae in this book.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Juliania, Orthopterygium.

(ii)
FOR WOOD STRUCTURE
(Juliania), (Orthopterygium).

LITERATURE
(i) On General Anatomy
Copeland and Doyel 466, Engler and Gilg 643 A, Fritsch 724, Hemsley 951, Hutchinson
1113, Kershaw 1234, Wettstein 2416.
(ii) On Wood Structure
Heimsch 938, Kramer 1275, Record 1851, Record and Hess 1886, Webber 2377.

112. CORIARIACEAE
(FiG. 1 02 on p. 450)

SUMMARY
A family consisting of shrubs from temperate regions, belonging to the single
genus Coriaria. The wood exhibits the following characters. Vessels com-
monly grouped tangentially, sometimes ring-porous, perforations simple;
intervascular pitting alternate, small; pits to parenchyma similar, sometimes
oblong; members very short. Parenchyma paratracheal, scanty, vasicentric
or confluent, storied. Rays up to 15 cells wide and without uniseriates.
Fibres with simple pits, moderately to very short.

LEAF
The lamina has been most fully investigated (Wiesner 2423, p. 904) in
Coriaria myrtifolia Linn, which exhibits the following characters, Dorsi-
ventral. Cells of the upper epidermis polygonal, somewhat angular; those
of the lower epidermis more irregular, provided with pitted anticlinal walls
and covered externally by striated cuticle. Stomata present on both surfaces,
but most numerous on the lower side; rubiaceous, but with well-developed
 CORIARIACEAE 467
cuticular striations at right angles to the pore. Mesophyll including 2 layers
of palisade cells. Vascular bundle of the midrib surrounded by collenchyma.
Petiole of Coriaria sinica Maxim (Fig. 102 i) grown at Kew exhibiting, in
transverse sections through the distal end, a solitary, open, arc-shaped vas-
cular strand, not supported by pericyclic sclerenchyma. Infrequent cluster
crystals present in the cortical region of the petiole in the same species.
Solitary monoclinic crystals, as well as crystalline concretions with a corroded
appearance, present in all parts of the mesophyll of C. myrtifolia. All cells in
the same species coloured blue on treatment with ferric chloride solution.
Needles of gypsum deposited on the addition of dilute sulphuric acid. The
mesophyll is coloured brownish-red, and a similarly coloured precipitate
formed on applying caustic soda solution, especially following a preliminary
treatment with alcohol.

Axis
YOUNG STEM (Fig. 102 K)
The following details referparticularly to Coriaria sinica Maxim grown at
Kew. Cork arising in the sub-epidermis. Primary cortex narrow, consisting
of rounded, somewhat spongy, parenchymatous cells. Pericycle containing
a few large strands of fibres. Primary phloem forming well-developed
strands, containing no sclerosed elements. Xylem in the form of a cylinder,
traversed by broad primary medullary rays, the latter being lignified between
the xylem groups. Vessels mostly in irregular clusters, the individual elements
varying considerably in size and shape as seen in transverse sections, up to
about 100 fjL in radial diameter; perforations simple, somewhat oblique. Pith
fairly broad. Infrequent cluster crystals present in the pith, and occasional
solitary ones in the ray cells.

WOOD 1

Vessels small (mean tangential diameter less than 100 p)\ mostly solitary
or mostly in multiples and clusters that often extend tangentially occasionally
;

ring-porous. Perforations simple. Intervascular pitting alternate and small;

often with coalescent apertures; pits to ray and wood parenchyma similar,
sometimes with a few oblong pits. Mean member length about 0*2 mm.
(1206). Parenchyma paratracheal, about the vessel groups and often linking
them; rather sparse in species v:ith mostly solitary vessels, forming broad,
irregular confluent bands in others. Fusiform cells present. Storied. Rays
up to 15 or more cells wide and often more than 4 mm. high; uniseriates
absent composed almost entirely of square to upright cells with sheath cells
; ;

and crystals. Fibres with simple pits; mean length about 0-6-0-8 mm.

TAXONOMIC NOTES
In the Engler system the Coriariaceae are included in the Sapindaies and
placed near the Anacardiaceae. In the Bentham and Hooker system they are
also placed next to the Anacardiaceae. They resemble the Anacardiaceae in
containing abundant tannin, but differ from them in several important details,
such as the absence of resin canals and the possession of rubiaceous stomata.
1
Based entirely on the literature.
 468 CORIARIACEAE
Hutchinson (1113) treats the Coriariaceae as the sole family in the Coriariales,
which he believes to have no connexion with the Anacardiaceae.
The wood structure lacks two of the features most characteristic of the
Anacardiaceae, the septate fibres and radial intercellular canals; neither of
these features, however, is constant throughout that family. The type of ray
in the Coriariaceae is very different from that found in the Anacardiaceae.
Heimsch (938) states: 'It seems doubtful, because of the extreme specializa-
tion of the wood rays, that these families*, i.e. Corynocarpaceae and Coria-
riaceae, 'belong with those forming the nucleus of Wettstein's Terebinthales.'

ECONOMIC USES
The leaves of Coriaria myrtifolia Linn, are sold as a source of tannin. In
this respect they strongly resemble those of sumach (Rhus coriaria Linn.).
(See Anacardiaceae, 'Economic Uses*.)

GENUS DESCRIBED
(Coriaria.)*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Casparis 345, Hutchinson 1113, Wiesner 2423.

(ii) On Wood Structure

Dadswell and Record 533, Heimsch 938, Kanehira 1206, Record 1843, 1851, Record
and Hess 1886.

113. MORINGACEAE
(FiG. 104 on p. 470)

SUMMARY
The family consists of trees with an Acacia-like habit belonging to the single
genus Moringa, of which there are only a few species. They occur in India,
North Africa, &c. The following description of the leaf and stem structure
is largely based on that by Durin (618). One of the most significant anatomical

features is the presence of myrosin cells, which occur in the mesophyll of

the leaf as well as in the cortex and phloem of both stem and root. Vessels
of the mature wood medium-sized, with numerous multiples of 3 or 4 cells,
perforations simple, pits to parenchyma large and often simple; members
moderately short. Parenchyma paratracheal vasicentric or slightly aliform,
strands of 2-4 cells, storied. Rays 2-3 cells wide, about 8 per mm., homo-
geneous, storied. Fibres with large simple pits on the radial walls, storied,
moderately short.
LEAF
Compound. Leaflets exhibiting the following characters. Dorsiventral.
Hairs unicellular, with blunt tips. Cells of the epidermis said to be mucila-
ginous in M. aptera Juss. Stomata ranunculaceous. Mesophyll partly
composed of well-developed palisade tissue which forms a continuous layer
from one margin to the other in M, pterygosperma Gaertn., but is interrupted
 MORINGACEAE 469
by collenchyma above the main vein in M. concannensis Nimmo. Midrib in
M. pterygosperma containing a crescentic strand of xylem and phloem, accom-
panied above and below by collenchyma. Starch, myrosin, and cluster
crystals of calcium oxalate abundant.

Axis
YOUNG STEM
The following description applies mainly to M. pterygosperma Gaertn.
except where stated. Cork narrow. Cortex parenchymatous, but including
strands of sclerenchyma towards the phloem (probably pericyclic scleren-
chyma). A continuous ring of fibres also occurs, except in very young stems,
nearer the periphery of the cortex. Phloem consisting of very small cells.
Xylem in the form of a cylinder traversed by rays up to 3 cells wide largely
;

composed of radial groups of vessels and fibres respectively. Vessels with

simple perforations. Pith chiefly consisting of parenchymatous cells, but
including a large, central mucilage canal, lined with epithelial tissue, and
sometimes accompanied by a second, smaller canal. Starch and myrosin
abundant in the cortex. Cluster crystals also numerous in the parenchyma-
tous tissues, especially in the portions of the rays between the phloem groups.
The general structure is similar in other species of Moringa apart from small
variations in the amount of starch, the distribution of sclerenchyma, and other
minor features.

WOOD (Fig. 104 A and D)

Vessels medium-sized (100-200 ^ mean tangential diameter); commonly
with numerous multiples of 3 or 4 cells and occasionally more; 1-5-12 per
sq. mm. Perforations simple. Intervascular pitting alternate, large; pits to
wood and ray parenchyma large, simple or slightly bordered and sometimes
elongated horizontally. Contents not observed. Mean member length 0-25-
0-3 mm. Parenchyma paratracheal, vasicentric or sometimes slightly aliform.
In some species the parenchyma includes areas of very thin-walled spongy
tissue (Lucken), which may also include parts of the rays. Sometimes con-
taining single crystals in the ordinary cells; strands of 2-4 cells, with a few
fusiform cells in some species. Storied. Rays 2-3 cells wide and low;
uniseriates very few in some species, composed of procumbent cells; about
8 per mm. homogeneous (Kribs's Types I-II). Cells sometimes containing
;

single crystals. Storied. Fibres with large lenticular, simple pits, numerous
on the radial walls but rare on the tangential. Walls thin to moderately thin.
Markedly 'bauchig'. Storied. Mean length 0-75-1*0 mm. Intercellular
canals of the vertical traumatic type reported by Record (1801). Growth
rings. The seasonal development of the wood has been investigated by
Coster (481).

ROOT
Tuberous when young, but becoming branched when mature. Cork broad
even in young roots, consisting of rectangular cells in radial rows. Cortex
parenchymatous, but including numerous groups of sclerenchymatous ele-
ments in the outer part. The xylem of M. concannensis Nimmo occurs
470 MORINGACEAE
in individually distinct vascular bundles, but is more continuous in M.pterygo-
sperma Gaertn. ; including vessels of various sizes, irregularly distributed, or,
less frequently, exhibiting a radial arrangement. Pith generally absent from
M. pterygosperma, but a small amount present in M. concannensis. Starch and

'E
FIG. 104. MORINGACEAE, A and D; CORYNOCARPACEAE, B and E;
CONNARACEAE, C and F
A, Moringa pterygosperma Gaertn. B, Corynocarpus laevigata Forst. C, Manotes macrantha Schellen-
berg. D, Moringa pterygosperma Gaertn. E, Corynocarpus laevigata Forst. F, Manotes macrantha
Schellenberg.

my rosin abundant in the cortex; cluster crystals widely distributed in the

parenchymatous tissues.

TAXONOMIC NOTES
The possession of myrosin cells indicates the existence of affinities with the
Capparidaceae, Resedaceae, and Cruciferae, and thus supports the position
 MVRINGACEAE 471
to which the family is assigned in the respective systems of Engler and
Hutchinson.
ECONOMIC USES
M.
pterygosperma Gaertn. is known as the Horse-radish Tree, and is
commonly cultivated in India and adjacent countries as well as in the West
Indies. The presence of myrosin gives the roots the flavour of horse-radish.
The fruits are edible, and the seeds yield an oil which has been used for
lubricating delicate machinery.

GENUS DESCRIBED
Moringa.
LITERATURE
(i) On General Anatomy
Durin 618, Pax 1661.
(ii) On Wood Structure
den Berger 182, Coster 481, Janssonius 1154, Record 1801, 1809, 1851.

114. CONNARACEAE
(FiG. 104 on p. 470; Fro. 105 on p. 472)

SUMMARY
(i) GENERAL
All members of
this tropical family are woody, but they exhibit a con-
siderable diversity of habit. The genera Ellipanthus, Hemandradenia, Jolly-
dora, Schellenbergia, and certain species of Cnestis and Connarus are trees.
Trees belonging to Jollydora are palm-like in habit. Most other genera are
erect shrubs, climbers, or lianes. The following description of the stem and
leaf structure is largely based on that by Schellenberg (2029). The hairs
include the following types, (i) Unicellular with one or two arms, (ii) Sym-

podial, branched hairs,

(iii) Tufted. Glandular hairs are mostly confined to
the floral organs. The epidermis consists partly or wholly of mucilaginous
cells in several genera. Stomata are confined to the lower surface of the leaf,
but the arrangement of the adjacent cells is somewhat variable, thus enabling
certain groups of genera to be distinguished from one another (see 'Leaf ').
Secretory cells and cavities of various types have been reported from the
leaf and axis of certain members of the family. The cork arises superficially
and the pericycle contains a composite and continuous ring of sclerenchyma
in the few species which have been examined. The phloem of the axis
contains tannin-sacs in Connarus. Crystals, where present, are stated to be
exclusively solitary, and prismatic or pyramidal in shape.

(ii) WOOD
large, almost exclusively solitary or with numerous
Vessels medium-sized to
multiples, occasionally with spiral thickening, perforations simple, inter-
vascular pitting alternate, pits to parenchyma typically large and simple;
members of medium length. Parenchyma typically absent or as a few cells
round the vessels, sometimes in long crystalliferous strands among the fibres.
 FIG. 105. CONNARACEAE
A-C, Two-armed and one-armed trichomes of Connarus blanchetii Planch. D~F, Branched hairs of
species of Connarus: D-E, Connarus fulvus Planch,; F, Connarus pachyneurus Radlk. the trichome in
;

E is seen from above, that in D and F from the side. By Solereder.

FIG. 106. LEGUMINOSAEPAPILIONACEAE

A, Transverse section through an intramural gland of the leaf in Psoralea hirta L. B, Transverse
section through a secretory cavity in the primary cortex of Lonchocarpus spruceanus Benth., charac-
terized by the papillose differentiation of the epithelium. C, Peculiar secretory cavity of Lonchocarpus
spruceanus with bracket-epithelium. By Solereder.
 CONNARACEAE 473
Rays exclusively uniseriate or occasionally biseriate, very numerous, com-
posed entirely of upright cells, or of procumbent cells or of both. Fibres with
;

simple pits, septate, occasionally crystalliferous moderately long; thin- walled

;

groups often present, with a parenchyma-like distribution. Vasicentric

tracheids, latex tubes, gum cysts, and intercellular canals present in
some species. Included phloem of the 'concentric' type present in Rourea.

LEAF
DorsiventraL Hairs (Fig. 105 A-F) mostly unicellular or occasionally
bicellular, but exhibiting a considerable diversity of external form. Uni-
cellular, 2-armed in Connarus and Vismianthus. Horizontal hairs supported
by short pedestals recorded in Bernardinia, Burttia, and most species of
Connarus. Hairs of Jolly dora similar but multicellular. Sympodially branched,
tree-like hairs, often appearing stellate when viewed from above, recorded in

many species of Connarus. Stellate hairs, consisting of clusters of 4 or

occasionally more members, also occur in Agelaea. Short, pin-like trichomes
present amongst longer ones in Manotes (accompanied by glandular hairs in
this genus), Paxia (forming a golden-yellow clay-like coating on young
leaves and stems of P. calophylla Gilg), Roureopsis, Spiropetalum. Glandular
hairs, mostly confined to the floral organs, recorded in Cnestis, Connarus,
Jollydora (sometimes with long stalks, and resembling those of tobacco),
Pseudoconnarus, Rourea (pro parte). Epidermis papillose on the lower surface
in species of Byrsocarpus, Cnestis, Pseudoconnarus (very numerous), Santa-
hides', including a varying proportion of mucilaginous cells, which are
frequently large and penetrate into the mesophyll, in species of Bernardinia,
Byrsocarpus* Castanola, Cnestis, Rourea', mucilaginous cells typically absent
from Agelaea, Ellipanthus, Hemandradenia, Jollydora, Pseudellipanthus, and
usually lacking in Connarus. Epidermis appearing to consist partly or wholly
of several layers owing to the formation of secondary divisions in species of
Byrsocarpus, Castanola, Cnestidium, Jaundea, Paxia, Roureopsis, Santaloides,
Spiropetalum. Epidermal cells with vertical divisions recorded in Paxia,
Roureopsis, Spiropetalum. Stomata confined to the lower surface, except
rarely on the upper side in species of Connarus. Arrangement of the neigh-
bouring cells very variable. Stomata rubiaceous in Bernardinia, Byrsocarpus,
Cnestis, Jaundea, Jollydora, Paxia, Rourea, Roureopsis, Santaloides, Spiro-
petalum, Taeniochlaena', surroanded by a circle of small cells in Cnestidium,
Connarus, Ellipanthus, Hemandradenia, Manotes, Pseudellipanthus, Pseudo-
connarus. Subsidiary cells in the first of these 2 groups of genera sometimes
become subdivided, after which the stornata in both groups of genera resemble
one another. Stomata caryophyllaceous in Jollydora', surrounded by 3 sub-
sidiary cells in Agelaea and Castanola. For ecological specializations and
other details of stomata see Schellenberg (2029). A hypoderm of thickened
cells recorded beneath the upper epidermis in Agelaea oligantha Gilg (locally)
and Connarus spp. Mesophyll not usually exhibiting any special characters;
palisade tissue consisting of cubical cells in species of Jollydora. Solitary stone
cells frequent in the mesophyll above and below the midrib in most genera.
Sclerosed elements also recorded on the inside of the lower epidermis in
Rourea ligulata Baker. Secretory cavities recorded in Castanola, Connarus,
Vismianthus those of Castanola consisting of spaces filled with mucilage in
\
474 CONNARACEAE
the spongy tissue, and, in Connarus and Vismianthus, of cavities filled with
brown resinous material in all parenchymatous tissues. Vascular bundles of
the veins usually accompanied by sclerenchyma, frequently extending from
the vascular bundle to the upper surface of the leaflet. Bundles less frequently
surrounded by ordinary mesophyll cells, but this occurs in Jollydora and
Pseudoconnarus and also in certain species of Paxia, Roureopsis, Santaloides,
Spiropetalum, Taenochlaena. Small cells, each containing a solitary crystal,
present above and below the bundles, especially in the Eucnestis section of
Cnestis (but not in the sub-genus Ceratocnestis) and in Manotes. Solitary
crystals only have been recorded in the family, notably in species of Agelaea
and Cnestis. Small crystals also stated to occur in the mucilaginous lining of
the epidermal cells in certain species of Rourea. Secretory cavities, see
*MesophylP. For anatomy of the leaf joint (pulvinus) see Sperlich (2167) and
Funke (734).

Axis
YOUNG STEM
Cork arising in the sub-epidermis in species of Connarus and Rourea,
composed of cells with fairly wide lumina. Primary cortex containing
mucilage cavities formed from cells with mucilaginous membranes in Rourea
induta Planch. Pericycle containing a composite continuous ring of scleren-
chyma in species of Connarus and Rourea. Secondary phloem including

tangential groups of tannin sacs with wide lumina in Connarus fulvus Planch. ;
provided with groups of fibres in Rourea induta. Xylern in the form of a
continuous cylinder traversed by narrow rays; vessels with simple perfora-
tions. Crystals said to be exclusively solitary.

WOOD (Fig. 104 c and F)

Vessels large (mean tangential diameter more than 200 p) in Byrsocarpus,
ConnaruS) and Manotes, medium-sized in some specimens of these and in
the other genera; sometimes nearly all solitary, particularly where the vessels
are of the liane type, but otherwise commonly with numerous radial multiples,
which are sometimes of 4 or more cells in Connarus and produce a radial
pattern in Byrsocarpus] in addition there may be long clusters of 1-3 rows of
small angular vessels in Cnestidium and Connarus (938); mostly 7-15 per sq.
mm.; semi-ring-porous in some species of Connarus spiral thickening
;

reported, at any rate in the smaller vessels or at the tips of the members, in
Connarus (938, 1868) and Ellipanthus (1154). Perforations simple. Inter-
vascular pitting alternate, moderately small to moderately large; Heimsch
(938) states that some opposite pit-pairs are present in all species; pits to
wood and ray parenchyma mostly large and simple, varying from round to
horizontally elongated and sometimes slightly angular. Solid deposits and
tyloses present in some specimens, the tyloses sclerosed in Byrsocarpus.
Mean member length 0*5-0*7 mm. Parenchyma typically scanty para-
tracheal, e.g. in Ellipanthus, or absent. Williams (2430) describes the paren-
chyma of Connarus as 'metatracheal; in widely and irregularly spaced
concentric lines or bands', but this is not typical of the material examined by
the author. With long, fibre-like strands of chambered crystals scattered among
the fibres in all the genera except Byrsocarpus and Rourea (938). Rays
 CONNARACEAE 475

exclusively uniseriate or with a few biseriates in addition, though Solereder

records rays up to 5 cells wide in Cnestidium lasiocarpum Bak.; about 19 per
mm. ; the type of cell varying from square or upright only, e.g. in some species
of Connarus and Rourea (Kribs's Heterogeneous Type III), to procumbent
only, e.g. Ellipanthus (Kribs's Homogeneous Type III); frequently filled with
gummy contents in echelon in Ellipanthus. Fibres with simple pits, equally
;

numerous on both radial and tangential walls septate usually with moderately
; ;

thin walls. Often with groups of thinner-walled fibres, which also differ in
contents, that in transverse sections resemble bands or patches of parenchyma.
According to Janssonius these cells in Ellipanthus are distinctly shorter than
the other fibres. Crystalliferous, with the crystals separated by septa, in
Cnestidium (938) and Connarus. Mean length 0-7-1-0 mm. Vasicentric
tracheids present in, at least, some species of Byrsocarpus, Cnestidium (938),
Connarus, Manotes, and Rourea; usually intermediate in length between the
vessel members and the fibres and straight to irregular in shape; sometimes
with spiral thickening (938). Structures resembling latex tubes are described
by Heimsch (938) in the rays of Connarus and also reported by Record and
Hess (1886) for Cnestidium. Intercellular canals of the vertical type noted
by Heimsch (938) in Connarus martii Schellenb.; Heimsch also records the
occurrence of scattered secretory cavities, which may be cyst-like or elongated
vertically, in other species of Connarus and in two species of Cnestidium.
Included (interxylary) phloem of the 'concentric* type (c.l. circumvallatum)
present in Rourea pulchella Planch., with successive layers of xylem and
phloem separated by broad bands of conjunctive tissue containing numerous
layers of stone cells.

TAXONOMIC AND PHYLOGENETIC NOTES

Schellenberg (2029) has discussed the possible relationships between the
Connaraceae and other families, and also the phylogenetic relationships
between the genera within the Connaraceae. Anatomical features are not
taken into consideration to any great extent in the discussion.
Heimsch (938) considers that the wood of the Connaraceae is structurally
of an advanced type and similar to most other families of the Sapindales,
though he points out that the evidence of the wood structure alone cannot
decide whether this indicates a true relationship or merely parallel develop-
ment. He suggests that the Connaraceae may belong to Hutchinson's
Tinnatae', standing near the Sapindaceae rather than the Rosaceae or
Leguminosae.
ECONOMIC USES
The
family does not yield many economic products. Connarus africanus
Lam. is stated to possess anthelminthic properties, while the seeds of species
of Bernardinia, Byrsocarpus, Cnestis, Connarus, Rourea have been used locally
as poisons. The fruit of Cnestis natalensis (Hochst.) Planch, et Sonder is
edible, and known locally as wild pear.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Agelaea, Bernardinia, Burttia, Byrsocarpus, Castanola, Cnestidium, Cnestis,
476 CONNARACEAE
Connarus, Ellipanthus, Hemandradenia, Jaundea, Jollydora, Manotes, Paxia,
Pseudellipanthus, Pseudoconnarus, Rourea, Roureopsis, Santaloides, Spiro-
petalum, Taeniochlaena.

(ii)
FOR WOOD STRUCTURE
Byrsocarpus, (Cnestidium), (Cnestis), Connarus, Ellipanthus, (Jollydora),
Manotes, Rourea.
LITERATURE
(i) On General Anatomy
Courchet 485, Funke 734, Schellenberg 2027, 2028, 2029, Sperlich 2167.
(ii) On Wood Structure
Heimsch 938, Janssonius 1154, Record 1843, 1851, 1864-72, Record and Hess 1886,
Stone 2202, 2207, Williams 2430.

115. LEGUMINOSAE
115 A. MIMOSACEAE
(Fio. 107 on p. 478; FIG. 108 on p. 482; FIG. 118 on p. 536)
SUMMARY
(i) GENERAL
A tropical and sub-tropical family, consisting mostly of trees and shrubs,
but including lianes and a few herbs. The leaves of certain members of the
family exhibit 'sensitive movements'. Distinctive features are not numerous.
Both glandular and non-glandular hairs of various types occur, the former
being universally present. Uniseriate hairs, each consisting of short basal cells
and an elongated distal cell, which are common amongst the Papilionaceae,
have not been recorded in the Mimosaceae. Two-armed hairs, which occur
in certain genera of Papilionaceae and Caesalpiniaceae, also appear to be
absent. Glands occur on the petioles of most members of the family. The
epidermis of the leaf is papillose or sub-papillose in a considerable number
of genera. Vertical and horizontal divisions of the epidermal cells have been
recorded in a few instances, but true hypoderm is unknown. The stomata,
in nearly all recorded instances, are rubiaceous. The structure of the meso-
phyll varies considerably according to the type of leaf; the larger pinnules
being dorsiventral and the smaller ones centric. The centre of the mesophyll
in a number of genera contains relatively little chlorophyll, the cells being
filled with brown tanniniferous contents. Fibres, connected to the scleren-

chyma of the veins, also occur in the mesophyll of some species. The crystals
are usually solitary rhombohedra, rod-shaped, and sometimes resemble
styloids. When present in the axis they are generally situated in chambered
fibres. Secretory elements. Cells with variously coloured contents occur
in the leaf of a number of genera. Elongated secretory sacs have also been
recorded in the leaf in a few instances, but they are much more common in,
and characteristic of, the soft phloem of the young stem. Tannin is very
common in the tissues in all parts of the plant. Its occurrence, distribution,
and possible function have been discussed by Byl (326) with special reference
to Acacia. Phyllodes are common, and in these the vascular bundles are
arranged in a ring which, in most instances, is compressed so as to conform
to the flattened shape of the phyllode. According to Shirley and Lambert
 LEGUMINOSAEMIMOSACEAE 477
(2091) they usually bear stomata in equal numbers on both surfaces. For
details concerning the structure of the phyllodes of Acacia see the articles by
Buscalioni and Catalano (318, 319), Wood (2458), Peters, P. (1702), Peters, T.
(1703), and Boke (217). The assimilatory stems of the long-rooted, umbirella-
shaped acacias such as A. seyal Del, which grow in regions of restricted
rainfall, are also noteworthy on account of their specialized anatomical struc-
ture. (For further details see 'Young Stem'.) Transverse sections through
the distal end of the petiole in the small number of species which have been
investigated exhibit a continuous or interrupted ring of bundles, accompanied
by a few lateral ones. In the young stem the cork arises superficially in the
few species investigated. is sometimes developed instead of
Aerenchyma
cork in Mimosa and Neptunia oleracea Lour. The pericycle is
cinerea Veil,
generally characterized by a composite and continuous ring of sclerenchyma.
Centric medullary bundles have been recorded in Elephantorrhiza burchellii
Benth. Winged and grooved stems occur in some, but by no means all, of
the lianes in this family, whilst true anomalous structure occurs in Entada,
where strands of sieve tubes develop in the parenchymatous groundwork of
the xylem. The well-known of the Acacias is formed from parenchyma-
gum
tous elements in the pericycle, phloem, and wood.

(ii) WOOD
Vessels typically mostly solitary but with a few small multiples and
irregular clusters; with a rather vague oblique or tangential pattern in some
species ring-porous or semi-ring-porous in a few species perforations simple,
; ;

intervascular pitting alternate and small, pits to parenchyma similar; pits

vestured; members of medium length to very short. Parenchyma usually
abundant, typically paratracheal, in round or diamond-shaped sheaths, and
commonly also with diffuse strands which usually contain 1 1 or more cham-
bered crystals paratracheal parenchyma occasionally confluent or in regular
;

broad bands; terminal bands present in many species but seldom conspicuous ;

chambered crystals common in both the diffuse and the paratracheal paren-
chyma; strands usually of 2-4 cells, fusiform parenchyma cells common in
some Rays 1-9 (mostly 2-5) cells wide or exclusively uniseriate;
species.
cells typicallynarrow tangentially homogeneous with few uniseriates in about
;

two-thirds of the genera with multiseriate rays with some tendency to echelon
;

or storied arrangement in all species, but seldom distinctly storied. Fibres

with small, simple pits; septate in some genera; moderately long to moderately
short. Intercellular canals, of the vertical traumatic type, very rare.

LEAF
Dorsiventral, isobilateral or centric. Hairs including both glandular and
non-glandular types. Two-armed and uniseriate hairs consisting of short
basal cellsaccompanied by an elongated distal cell not common, but hairs
consisting of an elongated cell and 2 basal cells sunk in the epidermis
recorded by Kienholz (1236) in Sophora iomentosa Linn.

I. Non-glandular hairs
(a) Unicellular or uniseriate, in the latter case with thin division walls,
(i)
Uniseriate but with a bulbous base in Albizzia sp. (ii) Unicellular,
478 LEGUMINOSAEMIMOSACEAE
one-armed, but with a swelling in the place where a second arm might
be expected in Inga and Xylia. (iii) Unicellular, bracket-shaped,
hooked in species of Calliandra. (iv) With the base surrounded by a
rosette of cells having cystolith-like thickenings in Affonseajuglandifolia
St. Hil.

FIG. 107. LEGUMINOSAE

A, Shaggy hair of Mimosa obtusifolia Willd. B, Shaggy hair of Mimosa furfuracea Benth., resembling
a fir-tree. To the base of both of these sclerenchymatous cells are attached, which, penetrating into
the mesophyll, anchor the hair firmly, and are accompanied by crystal-cells (crystals shaded). C,
Shaggy hair of Mimosa incana Benth., resembling a fir-tree. D-E, Branched glandular hairs of
Mimosa spruceana Benth.; E, Gland seen from above. F-G, Peltate gland of Mimosa punctulata
Spruce. By Solereder.

(b) Branched shaggy hairs (Fig. 107 A-C) of various types in numerous
species of Mimosa, possibly of specific diagnostic value.
(c) Tufted hairs recorded in species of Pentaclethra and Prosopis.

II. Glandular hairs

(a) Hairs with uniseriate stalks and ellipsoidal, multicellular heads of
variable size recorded in species of Acacia, Adenanthera, Affonsea,
Albizzia, Calliandra^ Desmanthus, Elephantorrhiza, Entada, Enterolobium,
Gagnebina, Inga, Leucaena, Lysiloma, Mimosa, Neptunia, Parkia, Pipta-
 LEGUMINOSAEMIMOSACEAE 479
denia, Pithecolobium, Prosopis, Schrankia, Serianthes, Stryphnodendron,
Xylia.
(b) Peltate, with heads of various sizes in species of Acacia and Mimosa
(Figioyp-G).
(c) Branched or palmately divided glandular hairs present on the lower
side of the leaf in Mimosa spruceana Benth. (Fig. 107 D-E).
(d) Glandular shaggy hairs with multi- or uni-seriate stalks and spherical
or obconical heads in a number of species of Mimosa.

Glands, consisting of groups of parenchyma cells situated above bundle-

endings and surrounded by abundant crystals, present in the petiole of most
members of the family. The occurrence and distribution of glands in
Australian Acacias have been described by Hardy (889). They are to be found
on the upper edge of the petiole, rachis, or phyllode, and are most numerous
in the Bipinnatae, where there is usually a gland situated immediately below
the base of each pair of pinnae. In the Phyllodineae they occur relatively high
up (towards the distal end) in species with relatively broad phyllodes, but
more towards or below the base in species with long phyllodes. Certain
species belonging to the same group bear 2 or more glands, and several have
an auxiliary nerve connecting the marginal gland obliquely to the midrib, but
this inconstant feature is said to be restricted to the Uninerves. Glands are
absent from or most inconspicuous in members of the Calamiformes and
other Acacias from the dry north-west districts of Victoria. Cuticle usually
verrucose or granular, rarely smooth. Leaf surface sometimes covered with a
white deposit of a wax-like substance in species of Acacia and Mimosa or by
a resinous secretion from glandular hairs in Acacia sp. sometimes having a
;

varnished appearance owing to the structure of the cuticle itself in species of

Mimosa. Epidermis with short, hemispherical papillae on the lower side of
the leaf in species of Acacia, Calliandra, Enterolobium, Mimosa, Pithecolobium,
Sophora, Stryphnodendron, with mammiform papillae in Albizzia, Entero-
lobium, Mimosa, Pithecolobium, occasionally with finger-shaped papillae in
Albizzia and Pithecolobium. Epidermis sub-papillose, particularly on the
lower surface, in species of Adenanthera, Albizzia, Calliandra, Elephantorrhiza,
Entada, Enterolobium, Gagnebina, Leucaena, Lysiloma, Piptadenia, Pitheco-
lobium, Plathymenia, Prosopis, Serianthes, Stryphnodendron. Outer walls of the
epidermal cells often considerably thickened; with ridge-like external projec-
tions either on the whole surface, or confined to the region of the stomata, in
species of Acacia and Mimosa. Inner walls of the epidermal cells frequently
mucilaginous in species of Acacia, Adenanthera, Albizzia, Calliandra,
Desmanthus, Dichrostachys, Elephantorrhiza, Entada, Leucaena, Lysiloma,
Mimosa, Neptunia, Parkia, Piptadenia, Pithecolobium, Schrankia, Serianthes,
Stryphnodendron. Epidermal and adjacent cells silicified in species of
Calliandra and Pithecolobium. Epidermal cells usually with vertical divisions
in Enterolobium with vertical and horizontal divisions in Albizzia ferruginea
;

Benth. Hypoderm not recorded. Stomata often confined to the lower

surface, particularly amongst the Adenanthereae, Ingeae, and Parkieae;
uniformly distributed on both sides of the leaf in the Acacieae, the Eumi-
moseae, as well as in Dichrostachys and Neptunia', confined on the lower side
to the bases and on the upper side to the tips of the leaflets in a few species of
480 LEGUMINOSAEMIMOSACEAE
Mimosa and Piptadenia, this peculiar distribution possibly being correlated
1
with the 'sleep-position assumed by the leaflets. In Mimosa cruenta Benth.
stomata occur all over the upper surface, but on the lower side they are con-
fined to one longitudinal half of each leaflet. They are absent altogether from
the lower side of the basal leaflet. These features correspond with the 'sleep-
position' of the leaves of the species in question, the parts of the epidermis
which bear stomata being covered, while the parts free from stomata are not
covered during 'sleep'. Stomata usually rubiaceous except where this is
obscured by transverse or oblique secondary division of the subsidiary cells,
e.g. in species of Acacia and Pithecolobium, Mesophyll structure varying
according to the size and shape of the leaflets dorsiventral in the large pinnules
;

of lnga\ centric in small, narrow leaflets; with 3-4 layers of palisade cells on
the adaxial side and of arm-palisade cells on the abaxial side in Acacia Senegal
Willd. according to Sabnis (1977). The vertical phyllodes of Acacia are also
centric. Central portion of the mesophyll often differentiated from the
remainder by being relatively free from chlorophyll, the cells instead often
containing brown tanniniferous materials in species of Acacia, Calliandra,
Elephantorrhiza,Leucaena, Lysiloma, Mimosa, Pithecolobium Prosopis. Groups
,

of water-storage cells present in the mesophyll of Sophora tomentosa L. Fibres

connected with the sclerenchyrna of the veins, but much more abundantly
developed in some species than in others, recorded in the mesophyll of
Affonsea, Calliandra, Inga, Parkia, Pithecolobium. Spool-shaped stone cells
recorded by Kienholz (1236) below the upper and lower epidermis in Prosopis
sp. Smaller veins usually embedded in the mesophyll, but accompanied and
generally surrounded by sclerenchyrna vertically transcurrent by thin- walled
;

tissue in species of Calliandra, Plathymenia, Xylia, and by sclercnchymatous

tissue in Stryphnodendron coriaceum Benth. Many of the veins said by Kien-
holz (1236) to be sheathed by colourless parenchymatous cells in Prosopis sp.
Enlarged terminal tracheids occur in Adenanthera and Piptadenia. Vascular
bundles of the phyllodes arranged in a ring, compressed to correspond with
the flattening of the petiole, the xylems of the opposed pairs of bundles being
directed towards one another, and the phloem groups of each connected to
the epidermis by a small mass of sclerenchyma. Petiole in transverse sections
through the distal end, exhibiting a continuous or interrupted ring of bundles,
accompanied by accessory bundles in the wings in species of Acacia (Fig. 1 18 A),
Entada, Inga (Fig. 118 c), Mimosa, Pithecolobium. Main vascular strand
dorsally flattened in Pithecolobium dulce Benth. consisting of a ventral arc and
;

a separate dorsal portion in 'Prosopis jacari Hort.' (Fig. n8B). A

large
accessory, centric, medullary bundle with central phloem observed in Inga
punctata Willd. Main vascular strand of the petiole supported by 'pericyclic*
fibres in all of the species examined at Kew. Further details concerning

petiole structure have been recorded by Morvillez (1560) and Watari (2364).
Crystals predominantly solitary, often rhombohedral, less frequently re-
sembling styloids; clusters recorded only in the mesophyll and accompanying
the veins in Mimosa and Piptadenia. Crystals in many species particularly
numerous below the bases of the hairs. Some of the mesophyll cells not
infrequently divided into chambers, each containing one crystal embedded in
a local thickening of the cell wall.
 LEGUMINOSAEMIMOSACEAE 481

Secretory elements.
I. Cells, (a) With colourless contents, in the spongy parenchyma, and
appearing as transparent dots recorded in certain species of Calliandra,
Pentaclethra, Pithecolobium, Prosopis. (b) With colourless or yellow contents,
and arranged in groups in the neighbourhood of the veins in species of
Pentaclethra and Pithecolobium. (c) Tubular or rounded, with yellowish con-
tents and mucilaginous walls, occurring in all parts of the mesophyll in certain
species of Prosopis belonging to the section Algaroba. (d) With undetermined
contents, observed in the unlignified tissues of the phloem in species of
Acacia, Inga, Mimosa, Pithecolobium, examined at Kew.
II. Sacs. Recorded in species of Pithecolobium, and probably occurring
elsewhere as well.
The anatomy of the leaves of members of the family which exhibit 'sensitive
movements* has been studied by Steckbeck (2190) and Funke (734). Steck-
beck concluded that the movements are controlled by crystals in the endo-
dermis, each of which is surrounded by a protoplasmic sac and provided with
intercellular protoplasmic connexions.

Axis
YOUNG STEM (Fig. 118 D-E)
Cork arising superficially in all recorded instances; sometimes in the second
or third layer of cells below the epidermis more rarely in the sub-epidermis.
;

Evidence of occasional deep-seated cork reported for Acacia (see 'Economic

Uses'). Cork cells slightly flattened or almost cubical; occasionally sclerosed
on one side in Acacia-, thickened on all sides in Inga\ sometimes with delicate
walls. Cork in Acacia armata R. Br. consisting of an outer, crushed layer of
tanniniferous cells,a central region of very regular thin-walled cells without
contents and an inner zone of irregular, densely tanniniferous cells crystals
;

abundant in all parts of the cork. Cork replaced by aerenchyma on the

bladder-like swellings of the stem of the floating plant Neptunia oleracea Lour.,
but normal cork is produced in the same position when more mature. Met-
calfe (1494) confirmed that the aerenchyma originates from a phellogen in the
outer part of the cortex and consists of cells with cellulose walls and living
contents. Aerenchyma also recorded on the stems and peduncles of Mimosa
cinerea Veil. Primary cortex consisting of thin- walled cells or of collenchyma,
often containing tannin composed of specially large cells in Parkia filicoides
;

Welw, Stone cells recorded in the primary cortex only in Albizzia anthel-
minthica Brogn. Pericycle usually containing a continuous and frequently
composite ring of sclerenchyma, the fibres often being mucilaginous or un-
lignified; strands of fibres less common. Secondary phloem sometimes
stratified into fibrous and soft portions. Distal ends of the rays, where traversing
the phloem, frequently broadened towards the outside; generally devoid of
sclerotic cells. Sieve tubes provided with scalariform sieve plates. Xylem in
the form of a continuous cylinder traversed by narrow rays, more interrupted
by broader rays in some genera. Vessels with simple perforations. Pith
generally consisting of lignified cells except in aquatic species of Desmanthus,
Dichrostachys, Mimosa, Neptunia. Medullary bundles, consisting of soft
phloem surrounded by xylem almost devoid of vessels, recorded in Elephanto-
4SM li
 D
B

FIG. 108. LEGUMINOSAEMIMOSACEAE

A, Acacia melanoxylon R, Br. B, Piptadenia africana Hook. f. C, Albizzia odoratisnma Be nth.
D, A. lucida Bcnth. E, Dichrostachys glomerata Chiov. Crystalliferous parenchyma strand. F,
Vackellia astringent Speg. G, Entada abyssinica Steud. H, Acacia nilotica Del. I, Piptadenia africana
Hook. f. J> Pithecolobium confertum Benth. K, Adinobotrys atropurpureus Dum.
 LEGUMINOSAEMIMOSACEAE 483
rrhiza burchellii Benth. Secretory elements. Cells with undetermined but
probably tanniniferous contents observed in the cortex of certain species of
Acacia, Inga, Mimosa, and Pithecolobium represented in the Kew slide collec-
tion, in the phloem and medullary rays of the same genera and species except
Mimosa, and in the pith of Acacia and Inga. Elongated secretory sacs,
resembling the tanniniferous sacs of the Papilionaceae, but filled with various
substances, also recorded by Solereder in Mimosa, their presence in other
genera also being implied. For tanniniferous cells see also 'Cork' and 'Cortex'.
Solitary crystals stated to occur in thickenings of the cell walls of the primary
cortex and in chambered fibres in the phloem; large or very large solitary
crystals observed in the cortex and phloem of several species. Groups of small
crystal cells also recorded in the pith of species of Adenanthera, Desmanthus,
Dichrostachys, and Neptunia. See also under 'Cork'. The gum of the acacias
is said to be formed by the transformation of parenchymatous elements in the

pericycle, phloem, and wood. Some details concerning the structure of young
stems of Australian acacias have been published by Shirley and Lambert
(2091).
Hagerup (866) has drawn attention to the interesting structure of the
assimilatory stems of umbrella-shaped acacias such as A. seyal Del. These
plants, with long tap-roots, grow in localities where water supplies are
restricted except during a brief annual period of high rainfall. The green,
primary cortex which is the chief assimilatory tissue, is protected by an
external layer of cork consisting of living cells with translucent contents. Only
the outermost layer of cork is composed of cells with thickened inner walls,
and this layer becomes disorganized and cast off in the form of a red powder
as the stem grows older. The next layer of cork cells then becomes thickened
and behaves in the same way. Air diffusion occurs through numerous lenticels.
Leaves are developed only during the wet season when they serve as accessory
assimilatory organs. The outer layer of cells of the thorns is filled with air,
and is thought to reflect the light on to the lateral assimilatory surfaces of the
branches, which are also illuminated below by reflection from the brightly
coloured ground. Hagerup points out that the plants are not unlike cacti
during the unfavourable season, whilst the development of leaves during
rainy periods makes them more like trees in a tropical rain forest.

WOOD (Fig. 1
08)
Vessels typically medium-sized (100-200 /* mean tangential diameter) to

large; small in Serianthes myriadenia Planch, and Vachellia spp., largest in

some species of Adinobotrys, Albizzia, Entada, Enterolobium, Fillaeopsis,
Parkia, Piptadenia, and Pithecolobium typically solitary with a few multiples
;

of 2 or 3 cells and, some irregular clusters; these clusters (Fig. 108 H), usually
of small vessels, are not present in every section, but a tendency to produce
them locally appears to be characteristic, particularly of Acacia, Inga, Penta-
clethra,Samanea, Tetrapleura, and Vachellia', such clusters often accompanied
or replaced by multiples of one large and several small vessels, e.g. in some
species of Acacia, Amblygonocarpus, Enterolobium, and Xylia\ with a tendency
to an oblique pattern in some species, e.g. of Albizzia and Calpocalyx,
Pithecolobium (1894) anc* * n occasional tangential rows in some species of
Acacia, Albizzia (1154), Lysiloma, Parkia, Piptadenia (2430), Prosopis (1894),
484 LEGUMINOSAEMIMOSACEAE
and Vachellia] mostly i'5~5 per mm., rather more numerous (mostly between
5 and 10 per mm.) in some species of Abarema, Acaciella, Cedrelinga, Chloro-
leucon, Cojoba, Dichrostachys, Faidherbia, Leucaena, Poponax, Prosopis, and
Vachellia', semi-ring-porous in some species of Acacia, Albizzia, Parkia,
Prosopis (1894), and Vachellia, sometimes with a distinct zone at the beginning
or end of the growth ring, of vessels that are smaller than those of the rest of
the ring; spiral thickening reported by Record (1864) in Prosopis. Perforations
simple. Intervascular pitting alternate, small; very small in Abarema,
Calliandra, Cojoba, Lysiloma, and Wallaceodendron, only moderately small in
Enterolobium, Parkia, and Plathymenia\ sometimes with coalescent apertures;
pits to parenchyma similar to the intervascular pitting; vestured. Solid
deposits present in most of the species; tyloses not observed. Mean member
length usually 0-2-0-4 mm. Greiss (816, 817) has investigated the effect of
different conditions of water-supply on the structure of the wood of Acacia
arabtca var. nilotica Forst. and the effect of such changes of structure on the
conductivity of the wood. Parenchyma usually abundant, and predominantly
paratracheal, sometimes very abundant and occupying more space than the
fibres, e.g. in Entada and Faidherbia typically as a sheath, several cells wide,
;

about the vessels, round, diamond shaped or distinctly aliform on cross-

section and becoming confluent locally where the vessels are close together;
with considerable variation within these limits in different parts of the ring
and in different specimens; more consistently rounded (vasicentric) in some
species of Abarema, Acacia, Acaciella, Albizzia, Cedrelinga, Chloroleucon,
Dichrostachys, Enterolobium, Inga, Piptadenia, Plathymenia, Prosopis, and
Pseudosamanea more consistently confluent in some species of Acacia, Adino-
;

botrys (Fig. 108

K), Amblygonocarpus, Aubrevillea, Enterolobium, Parkia,
Prosopis, and Vachellia (Fig. 108 F); in broad bands (4-seriate or more) in
Acacia albida Del., Acacia farnesiana Willd., Adinobotrys, Calliandra, Entada,
Faidherbia, and Poponax; scattered strands (diffuse), usually containing
chambered Acacia (a few species only), Adenanthera,
crystals, present in
Adinobotrys, Albizzia p.p. (Fig. 108 c), Calpocalyx p.p., Cathormion, Cedre-
linga, Chloroleucon, Cojoba, Cylicodiscus, Dichrostachys, Inga, Leucaena,
Lysiloma, Parkia, Piptadenia, Pithecolobium p.p., Poponax, Pseudosamanea,
Serianthes, Vachellia, Wallaceodendron, and Xylia\ sometimes with terminal
parenchyma, e.g. in Acacia p.p., Albizzia p.p., Amblygonocarpus, Calliandra,
Fillaeopsis, Parkia, Pentaclethra, Vachellia, and Xylia. Chambered crystals
present in (a) scattered strands, in the genera listed above, and (b). in the
paratracheal parenchyma, particularly on the margins, of nearly all the species
examined, rare or absent from Abarema, Acaciella, Adenanthera pavonina L.,
Albizzia antunesiana Harms., Enterolobium spp., Plathy menia reticulata
Benth., Pithecolobium p.p., and Serianthes myriadenia Planch.; the maximum
number of crystals per strand seldom fewer than 1 1 and often many more
(Fig. 1 08 E); the majority of the crystals in Vachellia large and contained in
thick- walled idioblasts; with the chambered strands often locally biseriate in
Wallaceodendron celebicum Koord., and, less commonly, in Serianthes
myriadenia (tangential section); cells sometimes containing gum-like deposits.
Often with pronounced intercellular spaces. Strands most commonly of
2-4 cells, fusiform parenchyma cells common in some species of Acacia,
Albizzia, Dichrostachys, Leucaena, Pithecolobium montanum Benth. (1154),
 LEGUMINOSAEMIMOSACEAE 485

Prosopis, and Samanea. Sometimes with a tendency to storied arrangement,

e.g. in Plathymenia. In Plathymenia the sheath of vasicentric parenchyma is
narrow but this is in turn surrounded by a sheath of septate fibres that, owing
to their wider lumina and thinner walls, are distinct from those of the ground
tissue. Rays mostly2 cells wide; exclusively uniseriate or with only a few
biseriate rays in Abarema, Acacia p.p., Cedrelinga, Chloroleucon, Cojoba,
Faidherbia, Pithecolobium p. p., Pseudosamanea, Serianthes myriadenia (Benth.)
Plane., and Wallaceodendron celebicum\ 4-7 cells wide in at least some
species of Acacia, Adinobotrys, Albizzia, Dichrostachys, Entada, Entadopsis,
Inga, Parkia, Piptadenia, Pithecolobium, Prosopis, and Vachellia', 8 cells wide
in some species of Acacia and Prosopis', woods with multiseriate rays usually
with very few or very low uniseriate rays, but uniseriate rays moderately
abundant in some species in which the maximum width is 2 or 3 cells,
e.g. CalltandraguildinguBtnth., Calpocalyx klainei Pierre, Enterolobium saman
Prain, Leucaena, Parkia singularis Miq., Samanea saman (Jacq.) Merrill,
Wallaceodendron celebicum, and Xylia dolabriformis Benth., and more rarely
in some woods with larger rays, e.g. Adinobotrys atropurpureus Dum. and
Albizzia odoratissima Benth.; mostly from 4 to 8 rays per mm. but more
numerous (up to 14 per mm.) in some of the woods with small rays; homo-
geneous (Kribs's Type II, sometimes I or III); Kanehira (1206) records
heterogeneous rays in Caesalpinia sappan L.; cells usually small (about 10 /LC-)
in tangential diameter (Fig. 108 B), but distinctly larger in Amblygonocarpus,
Calpocalyx, Entada, Fillaeopsis, Prosopis p.p., and Tetrapleura\ commonly
containing gum-like deposits, crystals extremely rare. Almost always with
some tendency to arrangement in echelon or stories unless the rays are
commonly 2 or more times the height of the carnbial initials storying least
;

marked in some species of Acacia, Calpocalyx, Inga, Prosopis, and Vachellia

and most distinct in some species of Adinobotrys, Parkia, Pithecolobium (1894),
and Plathymenia. Fibres with few, small, simple pits, more numerous on the
radial than on the tangential walls. Septate in Albizzia, Arthrosamanea,

Calpocalyx, Cathormion, Dugandia (1886), Havardia (1886), Inga, Leucaena

(1206, 1886), Pentaclethra, Piptadenia, Pithecolobium p.p., Plathymenia (see
also under 'Parenchyma*), Senegalia (1886), and Xylia. Walls usually
moderately to very thick, but thin in some species of Acacia (exceptional),
Albizzia, Entada, Enterolobium, Lysiloma, Parkia, Samanea, and Serianthes;
Janssonius (1154) states that the fibres next the paratracheal parenchyma are
usually thinner-walled than the others and have intercellular spaces, and
suggests this feature as one means of distinguishing the Mimosaceae from the
Caesalpiniaceae sometimes with a gelatinous inner layer and sometimes con-
;

taining starch or gum-like contents. Mean length o-8-i'75 mm. Inter-

cellular canals of the vertical traumatic type observed in Entada abyssinica
Steud. and reported (1886) in 'Senegalia glomerosa\ Growth rings and
seasonal development of the wood have been investigated by Coster (481) in
Acacia, Adenanthera, Enterolobium, Leucaena, and Pithecolobium and by
Chowdhury in Acacia (414) and Albizzia (415).
ANOMALOUS STRUCTURE
Winged or grooved stems occur in some, but by no means all, of the lianes
belonging to this family. True anomalous structure recorded only in species
486 LEGUM1NOSAEMIMOSACEAE
of Entada with secondary strands of sieve tubes embedded in the soft paren-
chyma forming the groundwork of the xylem. Asimilar groundwork of
parenchyma forms the greater part of the xylem in Entadopsis.

TAXONOMIC AND PHYLOGENETIC NOTES

See under Papilionaceae, p. 527.

ECONOMIC USES
Gum Arabic, now derived from Acacia Senegal Willd., was formerly
obtained from A. nilotica Del. and other species of Acacia. Edible products
are less numerous than in the Papilionaceae, but the pods of the Honey Locust
(Prosopis juliflora DC.) are used for food in the West Indies and Mexico. The
family is well known as a source of tannin derived from various barks such as
the Wattles (Acacia spp.) in Australia, South Africa, India, and the warm
parts of America. The Australian material has a particularly good record for
*
a high tannin content. The blackish- violet bark of the Black Wattle' (Acacia
decurrens Willd.) is one of the most important kinds. The anatomy of
Australian wattle barks has been studied by Welch, McGlynn, and Coombs
(2409) who give the following particulars. Epidermis persisting for a long
time, occurring even in barks of considerable thickness. Surface of the bark
of some trees coated with a whitish deposit of wax. Cork usually arising
superficially, but evidence of its deep-seated origin observed in some speci-
mens; composed of flattened cells with moderately thick walls and brown
contents. Periderm seldom more than 0*2 mm. thick. Cortical tissues bounded
internally by a more or less complete ring of irregularly-shaped, thick-walled
stone cells. Phloem including tangential bands of sieve tubes, which soon
become disorganized, situated between rows of parenchymatous cells. Phloem
also including groups of thick-walled fibres, up to 0-6 mm. in tangential
diameter and o-i mm. wide, each group partly or wholly sheathed by a single
row of short, thick- walled crystalliferous cells. Secondary phloem occasionally
containing concentrically arranged pockets filled with gum. Medullary rays
uniseriate or multiseriate, broadening considerably in the outer part of the
bark. Tannin present chiefly in the outer part of the medullary rays, in the
primary and secondary cortex and in the phloem parenchyma. Tannin con-
tent partly determined by the amount of fibre present in the phloem, a high
fibre content being correlated with a low yield of tannin. For further details

concerning the microscopy and properties of wattle barks see Wiesner (2423)
and Bodenstab (211). Brocardet (276) has described the microscopical struc-
ture and properties of a number of barks from Brazil which yield tannin.
These are derived from species of Acacia, Calliandra, Enterolobium, Inga,
Pithecolobium, and Stryphnodendron and known under variations of such local
names as Angico, Barbatimao, Cambuy, Winhatico, Inga, and Jurema.
This family produces a considerable variety of timbers and, though few of
them are of world-wide importance, some are moderately widely known and
many others have considerable local importance. Among the best known are
Australian Blackwood, Acacia melanoxylon R. Br., Kokko, or East Indian
Walnut, Albizzia lebbek Benth., and Pyinkado, Xylia dolabriformis Benth.
Timbers that are in demand locally are also obtained from other species of
 LEGUMINOSAEMIMOSACEAE 487
Acacia and Albizzia and from species of Enterolobium^ Lysiloma, Marmaro-
xylon, Piptadenia, Pithocolobium, and Prosopis.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acacia,* Adenanthera, Affonsea, Albizzia, Calliandra, Desmanthus, Dichro-
stachys, Elephantorrhiza, Entada,* Enterolobium, Gagnebina, Inga,* Leu-
caena, Lysiloma, Mimosa,* Neptunia, Parkia, Pentaclethra, Piptadenia,
Pithecolobium,* Plathymenia, Prosopis,* Schrankia, Serianthes, Sophora,
Stryphnodendron, Xylia.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Abarema, Acacia, Acaciella, Adenanthera, Adinobotrys, Albizzia, Ambly-
gonocarpus, Calliandra, Calpocalyx, Cathormion, Cedrelinga, Chloroleucon,
Cojoba, Cylicodiscus, Dichrostachys, Entada, Entadopsis, Enterolobium,
Fillaeopsis, Inga, Leucaena, Lysiloma, Mimosa, Parkia, Pentaclethra, Pipta-
denia, Pithecolobium, Plathymenia, Poponax, Prosopis, Pseudosamanea,
Samanea, Serianthes, Tetrapleura, Vachellia, Wallaceodendron, Xylia.

LITERATURE
(i) On
General Anatomy
Bodenstab 211, Boke 217, Brocardet 276, Buscalioni and Catalano 318, 319, Byl 326,
Funke 734, Hagerup 866, Hardy 889, Hart 910, Kienholz 1236, Metcalfe 1494, Morvillez
1560, Peters, P. 1702, Peters, T. 1703, Roncagliolo 1947, Sabnis 1977, Shirley and Lam-
bert 2091, Steckbeck 2190, Watari 2364, Welch, McGlynn and Coombs 2409, Wiesner
2423, Wood 2458.
(ii) On Wood Structure
Bailey 73, 78, Baker 104, Becking et al. 164, Beekman 167, Benoist 170, den Berger 179,
182, 183, Besson 186, Brown, F. B. H. 282, Burgerstein 310, 312, Chalk et al. 363, 364,
Chowdhury 411, 414, 415, Cooper and Record 461, Coster 481, Dadswell 525, Dixon 592,
Foxworthy 705, Giordano 786, Greiss 816, 817, Hess 960, Hopkinson 1083, Howard 1088,
Janssonius 1154, Jolly 1188, Jones 1191, Kanehira 1206, 1209, Kribs 1283, Lecomte 1334,
Louis and Fouarge 1392, Messeri 1493, Metcalfe 1497, Pearson and Brown 1679, Pfeiffer,
J. Ph. 1713, Record 1780, 1787, 1825, I $3 I '843, Record and Hess 1886, Record and
f

Mell 1894, Scott 2075, Stone 2202, 2206, 2207, Tang 2231, Williams 2430.

115 b. CAESALPINIACEAE
(FiG. 109 on p. 490; FIG. no on p. 494; FIG. in on p. 498; FIG. 118 on p. 536)

SUMMARY
(i) GENERAL
This mainly tropical family consists chiefly of trees and shrubs, and unlike
the Papilionaceae, includes but few herbs. The glandular and non-glandular
hairs are of various kinds, but the uniseriate type with short basal cells and
an elongated distal cell, so common amongst the Papilionaceae^ is rare. The
epidermis is often papillose and sometimes mucilaginous. The arrangement
of the subsidiary cells around the stomata is very variable, even within a
single species, the 2 main types being the rubiaceous and ranunculaceous.
This variability in the structure of the stomata is also a notable feature of the
Papilionaceae. Secretory elements. Secretory cells, with varied contents,
488 LEGUMINOSAE--CAESALPINIACEAE
are common in the mesophyll, but according to Solereder the tanniniferous
idioblasts,which are so common in the Papilionaceae, are rare in the Caesal-
piniaceae. Tanniniferous cells were, however, found to be common in the
cortex, phloem, and pith of young stems of various species grown at Kew (see
'Young Stem'). Secretory cavities, lined with epithelium, also occur in the
leaf of certain genera,and are sometimes present in the primary cortex, peri-
cycle, xylem, or pith of the axis as well. Secretory canals, which cannot easily
be distinguished in transverse sections from elongated cavities, sometimes
occur in the axis. The vascular bundles of the leaf veins are usually accom-
panied by sclerenchyma, and they are embedded in the mesophyll in some
species, but vertically transcurrent in others. The petiole often exhibits the
same type of vascular structure as that of the arboreal Papilionaceae, but more
complex types also occur. No single type of vascular structure is common to
all of the genera and species with pinnate leaves. In transverse sections
through the distal end of petioles with wings or well-developed adaxial
grooves, the main vascular strand consists of a ventral arc accompanied by a
separate dorsal strand between the arms, whilst there are small accessory
bundles in the wings. In petioles which are circular or oval in transverse
section the vascular system exhibits a considerable diversity of structure (see
'Petiole', p. 491). A sclerenchymatous ring usually surrounds the main petiolar
vascular strand and is bounded externally by a sheath of cells containing
solitary crystals. Both and clustered crystals are present in all parts
solitary
of the plants. Cluster crystals are particularly characteristic of the mesophyll,
thus serving to differentiate the Caesalpiniaceae from most Papilionaceae and
Mimosaceae, where the crystals are almost invariably solitary. The pericycle
of the young stem nearly always contains a composite and continuous ring of
sclerenchyma, or a continuous ring of fibres. The pericyclic sclerenchyma is
frequently accompanied on the outside by a sheath of cells containing solitary
crystals. Grooved stems occur in certain species of Cassia, whilst band-
shaped and winged stems are also to be found in various species of Bauhinia.
Anomalous secondary thickening is another characteristic feature of some
of the Bauhinias, and includes types with successive growth rings, or, less
frequently, with segmented xylem.

(ii) WOOD
Vessels typically mostly solitary, with a few small multiples and irregular
clusters; with an oblique or tangential pattern in a few species; ring-porous
or semi-ring-porous in a few species and very occasionally with spiral thicken-
ing; perforations simple, intervascular pitting alternate and small, pits to
parenchyma typically similar but subtending larger pits in a very few species;
pits usually vestured ; members of medium length to very short. Parenchyma
usually abundant; typically paratracheal, in round or diamond-shaped sheaths,
but in irregular confluent bands or more regular continuous bands in many of
the genera terminal bands present in most of the genera, sometimes broad
;

and conspicuous; broad bands containing intercellular canals present in a

few species; a few genera with diffuse strands which often contain 1 1 or more
chambered crystals; strands usually of 2-4 cells, fusiform cells common in
some species; storied in several genera. Rays 1-7 (mostly 2-3) cells wide or
exclusively uniseriate; some genera with narrow procumbent cells, but
 LEGUMINOSAECAESALPINIA CEAE 489
relatively fewer than in the Mimosaceae; homogeneous with few uniseriates
in about two-thirds of the genera with multiseriate rays; distinctly storied in
several genera. With or without echelon arrangement in the others. Fibres
with small simple pits septate in a few genera of medium length to moderately
; ;

short. Vertical intercellular canals scattered or in bands, normally present

in some genera, and traumatic canals present in some others. Included
phloem of the 'concentric' and 'disperse* types present in a few genera.

LEAF
Generally dorsiventral except in certain species of Hoffmanseggia and
Hymenaea, described as approximately isobilateral in 'Cassia obtusa Roxb.' by
Dastur and Saxton (543). Hairs glandular and non-glandular, but the
uniseriate type with short basal cells and an elongated distal cell so common
amongst the Papilionaceae has been recorded in the Caesalpiniaceae only from
Scorodophloeus zenkeri Harms.
I. Non-glandular
(a) Unicellular, of various lengths and types, (i) Thin-walled, superficially
granulated, shaped like a pruning-hook in Cassia armata Wats. curved
;

in C. obovata Collad (Sabnis 1977). (ii) Short, curved like a horn in

Cassia occidentals Linn, (iii) Spear-shaped, with tooth-like prongs in
Dialium platysepalum Bak. (iv) Bracket-shaped or hooked in species of
Apuleia^ Dialium Labichea. (v) Two-armed in Moldenhawera (two-
,

armed but bicellular in Bauhinia).

(b) Multiseriate, bristle-like in species of Cassia.

(c) Stellate or tufted in species of Cassia.
(d) Branched, resembling a fir-tree in Cenostigma.

II. Glandular
(a) Shaggy, with a glandular, spherical base in certain species of Cassia
(Fig. 109 A-B). Some of the epidermal cells of the glandular base are
themselves elongated to form simple, unicellular hairs in a few species
of the same genus.
(b) Spherical glands, similar to the bases of
the hairs described under (a),
recorded in species of Caesalpinia and Cassia.
(c) Large, club-shaped, visible to the naked eye, occurring on the leaf
margin in the Gleditschieae and in Cassia occidentals.
(d) Short-stalked, boat-shaped glands in many species of Bauhinia
(Fig. 109 c).
(e) Club-shaped glands with short stalks, smaller than any of the above,
recorded in Apuleia, Dialium, Sclerolobium, and Tachigalia.
(/) As (e) but consisting of a glandular base depressed below the leaf
surface, and bearing a laterally directed terminal cell in species of
Berlinia, Dicorynia, Heterostemon, Humboldtia (Fig. 109 D).
(g) Black glandular dots on the lower surface of the leaf in Caesalpinia
gilliesii Wall.
(h) Nectarial glands in Cassia sp.
(t) Pearl glands in Bauhinia anatomica Link.

Epidermis papillose or sub-papillose in species of Apuleia, Bauhinia,

49 LEGUMINOSAECAESALPINIACEAE
Caesalpinia, Cassia, Crudia, Cynometra, Delonix (Poinciana regia Boj.),
Detarium, Dialium, Dicorynia, Dimorphandra^ Hardwickia (sometimes on both
sides), Labichea, Macrolobium, Peltophorum, Pterolobium, Schizolobium, Scoro-
dophloeus, Wagatea; mucilaginous in species of Caesalpinia, Cassia, Cercidium,
Cercis, Crudia, Dialium, Dimorphandra, Haematoxylon, Hqffmanseggia, Pelto-
phorum, Scorodophloeus. Epidermal cells containing clustered crystals in
Apuleia and Eperua, and mixed solitary and clustered crystals in Saraca
triandra Bak, Brown tanniniferous masses recorded in the epidermis and

Fia 109. LEGUMINOSAECAESALPINIACEAE

A, Glandular shaggy hair of Cassia trachycarpa Vog. B, Glandular shaggy hair of Cassia aurivilla
Mart. C, Boat-shaped gland with intercellular secretory space of Bauhinta sp. D, External gland of
the leaf of Humboldtia sp. By Solereder.

hypoderm of Ceratonia siliqua Linn. Epidermal cells palisade-like in Sclero-

lobium divided by thin vertical walls in Copaifera (pro parte), Kingiodendron,
;

Oxystigma, Prioria. Epidermis many layered in Bauhinia confertiflora Benth.

Hypoderm recorded on both sides of the leaf in certain species of Cassia and
in Dicorynia paraensis Benth., but not in the closely related 7X floribunda
Spruce; confined to the lower side in species of Ceratonia, Cercidium, and
Parkinsonia*
Stomata usually confined to the lower surface, but recorded on the upper
side in species of Caesalpinia, Cassia, Cercidium, Erythrophleum (only in one
species). Arrangement of the subsidiary cells very variable even within a
species, distinct types having been observed side by side in a single leaf in
certain instances. Rubiaceous stomata commonly but not invariably present
in Afzelia, Amherstia, Apuleia, Berlinia, Brownea, Caesalpinia (pro parte),
Campsiandra, Cassia, Cenostigma, Copaifera, Crudia, Cynometra, Dialium,
Dicorynia, Delonix (Poinciana regia), Dimorphandra (pro parte), Diptychandra,
Eperua, Erythrophleum (one subsidiary cell smaller than the other), Haema-
toxylon, Hardwickia, Heterostemon, Hqffmanseggia, Humboldtia, Hymenaea,
Macrolobium, Mezoneurum, Moldenhawera, Peltogyne, Peltophorum, Prioria,
Pterolobium, Saraca, Schizolobium, Schotia (pro parte), Sclerolobium, Scorodo-
 LEGUMINOSAECAESALPINIACEAE 491

phloeus, Tachigalia, Tamarindus, Trachylobium, Wagatea. Subsidiary cells

generally absent from Caesalpinia (pro parte), Ceratonia, Cercidium, Cercis,
Dimorphandra, Gleditschia, Gymnocladus, Labichea, Parkinsonia, Peltophorum,
Poeppigia, Scholia (pro parte).
Mesophyll. Sometimes containing stone cells in Macrolobium and Saraca.
Sclerenchymatous fibres recorded in species of Cynometra, Heterostemon^
Saraca, and Schotia, often appearing as transparent dots (either in the whole
leaf or after cutting transversely), the bulbous bases of the fibres abutting on
the vascular bundles and filled with silica. Fibres unite the upper and lower
epidermis like girders in some species. Sclereids of an unspecified nature
recorded by Russell and Hedin (1974) and Russell (1971) in Pterygopodium
oxyphyllum Harms. Transparent dots also caused by the presence of groups
of silicified cells in Apuleia praecox Mart.
Vascular bundles of the veins usually accompanied by sclerenchyma ;

embedded in the mesophyll or vertically transcurrent. Embedded in Afzelia,

Berlinia, Caesalpinia, Cassia, Cercidium, Cercis (pro parte), Crudia, Cynometra,
Delonix (Poinciana regia), Dialium (pro parte), Dimorphandra (pro parte),
Gleditschia, Gymnocladus, Haematoxylon, Heterostemon, Hoffmanseggia, Macro-
lobium, Mezoneurum, Moldenhawera, Parkinsonia, Peltogyne, Peltophorum,
Poeppigia, Prioria, Pterogyne, Pterolobium, Saraca, Schizolobium, Scholia,
Tamarindus', vertically transcurrent in Apuleia, Brownea, Campsiandra,
Cenostigma, Cercis (pro parte), Copaifera, Didlium (pro parte), Dicorynia,
Dimorphandra (pro parte), Diptychandra, Eperua, Hardwickia, Humboldtia,
Hymenaea, Labichea, Peltophorum, Sclerolobium, Tachigalia, Trachylobium,
Wagatea.
Petiole. Examination of the limited number of slides in the collection at
Kew has revealed the following information concerning the structure to be
seen in transverse sections through the distal end. For a very detailed account
of the petiole structure in a number of Caesalpiniaceae see Morvillez (1558)
and Watari (2364). Main vascular strand cylindrical, but sometimes having
distinct dorsal and ventral portions, the latter usually being in the form of an

open arc and the dorsal part variously shaped according to the species. The
line of demarcation between these two types is not always clearly defined.

Xylem in both types of vascular strand more continuous in some species than
in others. Main vascular strand usually separated or tending to be separated
into dorsal and ventral portions as just described in Bauhinia fornicata Link.
(Fig. 118 j), Cassia grandis Linn., Cercis siliquastrum Linn., Gleditschia capsica
Desf., G. japonica Miq., Schotia latifolia Jacq., Tamarindus indica Linn.;
more perfectly cylindrical in Caesalpinia japonica Sieb. et Zucc., Gymnocladus
dioica K. Koch., Haematoxylon campechianum Linn., Saraca declinata Miq.
(Fig. 118 F). Vascular structure distinctive in Brownea coccinea Jacq. (Fig.
1 1 8 G). Additional accessory strands present in the wings in the above species

of Bauhinia, Cassia,, Gleditschia, Tamarindus, in the 'cortex* in Brownea and

Cercis and enclosed within the main vascular strand in Saraca. Vascular
tissue strongly supported by pericylic fibres, accompanied externally by cells
containing solitary crystals, in all of the species examined. Stone cells present
in the cortex and pericycle of Brownea.

Secretory elements of various types widespread, but according to

Solereder the tannin sacs, so characteristic of the Papilionaceae, are rare.
492 LEGUMINOSAECAESALPINIACEAE
There seems to be considerable confusion concerning the chemical nature of
the contents of the secretory cells in the family, and the subject needs further
investigation.

L Secretory cells, ellipsoidal when situated in the palisade tissue and causing
pellucid dots in the leaf; spherical when situated in the spongy mesophyll.
(a) With brown tanniniferous contents in Cassia and Erythrophleum (around
the vascular system).
(b) With pale-yellow contents in Caesalpinia (certain species mostly
included in the sections Coulteria, Guilandina, and Nugania), Dipty-
chandra, Mezoneurum, Pterolobium, Wagatea.
(c) Containing mucilage in certain species of Berlinia, Cassia, Cercidium,
Macrolobium, Peltogyne.
(d) Secretory cells with unidentified but apparently tanniniferous contents
present in the phloem and usually in the 'cortical' and 'medullary*
regions of the petiole in most of the rather limited material in the Kew
slide collection.

II. Secretory cavities, usually lined with a distinct epithelium.

(a) Containing yellow resin and causing transparent dots in the leaves in
Copaifera, Eperua, Hymenaea, Peltogyne, Prioria, Trachylobium.
(b) With brown tanniniferous opaque dots on the leaf in
contents, causing
species of Caesalpinia and Cenostigma.
(c) With unspecified contents in Daniella, Detarium, Hardwickia, Kingio-
dendron, Oxystigma, and, according to Russell and Hedin (1974), in the
parenchymatous tissues of the lamina and petiole in Gossweilerodendron
and Pterygopodium.

Crystals (see also under 'Epidermis* and 'Petiole'). Clusters common,

especially in the mesophyll, thus serving to differentiate the Caesalpiniaceae
from most Papilionaceae and Mimosaceae. Solitary crystals chiefly associated
with the veins; sometimes occurring as well in local swellings of the cell wall
of the palisade tissue in species of Eperua, Macrolobium, Tachigalia, Tama-
rindus.

Axis
YOUNG STEM
Cork not very fully investigated arising superficially in species of Bauhinia,
;

Brownea, Caesalpinia, Cassia, Ceratonia, Cercis, Gleditschia, Gymnocladus,

Haematoxylon, Saraca, Scholia, Tamarindus, and in the second or third layer
of the primary cortex in Gleditschia. Cork cells greatly flattened in Gleditschia
and Tamarindus, less so in Bauhinia and Gymnocladus, cubical in Cassia and
Ceratonia\ thin- walled in Bauhinia and Gleditschia, but thicker in Gymno-
cladus; thickened on the outer tangential wall in Tamarindus and on the inner
tangential wall in Ceratonia. Sclereids recorded by Planchon (1729) in the
cork of Erythrophleum. Outer part of the primary cortex occasionally collen-
chymatous; stone cells recorded in this region in species of Brownea, Erythro-
phleum, Gleditschia, Gymnocladus, Endodermis present as a well-defined
starch sheath in Cercis siliquastrum Linn. Pericycle almost invariably con-
taining a composite and continuous ring of sclerenchyma, except rarely in
 LEGUMINOSAECAESALPINIACEAE 493
Bauhinia. A continuous zone of fibres which becomes interrupted when older
has also been recorded in Erythrophleum. Secondary phloem frequently
containing fibres, e.g. in Brownea coccinea Jacq. and Tamarindus Mica Linn, ;
occasionally stratified into hard and soft portions, e.g. in Cassia grandis Linn.;
often containing solitary crystals, occasionally accompanied by clusters. Por-
tions of the rays passing through the phloem frequently broadened towards
the exterior. Xylem in the form of a continuous cylinder traversed by
narrow rays interrupted by broader rays in some species. Vessels with simple
;

perforations. Secretory elements. An oil, containing sulphur and smelling

of garlic, present in the form of a glucoside in the primary cortex of Scorodo-
is

phloeus zenkeri Harms. The glucoside accompanied by an enzyme occurs in

cells which are deeply stained when treated with iodine. Short oil cavities also
stated to occur in the primary cortex of Copaifera spp., and elongated secretory
cavities in the same position in species of Caesalpinia, Daniella, Detarium,
Hardwickia, Kingiodendron, Oxystigma, Scholia. Similar cavities recorded in
the pith and/or pericycle as well in Detarium, Erythrophloeum densiflorum
(Elm.) Merr., but according to Planchon (1729) not in other species of
Erythrophleum and Hardwickia. Fusiform secretory cavities recorded in the
,

cortex, xylem, and pith of certain species of Gossweilerodendron and in

Pterygopodium by Russell and Hedin (1974). Secretory canals occur in the
primary cortex of Eperua falcata AubL, and in the pith of species of Copaifera,
Daniella, Oxystigma, Prioria. Secretory cells, filled with material believed to
be tanniniferous, observed in the cortex, phloem, rays, and pith of Brownea
coccinea, Cercis siliquastrum, Saraca declinata Miq., Tamarindus indica and in
the phloem only in Cassia grandis and Haematoxylon campechianum Linn.
Cells with mucilaginous walls and contents which are stained pink by safranin,
also seen in the cortex and pith of Scholia latifolia Jacq. and in the pith of
Saraca declinata. Both solitary and cluster crystals common, solitary types
being especially frequent in the cortex in material represented in the slide
collection at Kew. See also under 'Phloem*.

WOOD (Fig. no)

Vessels typically medium-sized (100-200 p mean tangential diameter);
very small (25-50 ft) in Libidibia, Poeppigia, and Zuccagnia', moderately small
(50-100 p) in Apuleia, Baikiaea, Caesalpinia, Cercidium, Copaifera mopane
Kirk., Cynometra alexandri C H. Wright, Etaballia, Holocalyx, Parkinsonia,
Peltogyne, and Pseudocopaiva\ large (more than 200 pJ) in some species of
Abauria, Acrocarpus, Afzelia, Aldina, Berlinia, Cassia, Cordyla, Daniella,
Detarium, Dialium, Dinizia, Eperua, Erythrophleum, Koompassia, Mora, Pelto-
phorum, and Sclerolobium, typically solitary with a few multiples of 2 or 3 cells
and some irregular clusters; these clusters, usually of small cells, not always
present in every section, but a tendency to produce them locally appears to be
characteristic of most of the genera radial multiples more common and some-
;

times of 4 or more cells in Apuleia, Baikiaea, Bauhinia p.p., Burkea, Caesal-

pinia, Cercidium, Cercis, Copaifera p.p., Cynometra, Holocalyx, Libidibia,
Parkinsonia, and Zuccagnia, with an oblique pattern, distinct in Zuccagnia
(Fig. no B), and rather vague in Gleditschia\ clusters more pronounced and
forming broken lines in the late wood of Cercis and Gymnocladus (Fig. 1 10 c) ;

mostly between 1*5 and 5 per mm. 5-20 per mm. in some species of Brownea,
;
 FIG. no. LEGUMINOSAECAESALPINIACEAE
A, Diatiwn kingii Prain. B, Zuccagnia punctata Cav. C, Gymnocladus canadensis Lam. D, Eperua
falcata Aubl. E, Gleditschia japanica Miq. F, Eperua falcata Aubl. G, Gossweilerodendron balsamifera
Harms. H, Baitiaea plurijuga Harms. I, Holocalysc balansae Micheli. J, Afsselia bakeri Prain. K, A.
bakeri Prain. L, Cassia siamea Lam. M, Cynometra alexandri C. H. Wright. N, Tamarindus indica
Linn.
i.e. Intercellular canals
 LEGUMINOSAECAESALPINIACEAE 495
Burkea, Caesalpinia, Chidlowia, Copaifera, Cryptosepalum, Cynometra,
Dicymbe, Haematoxylon, Hardwickia, Hymenostegia, Melanoxylon, Parkinsonia,
Peltogyne, and Pseudocopaiva, 21-40 per mm. in some species of Apuleia,
Brasilettia, Browneopsis, Caesalpinia, Campsiandra, Mezoneuron, Peiranisia,
Poeppigia, and Pseudocopaiva, more than 40 per mm. in Baikiaea plurijuga
Harms, and some specimens of Cynometra alexandri; ring-porous or semi-
ring-porous in species of Asacara (1886), Caesalpinia, Cercidium, Cercis,
Gleditschia, Gymnocladus, Parkinsonia (1851), Trachylobium, and Zuccagnia
(1886); spiral thickening present in Cercis, Gleditschia, Gymnocladus, and
Zuccagnia (1864). Perforations simple. Intervascular pitting alternate, small;
very small in Bauhinia p.p., Caesalpinia, Copaifera p.p., Crudia, Cynometra,
Dimorphandra, Hardwickia, Holocalyx, Humboldtia, Intsia, Peltogyne, Saraca,
and Zuccagnia, minute in Browneopsis, Chidlowia, Delonix, Elizabetha y
Hymenostegia, Macrolobium, and Mora; moderately large in Aldina and
Martiodendron\ occasionally with marked striations due to coalescent aper-
tures, e.g. in Acrocarpus, Holocalyx, Parkinsonia, and Zuccagnia\ pits to
parenchyma and ray cells similar to the intervascular pits, but subtending
occasional larger oblong pits in the parenchyma walls in Bauhinia^ Burkea,
Cercidium, and Copaifera p.p. Record and Hess (1886) note unilaterally com-
pound pitting in Batesia, Cynometra, Eperua, Jacqueshuberia, and Parkinsonia.
Pits vestured except in the Bauhinieae, i.e. Bauhinia and Cercis. Solid
all the species, ty loses rare, observed or reported in
deposits present in nearly
Bauhinia malabarica Roxb., Gleditschia formosana Hay. (1209) and Melan-
oxyhn brauna Schott. (1894). Mean member length O-2-O-5 mm. Paren-
chyma usually moderately abundant and predominantly paratracheal ; most
typically as a sheath, several cells wide, about the vessels, round, diamond
shaped or distinctly aliform in cross-section and often locally confluent where
the vessels are close together (Fig. no K, and N); with considerable varia-
tion within these limits in different parts of the ring and in different specimens ;
commonly rounded (vasicentric) in Batesia, Brachystegia p.p., Brasikttia,
Ceratonia, Cercidium^ Cercis> Chamaesenna, Conzattia, Copaiferap.p., Dicymbe,
Dimorphandra, Englerodendron, Eperua, Gymnocladus, Heterostemon, Intsia
p.p., Parkinsonia, Peltophorum p.p., Prioria Pseudocassia, Pterogyne p.p.,
>

Schizolobiuni) Scholia^ Sindora, Tachigalia, Vouapa> and Zuccagnia\ pre-

dominantly aliform and most typically diamond shaped in Acrocarpus, Afzelia,
Berlinia, Brachystegia (most species), Brownea, Bussea> Caesalpinia, Camp-
siandra, Cassia p.p., Copaifera p.p., Cordyla, Daniella, Detarium, Dinizia,
Erythrophleum, Goniorrhachis, Gossweilerodendron, Hardwickia,Hymenaea,
Koompassia p.p., Macrolobium, Melanoxylon, Mezo-
Intsia p.p., Isoberlinia,
neuron, Mildbraediodendron, Mora, Oxystigma, Pahudia, Peiranisia, Pterogyne
p.p., Recordoxylon; Saraca, Sclerolobium, Tamarindus, and Trachylobium; in
irregular confluent bands in Afzelia quanzensis Welw., Aldina, Amphimas,
Baikiaea, Bauhinia, Campsiandra, Cassia p.p. (Fig. no L), Chidlowia, Cyno-
metra, Dicorynia, Distemonanthus, Gleditschia, Haematoxylon, Holocalyx,
Koompassia, Peltogyne, and Peltophorum p.p. intermediate between the above
;

and aliform in Burkea, Cassia p.p., Cryptosepalum, Elizabetha, Humboldtia,

Kingiodendron, Mora, and Pseudocopaiva\ in more regular bands that are
difficult to classify in Abauria, Apuleia, Cassia p.p., Crudia, Cynometra
p.p. (Fig. no M), Dialium, Etaballia (accompanied by abaxially aliform),
 496 LEGUMINOSAECAESALPINIACEAE
Hymenostegia, Pterygopodium (containing ducts), Sciacassia, Stahlia, Storckiella,
and Zollernia; scanty paratracheal in Jacqueshuberia (1836); scattered strands
(diffuse), without crystals, present in Aldina, Campsiandra (sometimes absent)
Ceratonia, Dinizia, Gossweilerodendron, Holocalyx, Pterogyne, and Recor-
doxylon, and containing chambered crystals in Bauhinia p.p. Bussea, Cercidium,
,

Copaifera p.p., Crypterosepalum, Englerodendron, Hardwickia, Humboldtia,

Peltophorum, Saraca, Schotia, Sindora, and Tamarindus', regular broad bands
containing vertical intercellular canals present in addition to the above in
some species (see below); terminal parenchyma present in most species.
Chambered crystals present in (a) scattered strands, in the genera listed above,
and (4) in the margins of the paratracheal or banded parenchyma of many of
the genera; chambered crystals not observed in Batesia, Brasilettia, Brownea,
Browneopsis, Campsiandra, Ceratonia, Chamaesenna, Chidlowia, Dicorynia,
Didelotia, Dinizia, Elizabetha, Erythrophleum, Gleditschia, Gymnocladus,
Martiodendron, Peltophorum p.p., Poeppigia, Pterogyne, and Schizolobiwn*,
the maximum number of crystals per strand seldom fewer than 1 1 and often
many more; with 2-4 large crystals per strand in Etaballia; chambered strands
sometimes locally biseriate (radial section) in Prioria copaifera Gris.; with
sclerotic cells in Martiodendron excelsa (Benth.) Gleason; cells sometimes

containing gum-like deposits. Record and Hess (1886) note 'inflated cells' in
Batesia, and gum cysts in Poincianella. Strands most commonly of 2-4 cells,
up to 8 cells in some species of Cynometra, Dicorynia, and Koompassia,
fusiform parenchyma cells moderately common in some species of Copaifera,
Dinizia (diffuse cells only), Kingiodendron, Macrolobium, and Pterogyne, not
observed in woods with heterogeneous rays. Storied in many of the genera,
e.g. in some species of Afzelia, Aldina, Apuleia, Baikiaea, Brachystegia,
Caesalpinia, Cercis, Copaifera, Cordyla, Cynometra, Daniella, Dialium,
Dicorynia, Distemonanthus, Etaballia, Haematoxylon, Holocalyx, Hymenaea,
Kingiodendron, Mildbraediodendron, Oxystigma, Peltogyne, Prioria, Pterogyne,
Pterygopodium, Tamarindus, Zollernia, and Zuccagnia. In Poeppigia procera
Presl., scattered among the septate fibres in groups and bands, there are thin-
walled, septate parenchyma strands these strands are distinctly shorter than
;

the surrounding septate fibres, the lumina wider and the pits rounded instead of
slit-like. With marked intercellular spaces in Aldina insignis. Rays mostly

2-3 cells wide, exclusively uniseriate or with only a few biseriate rays in some
species of Bauhinia, Brachystegia, Brownea, Browneopsis, Cassia, Chidlowia,
Crudia, Cynometra, Dicymbe, Dimorphandra, Elizabetha, Englerodendron,
ErythrophleumfordiiOliv. (1206), Etaballia, Heterostemon, Humboldtia> Ibadja
(1615), hoberlinia, Jacqueshuberia (1836), Loesenera, Macrolobium, Oxystigma,
Poincianella, and Tachigalia-, 4-7 cells wide in some species of Acrocarpus,
Amphimas, Burkea, Ceratonia, Cercidium, Cercis (2158), Copaifera, Cynometra
(1154), Daniella, Detarium, Dicorynia, Distemonanthus, Eperua, Gleditschia,
Gossweilerodendron, Gymnocladus, Haematoxylon, Hardzvickia, Hymenaea,
Melanoxylon, Parkinsonia, Peltogyne, Schotia, Schizolobium, and Trachy-
lobium\ of 2 distinct sizes in Eperua\ less than i mm. high, except in Eperua
and Prioria; woods with multiseriate rays often with few uniseriates, but
genera with moderately numerous uniseriates more common than in the
Mimosaceae, e.g. Bauhinia, Berlinia, Campsiandra, Cassia, Cercis, Cynometra t

Daniella, Detarium, Elizabetha, Eperua, Gossweilerodendron, Haematoxylon,

 LEGUMINOSAECAESALPINIACEAE 497
Hymenostegia, Krugiodendron y Mora, Parkinsonia, Peltophorum, Poeppigia,
Prioria, Saraca, Scholia, Tamarindus, Trachylobium, Vouapa, and Zuccagnia;
mostly from 4 to 12 rays per mm.; 13 to 20 per mm. in some species of
Apuleia, Bauhinia, Brachystegia, Browneopsis, Cercis, Cynometra, EUzabetha,
Heterostemon, Holocalyx, Hymenostegia, Ibadja (1615), Libidibia, Poeppigia^
Stahlia, Storckiella, and Tamarindus, fewer than 4 per mm. in Eperua,
Gleditschia, Gymnocladus, and Hymenaea; homogeneous (Kribs's Types I, II,
and III) in Afzelia, Aldina, Amphimas, Baikiaea, Batesia, Berlinia p.p.,
Brachystegia, Brasilettia, Burke a, Bussea, Caesalpinia, Campsiandra, Cassia,
Cercidium, Chamaesenna, Chidlowia, Conzattia, Copaifera, Cordyla, Cynometra
(except C. ramiflora (1154)), Dialium, Dicymbe p.p., Didelotia Dimorphandra,
y

Dinizia, Erythrophleum, Gleditschia, Goniorrhachis, Gossweilerodendron,

GymnocladuSy Haematoxylon, Hardwickia, Hymenaea, Intsia, Libidibia, Martio-
dendron, Melanoxylon, Mezoneuron, Oxystigma p.p., Pahudia, Parkinsonia,
Peiranisia, Peltogyne, Peltophorum, Poeppigia, Poincianella, Pseudocassia,
Pterogyne, Schizolobium, Sciacassia, Sclerolobium, Stahlia, Storckiella, Tachi-
galia, Tamarindus, and Zuccagnia\ heterogeneous (Kribs's Types II and III),
with 1-2 marginal rows of upright cells in the other genera and with up to
4 rows in Eperua\ procumbent cells small in tangential diameter (less than
10 IJL) in some genera, e.g. Brachystegia, Bussea, Caesalpinia^ Cassia, Cerci-
dium, Cercis, Cynometra, Dialium, Dinizia, Distemonanthus, Erythrophleum,
Gleditschia (Fig. HOE), Goniorrhachis, Gossweilerodendron, Gymnocladus p.p.,
Haematoxylon, Holocalyx, Hymenaea, Koompassia, Mora p.p., Parkinsonia,
Peltogyne, Poeppigia, Pterogyne, Schizolobium, Tamarindus, and Zuccagnia\
containing gum-like deposits in most species; crystals rare, observed only in
Abauria, Acrocarpus, Caesalpinia p.p., Cordyla, Cynometra p.p., Humboldtia,
Krugiodendron, Pterogyne, and Zuccagnia. Many genera without any distinct
tendency to arrangement in echelon or stories, but distinctly storied in some
species of Afzelia, Aldina, Amphimas, Apulea, Baikiaea, Bauhinia, Brachy-
Cordyla, Crudia, Cynometra, Daniella,
stegia, Brasilettia, Caesalpinia, Cercis,
Delonix, Dialium, Dicorynia, Distemonanthtis, Erythrophleum, Etaballia,
Goniorrhachis, Haematoxylon (1851), Holocalyx, Koompassia, Lecointea, Libi-
dibia, Martiodendron, Melanoxylon, Mildbraediodendron, Pahudia, (1851),
Peltogyne, Poeppigia, Poincianella, Pseudocopaiva, Pterogyne, Stahlia, Storcki-
ella, Tamarindus, and Zollerma. In several genera in which the other elements
are storied, the rays are commonly 2 or 3 stories high, and in consequence are
not themselves distinctly storied. Fibres with few small, simple pits, more
numerous on the radial than on the tangential walls. Septate in Batesia,
Brasilettia, Bussaea, Cassia, Ceratonia, Distemonanthus, Peltophorum, Poeppigia,
Poinciana (1886), and Schizolobium. Walls usually moderately to very thick,
but thin in some species of Acrocarpus, Afzelia, Cercidium, Intsia, Prioria,
Saraca, and Schizolobium commonly with a gelatinous inner layer and often
;

with dark gum-like contents. Bands of thin-walled fibres resembling paren-

chyma on cross-section present in Poeppigia ( 1 886). Mean length 0-7-1 -4 mm.
Intercellular canals occur regularly in bands in some species of Copaifera,
Detarium, Eperua (Fig. no D), and Sindora and scattered in Daniella, Goss-
weilerodendron, Kingiodendron, Prioria, and Pterygopodium Record (1851) notes
;

canals of 'normal vertical type* in Oxystigma, though absent from O. stapfiana

A. Chev. (461); vertical traumatic canals have been recorded (1851) in Berlinia,
4594 Kk
 FIG. in. LEGUMINOSAECAESALPINIACEAE
A, Winged and undulated stem of Bauhinia sp. (H. Schenck, collection of woods, no. 167). B,
Bauhinia rubiginosa Bong. C, of Bauhinia langtdorffiana Bong. A and B, natural
Split zylem-mass
size. C, approximately natural size. After H. Schenck.
 LEGUMINOSAECAESALPINIACEAE 499

Cercidiopsis, Hardmckia, Hymenaea, Macrolobium, and Peltogyne. Included

(interxylary) phloem occurs in certain genera. See 'Anomalous Structure*
below. Growth rings and the seasonal development of the wood have been
investigated by Coster (481) in Bauhinia, Cassia, Peltophorum, Poinciana, and
Tamarindus. Besson (186) records a very high silica content in Dicorynia and
Dialium and a high ash content in the latter.

ROOT
For information concerning bacteria in the roots of Gleditschia triacanthos
Linn, see Friesner's (720) account. Fusiform secretory cavities recorded
by Russell and Hedin (1974) in the cortex and xylem of Gossweilerodendron
and Pterygopodium. Well-developed cork, a broad region of phloem and peri-
cycle, strands of fibres, and conspicuous rays in the phloem recorded by
Dastur and Saxton (543) in the older parts of the long tap-roots of Cassia
auriculata Linn.
ANOMALOUS STRUCTURE
Apart from species of Cassia with grooved stems, anomalous structure is

confined to certain species of Bauhinia. The following types are recorded by

Solereder.
A. Band shaped and winged stems (Fig. 1 1 1 A-B) in 'Bauhinia blumenaviana
H. Schenck. and other species of Bauhinia'. Stems becoming band
shaped during thickening, owing to the cambium being confined to 2
opposed arcs. Wings and ribs arising subsequently on the broad sides
of older stems. Perimedullary part of the wood forming a cruciate ring,
denser in structure than the remainder.
B. Successive rings of growth recorded in B. angulosa Vig., B. rubiginosa
Bong., and B. vahlii W. et A.
C. Xylem becoming fissured, firstly owing to dilation of the thin- walled
xylem parenchyma, each segment of wood subsequently becoming sur-
rounded by a cambium which produces xylem internally and phloem
externally. Secondary fissures also occur. The central xylem mass
sometimes becomes broken. This type recorded with certainty only in
Bauhinia langsdorffiana Bong. (Fig. me).
Handa (885, 886) has described several types of anomalous thickening in
Bauhinia. In B.japonica Maxim, the three following types occur, (i) Dilation
tissue, originating in the pith and xylem, splits up the central area of the stem.
Vascular bundles provided with cambium then arise in the dilation tissue.
This anomaly occurs towards the base of the stem. (2) The axial and peri-
axial portions of the xylem become separated by dilation parenchyma in
which small vascular bundles, often arranged in 2 rows, subsequently arise.
The bundles of the inner row are normally, and of the outer row inversely,
orientated. This occurs immediately above the stem base* (3) Numerous
vascular bundles are differentiated in dilation parenchyma situated in the pith
and inner part of the xylem of the roots, especially those which are tuberous.
Thick axes of JB. championi Benth. exhibit a cleft xylem mass and i or 2 suc-
cessive rings of xylem and phloem. Secondary bundles arise in the dilation
tissue in the pith. Each segment of xylem becomes surrounded by an almost
continuous ring of cambium, the activity of which combined with that of the
500 LEGUMINOSAECAESALPINIACEAE
secondary bundles causes the xylem mass to become considerably dissected.
Later on successive rings of xylem and phloem are produced from meristems
which probably originate in the pericycle. For other work on the stem struc-
ture of Bauhinia see Loffler (1388) and Wagner (2343). Included phloem of
the 'concentric' type (c.L circumvallatwri) observed in some species of Koom-
passia, e.g. K. excelsa Ridl. and K. malaccensis Benth. and reported by Record
( 88 ) in one
1 1 rather doubtful species of Dicymbe.

TAXONOMIC AND PHYLOGENETIC NOTES

see under Papilionaceae, p. 527.

ECONOMIC USES
Edible fruits derived from members of the Caesalpiniaceae include the
Carob or Locust Bean (Ceratonia siliqua Linn.), Tamarinds (Tamarindus
indica Linn.), the Kentucky Coffee Tree (Gymnocladus canadensis Lam.).
Tannin occurs in many species and has been extracted for use from Caesal-
pinia and Cassia. Copaiba balsam is derived from species of Copaifera. Senna
pods and leaves, used in medicine, are obtained from species of Cassia, The
most important kind is Alexandrian Senna (C. acutifolia Del.) which is con-
sidered to be superior to Arabian or Tinnevelly Senna (C. angustifolia Vahl.).
The leaves of other species of Cassia are sometimes used as substitutes. The
bark of Erythrophleum guineense G. Don, used as an arrow poison in West
Africa, has also been employed as an anaesthetic in dentistry.
Senna leaves can be recognized by the following characters the thick- walled
:

unicellular trichomes the cells of the epidermis which are sometimes hori-
;

zontally divided and mucilaginous; the mostly rubiaceous stomata on both

surfaces the isobilateral mesophyll with a single layer of palisade cells towards
;

each surface; the spongy tissue in the centre of the mesophyll containing
cluster crystals of calcium oxalate; the vascular bundles accompanied by
fibres above and below, with cells containing solitary crystals situated externally
to the fibre groups. Various attempts have been made to devise methods of
distinguishing the powdered leaves of C. acutifolia from those of C. angusti-
folia. Levin (1360) found that the average vein-islet number is 26 for C.
acutifolia and 21 for C. angustifolia. Saber (1976) found that the epidermal
area per gramme was not sufficiently constant to serve as a basis for separating
the 2 species, but George (755) was more successful when using variations in
palisade ratios. Gilg and Schuster (780) state that the leaflets of Cassia
auriculata Linn, have been substituted for those of genuine senna. The sub-
stitute leaflets have a dorsiventral mesophyll with 2 layers of palisade cells.

Voigt (2339) records that powdered senna has been found with an admixture
of Belladonna,
The most characteristic anatomical features of the bark of Erythrophleum
guineense include the very numerous large groups of sclerenchymatous cells ;

the stone cells and fibres in the pericycle; the tanniniferous cells.
The woods of this family are often very durable and many are distinctively
coloured; some indeed have been important sources of dyes, e.g. Logwood,
Haematoxylon campechianum L., and Brazilwood, H. brasiletto Karst.
Several of the timbers are widely known, e.g. West Indian Locust, Hymenaea
courbaril L., Purpleheart, Peltogyne spp., Bubinga, Copaifera spp., and
 LEGUMINOSAECAESALPINIACEAE 501
'Rhodesian teak*, Baikiaea plurijuga Harms. The timbers of many other
species or genera are of local importance.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Afzelia, Amherstia, Apuleia, Bauhinia,* Berlinia, Brownea,* Caesalpinia,*
Campsiandra, Cassia,* Caulotretus, Cenostigma, Ceratonia, Cercidium,
Cercis,* Copaifera, Crudia, Cynometra, Daniella, Delonix, Detarium,
Dialium, Dicorynia, Dimorphandra, Diptychandra, Eperua, Erythrophleum,
Gleditschia,* Gossweilerodendron, Gymnocladus,* Haematoxylon,* Hard-
wickia, Heterostemon, Hoffmanseggia, Humboldtia, Hymenaea, Kingioden-
dron, Labichea, Macrolobium, Mezoneurum, Moldenhawera, Oxystigma,
Parkinsonia, Peltogyne, Peltophorum, Poeppigia, Poinciana, Pomarea, Prioria,
Pterogyne, Pterolobium, Pterygopodium, Saraca,* Schizolobium, Schotia,*
Sclerolobium, Scorodophloeus, Tachigalia, Tamarindus,* Trachylobium.
* Kew
Represented in the slide collection,

(ii) FOR WOOD STRUCTURE

Abauria,
Acrocarpus, Afzelia, Amphimas, Apuleia, Baikiaea, Batesia,
Bauhinia, Berlinia, Brachystegia, Brasilettia, Brownea, Browneopsis, Burkea,
Bussea, Caesalpinia, Campsiandra, Cassia, Ceratonia, Cercidium, Cercis,
Chamaesenna, Chidlowia, Conzattia, Copaifera, Cordyla, Crudia, Cynometra,
Daniella, Delonix, Detarium, Dialium, Dicorynia, Dicymbe, Didelotia,
Dimorphandra, Dinizia, Distemonanthus, Elizabetha, Eperua, Erythro-
phleum, Gleditschia, Goniorrhachis, Gossweilerodendron, Gymnocladus,
Haematoxylon, Hardwickia, Heterostemon, Holocalyx, Humboldtia,
Hymenaea, Hymenostegia, (Ibadja), Intsia, Isoberlinia, Kingiodendron,
Koompassia, Lecointea, Libidibia, Loesenera, Macrolobium, Martiodendron,
Melanoxylon, Mezoneuron, Mildbraediodendron, Mora, Oxystigma, Pahudia,
Parkinsonia, Peiranisia, Peltogyne, Peltophorum, Poeppigia, Poincianella,
Prioria, Pseudocopaiva, Pterogyne, Pterygopodium, Recordoxylon, Saraca,
Schizolobium, Schotia, Sciacassia, Sclerolobium, Sindora, Stahlia, Storckiella,
Tachigalia, Tamarindus, Trachylobium, Vouapa, Zollernia, Zuccagnia.

LITERATURE
(i) On
General Anatomy
Dastur and Saxton 543, Friesner 720, George 755, 2522, Gilg and Schuster 780, Handa
885, 886, Levin 1366, LoefHer 1388, Maplethorpe 1437, Morvillez 1558, Planchon 1727,
1729, Russell 1971, Russell and He"din 1974, Saber 1976, Sabnis 1977, Voigt 2339,
Wagner 2343, Watari 2364, Zemke 2505.
(ii) On Wood Structure
Bailey 78, Becking ft al. 164, Benoist 170, den Berger 179, 182, 183, Besson 186, B.
Hond. Forestry Department 274, Brown, F. B. H. 280, Brown H. P. 289, Burgerstein 310,
312, Cooper and Record 461, Coster 481, Desch 568, 569, Dixon 592, Fanshawe 2519,
Foxworthy 705, Handa 885, Howard 1088, Janssonius 1147, 1x54, Jentsch 1176, Jones
1191, Kanehira 1206, 1209, Kramer 1275, Kribs 1283, Lecomte 1334, Martin-Levigne
1450, M6niaud 1492, Nicoloff 1593, Normand 1612, 1615, 1616, 1621, Pearson and
Brown 1679, Pereira 1687, Pfeiffer, H. 1711, Pfeiffer, J. Ph. 17x3, Record 1783, 1792,
1797, 1827, 1834, 1836, 1843, 1851, 1864, 1881, Record and Hess 1886, Record and Mcll
1894, Riera 1937, Scott 2075, Stone 2202, 2206, 2207, Torres 2269, Wagner 2343, Wallis
2348.
 502 LEGUMINOSAEPAPILIONACEAE
115 C. PAPIHONACEAE
(Fie. 1 06 on p. 472; FIG. 112 on pp. 504 and 505; FIG. 113 on p. 508; FIG. 114 on p. 512;
FIG. 115 on p. 520; FIG. 116 on p. 522; FIG. 117 on p. 526)

SUMMARY
(i) GENERAL
A widely distributed family, consisting of trees, shrubs, herbs, and
large,
including some xeromorphs. Although the family is well defined by its floral
and fruit characters, there is a considerable range of anatomical variation
which is largely correlated with the wide diversity of habit shown by the
different genera and species. Other anatomical variations are xeromorphic in
nature. Some anatomical characters are common to a very wide range of
genera, but none of them are sufficiently distinctive to demarcate the Papilio-
naceae from all other families at a glance.
The hairs are of glandular and non-glandular types. The latter include
typical, uniseriate hairs each with short basal cells accompanied by an
elongated terminal cell. Other kinds of non-glandular hairs are two-armed,
hooked, branched, and multicellular and shaggy. The glandular hairs may
be club-shaped with long or short stalks; with specially large, spherical heads;
uni- or biseriate but bulbous towards the base. The epidermis of the leaf is
characterized by the common occurrence of angular folds in the anticlinal
walls; by the development of papillae, especially in the lower surface, in a
great number of genera; by being frequently mucilaginous. The arrangement
of the cells surrounding the stomata is very variable, no one type being
constant throughout a single tribe. The following are the main types, (a)
Rubiaceous, accompanied on either side by i or 2 cells lying parallel to the
ostiole. (b) Surrounded by a rosette of cells, (c) Cruciferous, (d) Ranun-
culaceous.
Secretory elements. Cells or sacs, with or without tanniniferous contents,
often stained brown in herbarium specimens, and sometimes containing sub-
stances such as protein, mucilage, &c., are a very common feature both in the
stem and leaf. Secretory cavities of various types also occur, whilst canals
have been observed in Derris and Myroxylon. The crystals, which are pre-
dominantly solitary, but very variable in size and shape, sometimes have a
characteristic appearance and distribution, especially in the leaf epidermis.
Crystalliferous cells frequently form a sheath along the outer boundary of the
pericyclic sclerenchyma. Amongst other cellular contents indigo is worthy
of special mention, although it has been recorded from only a few genera.
The petiole exhibits a considerable range of vascular structure, which appears
to be partly correlated with the habit of the plant.
The young stem is frequently assimilatory in xeromorphic species. Leafy,
wing-like expansions of the stem occur in certain genera, e.g. in Lathyrus spp.
Palisade chlorenchyma is common in assimilatory stems. The assimilatory
stems of Cytisus, Genista, Spartium, Ulex, and related genera have been the
subject of several special investigations, whilst Dr. C. L. Hare, while working
at Kew, has made numerous observations on stems which exhibit features of

special interest (see 'Young Stem'). The position in which the cork originates
ismost variable, ranging from the sub-epidermis to the pericycle, considerable
 LEGUMINOSAEPAPILIONACEAE 503
differences being found between members of a single genus, or sometimes
within a species. Cork cells typically thin- walled, but in woody genera such as
Cladrastis, Laburnum, Wistaria, with thickened tangential walls; some layers
of cork develop as typical stone cells in Ulex europeus Linn. The cork is
replaced by aerenchyma in a few species and is not formed at all in some of
the herbaceous members of the Loteae, Trifolieae, and Vicieae with assimila-
tory stems. The distribution of sclerenchyma in the pericycle is also very
variable, as it may be in separate strands or in a composite and continuous
ring. Pericyclic fibres often exhibit little lignification. Various types of stem
structure have been recognized in herbaceous species, depending on the
nature and position of the sclerenchyma in relation to the vascular bundles.
Herbaceous species are typically provided with vascular bundles separated
by conspicuous medullary rays, but in the woody ones the primary rays are
usually narrow (see 'Young Stem* below). Cortical vascular bundles occur
in a few species, especially amongst those with winged or grooved stems. Two
types of anomalous structure have been recorded. In the first of these there
are successive growth rings in the primary cortex, pericycle, or, more rarely,
in the phloem. The second type, which is less common, consists in the
development of interxylary phloem. The most noteworthy and familiar fact
about the root system is the almost universal occurrence of nodules contain-
ing nitrogen-fixing bacteria.

(ii) WOOD
Vessels. Radial multiples of 2 or 3 cells moderately common in most
species; with an oblique or tangential pattern in several genera; ring-porous
or semi- ring-porous in nearly a quarter of the genera; spiral thickening
moderately common; perforations simple, intervascular pitting alternate and
small, pits to parenchyma typically similar but simple or subtending larger
pits in some species; pits vestured; members moderately to very short.
Parenchyma usually abundant and occasionally forming the ground tissue;
most typically in either confluent or more regular bands, though aliform or
vasicentric in several genera; diffuse strands present in only a few genera;
terminal bands sometimes present chambered crystals common on the margins
;

of the banded or paratracheal parenchyma, typically with 8 and not more than
ii crystals per strand, but with more in a few genera; strands very commonly
of 1-2 or only i cell (fusiform), storied in most of the genera. Rays 1-12
(mostly 2-3) cells wide (except for a few species with rays 20 or more cells
wide), or exclusively uniseriate; some genera with narrow procumbent cells,
but relatively fewer than in the Caesalpiniaceae; homogeneous in about 45 per
cent, of the genera; with few uniseriates in about two-thirds of the genera
with multiseriate rays; distinctly storied in genera with low rays, usually
without echelon arrangement in the others, the rays being more than i story
high. Fibres with small simple pits; very rarely septate; of medium length
to very short. Intercellular canals of the vertical, scattered type present in
one genus. Included phloem of the 'concentric' and 'disperse' types present
in a few genera, mostly of the former type.

LEAF
Very variable in structure owing to the wide range of leaf types in the
 FIG. 112. LEGUMINOSAEPAPILIONACEAE
AC, Ordinary papilionaceous hairs. D, Simple hair of Lonchocarpus negrensis Benth., with bulbous
septate base. E, Branched hair of Erythrina indica Lam. F, Branched trichomes of Dipteryx rosea
Spruce. Gj, Anchor-like shaggy hair of Cranocarpus martii Benth. G, Head of the anchor, seen from
;

above. H, Glandular hair of the Phaseoleae. J, Uniseriate glandular hair of Pongamia glabra Vent.
K-L, Peltate glands of Pterocarpw ancylocalyx Benth. M, Section through a peltate gland of Centro-
lobiwn robustum Mart. N, External gland of the Phaseoleae. O, Bulbous hair of Fageliat, P-Q, Vicia
faba Linn.; P, Transverse section through a stipule with its glandular area; Q, The glands of the
glandular area strongly magnified. A, B, C, D, J, K, L and M after Kopff; H, N after Debold;
Q G G
and R after Haberlandt; E, F, t , s and O by Solereder.
 LEGUMINOSAEPAPILIONACEAE 505

family; usually dorsiventral and less frequently isobilateral. Hairs glandular

and non-glandular.
L Non-glandular types (Fig. 112 A-D)
with a variable number of short basal cells, accompanied by
(a) Uniseriate,
an elongated terminal cell. This type particularly prevalent, according
to Solereder, in the Dalbergieae, Galageae, Hedysareae, Phaseoleae,

Sophoreae, and Swartzieae, and recorded also in the genera Anagyris,

Cytisus, Gastrolobium, Genista, Lotus, Lupinus, Podalyria, Retama,
Trifolium, Ulex y
Vicia.

FIG. 112. (continued)

(b) Uniseriate, consisting of cells of more equal size. This type infrequent,
recorded, for example, in Ononis.
(c) Equally or unequally two-armed in species of Aspalathus, Astragalus,
Buchenroedera, Calycotome, Chorizema, Crotalaria (rare), Cyamopsis,
Dillwynia, Diplotropis, Erinacea, Erythrina, Genista (rare), Guelden-
staedtia,Hovea, Indigofera (common), Jacksonia, Lebeckia, Lessertia>
Lotononis, Mirbelia, Oxylobium, Priestleya, Pultenaea, Sphaerophysa,
Swamsona.
(d) Branched hairs: (i)
with a uniseriate stalk in Erythrina indica Lam.
(Fig. 112 E); (ii)unicellular in Dipteryx spp. (Fig. 112 F).
(e) Hooked hairs, with short basal cells, and a larger bent, terminal cell in
Barbiera, Canavalia, Centrosema, Clitoria, Desmodium,
Alysicarpus,
Leptodesmia, Lourea, Mecopus, Ougeinia, Periandra, Phaseolus,
Eleiotis,
Pictetia, Pseudarthria, Psoralea (terminal cell parallel to leaf surface),
Pycnospora, Uraria.
(/) Multicellular, shaggy hairs recorded only in Aeschynomene, Arachis,
Chapmannia, Cranocarpus, Discolobium, Geissaspis, Ormocarpum,
Smithia (bottle-shaped), Stylosanthes. Anchor-like shaggy hairs occur
in Cranocarpus martii Benth. (Fig. 112 G).

(g) Stellate hairs recorded in Erythrina.

II. Glandular types

(a) Club-shaped, with or without a distinct stalk, common in the Phaseo-

and in Alysicarpus, BarUera, Canavalia, Ctcer (long
leae (Fig. 112 H),
So6 LEGUMINOSAEPAPILIONACEAE
stalked), Chadsia, Desmodium, Eleiotis, Hallia, Hecastophyllum, Lepto-
desmia, Lonchocarpus, Lourea, Medicago (long stalked in certain species),
Melilotus, Millettia, Mundulea, Olneya, Ononis (long stalked), Parochetus,
Psoralea, Rhynchosia, Tephrosia, Trifolium, Trigonella.
(b) Peltate hairs of various types in species of Brongniartia, Centrolobium
(Fig. 112 M), Glycyrrhiza, Harpalyce, Petteria, Pterocarpus (Fig.
II2K-L).
(c) Especially large, spherical headed glands (Fig. 112 N) with a few short
basal cells, the secretion being formed in abundance between the cuticle
and cells of the head, in Atylosia, Cajanus, Cylista, Dunbaria, Eriosema,
Flemingia, Rhynchosia.
(d) Uni- or biseriate hairs, bulbous towards the base, recorded in Adesmia,
Atylosia, Eriosema, Fagelia (Fig. 112 o), Ormocarpum, Rhynchosia.

Glandular leaf teeth in Myroxylon pubescens H. B. et K. Extra-floral

nectaries on the stipules of Canavalia, Dolichos, Erythrina (on the spines
also), Vicia (composed of stalked glands mixed with clothing hairs (Fig. 112

.
Cuticle with verrucose thickenings, often surrounding peg-like projections
of the cellulose membranes of the adjoining epidermal cells and thus producing
the appearance of pseudo-pitting in Aotus, Aspalathus, Burtonia, Cyclopia,
Davtesia, Lupinus, Oxylobium Podalyria, Pultenaea. Cuticular projections,
y

sometimes resembling papillae but at others more like hairs, recorded in

species of Aotus, Brachysema, Chorizema Daviesia, Dillwynia, Eutaxia,
y

Gastrolobium, Gompholobium, Mirbelia^ Oxylobium, Pultenaea Sphaerolobium.

y

Epidermis. Anticlinal walls provided with angular folds in certain species

ofAnagyrtSy Anthyllts, Coronilla, Dorycnium, Euchresta, Hippocrepis, Hosackia,
Hymenocarpos, Lotus, Melilotus, Onobrychis, Ononis, Securigera, Trifolium,
Trigonella.
Lower epidermis papillose or sub-papillose in species of Abrus, Adeno-
carpus, Alysicarpus, Amphicarpaea, Anthyllis, Aotus, Argyrolobium, Barbiera,
Bossiaea, Bowdichia, Brachysema, Brongniartia, Burtonia (with coronulate
apices, the individual papillae connected by cuticular ridges), Calpurnia,
Calycotome, Camptosema, Chorizema, Cladrastis, Clitoria, Coelidium, Cytisus,
Dalbergia, Dalea, Daviesia, Derris, Desmodium, Dillwynia (in longitudinal
rows), Diphysa, Diplotropis, Discolobium, Dorycnium, Drepanocarpus, Eleiotis,
Erythrina, Eutaxia, Gastrolobium, Genista, Glycine, Gompholobium, Goodia,
Hardenbergia, Harpalyce, Hosackia, Hovea, Hypocalyptus, Indigofera, Ken-
nedya, Laburnum, Latrobea, Leptodesmia, Lespedeza, Liparia, Lonchocarpus,
Lotus, Lourea, Lupinus, Machaerium, Mecopus, Mirbelia, Mucuna, Muellera,
Myrospermum, Ougeinia, Oxylobium, Periandra, Phyllota, Piscidia, Poitaea,
Priestleya, Priotropis, Pseudarthria, Pterocarpus, Pterodon, Pueraria, Pultenaea,
Pycnospora, Rhynchosia, Robinia, Shuteria, Sophora, Sphaerolobium, Strongy-
lodon, Swartzia, Sweetia, Templetonia, Trifolium, Uraria.
Papillose upper epidermis recorded only in species of Dalbergta and
Spartium.
Epidermis often including a proportion of mucilaginous cells in species of
Adenocarpus, Adesmia, Aeschynomene, Ammodendron, Amorpha, Andira, Aotus,
Arachis, Argyrolobium, Aspalanthus, Atylosia, Borbonia, Bossiaea, Bowdichia,
 LEGUMINOSAEPAPILIONACEAE 507
Brachysertia, Brongniartia, Burtonia, Calycotome, Camptosema, Canavalia,
Chaetocalyx, Chapmannia, Chorizema, Cladrastis, Clitoria, Cratylia, Crota-
laria, Cyclopia, Cymbosema, Cytisus, Dalbergia, Dalea, Dichilus, Dillwynia,
Dioclea, Diphysa, Discolobium, Dolichos, Drepanocarpus, Erinacea, Eriosema,
Euchlora, Eutaxia, Galactia, Gastrolobium, Geissaspis, Genista, Goodia, Her-
miniera, Heylandia, Hovea, Kennedya, Laburnum, Latrobea, Lebeckia, Loto-
nonis, Melolobium, Mirbelia, Nissolia, Ormocarpum, Oxylobium, Periandra,
Petalostemon, Petteria, Phaseolus, Phyllota, Pictetia Piscidia, Platymiscium,
y

Platypodium Poecilanthe, Poiretia, Priotropis, Pterocarpus, Pultenaea, Rafnia,

y

Rhynchosia, Rothia, Rudolphia, Smtthia, Soemmeringta, Sophora, Spartium,

Sphaerolobium, Strongylodon, Stylosanthes, Swartzia, Sweetia> Templetonia,
Tephrosia, Ulex, Viborgia, Viminaria, Zornia.
Epidermal cells with gelatinous inner membranes not recorded from the
Loteae, Trifolieae, and Vicieae.
Epidermal cells vertically divided by thin walls in species of Andira,
Centrolobium, Cytisus, Dalbergia, Dipteryx, Geoffraea, Hecastophyllum, Petteria
(near the stomata), Platypodium> Poecilanthe, Pterodon. Upper epidermal
cells specially large in Argyrolobium emirense Baker (Lacoste 1312).

Hypoderm present on the upper side of the leaf in species of Andira,

Canavalia, Cratylia, Crotalaria, Dalbergia, Diplotropis, Hecastophyllum^
Lonchocarpus, Myrospermum, Myroxylon, Sophora, Zollernia; on the lower
side in a few species of Andira, Bowdichia, Centrolobium, Dillwynia, Eutaxia,
Geoffraea, Pterocarpus, Pultenaea, Swartzia, Sweetia\ beneath both surfaces
in species of Daviesia and Pterocarpus.
Stomata very variable, no single type being present throughout any one of
the tribes, nor is the distribution constant throughout many of the genera.
Present on both surfaces in all investigated Loteae and Trifolieae as well as in
species of Alysicarpus, Arachis^ Argyrolobium^ Canavalia, Crotalaria (pro
parte), Desmodmm (pro parte), Dolichos, Eriosema, Indigofera,
Dalbergia,
Mucuna (pro parte), Mundulea (very small in some species), Psophocarpus,
Rhynchosia, Sesbania, Smithia, Strongylodon, Vigna, Voandzeia, Zornia. Often
confined to the upper surface in Coelidium, Dillwynia, Diphaca^ Eutaxia,
Geoffraea, Pultenaea confined to the lower surface in Aeschynomene, Chadsia,
;

Clitoria, Crotalaria (pro parte), Derris, Desmodium (pro parte), Dioclea,

Dumasia, Fagelia, Leptodesmia, Lonchocarpus, Millettia, Mucuna (pro parte),
Strongylodon; parallel to one another but placed transversely to the direction
of the midrib or of the longitudinal axis of assimilating branches of Alhagi,
Anarthrophyllum, Carmichaelia, Daviesia, Latrobea.
Arrangement of surrounding cells as follows.

(a) Rubiaceous in Alysiocarpus, Arachis> Bowdichia, Cicer (some of the

stomata), Cranocarpus Dalbergia (most stomata), Dalhousiea, Des-
y

modium, Dillwynia, Diplotropis, Eleiotis, Eutaxia, Geissaspis, Hallia,

Hovea, Hypocalyptus, Jacksonia, Lathyrus (some stomata), Lens (some
stomata), Leptodesmia, Lespedeza, Lourea, Mecopus, Millettia, Ononis %
Ormosia, Ougeinia, the Phaseoleae, Pseudarthria, Psoralea, Pycnospora,
the Swartzieae (except Zollernia), Uraria, Vicia (some stomata),
(b) Rubiaceous but with 2 pairs of subsidiary cells parallel to the pore in
species of Aotus> Brachysema, Dillwynia, Oxylobium.
508 LEG UMINOSAEPA PI LION A CEAE
(c) Surrounded by 3 or more subsidiary cells in most Galageae, Hedysareae,
Podalyrieae and Sophoreae, and sometimes in Abrus, Crotalaria (Lacoste
1311), Euchresta, Gastrolobium, Jacksonia, Latrobea, Pterocarpus, Pul-
tenaea, Scorpiurus, and Sphaerolobium.
(d) Approximating to the cruciferous type in species of Borbonia, Crotalaria,
Lebeckia, Lotononis, Priotropis, Rafnia, Viborgia.
(e) Surrounded by a rosette of cells in species of Anarthrophyllum, Genista,
Lebeckia, Templetonia.

FIG. 113. LEGUMINOSAEPAPILIONACEAE

A, Transverse section of the leaf of Rhynchosia rufescens DC. with styloid-like crystals in the palisade-
tissue. B, Surface-view of the epidermis of the leaf of Stylosanthes procumbens Sw. C, Twin crystal-
containing cells in the epidermis of the leaf of Canavalia villosa Benth. By Solereder.

(/) Ranunculaceous in most Loteae and Vicieae (for Vicia see also (a)
above).
(g) Special type in Stylosanthes (Fig. 113 B).

Central layers of mesophyll occupied. by cells containing little chlorophyll,

and often, but not invariably, filled with tanniniferous contents which are
coloured brown in dried material in species of Alysicarpus, Arachis, Butea,
Calopogoniwn, Camptosema, Cajanus, Canavalia, Chapmannia, Cleobulia,
Cranocarpus, Cratylia, Cylista, Cymbosema, Derris, Desmodium, Dtoclea,
Dolichos, Dunbaria, Eleiotis, Eriosema, Erythrina, Fagelia, Flemingia, Galactia,
Glycine, Halimodendron, Hardenbergia, Indigofera, Kennedya, Leptodesmia,
Lespedeza, Lonchocarpus, Muellera, Mundulea^ Ougeinia, Periandra, Petalo-
stemon, Phaseolus, Physostigma, Pongamia, Pseudarthriaj Pueraria, Pycnospora,
Rhynchosia, Rudolphia, Spatholobus, Stylosanthes, Tephrosia, Teramnus\
aqueous tissue recorded in the corresponding position in species ofAlysicarpus,
 LEGUMINOSAEPAPILIONACEAE 509
Crotalaria, Psoralea (Sabnis 1977). Fibres or branched sclerenchymatous
idioblasts present in the mesophyll of a few species of Ammodendron, Andira,
Bossiaea> Bowdichia> Dillwynia> Ormosia> Platymiscium> Pultenaea> Swartzia.
Sclerosed parenchyma recorded in the mesophyll of Buchenroedera.
Veins variable in structure, with or without a sheath of sclerenchyma ;

smaller ones embedded or vertically transcurrent, the latter type recorded

particularly in certain Genisteae, Podalyrieae, Trifolieae, and Vicieae, as well
as in species of Alysicarpus, Chadsia, Rhynchosia, Tephrosia. Median veins
containing two opposite systems of bundles, with their xylem groups abutting
on one another in those species of Daviesia having approximately horizontal
leaves. An annular arrangement of bundles surrounding a pith-like tissue at
the centre of the leaves occurs in certain species of Lebeckia.
Petiole. Structure very variable, differences being correlated to some
extent with the general morphology of the leaf and the habit of the plant.
Some idea of the range of structure is indicated by the following information
concerning the appearance of transverse sections through the distal end of
the petioles represented in the slide collection at Kew. For recent detailed
descriptions of the vascular system throughout the length of the petiole the
work of Morvillez (1560) and Watari (2364) should be consulted.
I. Main vascular strand composed of separate bundles.
(a) In the form of a widely open arc in Arachis hypogaea Linn., Coronilla
vaginalis Lam., Lotus tetragonolobus Linn., Medicago saliva Linn. (3
bundles only), Melilotus officinalis (L.) Lam., Trifolium ochroleucum Huds.
(b) U-shaped, but ends not incurved in Baptisia australis R. Br., Cicer
arietinum Linn., Colutea persica Boiss., Lathyrus pratensis Linn., L.
sylvestris Linn, (wings well developed), Lespedeza formosa Koehne,
Vicia cracca Linn.
(c) U-shaped but with incurved ends in Astragalus frigidus A. Gray
(Fig. 1140), Chorizema cordatum LindL, Colutea arborescens Linn.,
Psoralea onobrychis Nutt.
(d) U-shaped with incurved ends and dorsiventrally compressed in
Caragana microphylla Lam., Vicia gigantea Bunge.
(e) Cylindrical and approximately circular in Caragana arborescens Lam.,
Chadsia (Dubard & Dop 611), 'Crotalaria axillaris Hort.' (according to
Lacoste (1311) cylindrical and arc-shaped strands occur in different
species of Crotalaria), Erythrina crista-galli Linn., Galega officinalis
Linn, (surrounding a hollow pith), Glycine soja Sieb. et Zucc., Glycyr-
rhiza glabra Linn., Halimodendron halodendron Druce, Lathyrus mariti-
mus Bigel (Fig. 1140), Lupinus mutabilis Sweet (pith hollow), Phaseolus
multiflorus Willd., Pisum elatius Bieb., Robinia pseudacada Linn.,
Wistaria floribunda DC., W. sinensis Sweet.
(/) Cylindrical but dorsally concave in Apios tuberosa Moench.
II. Main vascular strand continuous, i.e. consisting of a single vascular strand.
(a) U-shaped, but ends not incurved in Spartium junceum Linn, and
Thermopsis montana Nutt.
(b) U-shaped, but dorsally compressed and with strongly incurved ends in
Indigo/era gerardiana R. Grah. (according to Lacoste (1312) the petiole
structure varies considerably in different species of Indigofera),
 Sio LEGUMINOSAEPAPILIONACEAE
Myroxylon balsamum (L.) Harms var. pereirae (Royle) Harms (Fig.

1148), Petteria ramentacea PrsL, Piptanthus tomentosus Franch, Sophora

viciifolia Hance.
(c) Cylindrical and approximately circular in Abrus precatorius Linn, (de
Boer 214), Castanospermum australe A, Cunn., Cladrastis sinensis Hemsl.,
C. tinctoria Rafin, Denis malaccensis Prain (Fig, 114 A), Laburnum
alpinum Bercht et PrsL, L. anagyroides Med., Lonchocarpus sericeus
H. B. et K., Robinia viscosa Vent., Sophora japonica Linn.

Accessory bundles observed in the wings or 'cortex* in the same species of

Apios, Castanospermum, Derris, Glycine, Glycyrrhiza, Indigofera, Laburnum,
Lathyrus, Lespedeza, Lonchocarpus, Myroxylon, Phaseolus, Piptanthus, Robinia,
Sophora, Vicia, Wistaria. Pericyclic fibres usually present around the main
vascular strand, but none observed in Chorizema, Glycine, Melilotus, Phaseolus,
Spartium, Thermopsis.
Petioles whose vascular structure does not conform to any of the above
types include the following. Desmodium gyrans DC. with a triangular petiole
with a single bundle in each of the three angles. Platyosprion platycarpum
Maxim, with a cylindrical but dorsally concave vascular strand having a small
accessory bundle in the concavity. Mucuna pruriens DC. with a flattened
petiole supplied by a compressed circle of widely spaced vascular bundles
(Lacoste 1915). Small secretory canals observed in the outer cortical region
of the petiole in Derris malaccensis, and larger ones in Myroxylon balsamum
var. pereirae.

Secretory elements (see also 'Axis'). Cells or sacs containing tannin,

protein, or a gum-like substance, often coloured brown in dried material,
observed or recorded in various parts of the mesophyll or petiole in species of
the following genera, but probably present in many others as well, Alhagi,
Anthyllis, Aotus, Apios, Brachysema, Calophaca, Caragana, Castanospermum,
Chorizema, Cladrastis, Coromlla, Corynella, Cyclopia, Cytisopsis, Dalbergia,
Daviesia, Derris, Desmodium, Dillwynia, Diphaca, Dorycnium, Ebenus, Ery-
thrina, Eutaxia, Eversmannia, Glycine, Glycyrrhiza, Goodia, Hallia, Halimo-
dendron, Hedysarum, Helminthocarpum, Hosackia, Indigofera (very numerous
in some species), Latrobea, Lespedeza, Lonchocarpm, Lotus, Machaerium (flask
shaped, belonging to the epidermis but penetrating into the mesophyll),
Melilotus, Mirbelia, Muellera, Mundulea, Myroxylon, Olneya, Onobrychis,
Ormocarpum, Ornithopus, Oxylobium, Parochetus, Petalostemon, Phaseolus,
Phyllota, Poitaea, Pongamia, Psoralea, Pterocarpus, Pultenaea, Rhynchosia,
Robinia, Sabinea, Scorpiurus, Sophora (often flask shaped as in Machaerium),
Taverniera, Tephrosia, Wistaria.
Secretory cavities of various types, such as those shown in Fig. io6c,
usually appearing as transparent dots, recorded in Amicia, Amorpha,
Anthyllis (with bright yellow contents), Apoplanesia, Dalea, Derris, Dipteryx,
Eysenhardtia, Lonchocarpus, Marina, Millettia, Muellera, Myrospermum,
Myroxylon, Petalostemon, Piscidia, Poiretia, Pongamta, Psoralea, Zornta.
Secretory canals observed in the petiole of Derris malaccensis,
Crystals mostly solitary, particularly in the Loteae and Trifolieae, occurring
in the form of rhombohedra (especially accompanying the vascular bundles
of the veins) or styloids (Fig. 113 A), the latter being more characteristic of
 LEGUMINOSAEPAPILIONACEAE 511
the mesophyll or epidermis. Spherical crystals observed only in Dalea and
Discolobium ; small crystalline grains or prisms recorded in the mesophyll of
Dalbergia, Derris, Haematophyllum, Swartzia, Virgilia. Large rhombohedral
or rod-shaped crystals, sometimes accompanied by small prismatic or acicular
forms, are particularly characteristic of Australian species of Podalyrieae; but
crystals stated to occur exclusively in the form of small prisms or needles in
species of Cyclopia and Podalyria endemic to S. Africa, as well as in species
of Anagyris, Baptisia, Piptanthus, and Thermopsis from the northern hemi-
sphere. Large crystals also recorded in Bossiaea, Goodia, Hovea, Platylobium,
and Templetonia, and small cubes, prisms, or needles in Adenocarpus, Amphi-
thalea, Anarthrophyllum, Argyrolobium, Aspalathtis, Borbonia, Calycotome,
Coelidium, Crotalaria, Cytisus, Genista, Hypocalyptus, Lathriogyna, Lebeckia,
Liparia, Lupinus, Melobium, Petteria, Priotropis, Spartium, Ulex, Viborgia,
Rod-shaped crystals (styloids) recorded in the palisade tissue of Abrus,
Aeschynomene, Amphicarpaea, Anthyllis, Arachis, Atylosia, Bossiaea, Calo-
pogonium, Canavalia, Centrosema, Chapmannia, Clitoria, Cologania, Coronilla,
Coursetia, Cytisopsis, Dalbergia, Dalea> Daviesia, Derris, Desmodium, Dioclea,
Diphysa, Drepanocarpus, Eriosema, Erythrina, Eutaxia, Eysenhardtia, Flemin-
gia, Galactia, Geoffraea, Glycine, Grona, Hammatolobium, Herminiera, Hovea,
Indigofera, hotropis, Lathyrus, Latrobea, Lonchocarpus, Lotus, Lourea,
Machaerium, Millettia, Mucuna, Muellera, Olneya, Ononis, Orobus, Pachy-
rhizus, Petalostemon, Petteria, Phaseohts, Pictetia, Platylobium, Poecilanthe,
Poitaea, Pongamia, Psoralea, Pueraria, Pultenaea, Rhynchosia, Robinia,
Rudolphia, Sophora, Spatholobus, Strongylodon, Swartzia, Templetonia,
Tephrosia, Teramnus, Trifolium (a few species), Vicia, Vigna.
Variously, and sometimes characteristically arranged, rod-shaped crystals
occur in the epidermis of certain species of Abrus^ Arachis Canavalia (in
y

isolated pairs of epidermal cells, Fig. 113 c), Chapmannia, Cleobulia, Crano-
carpus, Cratylia, Dalea, Dioclea, Dolichos, Drepanocarpus, Eriosema, Erythrina,
Galactia, Geoffraea, Glycine; Millettia, Petalostemon, Psoraka, Rudolphia,
Sophora, Stylosanthes (curious distribution shown in Fig. 113 B), Swartzia.
Especially large crystals accompany the veins in Brongniartia, Harpalyce,
and Sabinea. Large prismatic crystals, occupy the whole lumen of palisade-
like cells immediately below the two-armed hairs in Indigofera lespedezioides
H. B. et K. Sphaero-crystalline masses of unknown chemical composition,
recorded in the epidermis, particularly in herbarium material, of Anagyris,
Argyrolobium, Aspalathus, Chorizema, Crotalaria^ Cyclopia, Dillwynia,
Eutaxia, Hovea, Latrobea, Phyllota, Piptanthus, Podalyria, Pultenaea,
Thermopsis.
Saponin in shapeless masses, stated to occur in the epidermis of Aspalathus,
Indigo, or bodies resembling this substance, recorded in Crotalaria, Hel~
minthocarpum, Hosackia, Hymenocarpos, Indigofera, Lathyrus, Lens, Lotus,
Melobium, Petalostemon Priotropis. For further details concerning the leaf
9

structure in numerous species see Leupin's (1364) thesis.

Axis
YOUNG STEM 114 D, E, F, H, and i)
(Fig.

Epidermis of Chorizema cordatum Lindl. persistent, with very thick

cuticle.
 FIG. 114. LEGUMINOSAEPAPILIONACEAE
A, Derm malaccensis Prain. x 18. B, Myroxylon balsamum (L.) Harms var. pereirae (Royle)
Petiole
Harms. Petiole X 18. C, Astragalus frigidus A. Gray. Petiole x 18. D, Lathyrus sylvestris Linn.
Stem X7. E, Apios tuberosa Moench. Young Stem X33 F, Cytisus albus Link. Stem X 18. G,
Lathyrus maritimus Bigel. Petiole x 13. H, Spartium junceum Linn. Stem x 18. I, Caragana arbore-
scens Lam. Young stem X 33.
 LEGUMINOSAEPAPILIONACEAE 513
Cork arising in very varying positions, even amongst different members of
a single genus.

(a) Arising in the epidermis in Abrus, Coursetia, Cytisus, Erinacea, Indigo-

fera (pro parte), Loddigesia, Mundulea, Myroxylon, Oxylobium, Platy-
osprion, Pseudarthria, Pycnospora, Robinia, Ulex, Uraria, Virgilia.
(b) Arising in the sub-epidermis or between this and the sixth cell layer in
species of Aeschynomene, Amorpha, Amphitaka, Argyrolobium, Baphia,
Buchenroedera, Calopogonium, Calycotome, Canavalia, Centrolobium,
Centrosema, Chadsia, Coelidium, Cordyla, Goronilla, Cranocarpus,
Cratylia, Cytisus, Dalbergia, Dalea, Desmodium, Dillwynia, Diphysa,
Eutaxia, Eysenhardtia, Gastrolobium, Geissaspis, Geoffraea, Goodia,
Hallia, Hovea, Hypocalyptus, Indigofera (pro parte), Laburnum, Lathrio-
gyna, Liparia, Lotononis, Phaseolus, Pictetia, Podalyria, Priestleya,
Priotropis, Pterocarpus, Pueraria, Pultenaea, Robinia, Sabinea, Smithia>
Soemmeringia, Sophora, Spartium, Stylosanthes, Thermopsis, Wistaria.
(c) Arising in the middle of the cortex in Anagyris, Atylosia, Camptosema,
Cleobulia, Clitoria, Cylista, Dalhousiea, Eriosema, Erythrina, Galactia,
Indigofera, Piptanthus, Psoralea, Rhynchosia, Rudolphia.
(d) Arising in the innermost layer of cortex in species of Cicer^ Colutea,
Dioclea, Galega, Glycine, Halimodendron (mucilaginous cork), Lotus,
Trifolium,
Arising in positions varying from the middle of the cortex to the endo-
(e)
dermis in species of Hippocrepis, Lathyrus, Medicago, Onobrychis,
Ononis, Orobus. *

(f) Arising internally to the groups of pericyclic fibres in Adenocarpus,

Adesmia, Aspalathus, Borbonia, Colutea, Cytisus, Halimodendron,
Lebeckia, Petteria, Sphaerophysa, Ulex, Viborgia.

Aerenchyma is formed instead of cork in Lotus uliginosus Schkuhr,

Sesbania aculeata Poir., and S. marginata Benth., and is stated to occur also
in Phaseolus when grown in water cultures.
Outer part of the primary cortex often collenchymatous and containing
stone cells. A
large part of the cortex, in species with reduced leaves, composed
of chlorenchyma, frequently exhibiting palisade structure and accompanied
by abundant stomata, the latter sometimes restricted to grooves. Species with
reduced leaves also frequently characterized by strands of fibres in the cortex
or attached to the vascular system. In the old stems the cortical fibres may
become exfoliated by a deep-seated phellogen, e.g. in the prominently ribbed
*
stem of Caragana arborescens Lam. See also Stems with special features'
on p. 516.
Pericycle containing either isolated strands of fibres or a composite and
continuous ring of sclerenchyma.

(a) Isolated strands of fibres, occasionally with stone cells between them,
stated to occur in most Hedysareae and in at least certain species of
Adenocarpus, Alysicarpus, Ammodendron, Amphithalea, Anagyris, Apios,
Arachu, Argyrolobium* Aspalathus, Astragalus, Baphia, Baptisia, Brongni-
artia, Buchenroedera, Calpumia, Calycotome, Caragana, Carmichelia,
Chadsia (pro parte), Chorizema, Cicer, Cladrastis, Coelidium, Corynella,
4594 L1
 5X 4 LEGUMINOSAEPAPILIONACEAE
Courestia, Crotalaria (sometimes), Cyclopia, Dalbergia, Dalea, Desmo-
dium (several species), Dichilus, Dillwynia, Diphaca, Diphysa, Dolichos
(pro parte), Drepanocarpus, Dumasia, Erinacea, Euchlora, Euchresta,
Eutaxia, Eysenhardtia, Gastrolobium, Genista, Geoffraea, Goodia, Hali-
modendron, Hecastophyllwn, Hypocalyptus, Indigofera, Laburnum,
Lathriogyna, Latrobea, Lebeckia, Lessertia, Loddigesia, Lotononis,
Lupinus, Machaerium, Melolobium, Mirbelia, Mundulea (pro parte),
Oxylobium, Petalostemon, Petteria, Piptanthus, Platymiscium, Platy-
podium, Podalyria, Poecilanthe, Priotropis, Psoralea, Pterocarpus,
Rafnia, Rhynchosia (pro parte), Rothia, Sabinea, Sesbania (pro parte),
Smithia, Sophora (fibre groups becoming crescent shaped and almost
y
continuous in 'S.flavescens Hort (Schilling 2032)), Spartium, Sphaero-
physa, Strongylodon, Tephrosia, Teramnus, Thermopsis, Tipuana, Ulex,
Viborgia, Vigna, Zornia.
(b) A ring of fibres present in Colutea (pro parte), Desmodium (pro
parte),
Galega, Glycyrrhiza, Harpalyce.
(c) A composite and continuous ring of sclerenchyma occurs in most of the
Phaseoleae and in Abrus, Aldina, Amorpha, Andira, Barbiera, Borbonia
(almost continuous), Bowdichia, Brachysema, Calophaca, Canavalia,
Centrolobium, Chadsia (pro parte), Cladrastis sinensis (bounded externally
by a sheath of cells containing large, solitary crystals), Colutea (pro
parte), Crotalaria (pro parte), Dalhousiea, Derris, Dioclea, Diplotropis,
Dipteryx, Dolichos (pro parte), Eriosema, Fagelia, Glycine, Hovea,
Leptodesmia, Liparia, Lonchocarpus, Millettia, Muellera, Mundulea (pro
parte), Myrospermum, Myroxylon, Olneya, Ormosia, Ougeinia, Piscidia,
Platylobium, Pongamia, Priestleya, Pterodon, Pultenaea, Rhynchosia (pro
parte), Robinia, Sesbania (pro parte), Swartzia, Sweetia, Virgilia,
Voandzeia, Wistaria, Xanthocercis, Zollernia.

Secondary phloem containing compact, sometimes stratified, strands of

the latter occasionally exhibiting a mucilaginous
fibres, layer. Chambered
fibres containing rhombohedral crystals or
styloids sometimes present as
well Medullary rays, where passing through the phloem, usually more than
4 cells wide, never sclerosed, broadened towards the exterior.
Vascular bundles in herbaceous species usually
individually distinct in
transverse sections, but exhibiting considerable differences in the nature of
the interfascicular tissue. Some of the arrangements are as follows.

(a) The groups of fibres belonging to each of the separate vascular bundles
are connected to one another by sclerenchymatous parenchyma in most
species of Trifolium.
(b) Separate bundles united by sclerenchymatous parenchyma situated on
the inside of the cambium in species of Coronilla,
Onobrychis, Orni-
thopus, Scorpiurus, Securigera.
(c) As (6), but prosenchymatous elements present amongst the sclerenchy-
matous parenchyma in species of Astragalus, Astrolobium, Biserrula,
Coronilla, Galega, Hippocrepis, Lotus, Medicago, Melilotus, Ononis,
Phaca, Tetragonolobus, Trigonella.
(d) Interfascicular tissue on the inside of the cambium wholly sclerenchy-
matous in Dorycnium.
 LEGUMINOSAEPAP1LIONACEAE 515

Xylem generally constituting a closed cylinder in woody genera and

species ; with or without distinct growth rings. Vessels frequently blocked
with a dense black secretion in Arachis hypogaea Linn, and Derris malaccensis
Prain.
Cortical vascular bundles present in species of Borbonia, Bossiaea,
Cytisus, Derris, Genista, Lathyrus, Mucuna, Orobus, Pimm, Retama, Strongy-
lodon, Viborgia, Vicia, especially in winged or grooved stems.
Secretory elements, (a) Tanniniferous cells, coloured brown in dried
material, and sometimes containing protein, mucilage, and other materials as
well, often in groups situated at the margin of the pith, in the primary or
secondary phloem, or, more rarely, in the primary cortex.
(i)
In the pith and phloem in species of Amorpha, Amphicarpaea, Apios,
Atylosia, Butea, Cajanus, Calopogonium, Camptosema, Canavalia,
Centrosema, Cleobulia, Clitoria, Cothlianthus, Cologania, Cratylia,
Cylista, Cymbosema, Desmodium, Dioclea, Dolichos, Dorycnium, Dun-
baria, Eriosema, Eysenhardtia, Fagelia, Flemingia, Galactia, Grona,
Hardenbergia, Hedysarum, Kennedya, Lonchocarpus, Mucuna, Ono-
brychis, Ornithopus, Periandra, Phaseolus, Platycyamus, Pueraria,
Robinia, Sesbania, Shuteria, Spatholobus, Teramnus, Tetragonolobw,
Vigna.
(ii) Secretory cells in the cortex, phloem, and pith in species of Dalbergia
(frequency varying considerably in different species), Derris,
Erythrina, Indigo/era, Lespedeza, Mundulea, Pterocarpus, Rhynchosia
(very numerous), Wistaria.
(iii) In the pith in Adesmia, Aeschynomene, Anthyllis, Arachis, Cladrastis,
Coronilla, Crotalaria, Desmodium, Diphaca, Hippocrepis, Leptodesmia,
Lespedeza, Lonchocarpus, Lotus, Nissolia, Ornithopus, Ougeinia,
Pseudarthria, Securigera, Zornia.
(iv) In the pith and primary cortex in Adesmia, Apios tuberosa Moench
(numerous, small, arranged in a compact zone below the epidermis),
Calophaca, Chadsia, Dolichos, Erythrina, Hosackia, Scorpiurus,
Stylosanthes.
(v) Accompanying the pericyclic fibres in Amorpha, Chapmannia, Des-
modium, Discolobium, Ebenus, Eversmannia, Eysenhardtia, Glycyrrhiza,
Hedysarum, Indigofera, Leptodesmia, Lessertia, Lotus, MiUettia, Mun-
dulea, Olneya, Onobrychis, Ornithopus, Petalostemon, Psoralea, Robinia,
Taverniera, Uraria.
(vi) In the primary cortex in Alhagi, Alysicarpus, Anthyllis barba-jovis
Linn, (especially large, arranged in longitudinal series and forming a
definite zone), Apios, Astragalus, Clitoria, Cordyla, Coronilla, Dalea,

Hedysarum,
(vii) In the sub-epidermis in Alhagi, Taverniera.
(viii)In the epidermis in Lathyrus pratense Linn.
(ix) In the primary phloem in Alhagi, Eversmannia, Hammatolobium,
Psoralea, Rhynchosia.
(x) In the cortex and phloem in species of Castanospermum and
modium.
Si6 LEGUMINOSAEPAPILIONACEAE
(xi) Distribution of tanniniferous cells in Glycyrrhiza glabra Linn,
observed to be somewhat distinctive as follows, (a) As an almost
continuous, compact layer of small cells in the epidermis and sub-
epidermis, (b) As
occasional scattered cells in the outer part of the
cortex, (c) As
longitudinal series of elongated cells in the phloem,
solitary or in groups of 3-6. (d) As scattered longitudinal series in the
pith.
Similar secretory cells, but devoid of tanniniferous contents and there-
(b)
fore not stained brown, occur in a number of genera.

Secretory cavities (see also 'Leaf') of various types such as those illu-
strated in Fig. 106 also occur notably in the Galageae, e.g. in the phloem and
pith of Robinia viscosa Vent. similar cavities also observed in the phloem and
;

pith of Fagelia sp., in the phloem of Canavalia sp., and in the cortex of Derris
malaccensis. Solereder records the existence of secretory cavities situated in
lumps on young branches of certain species of Dalea. Cells or lacunae con-
taining mucilage also recorded in the secondary cortex and/or pith of species
of Alhagi and Halimodendron.
Small secretory canals observed in the outer cortex and at the periphery
of the pith in Derris malaccensis; larger ones also occur in the outer part of the
cortex in Myroxylon balsamum var. pereirae. Secretory canals lined with
epithelium recorded by Holm (993) in Apios.
Solitary crystals observed at Kew in the cortex in species of Anthyllis,
Apios, Caragana, Castanospermum, Chorizema, Cladrastis, Dalbergia, Derris,
Desmodium, Erythrina, Galega, Glycine, Glycyrrhiza, Indigofera, Lathy rus,
Lespedeza, Lonchocarpus, Medicago, Myroxylon, Ononis, Oxytropis, Platyos-
prion, Psoralea, Robinia, Sophora, Trifolium, Vicia, Wistaria', in the phloem
in species of Anthyllis, Castanospermum, Cladrastis, Dalbergia, Desmodium,
Erythrina, Glycyrrhiza, Halimodendron, Platyosprion, Robinia, Sophora in the ;

pith in species of Castanospermum, Indigofera, Myroxylon, Platyosprion,

Robinia, Sophora, Wistaria. Cluster crystals observed only in the cortex of
Erythrina crista-galli Linn., and distinctive I-shaped crystals in the pith of
Robinia viscosa. Crystals of the same types as those described above under
'Leaf* (p. 510) also occur in the stem.

STEMS WITH SPECIAL FEATURES

Certain species, especially but not exclusively amongst the xeromorphic
members of the family, exhibit a considerable range of anatomical peculiarities,
the precise nature of which cannot be fully described here. The following
particulars concerning a selection of interesting species gives some indication
of the unusual types of stem structure which have been observed by Dr. C. L.
Hare in material grown at Kew.

(i) ASTRAGALUS AMBIGUUS BUNGE.

Xylem in young stems very loose in texture, divided into sectors by broad
medullary rays. Vessels small, very thick- walled, solitary or in short tangential
clusters, with reticulate thickening. Ground-mass of the wood composed
mostly of thin-walled parenchyma, interspersed with small patches of fibres.
 LEGUMINOSAEPAPILIONACEAE 517

(ii) CARAGANA ARBORESCENS LAM. AND C. MICROPHYLLA LAM.

Young stems prominently ribbed, with strands of fibres in the angles
between the ribs.Ribs and fibres later becoming exfoliated owing to the
development of a broad band of cork in the cortex, Xylem very close in tex-
ture; vessels in uniseriate tangential bands embedded in parenchymatous
ground tissue rays very narrow.
;

(iii)CARMICHAELIA AUSTRALIS R. BR.

Young stem constituting a typical phylloclade. Stomata numerous, arranged
transversely to the longitudinal axis of the stem. Cortex consisting of alternat-
ing radial bands of sclerenchyma and collenchyma. Stems becoming circular
when older, through the exfoliation of the outer tissues following the develop-
ment of deep-seated cork. Xylem in the form of wedge-shaped masses,
separated by broad medullary rays. Vessels mostly in tangentially extended
clusters; walls with delicate spiral (tertiary) thickening.

(iv) CYTISUS, GENISTA, ULEX, AND RELATED GENERA

The xeromorphic assimilatory stems of different species of Cytisus and
related genera can often be distinguished by variations in the arrangement of
the ribs, sclerenchyma, assimilatory tissue, as well as by differences in the
occurrence and distribution of hairs. As an example of the type of structure
in Cytisus the following particulars of C. albus Link, are given. Exterior of the
stem with 18-20 prominent ribs, the distal ends of which are tangentially
expanded as seen in transverse sections. Grooves between the ribs lined with
hairs. Palisade chlorenchyma forming an almost continuous zone except
where interrupted by wedge-shaped strands of thick- walled fibres within each
of the ridges. Cytisus scoparius (L.) Link, similar, but provided with only
5 narrow ridges, hairs being absent from the grooves between them. Genista
cinerea DC. similar to Cytisus albus, but with cortical vascular bundles in some
of the ridges. Ribs less pronounced and grooves more shallow in Genista
hispanica Linn, than in G. cinerea. Stomata of G. hispanica raised, with a
characteristic pear-shaped vestibule almost closed by cuticular ridges.
Chlorenchyma in the same species not arranged as a palisade. Young stems
of Ulex europeus Linn, with 8-10 ribs separated by narrow grooves. Stem
clothed with twisted, uniseriate hairs. Each stomata with its long axis parallel
to that of the stem. Cortical vascular bundles absent from the ridges.
The anatomy of the spines of Ulex europeus has been described by Skipper
(2119), from whose paper the following details are taken. Pith consisting of
thick- walled cells, surrounded by a ring of vascular bundles. Cortex grooved,
the grooves being equal in number to but alternating with the bundles, Ridges
between the grooves supported by a strip of sclerenchyma, consisting of long,
thick-walled fibres with simple pits. Ridges flanked by palisade-like chloren-
chyma. Stomata mostly confined to the region of the palisade cells.

(v) LATHYRUS AND VICIA

Stem of Lathyrus maritimus Bigel flattened on one side and provided with
2 wings each containing a vascular bundle. Remaining vascular bundles
widely spaced and arranged in a flattened circle around the hollow pith.
Secretory tissue absent. Structure similar in L. pratensis Linn,, but abundant
518 LEGUMINOSAEPAPILIONACEAE
secretory cells present in the epidermis. Wings in L. sylvestris Linn, much
longer than in the above 2 species, with several vascular bundles in each wing.
The 2 wings of Vicia cracca Linn, each contain a single bundle.
According to Parsa (1657) the stem of Lathyrus szowitsii Boiss., a species
endemic to Iran, is provided with exceptionally long wings with swollen
distal ends. Each wing contains several small vascular bundles, whilst the
swollen ends are supported by well-developed strands of sclerenchyma.
Apart from the preceding records and observations, the assimilatory stems
of various members of the Papilionaceae have also been the object of several
previous investigations, Cytisus, Genista, Lebeckia, and Spartium having been
most thoroughly examined. Some of the earlier work has been summarized
by Solereder, but more recent investigations include those by Pellegrin (1682),
Jolivet (1187), an d Taylor (2239). Less extensive observations have also been
recorded by Goodlatte (795), Buchegger (303), Gravis (804), Evenari (Schwarz)
(665). Pellegrin found the distribution of cortical bundles, the nature and
arrangement of the ribs, and the nodal anatomy to be taxonomically important,
but it was seldom possible to distinguish species by means of the stem struc-
ture alone. Jolivet likewise found the genera to be more or less clearly defined,
but was unable to separate species with precision. Taylor, working mainly
with Cytisus and Genista, found that the species could be roughly grouped by
the appearance of the stem in transverse section. He divided them into five
classes based on the number and shape of the ribs and variations in the
distribution of fibres, cortical bundles, and assimilatory tissue. It is note-
worthy that the earlier French work was apparently unknown to Taylor when
his researches were in progress.
Lacoste (1312) has described the phyllode of Phylloxylon as having numer-
ous stomata in the epidermis, palisade chlorenchyma beneath both surfaces,
while the vascular bundles are widely spaced and arranged in a flattened ring.
The centre is occupied by a pith-like tissue.
The stems of 'weeping' varieties of Sophora japonica Linn, and Caragana
arborescens Lam. were found by Low (1397) to have fewer phloem fibres and less
fully developed prosenchymatous elements in the xylern than the erect forms
of the same species. The difference was first apparent during the second
year's growth,
X Laburnocytisus adamii Schneid. (syn. Laburnum adamii Schneid.) is
generally regarded as a periclinal chimaera in which the skin of tissue derived
from Cytisuspurpure a Vent, covers more deeply seated tissue derived homLabur-
num anagyroides Med. (syn. L. vulgare Bercht. et Prsl). Anatomical evidence,
which he believedto support this interpretation, was obtained by Buder (305).
Since the epidermal cells of Cytisus purpurea are richer in tannin than those
of Laburnum anagyroides the presence of tannin could be used for the detec-
tion of 'purpurea' cells in the graft hybrid. When sections of the leaves or
petiole were placed in a solution of potassium bichromate the dark-brown
9

'purpurea epidermis was clearly visible to the naked eye, and contrasted strongly
*
with the uncoloured vulgare' tissues beneath. In older stems the mode of cork
formation also helped to distinguish the tissues of the two parents. In C.
purpurea cork arises in the epidermis, in L. anagyroides its origin is more deep-
seated. In L. adamii cork originates in both positions, but cork formation
does not persist for so long in the 'purpurea' epidermis as in the more deep-
 LEGUMINOSAEPAPILIONACEAE 519
seated 'vulgare* tissues, so that eventually the outer 'purpurea* tissues are cast
off and an old stem is thus derived wholly from L. anagyroides. Buder's

thorough anatomical investigation failed to reveal any facts which are con-
trary to the concept that L. adamii is a periclinal chimaera.
According to Holm (993) the tuber at the base of the stem of Apios tuberosa
Moench. includes numerous small vascular strands in the pith, each contain-
ing secretory canals of the same type as those in other parts of the plant.

WOOD (Figs. 115, 1 1 6, and 117)

Vessels medium-sized (100-200 p mean tangential diameter) in most
genera; very small (less than 50/1) in Anagyris, Anthyllis, Apoplanesia,
Ateleia, Behaimia, Belairia, Benthamantha, Brya, Calycotome, Cytisus,
Drepanocarpus, Halimodendron, Lespedeza, Medicago, and Poecilanthe moder- ;

ately small (50-100 IJL) in Aeschynomene, Desmodium, Indigo/era, Lupinus,

Piptanthus, Piscidia, Podalyria, Sabinea, Schefflerodendron, Spartium, Swartzia
p.p., and Zollernia; large (more than 200 /x) in some species of Andira,
Bowdichia, Cashalia, Castanospermum, Clathrotropis, Dalbergia, Dussia,
Erythrina, Hymenolobium, Kunstleria, Millettia, Monopteryx, Ostryocarpus,
Platycyamus, Platymiscium, and Pongamia\ very large in some of the lianes,
e.g. reaching a diameter of 600 p in species ofEntada, according to Solereder ;

occasionally of 2 distinct sizes, e.g. in Butea superba Roxb. the larger vessels
are up to 425 (JL in diameter and the smaller not more than 175 /z; in other
species, particularly those with an oblique or tangential pattern, the larger
vessels may themselves be small and be set in a matrix of extremely small,
angular storied vessels that are comparable with tracheids in cross-section,
e.g. in Diocledy Halimodendron, Spartium (Fig. 116 D), Ulex, and Wistaria] the
tendency for the vessels to be mostly solitary, but with a few multiples and
clusters, noted as characteristic of many of the genera of the Mimosaceae and
Caesalpiniaceae, is not particularly noticeable in the Papilionaceae; in woods
without any definite radial pattern radial multiples are usually moderately
abundant, but do not commonly exceed 3 cells except in some species of
Belairia, Dalbergia, Medicago and Pterocarpus\ irregular clusters are common
y

in some species of Coursetia, Crotalaria, Dalbergia, Dalea, Daviesia,

Platymiscium, Pterocarpus, Pueraria, Retama (1493), and Sophora; with an
oblique pattern in some species of Anthyllis Calycotome, Cytisus, Dalea,
y

Genista, Laburnum^ Parosela, Pickeringia, Sophora, Spartium (Fig. 1160), and

Ulex, and with a tangential pattern in Anagyris, Halimodendron, Lupinus,
Piptanthus, Podalyria, Robinia, and Virgilia; clusters common in nearly all
these woods with a distinct vessel pattern; apart from the woods with very
numerous vessels in oblique or tangential rows, most species have either
fewer than 5 vessels per sq. mm. or between 20 and 40 per sq. mm. fewer than
;

5 per mm. in species with large vessels and also in Aeschynomene, Afrormosia,
Alexa, Amerimnon, Andira, Baphia, Butea, Canavalia, Cashalia, Couma-
rouna, Crotalaria, Derris, Dipteryx, Fordia, Inocarpus, Machaerium, Millettia,
Muellera, Ormosia, Ougeinia, Piscidia, Poecilanthe, Pongamia, Pterodon,
Tipuana, Torresia, and Zollernia; 5-20 per sq. mm. in Desmodium (Janssonius
1154), Haplormosia, Pericopsis, Pueraria, Rothia, Schefflerodendron, Sophora
p.p., Spatholobus, Swartzia, Sweetia, and Wistaria] more than 40 per
mm. in Brya, Eysenhardtia, and Medicago; ring-porous or semi-ring-porous
520 LEGUMINOSAEPAPILIONACEAE
in at leastsome species of Ammodendron, Amorpha, Anthyllis, Caiycotome,
Cascaronia (1864), Cladrastis, Cytisus, Dalbergia, Dalea, Edwardsia (1886),
Eysenhardtia, Genista, Gourliea, Halimodendron, Indigofera (1851), Laburnum,
Lespedeza, Parosela, Pickeringia, Podalyria, Pterocarpus, Robinia, Sophora,

FIG. 115. LEGUMINOSAEPAPILIONACEAE

A, Pericopsis mooniana Thw. B, Millettia pendula Benth, C, Afrormoria laxiflora Harms. D,
Erythrina stricta Roxb. E, Butea superba Roxb.

Spartium, Ulex, Virgilia, and Wistaria (1851); spiral thickening, often limited
to the smaller vessels, observed or reported in Amphithalea (1851), Anagyris,
Argyrolobium (1851), Calycotome, Coelidium (1851), Cytisus, Erinacea (1851),
Genista, Halimodendron, Laburnum, Lathiogyna (1851), Lebeckia (1851),
Liparia (1851), Lotononis (1851), Petteria (1851), Pickeringia, Platylobium
(1851), Podalyria, Priesthya (1851), Robinia, Sophora, Spartium, Ulex,
and
Wistaria; Solereder refers also to spirally thickened 'tracheids or vessels of
 LEGUMINOSAEPAPILIONACEAE 521
narrow lumen* in species of Adenocarpw, Ammodendron, Anthyllis, Car-
agana, Carmichaelia, Cladrastis, Colutea, Coronilla, Cyclopia, Dorycnium,
Edwardsia, Lotus, and Sweetia. Perforations simple. Intervascular pitting
alternate, typically small;minute in Brya, Dipteryx, Lespedeza, Myrocarpus,
Piscidia, Schefflerodendron, Toluifera, and Zollernia\ pits moderately large
in a few species, e.g. Butea superba Roxb., Centrolobium paraeme Tul.,
Drepanocarpus inundatus Mart., Pericopsis mooniana Thw., and Spatholobus
roxburghii Benth. and large in Hymenolobium and Kunstleria occasionally with
;

conspicuous striations due to coalescent apertures, e.g. in Rothia sp. and

Dalea spinosa Gray; pits to parenchyma and ray cells usually similar to the
intervascular pits, but sometimes either simple or subtending larger paren-
chyma pits in Gliricidia, Hebestigma, Hymenolobium, Kunstleria, Pericopsis,
Piscidia, Platymiscium p.p., Robinia, Sabinta, and Spatholobus y and, according
to Solereder, in Brya', sometimes elongated in Dioclea', pits vestured. Solid

deposits common, tyloses abundant in some species of Amorpha, Coursetia,

Dalbergia, Gliricidia (1886), Hebestigma, Lennea, Machaerium, Notodon (1886),
Olneya (1886), Robinia, and Sabinea\ those of Sabinea florida DC. containing
crystals. Mean member length o- 1-0-4 mm. Parenchyma usually moderately
to very abundant and either predominantly paratracheal or in moderately
regular bands that tend to be replaced by definitely paratracheal forms where
the parenchyma is less abundant round or diamond-shaped sheaths, such as
;

are characteristic of the Mimosaceae and Caesalpiniaceae, occur in some

genera but are less common than the confluent or other banded types. Very
abundant, in broad, moderately regular bands 4 or more cells wide in some
species of Bergeronia, Butea, Clitoria, Desmodium p.p. (1154), Erythrina,
Fordia, Inocarpus, Lonchocarpus, Millettia (Fig. 1156), Muellera, Piscidia,
Pongamia, and Spatholobus; in narrower bands in Alexa p.p., Baphia (Fig.
1 16
j), Brya, Dalbergia p.p., Drepanocarpus, Geoffroea, Haplormosia, Machae-
rium, Platypodium, Pterocarpus p.p., Schefflerodendron, Swartzia p.p., and
Zollernia\ confluent, forming irregular bands or the matrix for oblique or
tangential bands of vessels, in species of Afrormosia (Fig. 1150), Alexa p.p.,
Anagyris, Andira, Anthyllis, Belairia, Bergeronia, Calycotome, Canavalia,
Cashalia, Clathrotropis, Coursetia, Cytisus, Dalbergia, Dalea, Derris, Dussia,
Ecastophyllum, Erythrina, Etaballia, Genista, Gliricidia, Gourliea, Halimo-
dendron, Hebestigma, Hymenolobium, Ichthyomethia, Laburnum, Lonchocarpus,
Lupinus, Ostryocarpus, Parosela, Pickeringia, Piptanthus, Piscidia, Platycyamus,
Podalyria, Poecilanthe, Pongamia, Pterocarpus, Retama (1493), Sabinea,
Spartium, Swartzia p.p., and Ulex intermediate between aliform and con-
m

fluent in Dalbergia p.p., Desmodium p.p., Diphysa, Indigofera, Inocarpus

(1154), Medicago, Paramachaerium, Pericopsis, Platymiscium, Pterocarpus p.p.,
and Tipuana\ aliform, sometimes confluent at the end of the growth ring, in
Amherstia, Ateleia, Bowdichia, Castanospermum, Coumarouna, Diplotropis,
Harpalyce, Kunstleria, Lespedeza, Monopteryx, Myrospermum, Ormosia,
Ougeinia, Sophora p.p., and Torresia\ in predominantly rounded sheaths
about the vessels (vasicentric) in Amorpha, Apoplanesia, Behaimia, Brya,
Cladrastis (confluent in outer part of ring), Crotalaria, Dalea, Dioclea,
Eysenhardtia, Indigofera, Myrocarpus, Myroxylon, Rhynchosia, Robinia (con-
fluent in outer part of ring), Rothia, Sesbania, Sophora, Toluifera, and Virgilia;
in a narrow layer on the abaxial sides of the vessels in Centrolobium and
5*2 LEGUMINOSAEPAPILIONACEAE
Sweetia\ scattered strands (diffuse), which are sometimes crystalliferous,
present in Amerimnon, Andira, Bowdichia, Brya (very conspicuous),

FIG. 116. LEGUMINOSAEPAPILIONACEAE

A, Andira inermis H. B. et K. B,. Pterocarpus stevensonii Burtt Davy. C, Millettia atropurpurea Benth.
D, Spartium junceum Linn. E, Datbergia nigra Allem. F, Cladrastis amurenns Benth. G, Sarothamnus
scoparius (L.) Koch. H f Dalbergia melanoxylon Guill. I, Laburnum anagyroides Med. J, Baphia nitida
Lodd.

Centrolobiwn, Cladrastis, Dalbergia (a few species only), Diphysa, Genista,

Machaerium, Myrocarpus, Pterocarpus (a few species), Tolutfera, and Ulex\ in
 LEGUMINOSAEPAPILIONACEAE 523
Pueraria there are both vasicentric sheaths (paratracheal) and short,
apparently apotracheal bands; in Wistaria the parenchyma mixed with very
small vessels tends to form the ground tissue and in Aeschynomene parenchyma
constitutes the greater part of the tissues and forms a ground-mass in which
groups of fibres and vessels appear on cross-section as aliform patches
(Fig. 1 17 A); terminal parenchyma present in some genera. Chambered
crystals present in the marginal cells of the paratracheal or banded parenchyma
of most of the species but not observed in Alexa, Ammodendron, Amorpha,
Anagyris, Apoplanesia, Bowdichia, Calycotome, Canavalia, Cladrastis, Crota-
laria, Cytisus, Dalea, Daviesia, Diphysa, Dussia, Eysenhardtia, Genista,
Halimodendron, Hebestigma, Kunstleria, Laburnum, Lespedeza, Lupinus,
Medicago, Monopteryx, Paramachaerium, Parosela, Pericopsis, Pickeringia,
Piptanthus, Podalyria, Poecilanthe, Pueraria, Rothia, Sabinea, Scheffleroden-
dron, Sesbania, Sophora, Spartium, Torresia, Ulex, Virgilia, and Vouacapoua\
chambered crystalssometimes occurring also in diffuse parenchyma,
where this ispresent (see p. 524); the maximum number of crystals per
strand typically about 8 and not more than 1 1, but 12 or more in some genera,
particularly those in which the ordinary strands are commonly of 4 cells and
also in some woods with strands of not more than 2 cells, e.g. Bergeronia,

Brya, Muellera, and Zollerma. In Aeschynomene elaphroxylon Guill. et Perr.

the crystals are usually limited to a layer of cells adjoining the adaxial side
of the fibre groups and the chambered strands are sometimes locally biseriate.
Cells sometimes containing gum-like deposits. Record and Hess (1886) note
small patches of sclerotic cells in Clathrotropis. Strands most commonly of
1-2, or only i cell, but of 2-4 cells in a number of genera, e.g. Afrormosia,
Alexa, Andira, Bowdichia, Centrolobium, Diplotropis, Dipteryx, Haplormosia,
Hymenolobium, Lonchocarpus, Lupinus, Millettia, Myrocarpus, Myroxylon,
Ormosia, Pericopsis, Piscidia, Platymiscium p.p., Pterocarpus p.p., Robinia,
Rothia, Schefflerodendron, Sesbania, Spatholobus, Swartzia, Sweetia, Torresia,
Virgilia, and Wistaria; strands of more than 4 cells present in occasional
genera, e.g. Kunstleria, Monopteryx, and Pueraria fusiform parenchyma cells
;

common in Anagyris, Baphia, Butea p.p., Cladrastis, Dalbergia, Desmodium

(1154), Diphysa, Erythrina, Geoffraea, Gourliea, Laburnum, Paramachaerium,
Parosela, Platymiscium p. p., Podalyria, Pterocarpus p.p., Sophora, and Torresia,
and with the parenchyma cells almost exclusively fusiform in Aeschynomene,
Calycotome, Cytisus, Genista, Indigofera, Spartium, and Ulex. Strands
typically distinctly storied, but the stories rather vague in or absent from Alexa,
Amherstia, Amorpha, Anthyllis, Apoplanesia, Bowdichia, Butea p.p.,
Cashalia, Centrolobium p.p., Crotalaria, Diplotropis, Harpalyce, Hebe"
stigma, Hymenolobium, Lespedeza, Monopteryx, Ormosia, Pericopsis, Piscidia,
Platymiscium p.p., Pongamia, Pterygopodium, Pueraria, Rothia, Sabinea,
Schefflerodendron, Sesbania, Spatholobus, Swartzia, Torresia, Vouacapoua,
Virgilia, Wistaria, and Zollernia. Many of the woods with strands pre-
dominantly of 2 cells exhibit secondary storying of the parenchyma owing to
the regular position of the central cross-wall. Distinctly storied strands have
obtuse-angled gable-ends on tangential section and the pitting is sometimes
markedly more abundant on these end walls, e.g. in Aeschynomene elaphroxy-
lon. This species was investigated in great detail by Beijer (168) in his work on
cell division in the cambium leading to storied structure. Janssonius (1154)
 524 LEGUMINOSAEPAPILIONACEAE
uses the presence of these two characters as a means of distinguishing the
Papilionaceae from the Mimosaceae and Caesalpiniaceae, but there appear to
be many exceptions. According to Solereder, the wood parenchyma in some
of the lianes is thin-walled and unlignified, especially in the later-formed
secondary wood, e.g. in species of Dioclea, Mucuna, Phaseolus, and Pueraria',
axialwood of denser structure occurs in the neighbourhood of the pith. Rays
of more than half the genera 2-3 cells wide exclusively uniseriate or with only
;

occasional biseriate rays in some species of Canavalia, Centrolobium, Couma-

rouna, Dipteryx, Drepanocarpus, Etaballia, Geoffraea, Gourliea, Inocarpus,
Kunstleria, Machaerium, Myroxylon (2430), Paramachaerium, Piscidia,
Platymiscium, Platy'podium Pterocarpus, Spartium, Swart zia p.p., and
,

Tipuana\ 4-10 cells wide in Afrormosia, Ammodendron, Anagyris, Andira,

Anthyllis, Baphia, Benthamantha, Butea, Cladrastis, Clitoria, Cytisus, Dalea,
Desmodium, Dioclea, Erythrina, Halimodendron, Ichthyomethia, Indigofera,
Laburnum, Lonchocarpus, Lupinus, Medicago, Millettia, Monopteryx, Parosela,
Piptanthus, Platy cyamus, Retama (1493), Robinia, Sophora, Spartium,
Spatholobus, and Ulex\ sometimes more than 10 cells wide in the larger rays
of Erythrina, Laburnum and in the inter fascicular rays of Pueraria and
,

Wistaria; with some very large rays, 20-30 cells wide in Aeschynomene and
Monopteryx and, according to Solereder, in some species of Sarothamnus\
of 2 distinct sizes, the small rays storied, in Aeschynomene, Daviesia, and
Millettia albiflora Prain, and, without storying of the small rays, in Mono-
pteryx up to more than i mm. in height in Aeschynomene, Andira, Baphia,
;

Clathrotropis, Clitoria, Coursetia, Erythrina, Fordia, Hebestigma, Laburnum,

Lennea, Lupinus, Medicago, Millettia p.p., Monopteryx, Ostryocarpus, Parosela,
Pickeringia, Piptanthus (small stem), Pueraria (interfascicular), Sophora,
Spatholobus, and Wistaria (interfascicular) woods with multiscriate rays
;

usually with few uniseriates, e.g. Afrormosia, Andira, Baphia, Bergeronia,

Butea, Cladrastis, Dalbergia p.p., Drepanocarpus, Erythrina p.p., Halimoden-
dron, Haplormosia, Hymenolobium, Laburnum, Lonchocarpus, Medicago,
Monopteryx, Muellera, Myrocarpus, Myroxylon, Ougeneia, Pericopsis, Platy-
cyamuSj Podalyria, Pongamia p.p., Robinia, Schefflerodendron, Sophora p.p.,
Spartium, Spatholobus, Torresia, and Vouacapoua\ mostly from 4 to
12 rays per mm. more than 12 per mm. in some species of Amorpha, Belairia,
;

Bergeronia, Brya, Calycotome, Dalbergia, Dalea, Dipteryx, Drepanocarpus,

Ecastophyllum, Etaballia, Geoffraea, Gourliea, Hebestigma, Inocarpus, Machae-
rium, Parosela, Platypodium, Podalyria, Pterocarpus, Sabinea, and Sivartzia\
with fewer than 4 rays per mm. in Bowdichia, Erythrina, Halimodendron, and
Pueraria', homogeneous (Kribs's Types 1, II, and III) in Alexa, Anagyris,
Andira, Anthyllis, Baphia, Benthamantha, Bergeronia, Brya, Butea, Cashalia,
Centrolobium, Clathrotropis, Coumarouna, Daviesia, Derris, Desmodium,
Dipteryx, Dussia, Erythrina p.p., Etaballia, Fordia, Geoffraea, Gliricidia,
Halimodendron, Haplormosia, Ichthyomethia, Inocarpus, Lennea, Lonchocarpus,
Machaerium, Millettia, Muellera, Ostryocarpus, P&ramachaerium, Pericopsis,
Piscidia,Platycyamus, Platymisdum, Platypodium, Pongamia, Pterocarpus,
Robinia, Rothia, Schefflerodendron, Sophora, Spartium, Swartzia, and Ulex\
moderately heterogeneous (Kribs's Type II and occasionally III) with
1-2 marginal rows of square or upright cells in most of the other genera, but
with 4 or more such rows in Hebestigma, Indigofera, Kunstleria, and Lupinus \
 LEGUMINOSAEPAPILIONACEAE 525
cells tending to be all square (radial section) in Medicago and Parosela\
procumbent cells small in tangential diameter (less than 10 ju,) in some genera,

e.g. Anthyllis, Cladrastis, Cytisus, Desmodium, Diplotropis p.p., Genista,

Halimodendron, Indigo/era, Laburnum, Machaerium, Myrocarpus, Myroxylon,
Pericopsis, Piscidia, Pongamia, Robinia, Schefflerodendron, and Sweetia sheath
;

cells present in some

species of Butea, Clitoria, Cytisus, Dioclea, Erythrina,
Halimodendron, Laburnum, Lupinus, and Spartium', commonly containing
gum-like deposits; crystals observed in only a few species, sometimes
abundant, e.g. in Parosela and Zollernia. Usually storied in woods with low
rays, but in most of the genera the rays are several times as high as the
parenchyma strands and exhibit no storying; distinctly storied in some species
of Aeschynomene, Amorpha, Cascaronia (576), Castanospermum, Centrolobium,
Cladrastis, Dalbergia, Dalea, Drepanocarpus, Laburnum, Machaerium, Para-
machaerium, Pericopsis, Pickeringia, Pterocarpus, Robinia, Schefflerodendron,
Sophora, Swartzia, Sweetia, and Zollernia,
The rays of a few genera exhibit some unusual features. Solereder refers
to the occurrence of spirally thickened tracheids in the secondary rays of
Cytisus ardoini Fourn. and C. sauzeanus Burn, et Briqu. the large rays of
;

Monopteryx uacu Spruce include horizontal strands of thick* walled fibres;

in Aeschynomene elaphroxylon most of the rays are uniseriate and storied, but
in old material there are also broader rays containing vascular strands, which
sometimes include 30 or more vessels, arranged, as seen in tangential section,
in multiples and clusters of 2-6 cells; these vessels have simple perforations
and numerous, rather large intervascular, pits, which in some cases are
elongated to an almost scalariform type Solereder refers to smaller, spirally
;

thickened vessels in the centre; he quotes Klebahn as having shown that the
vascular strands correspond to the vascular part of the rudiments of roots
formed between the broad rays and lenticels on the surface of the stem.
Fibres typically with few, small, simple pits, more numerous on the radial
than on the tangential walls pits numerous in a few species and occasionally
;

moderately large, e.g. in Alexa and Swartzia', with conspicuous bordered pits
in Kunstieria. Very occasionally septate in Ougeneia, Robinia, Sophora, and
Tipuana. In the majority of genera walls moderately to very thick, but
occasionally very thin- walled, e.g. Lespedeza, Rothia, and Virgilia; often with
a gelatinous inner layer. In a few woods the fibres form small islands on the
cross-section owing to the abundance of wood and ray parenchyma, e.g. in
Aeschynomene (Fig. 117 A), Butea (Fig. 115 E), Clitoria, Pueraria, Spatholobus,
and Wistaria; in Aeschynomene elaphroxylon the fibres form aliform patches
about the vessels, similar in shape and distribution to aliform parenchyma.
Mean length 0-6-1-7 mm. Intercellular canals of the vertical traumatic
type observed or reported (1886) scattered throughout the ring in Andira
and Humboldtiella. Included (interxylary) phloem, see under 'Anomalous
Structure* below.

ANOMALOUS STRUCTURE
Anomalous structure, consisting of successive layers of xylem and phloem
repeating the primary structure of the stem, separated by tangential bands
of conjunctive parenchyma, observed in Machaerium, Pueraria, and Wistaria
and reported by Solereder in Derris, Mucuna, Pachyrhizus, Rhynchosia,
 526 LEGUMINOSAEPAPILIONACEAE
Spatholobus, and Strongylodon\ of the 'disperse' type, the stem cleft into
irregular strands by dilation of the parenchymatous elements, in Kunstleria
ridleyi Prain. The formation of successive rings of xylem and phloem in the
primary cortex, pericycle, or, more rarely, in the phloem also recorded in

FIG. 117. LEGUMINOSAEPAPILIONACEAE

Wood of Aeschynomene sp. A, Transverse section through the wood. B, Part of a transverse section
showing the terminal surfaces of the transversely cut fibrous cells, which have wide lumina. C, Radial
longitudinal section. D, Tangential section. By Solereder.

species of Pueraria and Wistaria by Handa (88,2 A). Interxylary phloem

recorded in species of Dioclea, Mucuna> Phaseolus, and, according to Russell
(1793), in Mittettia. Stems of lianes mostly cylindrical, but band-shaped in
Machaerium and Rhynchosia phaseoloides DC. A method whereby the xylem
becomes split up by the development from the cambium of a contractile tissue
has been described by Ziegenspeck (2506).
 LEGUMINOSAE PAPILIONACEAE 527
ROOT
The
roots of the Papilionaceae are interesting mainly on account of the
almost universal occurrence on them of nodules or tubercles containing
bacteria capable of fixing atmospheric nitrogen. For this reason leguminous
crops are commonly grown as a green manure* A considerable volume of
literature concerning this interesting subject exists, but is outside the scope
of this book. In woody species such as Wistaria sinensis Sweet, the nodules,
according to Jimbo (i 181), consist of a chain of 1-4 swellings. Small lamellae
termed palettes' have been described by Severini (2080) on the roots of
Hedysarum coronarium Linn. One surface of these bodies becomes covered
with absorbing hairs, whilst the other is flat, or at most provided with a few
small papillae. The 'palettes' arise in the pericycle, and their number is con-
trolled by the nature of the soil. They are quite distinct from root-nodules,
and become calcified by the deposition of calcium carbonate within the living
cells. According to Feh&r (68 1) tannin occurs in smaller quantities in the

young root of Robinia pseudacacia Linn, than in the aerial part of the plant,
but short tanniniferous tubes are present beneath the bark. An oily substance
occurs in young and old roots as well as in the tubercles of the same species,
and tyloses develop in the xylem vessels in the third year. For further informa-
tion concerning roots of the Papilionaceae see: Bond (226), Dastur and
Saxton (542), Holm (993), Lindemuth (1373), Smith and Kersten (2149),
Staber (2178), Ward (2360).

TAXONOMIC AND PHYLOGENETIC NOTES

(FOR THE LEGUMINOSAE IN GENERAL)
(i)
FROM GENERAL ANATOMY
Dormer's (600, 601, 602) recent investigations, which refer mainly to the
Leguminosae, lead him to conclude that phyllotaxy and the course of the
primary vascular system provide characters of taxonomic and phylogenetic
interest. He expresses the view that the 'evolution of a typical herbaceous
habit becomes possible only when the tangential continuity of the vascular
system is maintained by nodal anastomoses between the bundles, so that there
is no necessity for the formation of a continuous cylinder of secondary
tissues'. These interesting views will need to be substantiated by further

investigations on a much wide/ range of material before their significance can

be fully assessed. This will undoubtedly take time, because the technique
involved does not permit large quantities of material to be examined very
rapidly. Still more recently Dormer (605) has extended his line of inquiry by
examining the phylogenetic significance of the pulvinus in the Papilionatae,
the pulvinate tribes and genera being regarded as more primitive than those
which are epulvinate.

(ii) FROM WOOD STRUCTURE

The wood anatomy in generalsuggests that the Mimosaceae are the least
highly specialized and the Papilionaceae the most highly specialized of the
three Leguminous families. For example, the following characters in the
Papilionaceae indicate a higher degree of specialization than in the Caesalpini-
aceae or the Mimosaceae: (a) very short cambial initials, with a consequent
5*8 LEGUMINOSAE PAPILIONACEAE
higher proportion of woods with storied parenchyma and numerous fusiform
cells, (b) vessel characters, e.g. the more common occurrence of ring-porous-
ness, pronounced patterns, and spiral thickening, and (c) the more common
occurrence of anomalous structure. The Mimosaceae, on the other hand,
appears at first sight to have the most highly specialized rays of the three
groups, in that the rays are always homogeneous and are typically composed
of small cells and in that uniseriates are typically few. The position may,
perhaps, be better expressed, however, by saying that, though the Mimosaceae
have achieved a more uniformly high level of ray development, the Caes-
alpiniaceae, and more especially the Papilionaceae, include both more highly
specialized rays as well as less highly specialized types. The proportion of
genera with septate fibres shows a suggestive decrease from the Mimosaceae
to the Papilionaceae, with the Caesalpiniaceae intermediate, but the signifi-
cance of this feature is not clear.
Finally it may be noted that there is less difference in general structure and
in degree of specialization between the Mimosaceae and the Caesalpiniaceae
than between the latter and the Papilionaceae, The principal features that
tend to distinguish the three families are given below.
Normand (1615) states that in its wood anatomy Ibadja walkeri A. Chev.
closely resembles Loesenera kalantha Harms and differs from both Berlinia and
Hymenostegia.
Hess (960) considers the wood anatomy supports the view that Daubentonia
texana Pierce and D. punicea (Cav.) DC. may be separate genera and that
Senegalia angustifolia (Lam.) Britt. et Rose should be separated from Senegalia
and made into a monotypic genus.
Points of Difference between Mimosaceae, Caesalpiniaceae^ and Papilionaceae
Vessels. Oblique or tangential pattern never more than vague in Mimo-
saceae, occasionally distinct in the Caesalpiniaceae, and very distinct in several
Papilionaceae. Spiral thickening absent from Mimosaceae, more common
in Papilionaceae than in Caesalpiniaceae. More commonly ring-porous in
Papilionaceae. Gummydeposits common in all three groups but most pro-
nounced in the Caesalpiniaceae.
Parenchyma. Round or diamond-shaped paratracheal typical of the
Mimosaceae and Caesalpiniaceae and occurring in some Papilionaceae;
diffuse, usually crystalliferous strands very common in Mimosaceae, common
in Caesalpiniaceae, relatively rare in the Papilionaceae; terminal most pro-
nounced in the Caesalpiniaceae, present in the others. Strands most commonly
of 2 or 4 cells in Mimosaceae and Caesalpiniaceae, of 1-2 cells in the
Papilionaceae. Fusiform cells commonin a few genera of Mimosaceae and
Caesalpiniaceae, common and sometimes predominant in several genera of
Papilionaceae. Crystalliferous strands of more than 1 1 cells in the Mimosaceae

and Caesalpiniaceae, usually of less than 1 1 cells in the Papilionaceae. Distinctly

storied inmost Papilionaceae, often with obtuse-angled gable-ends sometimes
;

storied in the Caesalpiniaceae, more rarely and more vaguely storied in the
Mimosaceae. Strands very short in most Papilionaceae.
Rays. Mostly 2-5 cells wide in the Mimosaceae, always homogeneous and
composed of small cells (less than 10 /x tang, diam.); similar rays
typically
occur insome Caesalpiniaceae and Papilionaceae but heterogeneous rays
 LEGUMINOSAE PAPILIONACEAE 529
occur in some genera of Caesalpiniaceae, and in still more genera of Papilio-
naceae; mostly 1-3 cells wide in the Caesalpiniaceae and Papilionaceae. Rays
that are not distinctly storied usually exhibit echelon arrangement in the
Mimosaceae and Caesalpiniaceae but not in the Papilionaceae.
Fibres not uncommonly septate in Mimosaceae and Caesalpiniaceae, rarely
septate in Papilionaceae. Intercelluar canals of the vertical type, both
normal and traumatic, more common in the Caesalpiniaceae. Anomalous
structure most common in the Papilionaceae.

PHYSIOLOGY
Certain of the Papilionaceae have provided subjects for physiological
investigation in relation to their morphology or anatomical structure. Thus
Boodle (234) demonstrated that the number of trifoliate seedlings of gorse
(Ulex europeus Linn.) was greater when grown on a rich soil than on sand.
Warington (2361) showed how, in Vicia faba Linn., the absence of boron
caused cambium cells to disintegrate with or without preliminary hyper-
trophy, the phloem and ground tissue to become disorganized, and the xylem
to be poorly developed or even to break down. Kraus and Mitchell (1279)
studied the effect of alpha-naphthalene acetamide on bean plants. They found
that this chemical causes the root system to become more fibrous, whilst the
initiation of roots from derivatives of ray cells is also stimulated. Vascular
bundles were found to arise from proliferated endodermal cells. Nightingale
and Farnham (1602) studied the effects of nutrient concentration on the
anatomy, metabolism, and abscission of buds of the sweet pea, and Jenkins
(1170) examined snap bean tissues affected with black rot.

ECONOMIC USES
The economic products obtained from the Papilionaceae are numerous. The
most familiar are the various food-yielding plants such as the many kinds of
peas and beans derived from species of Arachis, Cajanus, Cicer, Dolichos,
Glycine, Lens, Phaseolus, Vicia, and the fodder plants including the vetches,
lupins, lucernes, clovers, &c., derived from species of Lupinus, Medicago,
Melilotus, Onobrychis, Trifolium. One of the best-known fibre plants is Sunn
Hemp (Crotalariajuncea Linn.), but there are others of smaller or local impor-
tance such as the Spanish Broom (Spartium junceum Linn.). The blue dye
indigo, obtained from species of Indigofera, was at one time a product of
considerable importance. Gums are obtained from certain members of the
family, notably Tragacanth from Astragalus gummifer Lab. and Sarcocolla
from A. sarcocolla Dymock. Balsam of Tolu and Balsam of Peru, two oleo-
resins derived respectively from Myroxylon balsamum (Linn.) Harms and
M. balsamum var. pereirae, are used in medicine and perfumery. In recent
years the roots of various species of Derris and Lonchocarpus have assumed
considerable importance as the source of insecticides, chiefly because of their
content of rotenone and related substances. Numerous plants with alleged
medicinal properties occur in the family, but few of them are of great impor-
tance except liquorice (Glycyrrhiza glabra Linn.).
The anatomical features of some of these economic products are important
for purposes of identification. A selection is given on pp. 530-2.
4594 Mm
530 LEGUMINOSAEPAPILIONACEAE
TUBA ROOT (Denis spp.)
Roots up to cm. in diameter with greyish-brown bark, with shallow,
i

sinuous, longitudinal furrows. Older roots partly encircled by transverse

ridges. Xylem, on a freshly cut
transverse surface, cream to pale yellow;
vessels easily visible except at the centre. Cork consisting of up to about
12 layers of cells; outer cells dark reddish-brown; inner ones lighter in colour
and thin-walled. Groups of stone cells and numerous secretory cells with
the cork. Phloem stratified
shining, golden-brown contents present beneath
into tangential zones of hard and soft tissue, and broken up into wedge-shaped
strands with enlarged distal ends of the rays. Phloem fibres very thick- walled,
resin and rotenone,
average diameter about 10 \L. Secretory cells, containing
frequent between the groups of phloem fibres. Xylem including vessels of
2 distinct sizes. Larger vessels elliptical to circular in transverse section,
diameter no X 90-210 X 170 p, but usually about 180 X 155 /*, mostly solitary
or occasionally accompanied by 1-6 much smaller angular vessels generally
about 27 X 32 p in diameter. Small vessels also occur arranged in radial rows
or clusters of 3-6. Vessel members mostly 140-190 ^ long. Perforations
simple. Pits alternate, elliptical to polygonal,
about 6 /z broad, with horizontal,
slit-like borders; distinctly vestured. Ground tissue of the xylem consisting
of storied cells filled with starch, a few containing
mostly parenchymatous
resin or rotenone. Compact strands of fibres occur in the parenchymatous
elements about
ground-mass, each strand consisting of 2-50 very thick- walled
10 [L in diameter and surrounded by a sheath of cells each containing a solitary
embedded in of the inner tangential walls. Rays homo-
crystal thickenings
1-6 cells wide. (For another description of the root of Derris
geneous,
elliptica see Russell (1972).)

CUBE ROOT (Lonchocarpus utilis H. B. et K.)

Roots up to about 22 mm. in diameter, pale yellowish-brown, surface
slightly furrowed longitudinally and provided with chocolate-brown, trans-
surface pale cream in
versely elongated lenticels. Freshly cut transverse
colour, turning yellow when moistened with water. Cube roots can readily
be distinguished from Tuba (Derris) by the much less numerous phloem
fibres and by the relatively infrequent vessels, the latter being mostly solitary
or in occasional radial or oblique pairs, seldom exceeding 33 ju in diameter
to about 180
except in a few specimens where they rarely attain up ji.

it is not always easy to identify a particular

According to Panshin (1651)
species of rotenone-yielding plant from its microscopical structure alone, for
elements of the
although differences in the size and frequency of the different
secondary xylem are known to occur in closely related species, the variations
are no greater than those which may be found in sections of the stem and
roots taken from different parts of the same plant. Worsley (2473), after
examining 22 species of Papilionaceae for the presence of rotenoids,
found that
these substances occur in varying amount in Derris, Millettia, Mundulea, and
Tephrosia. He also concluded that rotenoids are waste products
of the plant's
metabolism except possibly in the seed. The structure of Millettia has been
described by Russell (1973). For information concerning the taxonomy of
plants which yield rotenoids see Krukoff and Smith (1290).
 LEGUMINOSAEPAPILIONACEAE 531

LIQUORICE (Glycynhiza glabra Linn, and related species of Glycynhiza)

The commercial product consists of the rhizome and root. It occurs in
pieces 1-2 cm. in diameter. Surface reddish-brown when unpeeled, but the
commercial product often has the outer tissues removed. The more impor-
tant microscopical features include the very broad rays by which xylem is
dissected into radiating groups; the cork consisting of up to about 15 layers
of golden to chocolate-brown cells with thickened tangential walls; the strands
of thick-walled fibres in the phloem and xylem, each strand usually approxi-
mately rectangular in transverse section and consisting of 10-50 but occa-
sionally including as many as 100 members often accompanied or sheathed
by cells each of which contains a prismatic crystal; the xylem vessels, either
solitary or in small clusters, the individual members being 40-130 (mostly
about 100) IJL
in diameter; the numerous starch grains mostly about 10 fi in
diameter. Senft (2079) has described and discussed the physiological function
of amorphous deposits in certain of the cells of Glycyrrhiza glabra.

BAPTISIA OR WILD INDIGO ROOT (Baptisia tinctoria R. Br.)

The woody rhizome and root are used in medicine as a laxative. The root
contains 2- to 4-compound starch grains, lignified phloem fibres, and brownish
cork cells. A rhizome and root, which had been substituted for those of
Baptisia tinctoria, were provisionally identified at Kew as Thermopsis sp. For
the anatomy of the root, stem, and leaf of Baptisia tinctoria see Holm (1030).

ANGELIM BARK (Andira and Hymenolobium sp.)

The anatomy of the so-called Angelim barks from Brazil, to which medicinal
properties have been attributed, has been fully described (Brocardet 276).

PISCIDIA ERYTHRINA BARK

Jaeger (1140) has described the structure of the bark of Piscidia erythrina
Lam., which is used to prepare fish poison. Characters mentioned as being
of particular diagnostic value are the resin cavities in the cortical parenchyma,
and the cells, in the same region, which contain bipyramidal quadratic
crystals, partly or wholly embedded in a lignified deposit within the cells.

ADESMIA BORONIOIDES
Adesmia boronioides Hook. f. from Patagonia is a small shrub, 18 inches
high, the leaves of which are covered with glands which secrete a viscid sub-
stance with a pleasant balsamic odour, which might be worthy of consideration
for use in perfumery. The small, resin-covered leaflets, especially those of the
young leaves when still not fully unfolded, have a characteristic appearance
which, in specimens from which the typical, yellow, papilionaceous flowers
are absent, might be mistaken at first sight for inflorescences. The stems are
green when very young, but reddish-brown and longitudinally striated when
slightly older. The microscopical features of the leaf include the infrequent,
simple hairs, each having a few short basal cells and a long terminal cell; the
approximately isobilateral mesophyll of the leaflets; the large masses of
feathery crystals of unidentified material in the mesophyll (the material is
dissolved during the preparation of balsam mounts) ; the glands embedded
532 LEGUMINOSAEPAPILIONACEAE
in the outer part of the mesophyll; the deeply crescentic group of widely
spaced vascular bundles, each supported externally by well-developed 'peri-
cyclic' fibres, visible in transverse sections of the rachis. The microscopical
features of the stem include the small glands in the outer part of the primary
cortex; the large, broad, tangentially elongated strands of pericyclic fibres;
the well-defined, primary vascular bundles with numerous vessels united by
interfascicular fibres to form a closed ring; the broad pith; the occasional,
elongated secretory cavities in the pith and cortex, those in the pith containing
yellowish-brown material.

MUMMY PEA (Pisum sativum van)

The structure of the so-called 'mummy pea' has been described by Compton
(456). The 'mummy pea* is a variety of Pisum sativum Linn, in which the
upper part of the axis is funnel-shaped and bears a dense cluster of leaves and
short inflorescences. The following is a quotation from Compton's paper:
'In the uppermost region of the swollen funnel-shaped axis, two continuous con-
centric rings of vascular tissue are present. The outer of these rings is directly
continuous with the single ring of bundles below, and the different vascular
elements of the xylem, phloem, etc. have normal orientation. The inner ring on the
other hand has an inverse orientation of tissues. In between the two rings of
vascular bundles is found a zone of parenchymatous tissue, histologically like a
medulla. No vascular connection was discovered between the bundles of the outer
cylinder and those of the inner, either at the node at which the latter takes its
origin, or at any point of its course.'

The structure is an example of ring fasciation and breeds true.

Other useful references to the microscopical structure of members of the
Papilionaceae of economic importance include the following. Winton (2443)
has described a method for the detection of lucerne (Medicago saliva Linn.),
red clover (Trifotium pratense Linn.), and alsike clover (T. hybridum Linn.)
when present in ground material used as cattle food. Anatomical details by
which it is claimed that species of Trifolium may be distinguished from one
another have been recorded by Vechot (2328). Articles dealing with the
anatomy of Arachis hypogaea Linn, have been published by Waldron (2344) and
Reed (1896), and on the Lima bean by Snell and Pollard (2155), on the coffee
bean (Cicer arietinum Linn.) by Holm (1003), on the lupin by Boas and
Merkenschlager (209), on the developmental anatomy of the root of Melilotus
alba Desr. by Bottum (242), whilst further general information concerning
leguminous crop plants may be found in the text-books of Hector (929) and
Hayward (927).
The histology si Astragalus molissimus Torr., which, when eaten, causes the
'Loco* disease of cattle in Kansas, has been described by Ritter (1904), with
a view to facilitating its identification in cattle food.

TIMBERS
This family produces a number of woods that have been known and prized
throughout the world for a very long time some of them are among the most
;

valuable woods known for inlay and cabinet work, musical instruments, and
carving.
 LEGUMINOSAEPAPILIONACEAE 533
The most important genus is Dalbergia, which is the source of the following
important timbers: Rosewood, e.g. Brazilian Rosewood, D. nigra Fr. All.,
Honduras Rosewood, D. stevensonii Standl., and Indian Rosewood, D. lati-
folia Roxb., Cocobolo, D. return HemsL, Tulipwood, Dalbergia sp., King-
wood, /). cearemis Ducke, and African Blackwood, D. melanoxylon Guill. et
Perr. Pterocarpus also produces several well-known timbers such as Padauk,
Narra, and Red Sanders. Among the other tirftbers of importance may be
mentioned Angelim or Partridge Wood, Andira inermis (Sw.) H. B. et K., Cocus
or Brown or Green Ebony, Brya ebenus DC., and Black Bean, Castanosper-
mum australe A. Cunn.
Though most of the woods are hard, heavy, deeply coloured, and durable,
the wood of the Ambatch tree, Aeschynomene elaphroxylon (Guill. et Perr.)
Taub., lies at the other extreme, with an air-dry specific gravity of about 0-2.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Abrus,* Adenocarpus,* Adesmia,* Aeschynomene, Alhagi, Alysicarpus,
Amicia, Ammodendron, Amorpha, Amphicarpaea, Amphithalea, Ana-
gyris, Anarthrophyllum, Andira, Anthyllis,* Aotus, Apios,* Apoplanesia,
Arachis,* Argyrolobium, Aspalathus, Astraglus,* Atylosia, Baphia, Baptisia,*
Barbiera, Borbonia, Bossiaea, Bowdichia, Brachysema, Brongniartia, Brya,
Buchenroedera, Burtonia, Butea, Cajanus, Calophaca, Calopogonium, Cal-
purnia, Calycotome, Camptosema, Canavalia, Caragana,* Carmichaelia,*
Castanospermum,* Centrolobium, Centrosema, Chadsia, Chaetocalyx, Chap-
mannia, Chorizema,* Cicer,* Cladrastis,* Clathrotropis, Cleobulia, Clianthus,
CHtoria, Cochlianthus, Coelidium, Cologania, Colutea,* Cordyla, Coronilla,*
Corynella, Cranocarpus, Cratylia, Crotalaria,* Cyamopsis, Cyclopia, Cylista,
Cymbosema, Cytisopsis, Cytisus,* Dalbergia,* Dalea, Dalhousiea, Daviesia,
Derris,* Desmodium,* Dichilus, Dillwynia, Dioclea, Diphaca, Diphysa,
Diplotropis, Dipteryx, Discolobium, Dolichos, Dorycnium, Drepanocarpus,
Dumasia, Dunbaria, Ebenus, Eleiotis, Erinacea, Eriosema, Erythrina,*
Euchlora, Euchresta, Eutaxia, Eversmannia, Eysenhardtia, Fagelia, Flemingia,
Galactia, Galega,* Gastrolobium, Geissaspis, Genista,* Geoffraea, Glycine,*
Glycyrrhiza,* Gompholobium, Goodia, Grona, Gueldenstaedtia, Halimoden-
dron,* Hallia, Hammatolobium, Hardenbergia, Harpalyce, Hedysarum,*
Helminthocarpum, Herminiera, Heylandia, Hippocrepis, Hosackia, Hovea,
Hymenocarpos, Hypocalyptus, Indigofera,* Jacksonia, Kennedya, Lablab,
Laburnum,* Lathriogyna, Lathyrus,* Latrobea, Lebeckia, Lens, Lepto-
desmia, Lespedeza,* Lessertia, Liparia, Loddigesia, Lonchocarpus,* Lotono-
nis, Lotus,* Lourea, Lupinus,* Machaerium, Marina, Mecopus, Medicago,*
Melilotus,* Millettia,* Mirbelia, Mucuna, Muellera, Mundulea, Myrosper-
mum, Myroxylon,* Nissolia, Olneya, Onobrychis, Ononis,* Ormocarpum,
Ormosia, Ornithopus, Ougeinia, Oxylobium; Oxytropis,* Pachyrhizus, Paro-
chetus, Periandra, Petalostemon, Petteria,* Phaseolus,* Phyllota, Phylloxylon,
Physostigma, Pictetia, Piptanthus,* Piscidia, Pisum,* Platycyamus, Platylo-
bium, Platymiscium, Platyosprion,* Platypodium, Podalyria, Poecilanthe,
Poiretia, Poitaea, Pongamia, Priestleya, Priotropis, Pseudarthria, Psopho-
carpus, Psoralea,* Pterocarpus, Pterodon, Pueraria, Pultenaea, Pycnospora,
 534 LEG UMINOSAEPAPILIONA CEAE
Rafnia, Rhynchosia, Robinia,* Rothia, Rudolphia, Sabinea, Scorpiurus,*
Securigera,* Sesbania, Shuteria, Smithia, Soemmeringia, Sophora,* Spar-
tium,* Spatholobus, Sphaerolobium, Sphaerophysa,* Strongylodon, Stylo-
santhes, Sutherlandia, Swainsonia, Swartzia, Sweetia, Taverniera,
Templetonia, Tephrosia, Teramnus, Tetragonolobus, Thermopsis,* Tipuana,
Trifolium,* Trigonella,* Ulex,* Uraria, Viborgia, Vicia,* Vigna, Viminaria,
Virgilia, Voandzeia, Wistaria,* Zollernia, Zornia.
* Kew slide
Represented in the collection.

(ii) FOR WOOD STRUCTURE

Aeschynomene, Afrormosia, Alexa, Amerimnon, Amherstia, Ammoden-
dron, Amorpha, Anagyris, Andira, Apoplanesia, Ateleia, Baphia, Behaimia,
Belairia, Benthamantha, Bergeronia, Bowdichia, Brya, Butea, Calycotome,
Canavalia, (Cascaronia), Cashalia, Castanospermum, Centrolobium, Cladras-
tis, Clathrotropis, Coumarouna, Coursetia, Crotalaria, Cytisus, Dalbergia,

Dalea, Daviesia, Derris, (Desmodium), Diphysa, Diplotropis, Dipteryx,

Drepanocarpus, Dussia, Ecastophyllum, Erythrina, Etaballia, Eysenhardtia,
Genista, Geoffraea, Gliricidia, Gourliea, Halimodendron, Haplormosia,
Harpalyce, Hebestigma, Hymenolobium, (Ichthyomethia), Indigofera, Ino-
carpus, Kunstleria, Laburnum, Lennea, Lespedeza, Lonchocarpus, Lupinus,
Machaerium, Medicago, Millettia, Monopteryx, (Mucuna), Muellera, Myro-
carpus, Myrospermum, Myroxylon, Ormosia, Ostryocarpus, Ougeinia,
(Pachyrhizus), Paramachaeriuin, Parosela, Pericopsis, Pickeringia, Piscidia,
Platycyamus, Platymiscium, Platypodium, Podalyria, Poecilanthe, Pongamia,
Pterocarpus, Pterodon, Pueraria, (Retama), Rhynchosia, Robinia, Rothia,
Sabinea, (Sarothamnus), Schefflerodendron, Sesbania, Sophora, Spartium,
Spatholobus, (Strongylodon), Swartzia, Sweetia, Tipuana, Toluifera, Torresia,
Ulex, Virgilia, Vouacapoua, Wistaria.

LITERATURE
(i) On
General Anatomy
Alexandrov 12, Anonymous 28, Beijer 168, Boas and Merkenschlager 209, Boer 214,
Bombacioni 223, Bond, G. 226, Boodle 234, Bottum 242, Bowman 250, Brocardet 276,
Buchegger 303, Buder 305, Compton 456, Dastur and Saxton 542, Dormer 601, 602, 604,
605, Dubard and Dop 6ix, Evenari (Schwarz) 665, Fehr 680, 681, George 756, Good-
latte 795, Gravis 804, Hagerup 864, Handa 882 A, Hayward 927, Hector 929, Hill, T. G.
973, Holm 993, 1003, 1030, Hunter mi, Jacobsohn-Stiasny 1137, Jaeger 1x40, Jenkins
1x70, Jimbo 1181, Jolivet 1187, Kraus and Mitchell 1279, Krukoff 1289, Krukoff and
Smith 1290, Lacoste 1311, 13x2, Leupin 1364, Lindemuth 1373, Low 1397, Manganaro
i43i,Metca!fe X494,Morvillez 1560, Nightingale and Farnham 1602, Pan shin 1651, Parsa
1057, Pellegrin 1682, Reed 1896, Reeve 1901, Ritter 1940, Russell 1972, 1973, Sabnis
1977, Schilling 2032, Senft 2079, Severini 2080, Skipper 2119, Smith and Kersten 2x49,
Snell and Pollard 2155, Staber 2178, Starr 2x88, Taylor 2239, Vechot 2328, Waldron 2344,
Walensky 2345, Ward, H. ML 2360, Warington 2361, Watari 2364, Winter 2442, Winton
2443, Worsley 2473, Zemke 2505, Ziegenspeck 2506.
(ii) On Wood Structure
Bailey 78, Beekman 167, Beijer 168, Benoist 170, den Berger and Endert 183, Besson
186, Bienfait and Pfeiffer 197, Brit. Hond. For. Dept. 274, Brown, F. B. H. 282, Brown,
H. P. and Panshin 288, 289, Burgerstein 3x0, 312, Chalk and Chattaway 362, Chalk et

Janssonius 1x47, 1x54, Janssonius and Moll 1x56, Jentsch 1x77, Jolly 1x88, Jones
Kanehira X2OO, 1209, 1213, Kanehira et aL 12x4, Kribs 1283, Lecomte 1334, Leonhardt
 LEGUMINOSAEPAPILIONACEAE 535
and Fay 1360, Martin -Levigne 1450, M^niaud 1492, Messeri 1493, Metcalfe 1497, Miller
1537, Normand 1615, Panshin 1651, Pearson and Brown 1679, Pereira 1687, Record
1703, 1788, 1836, 1841, 1843, 1846, 1850, 1851, 1862, 1863, 1883, Record and Hess
1886, Record and Mell 1894, Riera 1937, Scott 2075, Stevenson 2199, Stone 2202, 2203,
2206, 2207, Tang '2230, 2231, Torres 2269, Tortorelli 2273, Uspensky 2317, Wallis
2348, Williams 2430.

116. KRAMERIACEAE
(FiG. 118 on p. 536)

SUMMARY
A small family of shrubs, ranging from Mexico to Chile, which comprises
the single genus Krameria. The shape of the leaf or leaflet, as seen in trans-
verse sections, varies considerably in different species, and might well prove
to be of considerable diagnostic value. The outer walls of the epidermal cells
of the lamina are usually heavily cutinized. Hairs, which vary in frequency
and length and occur on the leaves and young twigs, are invariably thick-
walled and unicellular. Stomata are present on both surfaces of the leaf,
whilst the mesophyll includes clusters of sclerosed, pitted, parenchymatous
cells at the centre. Cork in the young stem usually arises in the inner part of
the cortex or pericycle, and, in some species, cuts off the large strands of
fibres which occur in the pericycle. The xylem in young twigs forms a
continuous cylinder traversed by narrow rays. The vessels are invariably
small, but infrequent to numerous in different species. The hard ground
tissue of the wood is composed of fibres with bordered pits. The diameter of
the pith varies considerably in different species. Cluster crystals are common
in the parenchymatous tissues, whilst secretory cells, with presumably
tanniniferous contents, are common.
LEAF
Lamina of the leaf or leaflet varying in shape in different species isobilateral
;

in the species with a relatively large lamina, e.g. K. argentea Mart., K.

lanceolata Torr., and K. tomentosa St. Hil., but centric in smaller leaves,
e.g. in K. parvifolia Benth. (Fig. 118 K). Hairs invariably unicellular, thick-
walled, but varying in length and frequency in different species, e.g. very
numerous in K. greyi Rose et Painter, relatively short and infrequent in K.
cistoidea Hook. Outer walls 01 the cells of the epidermis strongly cutinized ;

usually arched outwards and sometimes tending to be papillose. Stomata

present on both surfaces those of K, tomentosa (Fig. 1 1 8 M) usually sur-
;

rounded by 4-6 cells slightly differentiated from the remainder of the epider-
mis, 2 of the surrounding cells being commonly but not invariably parallel to
the pore. Stomata not examined at Kew in surface view for other species, but
said by Kunz (1306) to be mostly rubiaceous and variously orientated in other
species as well. Mesophyll. Usually with a single layer of palisade cells
towards both surfaces and at the margins; 2 palisade layers observed in K,
cistoidea Cav. and K. triandra Ruiz, et Pav. but palisade tissue not very well
defined in the last of these species. Centre of the mesophyll consisting of
somewhat spongy tissue, including, in all species examined txzeptK.paucifolia
DC., small clusters of thick-walled, pitted, sclerosed cells. Vascular bundles
of the veins embedded in the mesophyll, each supported by small groups of
 FIG. 118. LEGUMINOSAEMIMOSACEAE,A-E;LEGUMINOSAECAES-
ALPINIACEAE, F~J; KRAMERIACEAE, K-O
A, Acacia decurrens Wttld. var. dealbata F. Muell. Petiole X n. B, 'ProsopisjacariHort.'. Petiole
X32, C, Inga punctata Willd. Petiole X 18. D, Entada scandens Benth. Stem X 12. E, Acacia
decurrens Willd. var. dealbata F. Muell. Stem X 7. F, Saraca declinata Miq. Petiole x 10. G,
Brownea coccinea Jacq. Petiole X 13. H, Haematoxylon campechianum Linn. Young stem X5$.
I, Gymnocladus dioica K. Koch. Stem X 6. J, Bauhinia forficata Link. Petiole x 32. K, Kramena
parviflora Benth. T.S. Leaflet x 18. L, K. isdna Linn. Stem X 18. M, K. tomentosa A. St. Hil.
Leaf epidermis x 100. N, K. tomentosa A. St. Hil. Petiole X28. O, K, argentea Mart. Petiole X33-
 KRAMERIACEAE 537
thick-walled fibres, but the amount and arrangement of the fibres varying in
different species. Transverse sections through the distal end of the petiole
of K. argentea (Fig. 118 o) exhibit a small crescent-shaped vascular strand
with very much incurved ends (almost cylindrical), supported by large
columns of pericyclic fibres; structure similar but with less supporting fibres
in K. tomentosa (Fig. 118 N); vascular arc more open and not supported by

pericyclic sclerenchyma in K. cistoidea; main vascular strand with a cylinder

of xylem accompanied by a crescent of phloem and supported by a broad arc
of very thick-walled fibres in K. ixina Linn. Very small additional strands
towards the wings present in some species. Cluster crystals common in all
unlignified tissues of the leaf of most species, but varying in frequency;
sphero-crystals seen in the mesophyll of K. cistoidea and K. triandra crystal-
;

sand recorded by Kunz (1306) in certain species. Secretory cells with

amorphous, probably tanniniferous, contents common in the mesophyll and
petiole.

Axis
YOUNG STEM (Fig. 118 L)

Epidermis composed of cells with thickly cutinized outer walls cells in

;

some species arched outwards or even tending to be papillose. Outer part of

the primary cortex differentiated as palisade tissue in K. argentea Mart.,
K. cistoidea Hook, (not very well differentiated), K. greyi Rose et Painter,
K. ixina Linn., and K. tomentosa St. Hil. Definite palisade tissue not observed
in the other species examined. Inner part of the cortex in all species paren-
chymatous. Cork generally arising fairly deep in the cortex or even in the
pericycle, the pericyclic fibres being cut off when the cork is sufficiently deep
seated; composed of cells which are thin- walled apart from the suberized
outer walls which are thicker and dome-shaped. Pericycle usually including
isolated, but in some species large, strands of thick-walled fibres. Phloem
devoid of lignified elements in the material examined. Xylem in the form of
a broad or narrow continuous cylinder, traversed by inconspicuous rays.
Vessels varying in size, frequency, and arrangement in different species up to
;

30 /t in radial diameter and mostly in radial groups in K. argentea; smaller

and less numerous in K. cistoidea] infrequent, mostly solitary, radial diameter
seldom more than 25 //. in K. parvifolia Benth. numerous, up to about 45 //,
;

in radial diameter in K. tomentosa mostly solitary, but locally tending to be in

;

radial chains, radial diameter seldom attaining 30 //. in K. triandra Ruiz, et

Pav. with spiral thickening or bordered pits and simple perforations in all of
;

the species examined. Moderately to very thick-walled fibres with bordered

pits form the ground tissue of the xylem in all species. Pith varying con-
siderably in diameter in different species, e.g. broad in K. argentea, small in
K. parvifolia] in most instances composed of or including a high proportion
of moderately thick-walled, pitted cells. Clustered crystals present in all
species; sphero-crystals as well as other clustered types observed in K.
cistoidea, and K. triandra. Asmall proportion of the crystals contained in
sclerenchymatous idioblasts in K, greyi, K. ixina, and K. parvifolia. Scattered
secretory cells, with amorphous contents believed to be tanniniferous, occur
in unlignified tissues, but vary in frequency in different species.
538 KRA MERIA CEA E
ROOT, see 'Economic Uses' below

TAXONOMIC NOTES
Krameria has been variously assigned to the Polygalaceae or to the Legumi-
nosae-Caesalpiniaceae. The taxonomic position of the genus is fully discussed
by Kunz (1306) who took both anatomical and exomorphic evidence into
consideration. He concluded that Krameria has no very close affinities with
either of the above groups, and favoured the idea that it should form the basis
of a distinct family.

ECONOMIC USES
Rhatany, used in medicine as an astringent, consists of the crown and larger
roots of K. triandra Ruiz, et Pav. The roots are reddish, usually straight but
sometimes slightly tortuous, mostly about the thickness of a finger, but
attaining a diameter of 3 cm. or more near the crown. They have a somewhat
irregular, slightly fibrous fracture, and the bark is easily detached from the
xylem which occupies about two-thirds of the diameter. Microscopical
features include the following. Cork narrow, consisting of about 8 layers of
cells, only the outer tangential walls being thickened and dome-shaped some ;

of the cork cells filled with yellowish-brown contents. Primary cortex in the
commercial article usually absent, having been cast off by the activity of the
phellogen. Secondary cortex fairly narrow, composed of thin-walled paren-
chyma. Phloem in the form of elongated, triangular strands exhibiting out-
wardly directed, somewhat sinuous apices in transverse sections, including a
high proportion of thick-walled, unlignified fibres with the lumina variously
shaped and often slit-like. Phloem strands separated from one another by the
expanded distal ends of the medullary rays. Vessels of the xylem numerous,
arranged in approximately concentric circles, circular to oval, usually solitary
and up to about 45 /x or more in radial diameter, with bordered pits having
horizontal apertures; perforations simple. Ground- tissue of the wood com-
posed of thick-walled fibres with conspicuous, bordered pits. Wood paren-
chyma arranged in slightly interrupted, uniseriate, concentric bands.
Numerous, mostly simple but sometimes loosely compound, variously shaped
starch grains present in the parenchymatous tissues. Solitary crystals and
crystal-sand sometimes present in association with the phloem fibres.

GENUS DESCRIBED
Krameria.*
* The above
description is based mainly on sections, now in the Kew slide collection, from
herbarium specimens of the following species: K. argentea Mart., K. cistoidea Hook., K.
cytisoides Cav., K. greyi Rose et Painter, K. ixina Linn., K. lanceolate, Torr., K. parvifolia
Benth., K, paucifolia DC., K. tomentosa St. HiL, K. triandra R. et P.

LITERATURE
On General Anatomy
Kunz 1306.
 (539)

117. ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE)

(Fic. 119 on p. 542; FIG. 1 20 on p. 544)

SUMMARY
(i) GENERAL
Trees, shrubs, or herbs with very few anatomical features common to the
whole family. The family is very widely distributed but many species occur
in north temperate regions. Hairs generally unicellular, in the form of simple
trichomes, or occasionally united to form tufts. Glandular hairs, and glan-
dular shaggy hairs also occur. Nectaries are present on the petiole, leaf
surface, and on the leaf teeth in certain species. Stomata are ranunculaceous.
Cork in the young stem arises superficially in some species but endogenously
in others. Pericycle with strands of sclerenchyma or a continuous scleren-

chymatous ring. The vascular bundles in the young stem are separated by
very narrow medullary rays in some genera, so that the xylem appears in
transverse sections to form a continuous ring around the pith. In other genera
the bundles are widely separated by broad primary rays and therefore appear
as individually distinct units. Calcium oxalate is usually secreted in the form
of solitary or clustered crystals. True styloids are known only in Quillaja.
Secretory elements. Mucilaginous cells are common in the leaf epidermis
and in the parenchyma of the leaf veins and stem (Eriobotrya and Neurada)*
lysigenous mucilage canals also occur in the pith of Neurada. Tannin is
abundant but not usually localized in special cells.
Characters such as the outline of the epidermal cells as seen in surface view;
the structure of the stomata; the number of palisade layers; the size, nature,
and distribution of crystals; the structure of the larger vascular bundles; the
nature and size of the hairs have been employed by Gyhr (854) for the
microscopical identification of the leaves of members of the Rosaceae with
medicinal properties. These characters are, however, of value only for
specific and to a certain extent for generic diagnosis. No satisfactory leaf
characters for the separation of the tribes were found by Gyhr, but, on
the other hand, it was much easier to divide the leaves into two classes accord-
ing to whether they were from herbaceous or woody plants.
When making use of the outline of .the epidermal cells as seen in surface
view for diagnostic purposes, it would be as well to bear in mind the variations
in this character which were found by Haberlandt (859) in sun and shade
leaves of an individual species during his researches on the Crataegomespili.
Minor variations between mesophytic and dune forms of Prunus (Starr 2188)
and between sun and shade leaves of Rubus spp. (Bird 199) have also been
recorded. Variations in the stem structure which may occur within an indivi-
dual species of Crataegus, Prunus, or Pyrus have also been described by
Aubertot (48).

(ii) WOOD
Vessels in temperate species mostly small and numerous, with a tendency
to ring-porousness, with oblique or radial arrangement in some species, often
spirally thickened, perforations typically simple, but with sporadic multi-
perforate plates in some species, intervascular pitting alternate, small, pits to
540 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE)
parenchyma similar; members of medium length. Parenchyma typically
apotracheal, diffuse or in short bands, with some scanty paratracheal paren-
chyma in a few species. Rays mostly 2-5 cells wide, considerably more in
some genera and occasionally of 2 distinct widths; heterogeneous to homo-
geneous. Fibres typically with numerous distinctly bordered pits on radial
and tangential walls, of medium length to moderately long.

LEAF
Usually dorsiventral. Centric in Crataegus azarolus Linn. Hairs commonly
unicellular but showing considerable variations in structure, (i) With charac-
teristic,solid, wart-like projections in Kcrria and Neviusa. (ii) Forming
tufted or stellate groups in species of Potentilla (section Stelligerae), certain
Rubus spp., and Spiraea sorbifolia A. Br. Stalked capitate glands occur in
Alchemilla, Fragaria, Geum, Potentilla, Prunus, Rosa, Rubus, and Sanguisorba.
Non-capitate, multicellular hairs also recorded by Gyhr (854) in Sanguisorba.
Spines of Rosa and Rubus arise superficially. Nectaries on the petiole of
certain species of Prunus are stated by Gregory (815) and Knapheisowna
(1249) to contain sugar and tannin, and, according to Dorsey and Weiss (607),
to represent the suppressed members of an originally ternate leaf. Glandular
leaf teethrecorded in Crataegus, Cydonia, Pyrus, Sorbus, and secretory leaf
teeth, devoid of palisade epidermis, in Alchemilla, Fragaria, Kerria, Rubus,
Sanguisorba, Spiraea, Glandular spots present on the lower surface of the
leaf of Laurocerasus, and oil drops in the palisade cells of Prunus laurocerasus
Linn. Cuticle striated in Chaenomeles japonica Lindl. Elevations of cuticle
around the stomata recorded in Pyrus communis Linn. Cells of the epidermis
varying in outline as seen in surface view; provided with a stratified coating
of wax in Kerria japonica DC. Lower epidermis papillose in certain species
of Amelanchier, Cotoneaster, Prunus, Pyrus, Rosa rugosa Thunb., Sorbus, and
Spiraea. Papillose on both surfaces in Acaena adscendens Vahl. Inner walls
on the epidermal cells commonly mucilaginous. Variations in the gelatiniza-
tion in different species of Cliffortia have been described by Montcheff (1553).
Hypoderm occurs in certain species of Heteromeles, Osteomeles, Pygeum,
Rubus (section micranthobatus), and Sibiraea. Stomata nearly always con-
fined to the lower surface, except in Geum parviflorum Sm. (Betts 190),
Neurada procumbens Linn, (Sabnis 1977), and Rosa berberifolia Pal. (syn.
persica Michx. ?). Stomata situated in pits in Cliffortia (Montcheff 1553).

Hydathodes present in the leaf teeth of Agrimonia, Alchemilla, Crataegus,

Geum, Potentilla, and possibly other genera (see also under 'Glands'). Meso-
phyll generally with 2 or more layers of palisade cells, but i single layer
recorded by Gyhr (854) in certain species of Alchemilla and Geum. Normal
palisade stated to be absent from Prunus armeniaca Linn. Mesophyll con-
sisting wholly of short palisade cells in Neurada procumbens (Sabnis 1977).
Vascular bundles of the smaller veins with or without sclerenchyma in the
parenchymatous sheath surrounding them vertically transcurrent in Neurada
;

procumbens (Sabnis 1977). Petiole exhibiting the following types of structure

in transverse sections through the distal end. (i) An arc of vascular bundles
in Acaena (Morvillez 1556), Agrimonia (Fig. 119 F), Filipendula (Fig. 119 B),
Fragaria, Geum, Potentilla (Morvillez 1556), Poterium (Fig. 1190), Prunus
(pro parte), Quillaja, Raphiolepis, Rosa (pro parte), Rubus (Betts 189), (Holm
 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE) 541

1045), (Morvillez 1556), Sanguisorba, Sorbus (Morvillez 1556), Spiraea (sec-

tions aruncus and sorbaria) (Morvillez 1556), and Waldsteinia (Morvillez
1556). (ii) A A
solitary crescent-shaped bundle in certain species of melanchier,
Cotoneaster, Crataegomespilus, Cydonia, Dichotomanthes, Neviusa, Prunus
(Fig. 119 A) (pro parte and not including Laurocerasus), Pyracantha, Pyrus
(pro parte) (Fig. 119 E), Spiraea (section Holodiscus) (Morvillez 1556). (iii) A
principal solitary crescent-shaped bundle, accompanied by smaller or very
small subsidiary ones in or towards the wings, in certain species of Crataegus,
Eriobotrya, Kerria, Neillia, Photinia, Potentilla (Fig. 119 l) (pro parte) (Mor-
viilez 1556), Prunus (including P. laurocerasus), Pyrus (pro parte), Raphiolepis,
Sorbus (Fig. 1190), Stranvaesia. (iv) Several centric bundles, each surrounded
by an endodermis and consisting of a ring of xylem and phloem enclosing a
pith-like tissue, in at least some species of Alchemilla (Fig. 119 c). (v) With
8-9 bundles, i being centric in structure as in Alchemilla, the remainder being
hemicentric, each then consisting of an arc of xylem and phloem surrounded
by an endodermis in 'Sanguisorba canadensis Linn/. The principal strand of
Eriobotrya japonica Lindl. (Fig. 119 H) also centric (Morvillez 1556).
The Crataegomespili, or graft hybrids between Mespilus germanica Linn,
and Crataegus monogyna Jacq., exhibit a proportion of characters which are
intermediate between those of the parents but also exhibit a mosaic-like
combination of parental characters. See Haberlandt (858, 859, 860, 861) and
Weiss (2391).
Calcium oxalate generally present in the form of solitary or clustered
crystals. True styloids recorded only in Ouillaja. The type of crystal is
sometimes characteristic of groups of species, e.g. (i) Crystals simple, or in
clusters consisting of 3-4 components in Rubus, section Micranthobatus.
(ii) Crystals exclusively clustered in Rubus sections, Anoplobatus, Batho-

thamnus, Eubatus, Idaeobatus. Exclusively or predominantly clustered crystals

occur in Potentilla and Spiraea, and specially large rosette crystals in the
petiole of species of Prunus, excluding P. laurocerasus. Crystals stated by
Gyhr (854) to be absent from the leaf of Mespilus germanica.
Axis
YOUNG STEM (Fig. 119 j-o)
Cork originating in very different positions in various members of the
family, ranging from the epidermis or sub-epidermis in some species to the
pericyclic region in others. Cork cells variable in structure. Mostly cubical
with thin walls and tending to separate into concentric layers in Neillia
sinensis Oliv. Cortex. Inner portion frequently collenchymatous in woody

species occasionally containing groups of stone cells in Osteomeles and Pyrus,

;

and fibres in Crataegomespilus. Mucilage cells observed in the broad primary

cortex of Eriobotrya japonica Lindl., and recorded by Sabnis (1977) in the
cortex, rays, and pith of Neurada procumbens Linn. Palisade chlorenchyma
present in the cortex of Prinsepia. Endodermis well defined in species of
Alchemilla, Fragaria, Gillenia (Holm 1028), Kerria, Neviusa, Potentilla
(Priestley and Hinchliff 1751), Poterium, and certain Rubus spp. (Betts 189).
Pericycle of woody genera initially provided with primary bundles of fibres,
but groups of stone cells subsequently develop between them to form a broken
ring in certain species of Chaenomeles, Crataegus, Osteomeles, Sorbus, and
 E

c.e.

FIG. 119. ROSACEAE

A, Prunus persica Batsch. Petiole X 10. B, Filipendida ulmaria (L.) Maxim. Petiole X 9. C,
Alchemilla vulgaris agg. Linn. Petiole X to. D, Poterium officinalis A. Gray. Petiole X 9. E, Pyrus
communis Linn. Petiole X 26. F, Agrimonia eupatoria Linn. Petiole x 10. G, Sorbus aucuparia Linn.
Petiole X 13, H, Eriobotrya japonica Lindl, Petiole x 6. I, Potentilla rupestris Linn. Petiole X 13.
J, Raphiolepis umbellata Mak. Stem X 8. K, Rosa camna Linn. Stem X 9. L? Cotoneaster frigida
Wall. Stem X 6. M, Rubus deliciosus Torr. Stem X 9. N, Agrimonia eupatoria Linn. Stem x 9.
O, Kerria japonica DC. Stem X9.
c.*. Cutinized epidermis. . Endodermia. p. Thin-walled pith cells, t.p. Thickened pith cells.
 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE) 543
Stranvaesia. Isolated strands of fibres with intervening unlignified paren-
chyma present in certain species of Amelanchier, Cotoneaster, Crataegomespilus,
Crataegus, Cydonia, Dichotomanthes, Eriobotrya, Kageneckia, Kerria (Bow-
man 250), Lindleya, Neviusa, Osteomeles, Photinia, Potentilla, Poterium,
Prinsepia, Prunus, Pygeum, Pyracantha, Pyrus, Quillaja, Raphiolepis, Rosa,
Rubus, Sorbus, Stranvaesia, and Vauquelinia. Pericyclic sclerenchyma absent
from Neurada procumbens according to Sabnis (1977). Sclerenchymatous ring
continuous in at least some of the species of Agrimonia, Alchemilla, Fragaria,
Geum, Kerria, Potentilla\ and practically continuous in British species of
Filipendula. Vascular bundles separated by narrow primary rays, giving
the xylem the appearance of a closed ring in at least some of the species of
Agrimonia, Alchemilla, Cotoneaster (Fig. 119 L), Crataegomespilus, Crataegus,
Cydonia, Eriobotrya, Potentilla, Prunus, Pyracantha, Pyrus, Raphiolepis
(Fig. 119 j), Rosa (pro parte), and Stranvaesia. Vascular bundles sufficiently
separated by the primary rays to be clearly visible as distinct units in at least
some of the species of Agrimonia (Fig. H9N), Amelanchier, Cotoneaster,
Fragaria, Geum, Gillenia (Holm 1028), Kerria (Fig. 1190), Neviusa, Potentilla,
Poterium, Rosa (Fig. ngit) (pro parte), Rubus (Fig. H9M), and British
species of Filipendula. Rays somewhat twisted in appearance where traversing
the phloem as seen in transverse sections of certain species pf Prunus and
Pyrus. Vessels usually with simple perforations; reticulate plates occasional.
Fibres recorded in the secondary phloem in species of Cotoneaster, Crataegus,
Cydonia, Mespilus, Pyrus, Rosa, Sorbus, and Spiraea. Pith homogeneous in
Amelanchier, Cydonia, Eriobotrya, Neurada procumbens (Sabnis 1977), Pyrus
communis Linn., and heterogeneous in species of Crataegus, Mespilus, Rosa,
Rubus (Holm 1045), Sorbus, Spiraea, and Malus pumila Mill.; septate in
and becoming hollow in certain species of Geum, and in the British
Prinsepia,
species of Filipendula', very clearly differentiated into a thick- walled peri-
medullary portion and a central region consisting of much larger cells with
thin walls in Kerria japonica DC., Neillia sinensis, and Rubus deliciosus Torr.
For crystals see also 'Leaf. Noteworthy types include the large rosette
crystals in the primary cortex of species of Prunus (excluding P. laurocerasus
Linn.), Rosa hugonis Hemsl., Rosa var. 'Dorothy Perkins', in the pericycle of
Rubus deliciosus, and in the pith of Malus pumila, and Rosa hugonis. Solitary
crystals in special cells have been reported amongst the phloem fibres of
Pyrus communis and P. nivalis Jacq. as well as in the pericycle of Eriobotrya
japonica. Tannin very common, especially in parenchymatous tissues. The
arrangement of the tannin cells in the pith of Rubus has been used for
taxonomic purposes, according to Solereder. Mucilage canals recorded in
the pith of Neurada sp.

SPECIAL FEATURES
The mechanical elements are less well developed in 'weeping* than in erect
forms of Sorbus aucuparia Linn., but according to Low (1397) the differences
are not very well defined.
Anatomical differences between the stems of 'American Pillar* and 'Dorothy
Perkins* roses have been described in detail by Carlson (342).
The secretion of gum in Prunus may be sufficiently stimulated by parasitic
organisms or physiological disturbances to constitute a serious disease of
 FIG.- 120. ROSACEAE
A, Prunus lusitanica Linn. B, P. avium Linn. C, Rosa canina Linn. D, Pygeum africanum Hook. f.
E, Prunus maackii Rupr. F, Cptoneaster microphylla Wall. G, Prunus avium Linn. H, Rosa canina
Linn. I, Prunus laurocerasus Linn. J, Spiraea alpina Pall. K, Raphiolepis umbellata Mak. var.
 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE) 545
fruit-trees. The formation of these gum deposits has been described by
Butler (324).
The abscission of flowers or immature fruits of the apple has been studied
by McCown (1464). Although there is a constriction zone in the pedicels of
flowers and this persists during the life of the apple fruits, it plays no part in
the abscission of flowers or fruits. Before flowers or fruits fall off, a definite
abscission layer is formed. It is initiated independently in the pith and cortex.
The developmental anatomy of the stem apex of the almond (Prunus
amygdalus Batsch.) has been described by Brooks (278).
BARK
For anatomy of the bark of Quillaja saponaria Mol. see 'Economic Uses'
on p. 548.
WOOD (Fig. 120)
Vessels typically small, mean tangential diameter less than 100 \L, except
in some tropical species, often very small (25-50 ^), moderately large (200-
300 IJL)
in Eriobotrya, Hagenia, Laurocerasus p.p., Pygeum, and Rubus \
exclusively solitary or nearly so in the Pomoideae (except Osteomeles),
Amelanchier, Cotoneaster, Crataegus, Cydonia, Eriobotrya, Eriolobus, Malus,
Mespilus, Micromeles, Photinia, Pourthiaea, Pseudocydonia, Pyrus, and
Raphiolepis in some Rosoideae, Cercocarpus, Chamaebatia, and Purshia, and
;

in Quillaja (Spiraeoideae) with radial and oblique arrangement or pore

;

multiples of 4 or over in the Prunoideae, Laurocerasus, Nuttallia, Padus,

Prunus (Fig. 120 i), and Pygeum, and producing a flame-like pattern in Prunus
ilicifolia (Nutt.) Walp. (1886), in occasional clusters in Hagenia, Laurocerasus,
Padus p.p., Rubus p.p.; typically very numerous (40 per sq. mm.), except
where the vessels are large, between 30 and 40 per sq. mm. in Holodiscus,
Photinia, Pygeum, and Rubus p.p., between 5 and 20 per sq. mm. in Hagenia
and Laurocerasus p.p.; ring-porous or semi-ring-porous in some species of
Amelanchier, Cercocarpus, Chamaebatia, Cotoneaster, Fallugia, Holodiscus,
Laurocerasus, Lyonothamnus (1886), Nuttallia, Opulaster (2261), Padus,
Photinia, Physocarpus, Prunus, Purshia, Quillaja, Rosa, Sorbus, Spiraea, and
Vauquelinia (1886); spiral thickening occurs in about two-thirds of the genera
examined and is present in most species of Amelanchier, Cercocarpus, Chaeno-
1 2
meles, Chamaebatia, Cliffortia, Cotoneaster, Crataegus, Cydonia, Eriobotrya,
Exochorda, Heteromeles, Kage:ieckia, Laurocerasus, Lyonothamnus, Mcdus,
Mespilus, Nuttallia, Osteomeles, Padus, Photinia, Polylepis, Pourthiaea, Prin-
sepia (1206), Prunus, Pseudocydonia, Purshia, Pyrus, Raphiolepis, Rosa,
Sorbus, and Vauquelinia (164). Perforations usually exclusively simple, a few
scattered foraminate or reticulate plates occur in some species of Amelanchier,
Crataegus, Padus, Pourthiaea, Pyrus, Sorbaria, Sorbus, and Spiraea, and
according to Thompson (2254) in species of Cydonia and Potentilla. Inter-
vascular pitting typically alternate, never large, minute in Chaenomeles,
Eriobotrya bengalensis Hook, f., Kageneckia, Laurocerasus, Malus, Photinia,
Potentilla, Raphiolepis, Sorbaria, and Spiraea', Tippo (2261)
Pygeum,
describes the pitting as mostly alternate and opposite and occasionally opposite
or transitional; pits to ray cells similar to intervascular pits. The sieve-like
1
Recorded by Solereder.
2
Recorded by Solereder, but not observed in Cydonia oblonga Mill.
 546 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE)
structures in Prunusand Cerasus referred to by Jonsson (1192) according to
Bailey (78) are probably artefacts and not vestured pits. Tyloses rare;
deposits of gum present in some species, particularly in the Prunoideae.
Mean member length of mature material about 0-4-0-95 mm.; Tippo,
apparently including small stems, gives the range as 0*19-076 mm., with a
mean of 0-487 mm. Parenchyma usually apotracheal only, in scattered cells
or short uniseriate lines from ray to ray (Fig. 120 K) (short irregular meta-
tracheal bands in Pourthiaea) very sparse, with occasional scanty paratracheal
;

and diffuse cells (Fig. 120 j) in some species of Kerria, Laurocerasus, Padus,
Prunus, Rubus, and Spiraea. Kanehira (1206) notes some paratracheal paren-
chyma in Prinsepia. Crystals in chambered cells or idioblasts in some species
of Cercocarpus, Chaenomeles, Crataegus, Eriolobus, Malm, Pseudocydonia, and
Raphiolepis. Solereder records the presence of silica in Parastemon urophyllus
DC. Strands typically of 4 cells, occasionally up to 5 or 6. Rays multiseriate,
mostly 2-5 cells wide, up to 6-10 cells wide in some species of Hagenia,
Padus, Prunus and Spiraea, more than 10 cells wide in some species of
,

Kerria, Rubus, and Spiraea; over i mm. in height in Hagenia, Kerria, Rubus,
and Spiraea; sometimes of 2 distinct widths. Uniseriates rather few and com-
posed entirely of procumbent cells in woods with homogeneous rays;
moderately numerous and composed of both procumbent and upright cells
in most of the other genera, but composed only of square to upright cells in
Cotoneaster, Hagenia, Laurocerasus, Padus, Prinsepia (1206), and Stranvaesia;
uniseriates absent from Kerria. Typically 9-15 rays per mm., but only 3-5
per mm. in Kerria, Rosa, and Rubus. Homogeneous (Kribs's Type I) in
Cercocarpus, Crataegus, Cydonia, Malus, Micromeles, Padus, Polylepis, Pseudo-
cydonia, Pyrus, and Sorbus\ heterogeneous (Kribs's Type II B), with i or 2
marginal rows of square or upright cells in most of the other genera, but more
markedly heterogeneous (Kribs's Type II A) in Cotoneaster, Hagenia, Lauro-
cerasus, Prinsepia (1206), and Stranvaesia', composed entirely of square to
upright cells in some of the shrubby members, e.g. Kerria, Rosa, and Rubu$\
sheath cells occasionally present, e.g. in Hagenia and Kerria. Fibres with
numerous distinctly bordered pits, equally numerous on both tangential and
radial walls, except in the Pomoideae and some species of Spiraea, in which

they are less numerous on the tangential walls. Very fine septa and gum
1

plates occur in occasional fibres in genera of the Prunoideae, and in Spiraea

alpina Pall, and Photinia lindleyana Wight et Am. Walls thin to thick, often
radially flattened and with thicker walls towards the end of the growth ring;
spiral thickening observed or reported in some species of Amelanchier, Cerco-
carpus, Chaenomeles, Crataegus, Cydonia, Heteromeles, Lyonothamnus, Mespilus,
Osteomeles, Rosa, Stephanandra. Mean length 0-9-1-6 mm. Intercellular
canals* Traumatic vertical canals observed in one specimen of Laurocerasus
mackii (Rupr.) C. K. Schum. and reported (1801, 1886) in some species of
Prunus and Pygeum; radial canals reported (1801) in some specimens of
Pygeum. Growth rings. The seasonal development of the rings in Prunus,
Pyrus, and Rosa has been investigated by Coster (481).
1
Solereder (2158) records occasional fibres with simple pits in Kerria, Potentilla,
Rhodotypw, and Spiraea; Janssonius (1154) comments on the unusual difference between
species of the same genus with respect to fibre pitting. Such a difference, for example, occurs
between Spiraea alpina Pall, and S. salacifolia L., the borders being barely discernible in
the latter.
 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE) 547
ROOT
Roots of Rosa var. 'American Pillar* exhibit the following characters. Cork
composed of thin- walled cubical cells with amorphous contents. Phloem
strands, in transverse sections, shaped like tall triangles with outwardly
directed apices tending to be stratified into fibrous and unlignified portions.
;

Xylem exhibiting growth rings; vessels somewhat irregularly distributed,

mostly solitary, very variable in diameter but seldom exceeding 100 pin the
material examined; primary rays up to about 8 cells wide in one specimen, but
20 or more cells wide in others, amyliferous. Root structure similar in
another, undetermined variety of climbing rose, but provided with a higher
proportion of vessels of large diameter and less distinctly triangular phloem
strands.
Wedge-like thickenings of the radial and transverse walls of the cells

immediately external to the endodermis recorded in numerous genera by

Solereder, and further examined, especially in Gillenia, by Holm (1028). The
existence of a correlation between the anatomical structure of apple rootstocks
and the vigour of the scions grafted on them has been established by Beakbane
et al. (156-61). 'Vigour' in the scion is correlated with the relative proportions
of different tissue elements revealed in transverse sections through rootstocks,
the clearest connexion being with the proportion of wood ray tissue which is
present. Living and dead tissues in the wood were found to be present in
equal amounts in vigorous rootstocks, whereas in dwarfing stocks the propor-
tion of living was two or three times greater than that of dead tissue. The
vessels and xylem fibres were more numerous in the vigorous than in the
dwarfing stocks, and the size of the vessels, although not definitely propor-
tional to the vigour of the scion, tended to be larger in the vigorous stocks.
Further results are to be expected from a continuation of these researches.
See also under 'Economic Uses'. Differences in structure, of physiological
rather than taxonomic interest, between large adventitious* and 'fibrous
branch roots' recorded by White (2420) in the cultivated strawberry (Fragaria).
The developmental anatomy of the root of the pear (Pyrus communis
Linn.) has been described by Esau (659).

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY

The considerable range of petiole structure within the family is interesting,

the centric and hemicentric vascular strands in Alchemilla, Eriobotrya, and
Sanguisorba being especially noteworthy. Those of Alchemilla in particular
resemble some of those which occur in certain members of the Saxifragaceae,
but it would be unwise, without taking exomorphic and other anatomical
characters into consideration as well, to decide whether this provides evidence
of affinity or represents a parallel development. The range of petiole structure
in Spiraea (sensu laid) suggests that some members of this genus must be
somewhat remotely related to others. The rather scanty information available
suggests that Laurocerasus and Prunus should be regarded as distinct genera.

(ii)FROM WOOD STRUCTURE

The Prunoideae is the only tribe that can be distinguished from the others.
It is characterized by (i) fibres with less distinctly bordered pits, which are
548 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE)
less numerous on the tangential than on the radial walls, and frequent plates
of gum or fine septa, and (2) vessels that are commonly arranged in an oblique
or radial pattern on the cross-section and often in multiples. The group is
sometimes combined with the Chrysobalanoideae to form a separate family,
the Amygdalaceae; Record and Hess (1886), however, point out that 'from
the standpoint of the woods the Prunoideae belong with the Rosaceae and the
Chrysobalanoideae seem distinct enough to be segregated into a family of
their own*.
In the Pomoideae and also in Cercocarpus and Adenostoma of the Rosoideae
and Quillaja of the Spiraeoideae the vessels are almost exclusively solitary.
In the latter two tribes the vessels in the other genera are often grouped
tangentially or in clusters.
Spiral thickening of the vessels and distribution of parenchyma have been
suggested by Janssonius (1154) as a means of distinguishing between the
tribes, but have not proved to be valid characters for the larger number of
genera and species here investigated.
Lyonothamnus. This genus, mentioned by Dalla Torre et Harms under
Rosaceae as a genus of uncertain position, is referred by Britton and Shafer,
North American Trees, p, 408 (1908), to the family Cunoniaceae. Its wood
anatomy is quite consistent with its being a member of the Rosaceae, though
not of the tribe Prunoideae. It does not fit so well in the Cunoniaceae.
Prunus and Laurocerasus. Though it is not possible to separate all the
specimens of these 2 genera into 2 sharply defined groups there does appear
to be an appreciable divergence. Laurocerasus tends to have very definitely
heterogeneous rays, frequently with 4 or more marginal rows of upright cells,
and some vasicentric parenchyma, and some species (L. offidnalis Roem., L.
lusitanica Roem., and L. lyoni Britton) have a marked radial vessel pattern.
In Prunus the rays are more homogeneous, the marginal rows seldom being
more than 1-3 and their cells square rather than upright, the parenchyma
is diffuse only and often very sparse; some species have an oblique vessel

pattern, but this is less definite and more nearly tangential than in the extreme
forms of Laurocerasus. Examples of the tendency for the 2 groups to overlap
are provided by the following species: Laurocerasus maackii (Rupr.) C. K.
Schn., which has the structure typical of a Prunus, and Prunus annularis
Koehne, which might be a typical Laurocerasus.

ECONOMIC USES
Many species are cultivated as ornamental shrubs, whilst other members
of the family yield edible fruits such as the Loquat (Eriobotryajaponica Lindl.),
Strawberry (Fragaria vesca Linn.), Bitter Almonds (Prunus amygdalus Batsch.
var. amara Schneid.), Sweet Almonds (Prunus amygdalus Batsch. var. dulcis
Schneid.), Apricot (Prunus armeniaca Linn.), Plum (Prunus domestica Linn.),
Peach (Prunus persica Batsch), Pear (Pyrus communis Linn.), Apple (Malus
pumila Mill.), and Blackberry and Raspberry (Rubus spp.). Other economic
products include Quillaja bark (Quillaja saponaria Mol.), which is used as a
commercial source of saponin and for medicinal purposes. This bark can be
recognized by a narrow layer of cork cells with red-brown contents; tortuous
bundles of phloem fibres; abundant starch grains, usually 5-10 but sometimes
up to 20 in diameter; calcium oxalate crystals 170 p long and 30 ft wide.
fji
 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE) 549
The existence of several types of Quillaja bark, exhibiting variations in struc-
ture, has been reported, but these are probably from branches of different age
or from plants grown in various habitats. For further particulars see papers
by Holmes (1075, IO?6) and Cofman-Nicoresti and Tallantyre (440).
Beakbane and Thompson (159) have shown that the 'rubbery' condition of
certain parts of the wood of 'Lord Lambourjie' and other cultivated varieties
of apple-trees is due to lack of lignification of the wood fibres. In consequence
the affected stems and branches are abnormally flexible. In other cases
unusually flexible stems of apple and pear varieties and of some apple root-
stocks were found to be due to an abnormally large pith, or to a high propor-
tion of living cells in the wood. When the 'rubbery* condition is due to
imperfect lignification, this can be immediately demonstrated by treatment
with phloroglucinol and hydrochloric acid. The cause of the disease is at
present uncertain. (See also under 'Root'.)
Other recent work on the anatomy of rosaceous fruit-trees includes the
investigation by MacDaniels and Cowart (1407) concerning the structure and
development of the apple leaf, and that by Schneider (2046) dealing with the
structure of the phloem in peach and cherry.
The anatomy of species of Dryas, Geum, and Sieversia which are used in
folk medicine has recently been described by Schulthess (2048).
The woods of this family are not of great importance. The best-known
timbers are probably various cherries (Prunus spp.), particularly the American
Black Cherry, P. serotina Ehr., and the European Cherry, P. avium L., which
provide ornamental timbers. The wood of the Pear, Pyrus communis L., has
some special uses, e.g. as a substitute for boxwood for engraving.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Acaena, Agrimonia,* Alchemilla,* Amelanchier,* Chaenomeles, Coton-
easter,* Cliffortia, Crataegomespilus,* Crataegus,* Cydonia,* Dichoto-
manthes,* Eriobotrya,* Filipendula, Fragaria,* Geum,* Gillenia,* Hetero-
meles, Kageneckia, Kerria,* Laurocerasus,* Lindleya, Mespilus, Neillia,*
Neurada, Neviusa,* Osteomeles, Photinia,* Potentilla,* Poterium,* Prinsepia,
Prunus,* Pygeum, Pyracantha,* Pyrus,* Quillaja,* Raphiolepis,* Rosa,*
Rubus,* Sanguisorba, Sibiraea, Sorbus,* Spiraea,* Stranvaesia,* Vauquelinia,
Waldsteinia.
* * Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Adenostoma, Amelanchier, Cercocarpus, Chamaebatia, (Cliffortia),
Cotoneaster, Cydonia, Eriobotrya, Eriolobus, Exochorda,
Crataegus,
Fallugia, Hagenia, Heteromeles, Holodiscus, Kageneckia, Kerria, Lauro-
cerasus, Lyonothamnus, Malus, Mespilus, Micromeles, Nuttallia, Osteomeles,
Padus, Photinia, Physocarpus, Polylepis, Potentilla, Pourthiaea, Prunus,
Pseudocydonia, Purshia, Pygeum, Pyrus, Quillaja, Raphiolepis, Rosa, Rubus,
Sorbaria, Sorbus, Spiraea, (Vauquelinia).
550 ROSACEAE (EXCLUDING CHRYSOBALANOIDEAE)
LITERATURE
(i) On
General Anatomy (including Chrysobalanoideae)
Aubertot 48, Batrany 152, Beakbane etal. 156, 157, 158, 159, 160, 161, Betts 187, 188,
189, 190, Bird 199, Bowman 250, Brierley 269, Brooks 278, Butler 324, Carlson 342,
Cofman-Nicoresti and Tallantyre 440, Dorsey and Weiss 607, Esau 659, Foweraker 703,
Gleichgewicht 789, Gregory 815, Greguss 2522, Gyhr 854, Haberlandt 858, 859, 860,
861, Holm 1028, 1039, 1045, Holmes 1075, 1076, Knapheisowna 1249, Lepeschkin 1361,
Low 1397, MacDaniels and Co wart 1407, McCown 1464, Montcheff 1553, Morvillez 1556,
I 557 Price 1741, Priestley and Hinchliff 1751, Prodinger 1760, Rinde 1939, Sabnis 1977*
Schneider 2046, Schulthess 2048, Starr 2188, Watkins 2367, Weiss 2391, White 2420,
Woodcock 2460, Youngken 2496.

(ii) On Wood Structure (including Chrysobalanoideae)

Bausch 154, Beakbane et al. 156, 157, 161, Beekman 167, Benoist 170, den Berger 179,
182, den Berger and Endert 183, Besson 186, Bianchi 194, Bienfait and Pfeiffer 197, B.
H. For. Dept. 274, Brown and Panshin 288, 289, Burgerstein 310, 312, Chalk and Rendle
365, Coster 481, Cozzo 494, Dadswell and Record 533, Descole and O'Donnell 576, Fox-
worthy 705, Gonggrijp 794, Hale 870, Howard 1088, van Iterson 1126, Janssonius 1154,
Jones 1191, Jonsson 1192, Kanehira 1206, 1209, Lecomte 1334, Macduffie 1408, Malhorta
1430, Martin-Levigne 1450, NicolofT 1593, Pearson and Brown 1679, PfeifTer, J, Ph. 1713,
Record 1780, 1781, 1783, 1787, 1800, 1801, 1818, 1843, 1851, Record and Hess 1886,
Stone 2202, 2203, Tang 2230, 2231, Thompson 2254, Tippo 2261, Torres 2269, Uspensky
2317, Williams 2430, Yamabayashi 2478, Yatsenko Khmelevsky and Djaparidze 2488.

117 A. ROSACEAE (CHRYSOBALANOIDEAE)

(FiG. 121 on p. 552)

SUMMARY
(i) GENERAL
A tropical, chiefly American family, consisting of trees and shrubs. The
stomata are rubiaceous. Transverse sections through the distal end of the

petiole generally exhibit a closed ring of xylem and phloem accompanied by

two small adaxial vascular strands, but numerous modifications of the closed
ring are to be found in different genera or even in different species of a single
genus. The cork generally arises superficially. The pericycle exhibits
a ring of sclerenchyma which is usually continuous, but may be broken.
Stone cells, which occur in the pericycle, are generally characterized by being
thickened only on one side, thus appearing U-shaped in transverse section.
Silicified cell membranes and solid inclusions of silica are common.

(ii) WOOD
Vessels moderately to very large, almost exclusively solitary, few to
moderately numerous, typically in oblique lines, spiral thickening absent,
perforations exclusively simple, pits to ray cells always including some that
are larger than the intervascular pitting; members of medium length. Paren-
chyma apotracheal, predominantly in uni- to triseriate bands. Rays
exclusively or predominantly uniseriate; heterogeneous. Fibres with numer-
ous distinctly bordered pits; of medium length.

LEAF
Whole mesophyll usually consisting of palisade tissue except in Lecostemon.
Long, unicellular, arachnoid hairs with thin walls form a web-like covering
to the leaves of Couepia^ Licanta, Moquilea, Parinari\ stellate hairs occur in
 ROSACEAE (CHRYSOBALANOIDEAE) 551

Chrysobalanus, and peltate ones in Lecostemon. Glands present at the leaf

base of certain species of Parinari, and especially large and conspicuous ones
in Acioa, Chrysobalanus, Couepia (pro parte), Hirtella, Licania, Moquilea.
Cork warts recorded on the leaf of Couepia bracteosa Benth. Epidermis
composed of cells with mucilaginous walls in species of Chrysobalanus,
Grangeria, Hirtella, Parinari', papillose on the lower surface in Couepia.
Epidermal cells frequently palisade-like. Hypoderm present in Chrysoba-
lanus, Couepia, Grangeria, Hirtella, Lecostemon, Licania, Moquilea, Parastemon,
Parinari. Stomata recorded only on the lower surface except in Stylo-
basium\ rubiaceous. Mesophyll traversed by fibre-like 'spicular cells' in
Couepia, Lecostemon, and Licania. Vascular bundles of the veins (except in
Stylobasium) surrounded by sclerenchymatous fibres, which exhibit U-shaped
thickenings in transverse sections; vertically transcurrent in certain species of
Acioa, Chrysobalanus, Couepia, Hirtella, and Licania. Terminal tracheids
with wide lumina in Licania and Moquilea. Transverse sections through the
distal end of the petiole described by Morvillez (1557) as exhibiting an
abaxial closed ring of xylem and phloem which shows various modifications
towards the adaxial side in different genera and species. The ventral strand,
in all recorded instances, accompanied by 2 small separate adaxial bundles.
Modifications are as follows, (i) The main ring of xylem and phloem some-
what invaginated on the adaxial side in Hirtella. (ii) As (i) but with the
invagination expanded laterally at the base in Parinari. (iii) Main vascular
strand in Licania enclosing a solid medullary bundle with central xylem sur-
rounded by phloem, (iv) Similar to (iii) but with the xylem of the medullary
strand confined to the adaxial surface of the phloem in Moquilea guianensis
Aubl. (v) Main vascular strand consisting of 2 closed concentric rings of
xylem and phloem surrounding 2 small, solid bundles in Moquilea sclerophylla
Mart, (vi) Main vascular strand enclosing i large and several small solid
strands in Moquilea licaniflora Sagot. (vii) Main vascular strand enclosing a
greatly reduced central bundle consisting of an island of phloem in Chryso-
balanus icaco Linn. Transitional stages between these types also occur.
Silicified membranes common except in Lecostemon and Stylobasium',
silica bodies recorded in the epidermal cells and in the neighbourhood of the
leaf veins of all examined genera except Parastemon and Stylobasium, Similar
bodies also occur in the peltate hairs of Lecostemon in the mesophyll of the
;

leaf of some species of Couepia, Lecostemon, and Moquilea. Silica deposits in

certain species of Chrysobalanus and Hirtella sometimes enclose clustered
crystals. Secretory cavities recorded in Couepia bracteosa Benth. and in
Lecostemon spp.

Axis
YOUNG STEM
Cork, generally superficial in origin. Pericycle including strands of
primary fibres which subsequently become united by stone cells, the latter
showing horseshoe-shaped thickenings in transverse sections. Xylem in the
form of a continuous cylinder traversed by narrow rays. Vessels with simple
perforations. Idioblasts containing tannin present in the phloem of certain
species of Couepia, Moquilea, and Parinari. Silica bodies stated to occur in
the pericycle, medullary rays, and pith of all the genera examined except
 552 ROSACEAE (CHRYSOBALANOIDEAE)
Parastemon and Stylobasium. Similar bodies also recorded in the bark of
Hirtella americana Linn,

WOOD (Fig. 121)

Vessels mostly moderately large (mean tangential diameter 200-300 mm.),
very large in Chrysobalanus and Parinari] exclusively solitary or nearly so;
typically in oblique lines; few in most species (less than 5 per sq. mm.);
moderately numerous (5-20) in Licania and Chrysobalanus spiral thickening
;

absent; perforations exclusively simple; intervascular pitting alternate, never

FIG. 121. ROSACEAE CHRYSOBALANOIDEAE

A, Licania heteromorpha Benth. B, L. hypoleuca Benth. C, Parinari asperulum Miq.
D, P. mobola Oliv.

minute; some of the pits to ray cells and parenchyma always larger than the
intervascularpits and oblong; tyloses sometimes present, sclerosed in
Angelesia. Mean member length 0-6-0-8 mm. Parenchyma apotracheal
only, in numerous fine continuous bands, usually (locally 2) cells wide and
i

6-1 1 per mm.; 2-3 cells wide and usually slightly less numerous (less than
6 per mm.) in Acioa and the African species of Parinari, strands
typically of
up to 1 6 cells. Rays fine, exclusively uniseriate in Chrysobalanus, Couepia,
Grangeria, Hirtella, and Licania, and predominantly uniseriate, but with some
biseriate rays in the others; more than i mm.
high in Acioa, Couepia, Licania,
Parastemon, and some species of Parinari; 12-20 per mm. ; heterogeneous
Kribs's Type III; almost homogeneous in Parastemon. Silica is
reported in
the ray cells of Angelesia splendens Korth., Parastemon
inophyllum A. DC.
(794) and Parinari spp., and van Iterson (1126) states that the Chryso-
balanoideae is very rich in silica inclusions. Fibres with numerous
distinctly
bordered pits on the tangential walls, more numerous and often biseriate in
the wider cells bordering the vessels;
pits on the radial walls almost entirely
limited to areas in contact with rays; walls thick
(except in Couepia and some
species of Licania). Mean length 1-3-1 -4 mm.
 ROSACEAE (CHRYSOBALANOIDEAE) 553
TAXONOMIC NOTES
FROM WOOD STRUCTURE
The genera of this group are very uniform in structure and form a distinct
group that is easily separated from the rest of the Rosaceae; the most charac-
teristic features are the banded apotracheal parenchyma, the oblique pattern
of the vessels, the almost exclusively uniseriate rays, and the long parenchyma
strands of up to 16 cells.
There is no very distinct evidence of this group being at a different level of
specialization from the rest of the Rosaceae. As Tippo (2261) has pointed out,
both are more specialized than the Cunoniaceae.

ECONOMIC USES
The Coco Plum is The bark of the
derived from Chrysobalanus icaco Linn.
Caraipi tree of Para (Moquilea utilis Hook, f.) is mixed with clay to make
vessels for domestic use. The timbers are very similar throughout the group.
They are not widely used.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Acioa, Chrysobalanus, Couepia, Grangeria, Hirtella, Lecostemon, Licania,
Moquilea, Parastemon, Parinari, Stylobasium.

(ii) FOR WOOD STRUCTURE

Acioa, Angelesia, Chrysobalanus, Couepia, Grangeria, Hirtella, Licania,
Parastemon, Parinari.

LITERATURE, see Rosaceae, p. 550

118. SAXIFRAGACEAE
(FiG. 122 on p. 556)

SUMMARY
Herbs, sometimes tending to be succulent. The family occurs particularly
in temperate and cold regions. The leaf is dorsiventral or centric. Both
glandular and non-glandular types of hair are common. The leaf epidermis
sometimes includes a proportion of elongated cells with tanniniferous con-
tents. The stomata are usually ranunculaceous, but their distribution may
be of considerable specific diagnostic value. Hydathodes are common, and
in some instances secrete water containing calcium bicarbonate which becomes
deposited on the leaf surface as solid carbonate. Hydathodes which secrete
lime in this way are known as chalk glands. Crystals are rare, but clustered
where present. The occurrence in the petiole of one or more separate
bundles, each surrounded by an endodermis, is an interesting feature of
certain species of Saxifraga and somewhat recalls the petiolar structure in
y

Akhemilla (Family Rosaceae). The stem exhibits a vascular system com-

posed of individually distinct bundles in some species; in pthers the primary
rays are narrower and the xylem continuous. There is frequently a continuous
554 SAXIFRAGACEAE
ring of fibres in the pericycle. Cortical and/or medullary bundles have
been recorded. The nodes of Parnassia palustris Linn, are said to be polystelic.

LEAF
Dorsiventral in Astilbe, Chrysosplenium, Heuchera, Hoteia, Mitella, Saxi-
fraga (pro parte), Tellima, Tiarella, Zahlbrucknera', isobilateral or centric in
Saxifraga (pro parte, especially the section Porphyrion), and Vahlia. Hairs
glandular and non-glandular.
I. (i) Simple,
Non-glandular, uniseriate common in Saxifraga. (ii) Shaggy
;

in Saxifraga. Multicellular, multiseriate recorded in Astilbe, Boykinia,

(iii)

Heuchera, Peltiphyllum, Tiarella, Tolmiea. (iv) Fimbriate appendages, secret-

ing mucilage when young, in Parnassia.
II. Glandular, (i) Uniseriate with a unicellular or multicellular head in
Boykinia, Francoa, Heuchera, Peltiphyllum, Saxifraga, Suksdorfia, Tetilla,
Tiarella, Tolmiea, Vahlia, Zahlbrucknera. (ii) Glandular shaggy, with a multi-
cellular stalk and head in Astilbe, Hoteia, Mitella, Saxifraga, Tellima, Tiarella.

Epidermis composed of cells with straight or sinuous anticlinal walls.

Long vermiform cells with tanniniferous contents occur between and some-
times project above the ordinary epidermal cells in species of Chrysosplenium,
Lepuropetalon, Parnassia, Saxifraga (section Cymbalaria), Zahlbrucknera",
frequently recognizable in herbarium material as small brown striae. Stomata
present on one or both surfaces; ranunculaceous, e.g. in the Dactyloides,
Nephrophyllum, and Miscopetalum sections of Saxifraga', subsidiary cells
sometimes smaller than their neighbours, e.g. in Saxifraga, section Euaizoonia.
The distribution of stomata is probably of considerable specific diagnostic
value in Chrysosplenium and Saxifraga. Stomata confined to the upper sur-
face in some species of Saxifraga occurring on both surfaces at the tips of the
;

leaves in others; restricted to the leaf margins in a third group; confined to

the middle on the upper and to the margins on the lower surface in a fourth
group. Stomata in Chrysosplenium restricted to the lower side in the Alterni-
folia group, but occurring on both surfaces in the section Dialysplenium',
occasionally in groups in both sub-genera. Hryniewiecki (noo) records the
existence of guard cells with a distinctive type of structure in various members
of the family, but the utility of this character for diagnostic purposes is limited
by the difficulty of rapidly cutting sections of sufficiently good quality to show
what is required. Hydathodes recorded in Chrysosplenium, Heuchera,
Mitella, Saxifraga, Tellima, Zahlbrucknera, and probably occurring in other
genera as well, although apparently absent from Bergenia\ those in Saxifraga
mostly situated at the leaf margins, at the. tip of the leaf, or at the bases of the
leaf teeth, sometimes in depressions (e.g. in the sections Euaizoonia, Kabschia,

Porphyrion, Xanthizoori), more rarely on the leaf surface always above the ;

terminations of veins. Hydathodes sometimes secrete lime in the form of a

solution of calcium bicarbonate, from which solid calcareous deposits are
formed on the surface of the plant. Hydathodes which secrete calcareous
material in this way are known as chalk glands. The quantity of lime secreted
isgoverned by the nature of the soil. Further details concerning hydathodes
and chalk glands have been recorded by Kurt (1308). Mesophyll not pos-
 SAXIFRAGACEAE 555

sessing great diagnostic value owing to its variability in different species and
also in relation to the environment; not differentiated into palisade and
spongy portions in certain species of Saxifraga. Number of palisade layers,
where differentiated, ranging from i to 7. Petiole. Structure, in transverse
section, somewhat distinctive, especially in Saxifraga, owing to the presence
of at least one concentric or hemi-concentric bundle. Many species of Saxi-
fraga (Fig. 122 K) are provided with 3 such bundles each surrounded by a
separate endodermis, and in this respect resemble Alchemilla (Fig. 1190)
(Family Rosaceae). Three separate bundles also observed in species of
Heuchera (Fig. 122 i) but in this genus they are collateral. The thick, fleshy
petiole of Bergenia delavayi (Franch.) Engl. (Fig. 122 M) is supplied by
numerous, irregularly scattered bundles. For further details concerning the
vascular structure of the petiole see Morvillez (1559). Secretory cells with
unidentified but probably tanniniferous contents seen to be common in the
unlignified tissues of the petiole and recorded in species of Chrysosplenium,
Lepuropetalon, Parnassia (Thompson 2252), Saxifraga (Cymbalaria), and
Zahlbrucknera. Clustered crystals observed or recorded in a few species;
rare in Saxifraga.

Axis
STEM (Fig. I22N)
Cork usually arisingin the outermost layer of the pericycle ; but originating
in the sub-epidermis in Peltiphyllum and in the epidermis itself in Vahlia.
Cortex sometimes very lacunar, e.g. in Bergenia delavayi (Franch.) Engl.
Pericycle containing a broad continuous ring of fibres in the few species of
Heuchera and Saxifraga examined; fibres confined to separate strands at the
outer periphery of the phloem in Bergenia delavayi. According to Engler
(636) sclerenchyma is absent from the flowering stem of the Cymbalaria
section of Saxifraga and from certain species in other sections of the genus as
well. Phloem and xylem usually appearing in transverse sections as
individually distinct collateral bundles (Fig. I22N), but the primary rays
between them are sometimes rather narrow. There are considerable variations
in this respect even within the genus Saxifraga. For further details see Engler
(636). Vessels usually small; radial diameter not exceeding 15 ^ in Bergenia
delavayi up to about 30 JJL in Heuchera spp. Only simple perforations ob-
\

served. Pith observed to be very lacunar in species of Bergenia, Heuchera,

and Saxifraga. Medullary bundles recorded in a considerable number of
species of Saxifraga, particularly but not exclusively in the section Euaizoonia
as well as in Peltiphyllum and Rodgersia. Cortical bundles also reported
in Peltiphyllum. Stems polystelic at the nodes in Parnassia palustris Linn.
Secretory cells with tanniniferous contents recorded in the unlignified
tissues in species of Chrysosplenium, Lepuropetalon, Parnassia, Saxifraga,
Zahlbrucknera, and also observed in Heuchera.

TAXONOMIC NOTES
The Saxifragaceae in the Bentham and Hooker system included plants
which have been described in this book under Cunoniaceae, Grossulariaceae,
Escalloniaceae, and Hydrangeaceae. Engler likewise (636) included all of
these groups under Saxifragaceae, but he recognized the existence of a number
 FIG. 122. HYDRANGEACEAE, A, D, F-H, J; ESCALLONIACEAE, B-C and E;
SAXIFRAGACEAE, I, K, and M-N; GROSSULARIACEAE, L and O
A, Dentzta setchuensis Franch. Petiole x 13. B, Escalloma rubra Pers. Petiole X 12. C, Brexia
madagascariensis Thou. Petiole X 12. D, Hydrangea arborescens Linn. Petiole X 13. E, Escalloma
;

macrantha Hook, et Am. Stem x 13. F, Hydrangea petiolaris S. et Z. Petiole x 13. G, Philadelphia
pubescent Lois. Stem X 15. H, Hydrangea arborescens Linn. Stem X 13. I, Heuchera americana Linn.
Petiole X 7. J, Dichroafebrifuga Lour. Petiole X 7. K, Saxifragafortunei Hook. Petiole X 8. L, /?i'6
sangwneum Pursh. Petiole X 18. M, Bergema delavayi Engl. Petiole X 5. .2V, Saxifragafortunei Hook.
Stem Xi.i. O. Ribes zrossularia Linn. Stem Xi8.
 SAXIFRAGACEAE 557
of sub-families. Hutchinson treats the Saxifragaceae (sensu stricto) as if
evolved along a different line from the Cunoniales in which he includes the
other families mentioned above. It is thus clear that taxonomists are by no
means agreed concerning the interrelations of the groups concerned. It will
at once be appreciated that the Saxifragaceae, as treated in this book, constitute
a wholly herbaceous family, whereas the other groups in question are com-
posed of woody plants. For this reason alone one would expect to find
anatomical distinctions between them, but in practice the differences are not
of a kind which are usually to be found between herbs and woody plants
which are closely related to one another. It has been questioned by Arber
(see 2254) whether Parnassia truly belongs to the Saxifragaceae or has closer
affinities with the Hypericaceae. It is sometimes treated as a member of a

separate family, the Parnassiaceae.

ECONOMIC USES
Numerous members of the family are cultivated for ornamental purposes,
especially in rock-gardens.

GENERA DESCRIBED
Astilbe, Bauera, Bergenia,* Boykinia, Chrysosplenium, Colmeiroa, Dei-
nanthe, Francoa, Heuchera,* Hoteia, Lepuropetalon, Mitella, Parnassia,
Peltiphyllum, Rodgersia, Saxifraga,* Suksdorfia, Tellima, Tetilla, Tiarella,
Tolmiea, Vahlia, Zahlbrucknera.
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Briquet 271, Engler 636, Engler and Irmscher 644, Hryniewiecki iioo, Kurt 1308,
Morvillez 1559, Smith, H. 2150, Thompson, H. S.

119. CEPHALOTACEAE
(Fic. 123 on p. 558)

SUMMARY
is a
Cephalotus follicularis LabilL, the sole representative of the family,
perennial herb with an underground rhizome, the lower of whose basal leaves
are differentiated as pitchers similar to those of Nepenthes, the upper leaves
of the basal rosette being ordinary photosynthetic organs. The plant is con-
fined to marshes in King George's Sound, South-west Australia. The most
in the pitchers which
interesting anatomical feature is the occurrence of glands
are thought by some authorities to secrete substances capable of digesting
animal food. The morphology and anatomy of the plant has been recently
surveyed by Lloyd (1383), whilst an earlier account by Macfarlane (1411)
includes a very complete description of the anatomy of the whole plant.
Another important description was published by Schweiger (2063). The
particulars given below have been taken from these
sources.
 B

FIG. 123. HYDRANGEACEAE, A-C; ESCALLONIACEAE, D; CEPHALOTACEAE, E

A, Simple trichome of X Philadelphus insignis Carr. B, Stellate hair from the upper aide of the
leaf of Deutzia pulchra Vidal. C, Stellate hair from the upper side of the leaf of Deutzia gracilis Sieb.
et Zucc. D, Stem-gland of Escallonia viscosa Forbes. E, Gland of Cephalotus A-C, by Solereder;
D, after Thouvenin; E, After Goebel.

FIG. 124. CRASSULACEAE

A, of stoma in Sedum spurium Marsch. Bieb. B, C, Bladder-like epidermal cells of Crasntla
Type
falcata Wendl. D, Transverse section through the epidermis of the anterior portion of the leaf of
Sempervivum calcareum Jord. A after Strasburger; B, C after Areschoug; D
after Solma-Laubach.
 CEPHALOTACEAE 559
LEAF
(a) SCALE LEAVES ON ELONGATED PARTS OF THE Axis
Unicellular hairs present on both surfaces and along the margins; accom-
panied by glands similar to those on the foliage and pitcher leaves. Meso-
phyll poorly developed, traversed by a few, small vascular bundles.

(b) FOLIAGE LEAF

Unicellular hairs with solid tips and brownish contents towards the base,
present on the petiole and at the margins of the lamina. Glands (Fig, 123 E)
also occur on the petiole and lower surface of the lamina. Stomata situated
on both surfaces, but most numerous on the lower side and also occurring on
the petiole. Mesophyll approximately isobilateral, but palisade cells some-
what rounded and not quite equally developed towards both surfaces; central
portion composed of very lacunar spongy tissue. According to Macfarlane
(1411) the single bundle system which enters the base of the petiole 'splits
into one median and two lateral parts. The median part consists of four
bundles so placed that two of them are lateral, one inferior and one superior,
but with their xylems inclined towards each other, and their extra-phloem
sclerenchyma patches directed outwardly/

(c) PITCHER LEAVES

Epidermis on the outside of the pitcher composed of isodiametric cells,
perforated by stomata and bearing depressed glands and unicellular hairs.
Inner surface of the lid consisting for the most part of a so-called 'slippery
surface* composed of downwardly projecting, overlapping, striated projec-
tions from the epidermal cells. Glands (Fig. 123 E) similar to those on the
external surface also present. Glands in the interior of the pitcher flask-
shaped; those present on the brightly coloured, cushion-like projections,
which extend into the interior of the pitcher, being especially large. Hyda-
thodes also recorded on the projections. Glands said to be absent from the
lower part of the internal wall of the pitcher. Full details concerning the
vascular system of the pitcher are given in the special articles mentioned
above, but it is interesting to note that Lloyd says: The venation is derived
from two systems of bundles in the petiole, a dorsal of three veins, and a
ventral of two, these splitting near the pitcher into four, then six and branch-
ing further in spreading.'

Axis
RHIZOME
Epidermis young rhizome bearing long, unicellular hairs. Cork
in the
formation is Cortex fairly broad; amyliferous
initiated during the first year.

according to Macfarlane (1411) and tanniniferous according to Schweiger

(2063). Vascular system composed of a ring of phloem and xylem inter-
rupted by broad, foliar traces. Pith similar to the cortex. Brown deposits
present in the epidermis, cortex, and pith of old rhizomes.

ROOT
Epidermis pale coloured. Cortex composed of 2 or 3 layers of cells.
Endodermis consisting of flattened cells. Cork arising in the pericambium,
560 CEPHALOTACEAE
consisting, in mature roots, of 3 or 4 layers of cells, but according to Macfar-
lane (1411) the endodermis and cortical tissues do not become detached.
Schweiger (2063), on the other hand, describes an old root as covered
externally by a layer of cork bounded on the inside by rounded, mostly
tanniniferous cells. Stele triarch. Xylem vessels described by Schweiger as

having scalariform perforation plates, but this needs confirmation.

TAXONOMIC NOTES
The
similarity of the pitchers of Cephalotus to those of Sarracenia and
Nepenthes is probably due to parallel development. There are no close
taxonomic affinities between these plants.

GENUS DESCRIBED
Cephalotus.
LITERATURE
On General Anatomy
Arber 32, Diels 584, Lloyd 1383, Macfarlane 1411, Schweiger 2063.

120. CUNONIACEAE
(Fic. 125 on p. 562; FIG. 126 on p. 570)

SUMMARY
(i) GENERAL
Trees and shrubs mainly from Australasia. Some of the more important
anatomical characters include the occurrence of specially small stomata
whose guard cells are almost circular in outline; in twigs, the xylem with
uniseriate rays and small, often somewhat angular vessels, mostly with
scalariform perforation plates; the heterogeneous pith; the common occur-
rence of secretory cells with tanniniferous or mucilaginous contents.

(ii)
WOOD
Vessels moderately small,solitary or with some multiples, perforation
plates scalariform only or simple with a few scalariform plates, rarely simple
only; intervascular pitting scalariform to opposite, rarely alternate; pits to
parenchyma and horizontally elongated members moderately
typically large ;

long. Parenchyma apotracheal, diffuse or in bands 1-4 cells wide. Rays

up to 2-4 cells wide, markedly heterogeneous in most species. Fibres usually
with distinctly bordered pits; of medium length.

LEAF
Dorsiventral in all of the species examined. Hairs sometimes absent, but
chiefly represented by infrequent unicellular types. Tufted hairs recorded
in CalKcoma, and glandular shaggy ones in Ceratopetalum and Cunonia, the
latter occurring particularly on the stipules and leaf teeth. Cells of the
epidermis elongated and palisade-like in Codia montana Forst., Cunonia
(locally), Wtinmannia trichosperma Cav. Stomata particularly small and
 CUNONIACEAE 561
almost circular in outline in Cunonia and Platy tophus; guard cells of Belangera
with ridge-like humps which appear like small horns in a transverse section
of a leaf. Hypoderm, frequently consisting of 3 layers of cells towards the
upper surface, recorded in species of Anodopetalum, Callicoma, Codia
(resembling stone cells), Cunonia, Platylophus, Pullea, Weinmannia. A tanni-
niferous hypoderm towards the lower surface recorded by Betts (187) in
Weinmannia racemosa Linn. Cells of the epidermis and hypoderm provided
with mucilaginous inner walls in all of the species examined. Mesophyll
containing 'spicular cells' in Pancheria sp. Smaller veins vertically trans-
current or embedded. Petiole. Transverse sections through the distal end
supplied by an almost continuous, adaxially flattened, cylindrical strand in
Cunonia capensis Linn. (Fig. 126 c), but with the flat dorsal portion separated
from the ventral arc. Structure somewhat similar in Weinmannia tricho-
sperma (Fig. 126 B) but vascular strand continuous and rather different in
shape. Small additional strands present in the wings in Cunonia capensis.
Main vascular strand in both species strongly supported by a ring of peri-
cyclic fibres. Secretory cells with amorphous, presumably tanniniferous
contents present in the unlignified tissues of the petiole in Weinmannia
trichosperma', similar cells but with less definitely tanniniferous contents
observed in the cortical and medullary regions of the petiole in Cunonia
capensis. Solitary and clustered crystals common in the same 2 species.

Axis
YOUNG STEM (Fig. 126 F)
Cork superficial, arising in the epidermis or sub-epidermis. Cortex some-
what spongy and composed of small cells in Cunonia capensis Linn, and
Weinmannia trichosperma Cav. Pericycle containing a somewhat interrupted
or continuous, sometimes composite ring of sclerenchyma. Secondary
phloem including sclerosed cells in species of Callicoma, Cunonia, Geissois,
Pancheria, Weinmannia. Phloem and xylem constituting continuous
cylinders, traversed by narrow rays. Vessels observed to be angular and up
to about 40 IJL in radial diameter in Cunonia capensis and Weinmannia tricho-
sperma. Exclusively scalariform perforation plates recorded in Anodopetalum,
Caldcluvia, Callicoma, Codia, Cunonia, Platylophus, Weinmannia and mixed
simple and scalariform plates in Belangera and Ceratopetalum. Solereder
records the occurrence of fibres with bordered pits in species of Anodo-
petalum, Caldcluvia, Ceratopetalum, Cunonia, Pancheria, Platylophus, and
Weinmannia, and simple pits in Belangera. Pith of Cunonia capensis and
Weinmannia trichosperma quadrangular in transverse section, composed of
cells larger in transverse diameter than those in other parts of the stem, many
of them provided with pitted walls, mostly devoid of contents but some cells,
filledwith amorphous deposits, arranged in vertical columns. Other secre-
tory cells, sometimes with mucilaginous contents, observed in the cortex,
medullary rays, and pith of Cunonia capensis, and similar cells, but with
apparently tanniniferous contents, in the phloem as well as the same tissues
in Weinmannia trichosperma. Solitary and clustered crystals common in the

unlignified tissues.
4594 OO
 FIG. 125. CUNONIACEAE, A-G; EUCRYPHIACEAE, H-J*
ESCALLONIACEAE, I and K-L
A, Weinmannia blumei Planch. B, Platylophus trifoliatw D. Don. C, Ceratopetalum apetalum D
Don. D, C. apetalum D. Don. E, Schizomeria pulleana O. Schmidt, F, Cunonia
capensis Linn.
G, C. capensis Linn, H, Eucryphia lucida (Labill.) Bail!. I, Quintinia sieberi A. DC. J, Eucryphia
luctda (Labill.) Baill. K, Polyosma laete-virens Griff.
L, P. laete-virens Griff.
 CUNONIACEAE 563
WOOD (Fig. 125 A-G)
Vessels typically moderately small (50-100 \i mean tangential diameter),
slightly smaller in Anodopetalum and Callicoma (527) and rather larger in
some species of Ceratopetalum, Geissois, and Spiraeopsis solitary, apart from
\

the apparent pairs produced by overlapping ends, in Caldcluvia, Callicoma^

Cunonia, and Weinmannia, multiples of 2-3 cells common in the other genera,
particularly in Belangera (1886), Geissois, Schizomeria, and Spiraeopsis, with
multiples of 2-9 cells in Geissois and Schizomeria, with a tendency to a tan-
gential pattern in Platylophus 8-130 per sq. mm., fewer than 20 per sq. mm.
\

in Geissois, Schizomeria, and Spiraeopsis, about 100 per sq. mm. in Anodo-
petalum, Caldcluvia, Callicoma, and Platylophus (some specimens) semi-ring- ;

porous in Anodopetalum and Platylophus. Perforation plates scalariform only

and usually with fewer than 20 bars in Ackama, Caldduvia, Callicoma,
Cunonia, and Weinmannia, 1 with both simple and scalariform plates, the
latter often limited to the smaller vessels, in Anodopetalum, Belangera (2158),

Ceratopetalum, Geissois, Platylophus, Schizomeria, and Spiraeopsis, with simple

perforations only in some species of Spiraeopsis. Intervascular pitting pre-
dominantly scalariform in Ackama, Anodopetalum, Caldcluvia, and Cerato-
petalum, opposite to alternate in Geissois and Schizomeria, intermediate
between scalariform and opposite in the other genera pits to ray and wood
;

parenchyma typically large and horizontally elongated, sometimes exclusively

circular in Schizomeria. Tyloses observed in Caldcluvia, Platylophus,
Spiraeopsis, and Weinmannia. Mean member length 0-75-1-0 mm. Paren-
chyma apotracheal, as scattered cells (diffuse) in genera with solitary vessels,
e.g.Ackama, Caldcluvia, Callicoma, Cunonia (some specimens) (Fig. 125 G),
Platylophus, and Weinmannia, in numerous, irregular, uniseriate bands in
Anodopetalum, Cunonia, Geissois, and Spiraeopsis, those of Anodopetalum
almost entirely limited to the late wood; in wider and sometimes more
regular bands 2-5 cells wide in Ceratopetalum and Schizomeria (Fig. 125 D-E),
the short lines in Weinmannia and the broader bands in Schizomeria tending
to be associated with the abaxial sides of the vessels (Fig. 125 A). Frequently
with gummy contents chambered crystals observed or reported in Ackama
;

(525), Caldcluvia, Callicoma (2158), Ceratopetalum, Cunonia (2158), Geissois,

Spiraeopsis, and Weinmannia. Strands usually of 4-8 cells. Rays of 2 distinct
sizes in some genera, e.g. Callicoma and Cunonia, and with a tendency towards
this in other genera; 2-4, occasionally 5, cells wide, seldom more than i mm.

high except where there are 2 distinct multiseriate parts; exclusively uni-
seriate, according to Betts (188-9), m the shrubby Weinmannia racemosa
Linn. F. uniseriate rays numerous and composed of high upright cells in
;

Ackama, Callicoma, Cunonia, and Weinmannia, the cells less high in the other
genera and some of them procumbent in Anodopetalum, Ceratopetalum,
Platylophus, Schizomeria, and Spiraeopsis mostly 7-12 rays per mm*; more
;

numerous (12-22) in Caldcluvia, Callicoma, Geissois, and Platylophus',

heterogeneous (Kribs's Types I, II A and B), commonly with 4-10 marginal
rows of upright cells except in Anodopetalum, Ceratopetalum, Platylophus,
and Schizomeria p.p. Cells filled with gummy substance in many species,
Dadswell and Eckersley (527) note that Weinmannia lachnocarpa F. v. M. differs from
1

all other species of this genus in having simple perforation plates, alternate intervascular
pitting, fibres with indistinctly bordered pits, and in some other respects.
564 CUNONIACEAE
crystals reported (527) in the upright cells of Ackama. Fibres with distinctly
bordered usually equally numerous on all walls, but sometimes more
pits,
common on the radial walls, e.g. in Schizomeria and Spiraeopsis; Dadswell
and Eckersley (527) describe the borders as small and inconspicuous in
Geissois and Schizomeria. With thick walls in Ackama, Anodopetalum >
Cunonia, and Weinmannia. Septate and with simple pits in Belangera (1886).
Mean length i '0-1-5 mm. Pith flecks common in Geissois (527).

ROOT
The
occurrence of buttress roots has been recorded in Ackama and
Weinmannia by Francis (707).
TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The spongy structure of the cortex, the characteristic quadrangular
heterogeneous pith, and the tanniniferous secretions provide points of
similarity between the Cunoniaceae and Eucryphiaceae, and thus tend to
confirm the existence of affinities, which are suggested by the wood struc-
ture, between these two families. See also under 'Saxifragaceae*.

(ii)
FROM WOOD STRUCTURE
Dadswell and Eckersley (527) suggest that Weinmannia lachnocarpa F. v. M.
should be transferred to Geissois and note that Geissois and Schizomeria differ
rather markedly from the rest of the family.
Tippo (2261) makes the following statement about the relative degree of
specialization of the wood: 'The Cunoniaceae are on about the same level of
development as are the Brunelliaceae. The former are lower in that all the
species have tracheids, the vessel diameter is slightly smaller and two of the
genera have exclusively scalariform plates. On the other hand, the Cunoniaceae
have shorter vessel elements, the vessel member end-walls are less oblique
1
and the rays are of higher type.
Bausch (154) suggests that 'Platylophus seems to be more typical of the
Saxifragaceae', basing this on anatomy and chemical tests.

ECONOMIC USES
Coachwood, Ceratopetalum apetalum D. Don., and Red Els, Cunonia
capensis Linn., areused in Australia and South Africa respectively for cabinet-
work, and some of the other Australian genera provide wood that is used for
tool handles, turnery, and other purposes.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Anodopetalum, Belangera, Caldcluvia, Callicoma, Ceratopetalum, Codia,
Cunonia,* Geissois, Pancheria, Platylophus, Pullea, Weinmannia.*

(ii) FOR WOOD STRUCTURE

Ackama, Anodopetalum, (Belangera), Caldcluvia, Callicoma, Ceratopetalum,
Cunonia, Geissois, Platylophus, Schizomeria, Spiraeopsis, Weinmannia.
* Kew slide
Represented in the collection.
 CUNONIACEAE 565
LITERATURE
(i) On General Anatomy
Betts 187, Engler 638, Francis 707, Morvillez 1559.

(ii) On Wood Structure

Baker 104, Bausch 154, den Berger 179, 182, den Berger and Endert 183, Betts 188, 189,
Chalk et al 360, Dadswell and Eckersley 525, $27* Desch 574, Garratt 744, Howard 1088,
Janssonius 1154, Record 1843, 1851, 1867-68, Record and Hess 1886, Record and Mell
1894, Scott 2075, Tippo 2261.

121. ESCALLONIACEAE
(FiG. 122 on p. 556; FIG. 123 on p. 558; FIG. 125 on p. 562)

SUMMARY
(i) GENERAL
A
family of trees and shrubs which is widely distributed but occurs chiefly
in the southern hemisphere, particularly in the Andes. The most interesting
features recorded concerning the leaf are that the stomata are often provided
with pairs of small guard cells, nearly circular in outline, whilst a hypoderm
of 1-3 layers occurs in a number of genera. Crystals in both leaf and axis
are predominantly clustered, although solitary types, crystal-sand, and styloids
have also been reported. The nature and arrangement of sclerosed cells in
the pericycle of the young stem are very variable. Xylern. Vessels in
young stems of Brexia Escallonia, and probably of other genera, small,
y

angular; the perforations usually with scalariform plates except in Brexia.

(ii)
WOOD
Vessels small and numerous, typically solitary, sometimes with numerous
multiples and an oblique or radial pattern, occasionally ring-porous and with
spiral thickening, perforation plates scalariform, intervascular pitting typically
opposite, sometimes scalariform, pits to parenchyma similar; members of
medium length to very long. Parenchyma apotracheal, diffuse. Rays up
to 2-7 cells wide, markedly heterogeneous. Fibres with distinctly bordered
pits, of medium length to moderately long.

LEAF
Dorsiventral in all species investigated. Hairs of several distinct types
recorded or observed, (i) Unicellular, pointed, with moderately thick walls
in Escallonia. (ii) Glandular, with short sunken stalks and unicellular heads
in Abrophyllum. (iii) Glandular-shaggy, with multiseriate stalks of variable
length and spherical or peltate heads in Escallonia (Fig. 123 D); particularly
frequent on the leaf teeth of certain species, (iv) Peltate glands in Quintinia.
(v) Two-armed in Argophyllum. Cork warts, not to be confused with glands,
recorded on the leaf surface in one species of Roussea. Stomata with pairs of
small guard cells nearly circular in outline in Abrophyllum^ Argophyllum,
Escallonia, Itea, Quintinia, Roussea; rubiaceous in Quintinia ; with a double
front cavity in Brexia. Hypoderm of 1-3 layers occurring on the upper side
of the leaf in certain species of Argophyllum, Carpodetus, Escallonia, Polyosma,
Roussea. Mesophyll including a single layer of palisade cells in Abrophyllum
S 66 ESCALLON1ACEAE
and Argophyllum. Three vascular bundles enter the base in
Petiole.
Escallonia. Transverse sections through the distal end exhibit a single, open,
crescent-shaped bundle in Escallonia macrantha Hook, et Arn. and E. rubra
(Ruiz et Pav.) Pers. (Fig. 122 B), accompanied by subsidiary strands in the
wings in both species. Vascular structure in the petiole of Brexia madagas-
cariensisThouars (Fig. 122 c) with an abaxial crescent-shaped strand with
incurved ends, accompanied by an additional small cylinder of xylem in the
medullary region and 2 more dorsal ones. Secretory cells and cluster
crystals present in the unlignified tissues of the petiole of the same species
and secretory cells only in Escallonia macrantha. Clustered crystals also
recorded in Anopterus, Carpodetus, Escallonia^ Forgesia, Itea, Polyosma\
solitary and clustered ones in Brexia and Qumtlnia; crystal-sand in Abro-
phyllum.

Axis
YOUNG STEM (Fig. 122 E)
Cork superficial in most members of the family, but arising in the peri-
and composed of relatively thin-walled cells in species of Escallonia.
cycle,
Cortex containing slightly thickened sclerenchymatous cells in Quintinia
sieberi A, DC. with resin in the intercellular spaces in Roussea. Solereder
;

notes the following types of sclerenchyma in the pericycle in at least certain

species of the genera mentioned, (i) A
composite and continuous ring of
sclerenchyma in Abrophyllum. (ii) A
ring of stone cells in Polyosma, Quintinia,
Roussea. (iii) Aring of fibres in Phyllonoma. (iv) Isolated strands of fibres
in Anopterus and Itea. (v) A
composite and continuous ring or an interrupted
ring of sclerenchyma in different species of Brexia. An interrupted or some-
times almost continuous ring of fibres was observed in the pericycle of species
of Brexia and Escallonia examined at Kew. Abrophyllum differs from the
other genera described in having, in the pericycle, a ring of laticiferous sacs,
consisting of vertically elongated cells arranged in longitudinal rows and
filled with white friable contents. Phloem and xylem in Brexia madagas-
cariensis Thouars, Escallonia macrantha Hook, et Arn., and E. rubra (Ruiz,
et Pav.) Pers. constituting continuous cylinders traversed by narrow medullary

rays vessels small (radial diameter seldom exceeding 40 /LI), angular (less so in
;

Brexia than in Escallonia). This structure is probably characteristic of other

members of the family. Vessels exclusively with scalariform perforation
plates in most investigated genera; mixed simple perforations and scalariform
plates recorded in Brexia\ only simple perforations seen in B. madagas-
cariensis examined at Kew. Pith heterogeneous and somewhat spongy or

becoming hollow; component cells abundantly pitted in Escallonia rubra.

Secretory cells, with apparently tanniniferous contents, observed in the
cortex, phloem, and medullary rays and pith of species of Brexia and Escat-
lonia. Laticiferous sacs, see Tericycle'. Crystal^, of the same types as those
described under 'Leaf', occur also in the axis of the same genera; styloids also
recorded in the phloem of Escallonia.

WOOD (Fig. 125 i, K, and L)

Vessels moderately to very small (30-75 p mean tangential diameter);
typically solitary apart from the apparent tangential pairs produced by over-
 ESCALLONIACEAE 567
lapping ends, but with numerous radial multiples and a tendency to a radial
pattern in some species of Polyosma (Fig. 125 L), and with an oblique or
radial pattern in Itea (1154, 1206); numerous, varying from about 20 per
sq. mm. in some species of Itea (1206) and Polyosma to about 100 per sq. mm.
in Carpodetus and Escallonia p.p. semi-ring-porous and with spiral thicken-
;

ing in some species of Escallonia. Perforation plates typically scalariform,

with both simple and scalariform plates in Brexia (2158, 2261); with less than
20 fine bars in Carpodetus and Escallonia, up to 40-70 bars in Itea (1154),
Polyosma p.p., and Quintinia, and up to 125 bars in some species of Polyosma
(1154); with reticulate plates in some species. Intervascular pitting scalari-
form in Carpodetus and Itea (1154, 1206), round and usually opposite in the
other genera but with some transitional forms pits to ray and wood paren-
;

chyma large and oblong in Carpodetus and Itea (1154), small, round, and
similar to the intervascular pitting in the other genera; mean length o-5i-2,
mostly more than 0-9 mm. Parenchyma exclusively apotracheal, as scat-
tered cells (diffuse) with a tendency to form short uniseriate lines, e.g. in
Quintinia, and sometimes more abundant near the vessels, e.g. in Carpodetus ;

in regular, multiseriate apotracheal bands in Brexia (2261). With chambered

crystals in Carpodetus and Brexia (2261); sometimes filled with dark gum.
Strands of 4-8 cells. Rays very variable in width in different species, tending
to be of 2 distinct sizes; the larger rays up to 2-7 cells wide, widest in some
species of Escallonia and Polyosma\ exclusively uniseriate, at any rate in
small material, in Brexia (2261); slightly more than i mm. in height in
Polyosma and Quintinia; the uniseriate rays composed entirely of upright
cells; 16-24 rays per mm. in Carpodetus and Escallonia, less numerous (about
10 per mm.) in Polyosma and Quintinia\ heterogeneous (Kribs's Types I-II A
and III in Brexia), with 4 or more marginal rows of square or upright cells
and commonly with 10 or more rows in Quintinia\ with sheath cells in
Escallonia p.p. Sometimes with abundant gummy contents, e.g. in Quintinia
and according to Tippo (2261) with crystals in Brexia and Carpodetus. Fibres
with distinctly bordered pits, usually more numerous on the radial than on
the tangential walls ; walls moderately thick to thick ;
mean length i -0-2- 1 mm. ,

longest in Polyosma (1154).

TAXONOMIC NOTES
(i) FROM GENERAL
ANATOMY
The nature of the stomata and the general structure of the young stem
support the generally accepted view that the Escalloniaceae and Grossula-
riaceae are closely related. The stem of Escallonia and Ribes in particular are
very much alike. The distinctive structure of the petiole of Brexia suggests
that this genus may be rather remotely related to Escallonia and possibly to
1

other genera in the family. See also under 'Saxifragaceae .

(ii) FROM WOOD STRUCTURE

Tippo (2261) states:

'Anatomically the Escalloniaceae are very close to the Grossulariaceae. The

former perhaps may be considered to be a little higher, for i genus in the Escallo-
niaceae has vessel elements with simple and rarely vestigial scalariform perforation
568 ESCALLONIACEAE
plates. On the other hand, the Grossulariaceae are more specialized in regard to
certain other characters/
'The three families Hydrangeaceae, Grossulariaceae and Escalloniaceae are very
similar anatomically, but in general it may be said that the
Hydrangeaceae are
primitive, Grossulariaceae less so, and Escalloniaceae least primitive,*

Tippo (2261) considers the anatomy of the wood to be consistent with the
derivation of the Cunoniales (including the Hydrangeaceae, Escalloniaceae,
Cunoniaceae, Brunelliaceae, and Grossulariaceae) from the Magnoliales, and
with the idea of the Cunoniales having given rise to the Resales (including
the Rosaceae and perhaps the Calycanthaceae). He notes trends towards the
Resales in the Hydrangeaceae, Grossulariaceae, and Escalloniaceae and con-
siders that the last three groups could very well be placed in one family.

ECONOMIC USES
The
family includes a number of ornamental shrubs commonly cultivated
in gardens.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Abrophyllum, Anopterus, Argophyllum, Brexia,* Carpodetus, Escallonia,*
Forgesia, Itea, Phyllonoma, Polyosma, Quintinia, Roussea.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
(Brexia), Carpodetus, Escallonia, (Itea), Polyosma, Quintinia.

LITERATURE
(i) On General Anatomy
Engler 636.

(ii) On Wood Structure

den Berger 182, Desch 574, Janssonius 1154, Kanehira 1206, Record 1783, 1800, 1840,
1851, Record and Hess 1886, Tippo 2261, Welch 2401, Yamabayashi 2478.

122. EUCRYPHIACEAE
(Fie. 125 on p. 562; FIG. 126 on p. 570)

SUMMARY
(i)
GENERAL
A
family of trees and shrubs belonging to the single genus Eucryphia which
occurs in Australia, Tasmania, and Chile. The following description of the
leaf and young stem refers chiefly to Eucryphia glutinosa (Poepp. et Endl.)
Baillon grown at Kew. This species has rubiaceous stomata whilst, in the
young stem, the somewhat spongy cortex, the xylem with small vessels and
uniseriate rays, the quadrangular heterogeneous pith and the occurrence of
tanniniferous elements are useful diagnostic characters.
 E UCR YPHIA CEAE 569

(ii) WOOD
Vessels small, mostly solitary, but with some multiples, spiral thickening
sometimes present, perforation plates scalariform or scalariform and simple,
intervascular pitting transitional between scalariform and opposite, pits to
parenchyma often large, horizontally elongated and simple. Members of
medium length to moderately long. Parenchyma apotracheal, diffuse to
slightly banded, and sometimes in terminal bands. Rays exclusively uni-
seriate or 2-3 cells wide, heterogeneous. Fibres with bordered pits, of
medium length.

LEAF
Leaflets dorsiventral. Hairs infrequent, but where present occurring in
the form of short, unicellular, thick-walled trichomes. Stomata confined to
the lower surface, 30 p in diameter, mostly rubiaceous, with i or 2 pairs of
subsidiary cells lying parallel to the pore. Subsidiary cells somewhat variable
in shape, but many of them approximately rectangular. Mesophyll com-
posed of 2 or 3 layers of palisade cells and of a region of spongy tissue, each
occupying about half the thickness of the leaflet. Mesophyll cells, especially
those of the palisade tissue, containing yellowish plastids. Vascular bundles
of the smaller veins vertically transcurrent by sclerenchyma. Petiole. Trans-
verse sections through the distal end in Eucryphia glutinosa Baill. exhibit,
towards the abaxial side, a broad, crescent-shaped vascular strand, accom-
panied by a second bundle partially enclosed within but almost connected
with the incurved ends of the first, the whole vascular system thus resembling
a slightly interrupted, dorsally flattened cylinder (Fig. 126 A). Cortical
parenchyma around the vascular strand very spongy. Numerous solitary and
cluster crystals occur in the veins and, less frequently, in the mesophyll.

Axis
YOUNG STEM (Fig. 126 D)
Cork examined at Kew. Cortex somewhat spongy,
superficial in material
composed mostly of sometimes pitted cells, but including a few
small,
branched, sclerenchymatous idioblasts. Pericycle bounded externally by a
broad, almost continuous ring of thick-walled fibres with narrow lumina,
locally interrupted by stone cells, the latter more numerous in X E. inter-
media Bausch than in E. glutinosa Baill. (Fig. 126 D). Phloem and xylem
constituting continuous cylinders, traversed by narrow, almost exclusively uni-
seriate medullary Phloem consisting wholly of soft tissue. Xylem vessels
rays.
somewhat angular, seldom exceeding 30 p, in radial diameter; perforation
plates scalariform or scalariform and simple. Pith approximately quadrangular
in transverse section; very heterogeneous, composed of cells of much larger
transverse diameter than any of those elsewhere in the stem; most of the cells
thin-walled and devoid of contents, but others, often arranged in vertical
columns, provided with thicker, pitted walls and filled with dense, amor-
phous, probably tanniniferous contents. Apparently tanniniferous cells also
observed in the cortex, phloem, and medullary rays. Crystals not seen in
E. glutinosa, but occasional solitary ones present in the cortex of X E. inter-
media, and more numerous solitary and clustered crystals in another, un-
determined species of Eucryphia. Secretory cells, see 'Pith'.
570 E UCR YPHIA CEA E
WOOD (Fig. 125 H and j)
Vessels very to moderately small, varying from about 30 to 70 ft mean
tangential diameter, mostly about 60 p (154); mostly solitary but with some

FIG. 126. EUCRYPHIACEAE, A and D; CUNONIACEAE, B-C and F;

CRASSULACEAE, E and G-J
A, Eucrypkia glutinosa Baill. Petiole x 19. B, Weinmannia trichosperma Cav. Petiole X 33.
C, Cunonia capensis Linn. Petiole x 15. D, Eucrypkia glutinosa Baill. Stem x 19. E, Umbilicus
pendulinus DC. Petiole x 8. F, Cunonia capensis Linn. Stem x 8. G, Bryophyllum daigremontana
Hamet et Perrier. Petiole x 8. H, B. daigremontana Hamet et Perrier. Stem x 8. I, Sedum reflexum
Linn. Stem X 1 8. J, Crassula sp. Petiole x 8.

multiples and short chains, and with a slight tendency to tangential arrange-
ment in some species; usually 70-120 per sq. mm,, mostly about 100, reported
by Vestal (2329) to be sometimes up to 200 per sq. mm. with a tendency to
;
 E UCR YPHIA CEA E 571
be semi-ring-porous in a few species; spiral thickening present in some
species, absent from Eucryphia cordifolia Cav. (1886). Perforation plates
typically all scalariform, but sometimes scalariform and simple, the multi-
perforate plates usually with 20 or fewer bars but occasionally up to 40, and
sometimes reticulate. Intervascular pitting transitional between scalariform
and opposite. Pits to ray and wood parenchyma partly large, horizontally
elongated and simple, and partly round and bordered, sometimes unilaterally
compound. Thin- walled tyloses rarely present (154). Mean member length
about 0*8 mm. Parenchyma predominantly apotracheal, as scattered cells
(diffuse); tending to be more numerous and to form uniseriate lines in the
late wood; sometimes in terminal bands 1-4 cells wide; Bausch (154) states
that a small amount of paratracheal parenchyma is usually present. Cells
commonly with dark gummy contents. Strands usually of 4-8 cells. Rays
either exclusively uniseriate or up to 2-3 cells wide, and occasionally up to
4-6 cells wide in the middle (1886); 10-12 per mm.; heterogeneous (Kribs's
Types II B and III), with 1-2 marginal rows of square to upright cells. Cells
commonly filled with gummy or granular material. Fibres with distinctly
bordered equally numerous on both radial and tangential walls and in
pits,
single rows; Bausch (154) describes the pits as inconspicuously bordered in
some species. Walls usually thick, often differing markedly in the early and
late wood. Mean length about 1-4 mm.

TAXONOMIC NOTES
Bausch (154) in a study of the affinities of the family states: 'The family is
considered to be taxonomically most nearly related to the Cunoniaceae. This
conclusion is based on morphological characters and on similarity of anatomical
structure, especially on the occurrence of both simple and scalariform per-
forations in the vessels in the two families. Similarity in chemical properties
also points to the same relationship/
The features emphasized by Bausch in the wood anatomy do not necessarily
imply relationship, nevertheless his general conclusion appears to be valid.
Record and Hess (1886) also consider the wood anatomy to resemble that of
the Cunoniaceae. For other points of similarity to Cunoniaceae see that
family.

ECONOMIC USES
Leatherwood, Eucryphia lucida (Labill.) Baillon (syn. E. billardieri Spach.),
is reported (525) to be used to a small extent for tool handles.

GENUS DESCRIBED
Eucryphia.*
* Kew
Represented in the slide collection.

LITERATURE
(i) On General Anatomy
Bausch 154, Gilg 769-

(ii) On Wood Structure

Bausch 154, Dadswell and Eckersley 525, Record 1851, Record and Hess 1886, Record
and Mell 1894, Vestal 2329.
 (572)

123. GROSSULARIACEAE
(Fio. 122 on p. 556; FIG. 127 on p. 576)

SUMMARY
(i) GENERAL
A
family of shrubs, mostly from north temperate regions; often with
spines. The more important diagnostic characters include the small stomata
which are nearly circular in outline; the deep-seated origin of the cork; the
absence of pericyclic sclerenchyma from the young stem; in the xylem
the small, angular vessels which nearly always have scalariform perforation
plates.

(ii) WOOD
Vessels extremely small and very numerous, tending to form tangential
lines, often ring-porous; perforations typically scalariform; intervascular
pitting typically scalariform to opposite, pits to parenchyma similar; members
of medium length. Parenchyma absent or in very rare bands. Rays up to
11-14 or more cells wide, heterogeneous. Fibres usually septate, very to
extremely short. Vasicentric tracheids present.

LEAF
Dorsiventral. Hairs mostly unicellular, but glandular, shaggy trichomes
with multiseriate stalks of variable length and spherical or peltate heads also
occur on the leaves and stipules. Stomata provided with pairs of small guard
cells nearly circular in outline. Petiole examined in Ribes aureum Pursh.,
R. grossularia Linn., 1?. nigrwn Linn., and R. sanguineum Pursh. (Fig 122 L)
and the following structure observed. Three separate vascular bundles enter
the base of the petiole, but transverse sections through the distal end exhibit
a solitary crescent-shaped vascular strand, with the ends very much incurved
and almost in contact with one another. Vascular strand in all four species
supported by fibres in the pericyclic region. Secretory cells and clustered
crystals fairly abundant in the parenchymatous tissues of the petiole.

Axis
YOUNG STEM (Fig. 122 o)
Cork deep seated in origin; composed of thin- walled cells in at least certain
species. Pericycle devoid of sclerenchyma. Phloem and xylem in some
species, e.g. R. grossularia Linn., constituting almost or quite continuous
cylinders traversed by narrow medullary rays, but with the xylem dissected
by broader but lignified rays in other species, e.g. JR. aureum Pursh. Vessels
small and angular, e.g. up to about 36 in radial diameter in R. nigrum Linn,
//,

nearly always with scalariform perforation plates. Pith stellate in outline in

JR. aurea and R. nigrum but not in R. grossularia] commonly heterogeneous
and frequently spongy. Secretory cells with apparently tanniniferous con-
tents common and frequently abundant in the cortex, phloem, medullary
rays, and pith (usually arranged in vertical columns in the last of these
tissues).Clustered crystals common in the unlignified tissues, but relatively
infrequent in R. nigrum.
 GROSSULARIACEAE 573
WOOD (Fig. 127 F-G)
Vessels extremely small (less than 25 ft mean tangential diameter) to very
small (25-50 /x); with moderately numerous groups and tending to form a
distinct tangential pattern in some species; very numerous, 100-300 per

sq. mm. ; ring-porous in some species. Perforation plates typically scalariform,

with less than 20 thin bars, but Tippo (2261) notes both simple and scalariform

plates in one species. Intervascular pitting scalariform to opposite, rarely

alternate (2261); pits to wood and ray parenchyma similar in size and shape
to the intervascular pitting. Mean member length o-36-O'66 mm. (2261).
Parenchyma typically very rare or absent; Tippo (2261) reports the rare
occurrence of apotracheal bands 1-6 cells wide. Rays of 2 distinct sizes, the
larger up to 1 1-14 cells wide and occasionally up to 22 cells (2261) up to about ;

i mm.
high; uniseriate rays numerous, low, and composed entirely of square
or upright cells; rays 10-14 per mm.; heterogeneous (Kribs's Type HA),
with 1-3 marginal rows of upright cells, commonly with sheath cells. Fibres
usually septate, the septa rare in some species; pits with very narrow borders;
walls moderately thin to moderately thick; mean length about 0*5 mm.
Vasicentric tracheids present.

TAXONOMIC NOTES, see under Saxifragaceae and Escalloniaceae

ECONOMIC PRODUCTS
Gooseberries (Ribes grossularia Linn.), Black Currants (R. nigrum Linn.),
and Red Currants (R. rubrum Linn.) are well-known edible fruits, selected
varieties of which are commonly cultivated. Other species, e.g. R. sanguineum
Pursh., are cultivated on account of their ornamental flowers.

GENUS DESCRIBED
Ribes.*
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Engler 636, Morvillez 1559*

(ii) On Wood Structure

Baker 104, Chalk and Chattaway 358, Cozzo 494, Greguss 2522, Record 1843, 1851,
Record and Hess 1886, Tippo 2261, Welch 2401.

124. HYDRANGEACEAE
(Fic. 122 on p. 556; FIG. 123 on p. 558; FIG. 127 on p. 576)

SUMMARY
(i) GENERAL
A family of trees and shrubs from north temperate and sub-tropical regions.
One of the most interesting anatomical features is the common but not
universal occurrence of raphide sacs, which also contain mucilage, in both
leaf and axis. In the leaf they are usually to be found in the spongy mesophyll
574 HYDRANGEACEAE
where they lie parallel to the surface of the leaf, but they also occur in the
palisade tissue. In the axis they are present in the cortex, phloem, and pith.
A hypoderm of 1-3 layers is common in some of the genera. The cork in
the young stem is sometimes differentiated into alternating rows of cells which
are radially elongated and radially compressed respectively. A
characteristic
ring of stone cells is present immediately within the cork cambium, in certain
genera. In the xylem the vessels mostly have exclusively scalariform per-
foration plates, although simple ones are sometimes to be found amongst
them.

(ii) WOOD
Vessels small, numerous and solitary, commonly semi-ring-porous and
with spiral thickening, perforation plates typically scalariform with numerous
fine bars, rarely simple, intervascular pitting scalariform to opposite, pits to

parenchyma similar to the intervascular pitting, moderately to very long.

Parenchyma absent, diffuse or scanty paratracheal. Rays 2-9 cells wide,
heterogeneous. Fibres typically with numerous bordered pits, rarely septate,
with spiral thickening in 2 genera, very long.

LEAF
Dorsiventral in all the species investigated. Hairs. Simple, unicellular,
papillose trichomes observed in Philadelphia (Fig. 123 A), the frequency and
length varying in different species; very long unicellular trichomes present in
Jamesia americana Torr. et Gray; tufted in Broussaisia and Pileostegia (with
calcified walls); stellate (Fig. 123 B~C), calcified, unicellular types occur in

species of Deutzia. Leaf teeth sometimes glandular in Decumaria, Deutzia,

Philadelphus. Epidermis locally consisting of horizontally divided cells in
Carpenteria. Hypoderm of 1-3 layers on the upper side of the leaf recorded
in i species of Broussaisia and in Hydrangea (pro parte). Stomata rubiaceous
in certain species of Dichroa and Hydrangea ranunculaceous in Philadelphus.
;

Mesophyll including i layer of palisade tissue in Deutzia and Philadelphus.

Petiole. Transverse sections through the distal end exhibiting an open,
crescent-shaped vascular strand in species of Deutzia (Fig. 122 A), Jamesia,
and Philadelphus. Vascular strand crescent-shaped but less open and usually
somewhat dissected in species of Hydrangea (Fig. 1220) and Pileostegia.
Main strand consisting of an abaxial arc with a separate, flat, adaxial bundle
between the ends in Decumaria sinensis Oliv., Dichroa febrifuga Lour., and
Hydrangea petiolaris Sieb. et Zucc. (Fig. 122?). Additional strands present
in the wings in the same species of Decumaria, Dichroa, and Pileostegia and
in the medullary region in Hydrangea and Pileostegia. Secretory cells, with
unidentified amorphous contents, observed in the unlignified tissues of the
petiole in Decumaria, Deutzia, Dichroa, Hydrangea, Jamesia, and Pileostegia
but not in Philadelphus. Raphide sacs (see also 'Young Stem') containing
mucilage common; usually lying in the spongy mesophyll parallel to the
surface of the leaf, but sometimes amongst and parallel to the palisade cells;
occasionally visible as transparent dots in Decumaria. Similar secretory
sacs, but with grey, finely divided contents, observed in addition to ordinary
raphide sacs in Pileostegia viburnoides Hook. f. et Thorns. Clustered crystals
present in Carpenteria, Dichroa, Jamesia, Philadelphus.
 HYDRANGEACEAE 575

Axis
YOUNG STEM (Fig. 122 G-H)
Cork deep seated in origin in species of Deutzia, Hydrangea, Jamesia,
Pileostegia examined at Kew; consisting of rows of radially elongated cells
alternating with radially compressed cells in Jamesia, Philadelphus (radially
compressed cells sometimes sclerotic); some of the cells provided with horse-
shoe-shaped thickenings in Wkipplea. A ring of stone cells on the inside of
the cork cambium recorded in species of Broussaisia, Decumaria, Pileostegia,
Schizophragma; sclerenchymatous elements also observed in the same position
in species of Decumaria and Philadelphus. Pericycle devoid of sclerenchyma
in at least certain species of Decumaria, Deutzia, Hydrangea, Jamesia, Phila-
delphus, Pileostegia. Phloem usually in the form of a rather broad, con-
tinuous cylinder, consisting of unlignified tissue. Xylem also cylindrical and
traversed by narrow rays in certain species of Decumaria, Deutzia, and
Pileostegia, but dissected by broader, lignified rays in species of Deutzia,
Hydrangea (Fig. 122 H), Jamesia, Philadelphus (Fig. 122 G). Vessels small,
none seen to be more than about 50 /LC in radial diameter in the material
available for examination, frequently angular. Perforation plates exclusively
scalariform in Broussaisia, Decumaria, Deutzia, Dichroa, Fendlera, Hydrangea,
Jamesia, Philadelphus, Pileostegia, Platy crater, Schizophragma', simple and
sclariform types recorded in Cardiandra and Whipplea. Pith usually large in
proportion to the diameter of the stem, consisting mostly of thin-walled
tissue, without contents apart from a narrow perimedullary zone of cells with
thicker walls; tending to break down and become hollow, e.g. in Deutzia.
Pith of the type just described observed in species of Deutzia, Hydrangea,
Jamesia, Philadelphus', smaller and composed of spongy tissue in Hydrangea
petiolaris Sieb. et Zucc. and Pileostegia viburnoides Hook. f. et Thorns.
Raphide sacs (see also 'Leaf ') containing mucilage said to be very common
in the primary cortex, phloem, and pith; but none were observed in species
of Deutzia, Jamesia, and Philadelphus although seen in species of Decumaria,
Hydrangea, and Pileostegia, those of P. viburnoides much elongated and filled
with an orange secretion. The distribution of raphide sacs in the different
tissues varies according to the species. Secretory cells with unidentified,
amorphous contents observed in the parenchymatous tissues in species of
Decumaria, Deutzia (pro parte), Hydrangea, Pileostegia', none seen in Phila-
delphus. Elongated secretory sacs, resembling the raphide sacs but filled with
a finely divided greyish substance, observed in the phloem of Decumaria
sinensis Oliv.

WOOD (Fig. 127 A-E)

Vessels small(less than 100 /x mean tangential diameter), ranging from
25 fj,
some species of Deutzia to about 60 p in Philadelphus in the material
in
examined; Tippo (2261) gives the range as 22-130 /x; typically solitary, apart
from the apparent tangential pairs produced by overlapping ends, but with
moderately numerous multiples in Deutzia glabrata Komorow very numerous,
;

ranging from 40 to over 200 per sq. mm. ring-porous or semi-ring-porous in

;

some species of Deutzia, Fendlera (1864), Hydrangea, and Philadelphus and

with spiral thickening in some species of Deutzia, Hydrangea (1851), and
S?6 HYDRANGEACEAE
Philadelphia. Perforation plates typically all scalariform, with 20-90 fine
bars, but all, or predominantly, simple in Deutzia glabrata and some species
of Philadelphia (2478); Solereder, quoting Halle, notes both simple and

D E F G
FIG. 1*7. HYDRANGEACEAE, A-E; GROSSULAWACEAE, F-G
A, Philadelphus incanus Koehne. (Not entirely typical; most rays distinctly heterogeneous.) B,
Hydrangea bretschneideri Dippel. C, Deutzia scabra Thunb. D, Philadelphus incanus Koehne.
E, Hydrangea bretschneideri Dippel. F, Kibes glaciale Wall. G, R. glaciale Wall.

scalariform plates in Cardiandra and Whipplea. Intervascular pitting scalari-

form Dichroa (1154), Hydrangea, and Schizophragma, opposite in Deutzia
in

p.p. and Philadelphus, alternate in Deutzia glabrata; pits to ray and wood
 HYDRANGEACEAE 577

parenchyma similar to the intervascular pitting, large and oblong in Dichroa

(1154), Hydrangea, and Schizophragma, small and round in Deutzia and
Phtladelphus. Tyloses reported by Tippo in a single species. Mean member
length 0-56-1 '35, mostly above i-o mm. Parenchyma diffuse in Deutzia,
predominantly paratracheal (scanty), but with a few scattered cells in Phila-
delphus, absent or with a few cells round the vessels in Dichroa (1154),
Hydrangea, and Schizophragma. Strands of 6-8 cells. Rays tending to be of
2 distinct sizes; up to 2-3 cells wide and less than i mm. high in some species
of Deutzia and Hydrangea, up to 4 or more cells wide in other species and in
Philadelphus and Schizophragma, and up to 9 cells in Hydrangea p.p. (1206);
more than i mm. high in Deutzia p.p., Philadelphus, and Schizophragma]
uniseriate rays numerous and composed entirely of high upright cells rather ;

fewer in Philadelphus and Schizophragma; about 30 rays per mm. in Deutzia

and Hydrangea, distinctly fewer (about 10 per mm.) in some species of
Philadelphus and Schizophragma. Heterogeneous (Kribs's Type I), with 4-10
marginal rows of upright cells; usually with sheath cells. Fibres typically
with small to large bordered pits, often very numerous on both radial and
tangential walls; but with simple pits in Deutzia p.p., Dichroa, and some
species of Hydrangea (1206) and septate according to Kanehira (1206) in
Deutzia p.p., Dichroa (1154), and Hydrangea; walls moderately to very thick,
with spiral thickening in Deutzia and Philadelphus coronarius L. (2158).
Mean length i-i-i-6 mm.

TAXONOMIC NOTES, see under Saxifragaceae and Escalloniaceae

ECONOMIC USES
The family includes many well-known ornamental shrubs.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Broussaisia, Cardiandra, Carpenteria, Decumaria,* Deutzia,* Dichroa,*
Fendlera, Hydrangea,* Jamesia,* Philadelphus,* Pileostegia,* Platycrater,
Schizophragma, Whipplea.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Cardiandra), Deutzia, (Dichroa), (Fendlera), Hydrangea, Philadelphus,
Schizophragma, (Whipplea).

LITERATURE
(i) On General Anatomy
Engler 636, Morvillez 1559.

(ii) On Wood Structure

Greguss 2522, Kanehira 1206, 1209, Janssonius 1154, Record 1783, 1800/1840,
1864, Tippo 2261, Yamabayashi 2478.

Pp
 (578)

125. CRASSULACEAE
(Fio, 124 on p. 558; FIG. 126 on p. 570)

SUMMARY
A family from warm dry regions, consisting mainly of succulent herbs, but
tending to be miniature shrubs or trees in certain genera and species. Most
members of the family are remarkable for their xeromorphic structure, parti-
cularly the occurrence of water-storage tissue in the leaf and stem. Some are
believed to be capable of absorbing water directly from the air by special
hairs,epidermal cells, or adventitious roots (Marloth 1445 and Berger 176).
Kean (1224) concluded that the hydathodes (see below) of Rochea coccinea
(L.) DC. serve solely for the secretion and not for the absorption of water,
whilst the bladder hairs of Crassula falcata Wendl. (syn, Rochea falcata DC.)
can take up moisture from the atmosphere, Reiche (1907) has pointed out
that the vascular strands are widely and irregularly distributed throughout
the water-storage tissue of numerous species, and he likens the vascular
system of the Crassulaceae, embedded in aqueous tissue, to water plants whose
finely divided members are suspended in a fluid medium. Reiche terms
succulent plants which exhibit this structure 'innere Wasserpflanzen', The
stomata are nearly always surrounded by a girdle of 3 subsidiary cells and
occur on all parts of the surface of the leaf. The leaves are frequently centric,
and typical palisade tissue is rare. The fleshy structure of the stem is due to
the well-developed parenchymatous or collenchymatous tissues of the cortex
and pith. Secretory cells, with contents generally described as tannini-
ferous, are common in the same tissues and in the phloem. The xylem in
the stem is very seldom dissected by wide rays, and there is a considerable
range of variation in the distribution of the vessels. Cortical bundles,
which represent leaf traces, are common; medullary bundles and
anomalous structure have been recorded and described in a few genera and
species. A red anthocyan pigment, the colour of which is intensified by
bright light, is common in many members of the family, either throughout
the plant or localized in certain organs or parts of organs in different species.
Certain species of Bryophyllum, notably B. pinnatum (Lam.) S. Kurz (syn.
B. calycinum Salisb.) and B. daigremontianum R. Hamet et Perrier (syn.
Kalachoe daigremontiana R. Hamet et Perrier), are remarkable for the fact
that they reproduce vegetatively by means of foliar embryos which arise in
notches in the leaf margins. This method of reproduction has been the sub-
numerous anatomical and physiological investigations. For particulars
ject of
concerning a selection of these see Johnson (1186), Mehrlich (1479), Sobels
(2156), Yarbrough (2483, 2484).
LEAF
Usually centric or intermediate between dorsiventral and centric; typical
palisade tissue rare. Hairs usually infrequent, but several kinds recorded,
(i) Bladder-like hairs,
sometimes described as epidermal cells, present in
Crassula falcata Wendl. (Fig. 124 B-c) (covering the whole surface) and
Rochea coccinea (L.) DC. (confined to the leaf margins) (Lindinger 1374 and
Rehfous 1904); similar structures also observed in an unidentified species of
 CRASSULACEAE 579
Crassula examined at Kew.(ii) Glandular hairs, with short or long stalks and
which sometimes secrete mucilage, recorded in species ofAeonium, Cotyledon,
Echeveria, Kalanchoe (glands red in K. aromatica Perrier), Sempervivum.
(iii) Three-armed, pointed hairs present in Kalanchoe section stellatopilosae,

e.g.K. beharensis Drake according to Berger (176). (iv) Biseriate hairs forming
a cobweb-like surface to the leaf, together with transitions between these and
glandular shaggy types, present in Sempervivum arachnoideum Linn. Leaf
surface often covered by a bluish-white coating of wax secreted from the
epidermis. Epidermis (Fig. 124 D) usually composed of cells elongated
transversely to the longitudinal axis of the leaf; papillose in a few species of
Bryophyllum and Sedum. Papillae of a special type, each with a central
perforation of the cuticle, a central reduction of the outer wall, and a mucila-
ginous protoplast recorded by Sporer (2171) at the margins of the leaf of the
xerophytic Crassula pyramidalis Thunb, Epidermis occasionally 2-layered
in the same genera. Stomata present on all parts of the surface of the leaf;
surrounded by a girdle of 3 subsidiary cells, e.g. in Crassula muscosa (L.) Roth,
(syn. Tillaea muscosa Linn,) and Sedum spurium Marsch-Bieb (Fig. 124 A).
Hydathodes, which appear as small pits or spots on the leaf visible to the
naked eye, are variously distributed in different species, sometimes covering
the whole of both surfaces, at others confined to one surface or arranged in
rows near the leaf margin on both surfaces or only on the lower side. Hyda-
thodes recorded in species of Bryophyllum, Crassula, Kalanchoe, Sedum,
Umbilicus', appearing as red-coloured pores on the upper surface in Crassula
schmidtii Regel according to Weingart (2380). Transverse sections through
the distal end of the petiole (Fig. 126 E and j) exhibiting, in most of the few
species examined, an open arc or crescent consisting of a variable number of
collateral bundles of which the median one is larger or much larger than the
remainder. Median vascular strand accompanied by numerous smaller ones
in the cortical region in Bryophyllum daigremontianum R. Hamet et Perrier
(Fig. 126 G). Secretory cells, with apparently tanniniferous contents,
common in unlignified tissues, especially around the veins; only rarely mor-
phologically differentiated from neighbouring cells. Crystals common,
solitary, clustered, or in the form of sphaerites and crystal-sand.

Axis
STEM (Fig. 126 H-I)
Cork, usually consisting of thin-walled cells, arising in the epidermis in
Bryophyllum and Sedum, but sometimes sub-epidermal or even more deeply
seated in other genera; described by Heckel (928) and Jadin and Juillet (1139)
as becoming impregnated with resin and forming a thick layer capable of
reducing evaporation in certain species of Kalanchoe from Madagascar.
Heckel draws attention to the similarity between the resinous cork of these
species of Kalanchoe and that of Sarcocaulon (Family Geraniaceae). Cortex
well developed, fleshy; consisting wholly of parenchyma or with the outer
part collenchymatous. Centric, sometimes numerous cortical bundles (leaf
traces) with central xylem present in certain species of Aeonium, Greenovia,
Rochea; similar bundles also recorded by Dauphin^ and Hamet (549) and
Dauphine (547) in species of Cotyledon and Kalanchoe. Phloem poorly
developed, including narrow sieve tubes which are not easily seen. Xylem
580 CRASSULACEAE
nearly always in the form of a continuous cylinder, only rarely dissected by
wide rays, but exhibiting the following varied types of structure in species
with a well-developed axis, (i) Forming a closed ring, consisting wholly of
prosenchyma and without vessels (apart from those in the primary xylem) or
rays, situated on the outside of the primary groups of vessels in species of
Sedum and Umbilicus, (ii) Similar to (i) but small groups of vessels accom-
panied by elongated, unlignified, parenchymatous cells, included in the ring,
in species of Aeonium, Bryophyllum, Crassula, Echeveria, Sedum. (iii) As (ii)
but with the groups of vessels and accompanying unlignified parenchyma
larger and arranged in concentric circles, in species of Cotyledon, Crassula,
Kalanchoe, Rochea, and Sempervivum. (iv) Groundwork of the xylem in
Crassula argentea Thunb. consisting of unlignified tissue with vessels
irregularly distributed in it, the vascular tissues being in the form of bundles
separated from one another by primary rays. Vessels of the secondary xylem
with simple pits or reticulate thickening, bordered pitting not recorded;
perforations simple. Jeffrey and Cole (i 167) record that some of the so-called
vessels of the Crassulaceae resemble tracheids, since their terminal pores are
occluded. Growth rings not well defined. Cambial activity not very consider-
able even in species with relatively wide stems; secondary growth in thickness
stated to occur through cell divisions in the cortex and pith, e.g. in species of
Bryophyllum and Crassula. Medullary bundles first recorded in the flower
stalk of Greenovia by Hamet (875, 876), but subsequently noted by the same
author in Echeveria as well (877, 878), Medullary bundles of Echeveria
minutiflora Rose (syn. Thompsonella minutiflora Britton et Rose) differ from
those of other species of Echeveria and from Greenovia (i) in being more
remote from and not connected with the main vascular ring ; (ii) in developing
considerably later than instead of at the same time as the main vascular system.
Secretory cells, with contents presumed to be tanniniferous, common in the
unlignified tissues; sometimes sufficiently elongated to be described as sacs
and arranged in longitudinal series. Developmental anatomy. Maas(i4os)
studied the developmental anatomy of the stem of various species of Crassula^
and found considerable variations in different species.

ANOMALOUS STRUCTURE
The complex anomalous structure of old stems of Sedum populifolium Pall,
is described by Solereder as follows:
The young stem of this species, externally to the primary groups of vessels,
contains a ring of wood fibres, enclosing isolated vessels only. Subsequently the
cambium produces, in the first place, a ring of thin-walled tissue with groups of
vessels, which lie on the same radii as the primary vessels but this is followed by a
;

second ring of fibres, which only differs from the first in having more numerous
groups of vessels embedded in thin- walled tissue, and so on. The ring of thin-
walled tissue, which succeeds the first ring of fibres, soon becomes filled with con-
tents, and assumes the appearance of a pith its innermost layer of cells develops
;

into a cork cambium, and the cork-tissue derived from the latter shuts off all the
tissue lying internally to it
(pith, primary groups of vessels and the first ring of
fibres) from the outer woody tissue. The phenomena described are repeated. In
each period of vegetation a new ring of wood is developed, and begins with a zone
of thin- walled tissue; and similarly in the innermost cell layer of these zones
successive layers of cork are formed.'
 CRASSULACEAE 581
Anomalous structure also recorded in the rhizome of Sedum aizoon Linn, and
other species of Sedum possessing rhizomes of a similar type. For details see
Pfeiffer (1712). The middle part, but not the two ends, of the napiform roots
of Sedum section Telephium also exhibits anomalous structure described by
Solereder as follows:
'In transverse section this region shows a circle of concentric vascular bundles
with central wood. Originally the fibro-vascular system of the roots possesses
normal structure as Koch was the first to show. Subsequently, however, the ring
of cambium breaks up into a number of separate arcs, which extend round portions
of the original xylem and enclose them in an annular manner/

ROOT
Most members of the family are described by Molisch (1549) as having red
root tips, coloured by an anthocyan pigment, e.g. in certain species of
Echeveria, Kalanchoe, Sedum, Sempervivum, but red tips not observed by the
same author in certain species of Bryophyllum, Cotyledon, Crassula. Dried
and fresh adventitious roots form a reddish felt on the stemlets of Adromischus
spp. For the abnormal structure of the roots of Sedum section Telephium see
'Anomalous Structure* above.

ECONOMIC USES
Many members of this family are cultivated for ornamental purposes or
because of their curious habit. Gyr (855) has recently described the anatomy
of 6 species of Sedum which are used in folk medicine.

GENERA DESCRIBED
Adromischus, Aeonium, Bryophyllum,* Cotyledon, Crassula,* Echeveria,
Greenovia, Kalanchoe,* Rochea, Sedum,* Sempervivum,* Tillaea, Um-
bilicus.*
* Kew
Represented in the slide collection.

(So far as possible the synonymy in Berger's (176) monograph has been
adopted.)
LITERATURE
On General Anatomy
Berger 176, Cholodny 406, Dauphine" 546, 547, Dauphine and Hamet 549, Gyr 855,
Hamet 875, 876, 877, 878, Heckel 928, Jadin and Juillet 1139, Jeffrey and Cole 1167,
Johnson 1186, Kean 1224, 1225, 1226, 1227, Lindinger 1374, Maas 1405, Marloth 1445,
Mehrlich 1479, Meyer 1508, Molisch 1549, Pfeiffer 1712, Rehfous 1906, Reiche 1907,
Sobels 2156, Sporer 2171, Weingart 2388, Yarbrough 2483, 2484, Zemke 2505.

126. DROSERACEAE
(Fie. 128 on p. 582; FIG. 129 on p. 584)

SUMMARY
The morphology and biology of members of this widely distributed
family, many of which occur in boggy localities, have been described very
completely by Diels (586) and Lloyd (1383). There is, therefore, no need to
deal with them in great detail here, and it will suffice to give a brief resume
5** DRQSERACEAE
of such facts concerning their anatomy as have been recorded or observed.
The family consists of small shrubs with leaves bearing glandular hairs,
some of which secrete substances which ensnare and digest insects and
other small animal organisms. Since the morphology and anatomical struc-
ture of the plants differ somewhat, each of the genera is described separately
below.

FIG. 128. DROSERACEAE

A, Longitudinal section through a glandular tentacle of Drosophyllum lusitanicum Link. B, A similar
section through a shortly stalked gland of Drosophyllum. C, Longitudinal section through the glandular
disk of Dionaea muscipula Ellis. D, Surface-view of the same gland. E, Longitudinal section through
a glandular tentacle from the upper side of the leaf of Drosera rotundifolia Linn. F, Two-armed hair
of Aldrovanda vesiculosa Linn. G Glandular hair of Drosera rotundifolia. H, Four-rayed hair of
t

Aldrovanda. I, Stellate hair of Dionaea. A-C after Goebel, D

and J after Fraustadt, E after de Bary,
F and H after Caspary, G after Nitschke.
(m = Middle layer, t = Tracheids.)

DROSERA (Sundews)
Terrestrial herbs with a basal rosette of red leaves. Tentacles (Fig. 128 E)
present on the margin and upper surface of the leaf; each consisting of
a stalk of varied length, traversed longitudinally by fine tracheids terminating
in large groups in the heads. Outer part of the heads of the tentacles composed
of 2 external layers of glandular cells surrounding a suberized or cutinized
layer. Dome-shaped sessile glands (Fig. 128 B~D), often filled with a red
or purple fluid, also present on both surfaces of the leaf, as well as on the
stalks of the tentacles. Structures transitional between tentacles and sessile

glands recorded in certain species. Stomata present on both surfaces.

Mesophyll of D. rotundifolia Linn, composed of uniform, spongy tissue
consisting of round cells. Vascular bundles of the veins in the same species
embedded in the mesophyll and surrounded by a more or less definite sheath
of parenchymatous cells. Structure similar in D. binata Labill., but central
part of the mesophyll consisting of larger cells than those towards the 2 sur-
 DROSERACEAE 583
faces; chlorophyll granules much more numerous in the small outer than in
the large central cells. Transverse sections through the distal end of the
petiole of D. rotundifolia Lin. (Fig. 129 B) exhibit about 3 widely spaced
vascular bundles, not accompanied by sclerenchyma, but each including
well- developed phloem and a few small xylem vessels. Petiole of Z). binata
Labill. (Fig. 129 A) supplied by 2 much longer, cylindrical vascular bundles.
Scape of the same species (Fig. 129 F) with a narrow cortex, bounded in-
ternally by a broader, well-defined, mechanical ring of elongated, relatively
thick- walled cells ; a circle of vascular bundles tending to be centric in struc-
ture and including very few vessels ; a large, slightly spongy pith, composed
of thin- walled, somewhat elongated cells. In the root, according to Solereder,
the cortical parenchyma is composed of spirally thickened cells with wide
lumina in certain species the vascular bundles often being centric with central
;

phloem and arranged in a simple or multiple ring or irregularly distributed

in the ground tissue.

DROSOPHYLLUM
The following description applies particularly to D. lusitanicum (L.) Link.
General morphology similar to that of Drosera but taller, tending to be almost
shrubby. Tentacles and glands (Fig. 128 A~D) not unlike those of Drosera.
Mesophyll composed of uniform spongy tissue, but with very much larger
intercellular spaces than in either of the species of Drosera described. Vas-
cular bundles of the veins also larger than those of Drosera, the one in the
midrib being supported by a wide sheath of slightly collenchymatous cells
devoid of chloroplasts. Leaf base (Fig, 1290) composed of spongy tissue
similar to that of the lamina and supplied by 3 widely spaced, vascular
bundles. Outer part of the scape (Fig. 129 D) chlorenchymatous, somewhat
spongy. Vascular system consisting of a circle of small, widely spaced,
vascular bundles situated at the inner periphery of the chlorenchyma, and a
much more deeply seated circle of 3 vascular bundles separated from the
outer ring by a broad zone of collenchyma. It was impossible to decide
without further investigation whether the vascular system should be inter-
preted as consisting of a ring of small cauline bundles surrounding a group of
larger medullary ones, or whether the inner circle of larger bundles represents
the main cauline system, in which case the small outer ones would be regarded
as cortical.

DIONAEA (Venus's fly-trap)

Small terrestrial plants with a basal rosette of leaves whose distal ends are
differentiated as traps. The latter, which consist of two lobes of the lamina
united along the midrib, bear marginal cilia in the nature of emergences
which interlock when the leaf is closed. Outer surface of the trap provided
with stellate trichomes (Fig. 1281); inner surface covered with glands
having 2-celled stalks and multicellular heads made up of 2 tiers of cells.
Glands all morphologically alike, but differentiated physiologically into deep-
red digestive and colourless alluring types, the latter secreting substances
which attract insects. Inner surfaces also bearing jointed, sensitive bristles
which include no vascular tissue. Mesophyll composed of thin-walled
parenchyma; palisade tissue absent. Midrib traversed longitudinally by a
 K
FIG. 129. DROSERACEAE, A-D and F; HAMAMELIDACEAE, E and G-K
A, Drosera binata Labill. Petiole X 19^ B, Drosera rotundifolia Linn. Petiole X 33. C, Drosophyllum
lusitanicum Link. Leaf base X 31. D, D. lusitanicum Link. Inflorescence axis X 33. E, Liquidambar
;

styraciflua Linn. Petiole x 8. F , Drosera binata Labill. Infloresence axis X

x 19. G, Corylopsis spicata
S'. et Z. Petiole X 19. H, Fortunearia sinensis Render. Petiole X 32. I, Disantkus cerddifolius Maxim.
Petiole Xi8. J, Liquidambar styraciflua Linn. Stem X 15. K, Hamamelis japonica S, et Z. Stem Xi5.
 DROSERACEAE 585
double vascular bundle. Winged petiolar portion of the leaf described as
having a single vascular bundle. Molisch (1547) demonstrated the presence
of tanniniferous material in the epidermis and ground tissue of the leaf.
This is normally in solution, but can be precipitated in crystalline form by
treatment with hot water, dilute mineral acids, and other reagents.

ALDROVANDA
The genus represented by a single species A. vesiculosa Linn., a rootless,
is
with whorls of leaves attached to one another at their bases.
floating plant
Leaves somewhat resembling those of Dionaea, the lobed lamina with
marginal bristles being differentiated as a trap at the distal end of the flattened
petiole. Outer surface of the trap bearing short, 2-armed trichomes
(Fig. 128 F), together with almost sessile glands with swollen, multi-
cellular heads and a limited number of long, slender, sensitive bristles, con-
tact with which causes the trap to close. Transverse sections of the petiole
of Aldrovanda show polygonal air cavities below the epidermis, separated
from one another by parenchymatous lamellae; centre of the petiole occupied
by a single, somewhat reduced, vascular bundle. Vascular system of the scape,
according to Solereder, consisting of a ring of phloem and a little xylem
parenchyma, the latter enclosing an air passage.

GENERA DESCRIBED
Aldrovanda, Dionaea, Drosera,* Drosophyllum.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Diels 586, Lloyd 1383, Molisch 1547.

127. BYBLIDACEAE
SUMMARY
A small family of glandular herbs and small shrubs comprising the genera
Byblis from Australia and Roridula from Africa. Byblis, which is doubtfully
carnivorous, has been fully described by Lloyd (1383), who also considers
that Roridula is not carnivorous, although it was at one time thought to be so.
There are 2 species of Byblis, B. linifolia Salisb. and B. gigantea LindL, both
of which have rhizomes bearing the branched aerial portions of the plant with
linear leaves.

BYBLIS
Leaf triangular in transverse section with rounded angles, but nearly
cylindrical towards the apex; terminated by a knob with the function of a
hydathode supplied by well-developed tracheidal tissue. Stalked and sessile
glands present on all aerial parts of the plant; sessile glands on the leaf con-
fined to longitudinal furrows with glandless ridges between them. Stalked
glands scattered; provided with multicellular, umbrella-shaped heads; capable
of secreting abundant mucilage. Epidermis composed of thick-walled cells.
586 BYBLIDACEAE
Stomata situated in the furrows between the sessile glands; rubiaceous.
Mesophyll. Portions immediately below the epidermis consisting of chlor-
enchyma, which is not differentiated into spongy and palisade tissue central
;

part occupied by colourless ground parenchyma. Vascular bundles of the

leaf accompanied by mechanical tissue. In the stem Diels (585) records
the existence of a continuous starch sheath (endodermis), bounded internally
by a ring of sclerenchyma. Vascular system said to consist of about
9 bundles, with weakly bordered pits to the vessels in the xylem; perforations
simple. Solereder describes the vascular bundles as showing transitions from
the collateral to the centric type. Pith composed of large, lignified, pitted cells.
Young root said to be triarch,

RORIDULA
The following information concerning the structure of Roridula has been
taken from Solereder.

LEAF
Provided with tentacular glands of varying length; the latter consisting
of multiseriate stalks bearing ellipsoidal heads, composed of a central group
of isodiametric cells surrounded by a palisade-like secretory epidermis. Upper
surface of the leaf furrowed. Stomata confined to the lower surface. Meso-
phyll spongy; the lower part containing large lacunae. Vascular bundles of
the veins accompanied by sclerenchyma. In the stem a ring composed of
fibre-like cells adjoins the vascular bundles. In Roridula dentata L. the wood
fibres have simple pits the isolated vessels are spirally thickened, and with
;

bordered pitting where in contact with the rays; the perforation plates
scalariform with many bars. Rays 1-2 cells wide.

TAXONOMIC NOTES
The
genera Byblis and Roridula were included in the Droseraceae in the
system of Bentham and Hooker. On external morphological grounds both
genera have been placed by Hutchinson (1113), who followed Domin, in a
separate family, the Byblidaceae, believed to be related to the Pittosporaceae.
Byblis is also described under Byblidaceae in the Engler system, but Roridula
in the Roridulaceae. Byblis was at one time also referred to the Lentibu-
lariaceae. Roridula is distinguished by the scalariform perforation plates
in the vessels. Byblis, unlike the Droseraceae, has rubiaceous stomata. The
anatomical features thus tend to confirm the view that the resemblances
between the 2 genera and the Droseraceae are superficial.

GENERA DESCRIBED
Byblis, Roridula.
LITERATURE
OnGeneral Anatomy
Diels 585, Hutchinson 1113, Lloyd 1383.
 (58?)

128. HAMAMELIDACEAE
(Fie. 129 on p. 584; FIG. 130 on p. 590)

SUMMARY
(i) GENERAL
A
family of trees and shrubs which occur in the Far East, Madagascar,
Africa, and North America. The leaf is always dorsiventral, the hairs almost
exclusively non-glandular, and mostly tufted or stellate; stomata, confined
to the lower surface, are rubiaceous, with one or more subsidiary cells on
either side of the pore. Sclerenchymatous idioblasts of various types are
common in the mesophyll. The vascular bundles of the veins are frequently
surrounded by sclerenchyma, but in some species there is only an arc of
sclerenchyma adjacent to the phloem. Transverse sections through the distal
end of the petiole usually show a cylindrical or nearly closed vascular strand,
accompanied, in some genera and species, by additional bundles in the wings
on the adaxial side. The vascular structure of the petiole of Liquidambar
styraciflua Linn, is more complicated. Both solitary and clustered crystals
occur, the former sometimes appearing as transparent dots in the leaf. In the
young stem the cork usually arises in the sub-epidermis, and the pericycle
generally contains a composite and continuous ring of sclerenchyma. The
xylem and phloem, in internodes of young stems, constitute continuous
cylinders traversed by uniseriate or biseriate rays. The xylem includes
numerous small, angular vessels, seldom exceeding 25 p in diameter, and
provided with scalariform perforation plates with numerous bars. Small,
medullary, secretory canals occur in the stems ofAltingia and Liquidambar,
the 2 genera included in the Liquidambaroideae, and extend into the veins of
the leaf as well as into the root.

(ii) WOOD
Vessels extremely small to moderately small, often entirely solitary, some-
times with a little spiral thickening, perforation plates scalariform, with few
tomany bars, intervascular pitting scalariform to opposite, pits to paren-
chyma often large and horizontally elongated; members moderately to
extremely small. Parenchyma apotracheal, typically diffuse, but some-
times banded. Rays typically 2-3 cells wide, exclusively uniseriate in a few
genera, heterogeneous. Fibres with large, distinctly bordered pits, moderately
to very long. Intercellular canals of the traumatic type, vertical or radial,
very occasionally present.

LEAF
Always dorsiventral. Hairs almost exclusively non-glandular, mostly
tufted to stellate, often with thick walls and narrow lumina. Simple, uni-
cellular hairs occasional. Glandular leaf teeth, each containing a bundle
termination, composed largely of cells filled with tanniniferous mucilage and
provided with stomata on the upper surface, occur in Liquidambar styraciflua
Linn. Cork warts reported on both surfaces in Altingia excelsa Non Cuticle
strongly developed and granular in Bucklandia, Corylopsis, Dicoryphe,
Rhodoleia, Trichocladus. Epidermis stated to include mucilaginous cells in
588 HAMAMELWACEAE
Rhodoleia championii Hook. f. and R. teysmannii Miq. Hypoderm on the
upper side of the leaf known to occur only in Altingia exceha and A. chinensis
(Champ.) Oliver. Stomata confined to the lower surface; rubiaceous, with
i or more subsidiary cells on either side of the pore.
Subsidiary cells some-
times considerably elongated at right angles to the long axis of the stoma as
seen in surface view. Mesophyll including i layer of palisade tissue in
species of Corylopsis, Disanthus, Fothergilla, Hamamelis, Parrotia\ 2 layers in
species of Altingia, Bucklandia, Distylium, Eustigma, Liquidambar, Low-
petalum, Sycopsis, Trichocladus, and several layers in Rhodoleia. Spongy
tissue usually very lacunar. Mesophyll including the following types of
sclerenchymatous idioblasts. (i) Branched, but short and gnarled in Buck-
landia and Rhodoleia. (ii) Columnar, occasionally branched, mostly elongated
in the same direction as the palisade cells in Eustigma oblongifolium Gard.
et Champ, and Hamamelis virginiana L. (iii) Irregular sclerenchymatous
fibres with thick walls and narrow lumina in Dicoryphe stipulacea J. St. Hil.

(iv) Fibres similar to (iii) but with thinner walls and wider lumina in Disty-
lium, Loropetalwn, Sycopsis. Vascular bundles of the veins sheathed or
accompanied by an arc of sclerenchyma in Corylopsis, Dicoryphe, Distylium,
Eustigma, Fothergilla, Hamamelis, Loropetalum, Parrotia, Sycopsis, Tricho-
cladus, i.e. in the Hamamelidoideae; sclerenchyma associated with the veins
only poorly developed in Altingia, Bucklandia (sometimes lacking in this
genus), Liquidambar, Rhodoleia. The bundle system of the larger leaf veins
forms a ring of xylem and phloem in Altingia and Liquidambar, but is
crescent-shaped in the other genera. Petiole, in transverse sections through
the distal end, exhibiting a continuous, cylindrical vascular strand in Cory-
lopsis platypetala Sieb. et Zucc., C. spicata Sieb. et Zucc. (Fig. 129 G),
Disanthus cercidifolius Maxim (Fig. 129 i) (vascular ring sometimes not quite
closed), Fortunearia sinensis Rehder (Fig. 129 H), Fothergilla gardeni Murr.,
F. major (Sims) Lodd., Hamamelis mollis Oliv., H. virginiana Linn., Liquidam-
bar formosana Hance, Loropetalum chinense R. Br., Parrotia persica C. A.
Meyer, with a very slightly interrupted cylindrical strand in Bucklandia
populnea R. Br. vascular strand crescent-shaped but interrupted in Hamamelis
;

japonica Sieb. et Zucc. and Sinowilsonia henryi Heinsl. vascular strand con-
;

tinuous, crescent-shaped but with the ends almost in contact with one another
and somewhat prolonged towards the adaxial side in Sycopsis sinensis Oliv.
Additional strands present in the wings in the same species of Bucklandia and
Liquidambar. Vascular system, in the corresponding region of the petiole of
Liquidambar styraciflua Linn. (Fig. 129 E), more complex, consisting of an
interrupted circle of separate strands the abaxial ones of which are crescent-
shaped, accompanied by 2 small cylindrical strands in the wings and an
inversely orientated one in the medullary region. For further details concern-
ing the course of the vascular bundles in a few members of the family see
Morvillez (1561). Secretory cells, with unidentified but probably muci-
laginous or tanniniferous contents, occur in the unlignified tissues t>f the
petiole in all of the above genera
and species except Sinowilsonia henryi.
Secretory canals present on the inside of the xytem in both species of
Liquidambar. Crystals both solitary and clustered; specially large ones in
the palisade tissue appear as transparent dots in species of Corylopsis, Pother-
gilla, Loropetalum^ Parrotia, Trichocladus. Clustered crystals
in the mesophyll
 HAMAMELIDACEAE 589
are characteristic of Altingia, Bucklandia, Liquidambar, Rhodoleia, i.e. mem-
bers of the Bucklandioideae; solitary ones occur in the corresponding position
in Corylopsis, Dicoryphe, Eustigma, Fothergilla, Hamamelis, Loropetalum,
Parrotia, Sycopsis, Trichocladus, i.e. members of the Hamamelidoideae. This
distinction does not hold good in the axis. Small crystals accompany the
vascular bundles in Altingia and Liquidambar, but a few clustered and
numerous solitary ones occur in the corresponding position in Corylopsis and
Hamamelis.

Axis
YOUNG STEM (Fig. 129 J-K)
Cork precocious in all investigated genera except in Bucklandia and
Rhodoleia; originating in the sub-epidermis in Disanthus, Distylium, For-
tunearia, Fothergilla, Hamamelis, Liquidambar, Parrotia, Sinowilsonia, Sycop-
sis, Trichocladus; composed of thin-walled cells; somewhat spongy in
Distylium, Parrotia, Sycopsis. The development of phelloderm recorded or
observed in Corylopsis, Dicoryphe, Distylium, Hamamelis, Liquidambar,
Trichocladus. Primary cortex, especially the middle part, often collenchy-
matous sometimes including sclerotic cells in species of Altingia, Bucklandia,
;

Dicoryphe, Distylium, Liquidambar, Sycopsis. Pericycle with a composite,

continuous ring of sclerenchyma in all genera except Altingia and Liquidam-
bar (Fig. 129;) and sometimes in Loropetalum where it is interrupted.
Sclerenchymatous elements recorded in the secondary phloem in Dicoryphe,
Distylium, Fothergilla, Sycopsis. Xylem and phloem in the internodes con-
stituting a closed cylinder traversed by uniseriate or biseriate rays. Xylem
including numerous, small, angular vessels, seldom exceeding 25 p, in
diameter and provided with scalariform perforation plates with numerous
bars. Pith usually homogeneous, but heterogeneous in Liquidambar stryaciflua
Linn, and L. orientalis Mill., the component cells stated sometimes to be
elongated transversely to the longitudinal axis in Altingia, Corylopsis,
Hamamelis, Liquidambar', longitudinally elongated in Bucklandia and Rhodo-
leia. Secretory elements. Cells, with unidentified but probably tannini-
ferous or mucilaginous contents, present in varying numbers in the unlignified
tissues of all of the species represented in the Kew slide collection except
Sinowilsonia henryi. (For genera concerned see list of 'Genera described',)
A few (usually about 7 or 8) secretory canals occur in the pith very close to
the protoxylem groups in Altingia and Liquidambar; i.e. the 2 genera in the
Liquidambaroideae, each canal surrounded by small-celled epithelium, and
extending into the veins of the leaf and into the root. Medullary canals
recorded also in Ostrearia. Cortical secretory canals stated to occur in
Mytilaria (Harms 897).

WOOD (Fig. 130)

Vessels small, varying from about 25 to 90 \L mean tangential diameter,
largest (more than 50 p,) in Altingia, Bucklandia^ Rhodoleia, Sycopsis] Tippo
(2261) gives the mean for the family as 32 /A; exclusively solitary, apart from
the apparent tangential pairs due to overlapping ends, in Bucklandia, Distylium,
Eustigma (1206), Fortunearia, Fothergilla, Hamamelis, Loropetalum, Parrotia>
Parrotiopsis, Rhodoleia, Sycopsis, and Trichocladus, with some multiples of
590 HAMAMELIDACEAE
2 or 3 cells in the other genera; numerous, mostly 50-100 per sq. mm., but
considerably more than 100 per sq. mm. in some
species, e.g. Hamamelis
japonica Sieb. ; spiral thickening present in the tips of the vessel members of
Corylopsis and Rhodoleia (2158) and on the walls of Liquidambar formosana
Hance (1206); Record (1851) includes Hamamelis in a list of genera with

FIG. 130. HAMAMELIDACEAE

A, Liquidambar styraciflua Linn. B, Trichocladus crinitus
T ^ Pers.
r>
C, Hamamelis japonica
,._.!_,_,_. *...
Sieb. et Zucc.
^ r,,.. ,.
D, Liquidambar styradflua Linn. E, Altingia excelsc
?/$a J.S.G. F, Rhodoleia teysmanni Miq, G, Dtstylium
racemosum Sieb. et Zucc.

spiral thickening. Perforation plates all scalariform, mostly with fewer than
20 bars, but with more in Bucklandia, Liquidambar, Rhodoleia, and Tetra-
thyriwn\ the bars commonly anastomosing to give partially reticulate plates;
the perforations usually without borders (2261). Intervascular pitting scalari-
form to opposite, difficult to find in the woods with solitary vessels pits to ;

ray and wood parenchyma predominantly large and horizontally elongated,

simple or with distinct borders. According to Bailey, the sieve-like structures
reported by Janssonius in Altingia are not true vestured pits. Tyloses common
 HAMAMELIDA CEAE 591

(2261), present, for example, in Altingia, Liquidambar, Parrotiopsis, and

Sycopsis. Mean member length 0-8-2-4 mm5 longest
- in Bucklandia. Paren-

chyma apotracheal, sometimes scarce, e.g. in Liquidambar typically as

9
,

scattered cells (diffuse) (Fig. 130 B), but banded (metatracheal) in Distylium
(Fig. 130 G), Fortunearia, Loropetalum, Parrotia, and Sycopsis, the bands i cell
wide in most species but up to 3, or more rarely 4, cells wide in Distylium;
some authors (1154, 1679) refer to paratracheal parenchyma, but it is difficult
to distinguish this from diffuse parenchyma that happens to be contiguous
with some of the numerous vessels. Cells commonly filled with dark gum-like
substance; chambered crystal cells present in a few species. Strands of 8-16
cells. Rays tending to be of 2 distinct widths in a few genera, e.g. Corylopsis,

Eustigma, Sinowilsonia, and Tetrathyrium\ typically 2-3 cells wide, commonly

up to 4 cells wide in Altingia exclusively uniseriate or with only occasional
;

small biseriate parts in Bucklandia, Fothergilla, Hamamelis, Parrotiopsis (small

stem), and Tetrathyrium; usually less than i mm. high, but rather more in a
few species; uniseriates in woods with multiseriate rays, numerous to only
moderately numerous, usually composed of both upright and procumbent
cells, but sometimes almost entirely of upright cells; typically more than
12 rays per mm. and up to 25 per mm. in Trichocladus, but rather fewer than
12 per mm. in Altingia, Bucklandia, Fothergilla, Liquidambar, and Rhodoleia;
heterogeneous (Kribs's Types II A and B, and III, rarely I), typically with
4-10 marginal rows of upright cells, but seldom with more than 3 rows in
Liquidambar, and with more than 10 rows in Sycopsis and Trichocladus. Cells
usually filled with dark gummy substance; containing crystals in only a few
species, e.g. Altingia. Fibres with large, distinctly bordered pits, equally
numerous on both radial and tangential walls commonly thick- walled. Tippo
;

(2261) classifies these elements as true tracheids in the sense defined by

Bailey (74). Solereder refers to spiral thickening in Hamamelis chinensis R. Br.
Mean length 1-4-2-9 mm., longest in Altingia and Bucklandia. Intercellular
canals of the vertical traumatic type are recorded by various authors in some
species of Altingia and Liquidambar and occasional schizogenous canals are
noted by Janssonius (1154) in the rays of Altingia excelsa Nor.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
Harms (897) has drawn attention to the fact that the taxonomic subdivisions
of the family exhibit corresponding anatomical differences. This applies
particularly to the type of crystal in the mesophyll and to the distribution of
sclerenchyma around the vascular bundles of the veins. The presence of
medullary secretory canals in Altingia and Liquidambar serves to distinguish
these genera from others which have been examined with the exception of
Mytilaria.

(ii) FOR WOOD STRUCTURE

Tippo (2261). 'The secondary xylem of this family is very close in all
anatomical details to certain groups in the Magnoliales as described by
McLaughlin. The Magnoliales exhibit a considerable range of anatomical
structures, yet it may be said that on the whole the least specialized members
of that order are more primitive anatomically than are the Hamamelidaceae/
 592 HAMAMELIDACEAE
ECONOMIC USES
Storaxthe balsam produced when the bark of Liquidambar orientalis Mill,
is
is wounded. It is said to be chemically distinct from the material present in
the secretory canals in the same genus. It is used to a limited extent in
medicine as a remedy for skin complaints. The leaves and bark of Hamamelis
virginiana Linn, are used in medicine to prepare a cooling lotion which
has astringent properties. The leaves may be recognized by the stellate or
tufted hairs made up of unicellular components with thick walls and which
occur particularly below the veins on the lower surface of the leaf; the
rubiaceous stomata confined to the lower surface; the single layer of palisade
cells; the slightly branched, sclerenchymatous idioblasts in the mesophyll;
the mostly solitary crystals of calcium oxalate which occur in the mesophyll
and in chambered cells which accompany the fibrous sheaths around the
veins. Microscopical characters for the bark of the same species include: the
cork composed of broad layers of thin- walled cells alternating with narrower
zones of sclerotic cells the zone consisting of several layers of stone cells at
;

the inner periphery of the phelloderm; the groups of stone cells in the outer
part of the secondary phloem; the groups of fibres, also in the secondary
phloem, accompanied by chambered cells containing prismatic crystals; the
narrow medullary rays the absence of starch and the presence of tanniniferous
;

substances in the parenchymatous tissues.

Various members of the family are cultivated in gardens either for their
early flowers which come into bloom before the leaves develop, e.g. Hamamelis
spp., or because of the beautiful tints assumed by the autumn foliage,
e.g. Parrotia persica C. A. Meyer.
Useful timbers are produced by species of Altingia and Bucklandia from
the Indo-Malayan region and Liquidambar from North America.
The American Red or Sweet Gum, Liquidambar styraciflua Linn., is widely
used for a great variety of purposes, but particularly for furniture and
interior trim.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Altingia, Bucklandia,* Corylopsis,* Dicoryphe, Disanthus,* Distylium,*
Eustigma, Fortunearia,* Fothergilla,* Hamamelis,* Liquidambar,* Loro-
petalum,* Maingaya, Mytilaria, Ostrearia, Parrotia,* Rhodoleia, Sinowil-
sonia,* Sycopsis,* Trichocladus.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Altingia, Bucklandia, Corylopsis, Disanthus, Distylium, Eustigma, Fortune-
aria, Fothergilla, Hamamelis, Liquidambar, Loropetalum, Parrotia, Parro-
tiopsis, Rhodoleia, Sinowilsonia, Sycopsis, Tetrathyrium, Trichocladus.

LITERATURE
(i) On General Anatomy
Harms 897, Holm 1024, Morvillez 1561, Puri 1763, Starr 2188, Varossieau 2324.
 HAMAMELIDACEAE 593
(ii) On Wood Structure
Bailey 74, Beekman 167, den Berger 179, 182, den Berger and Endert 183, Brown and
Panshin 288, 289, Chowdhury 411, Desch 574, Greguss 2522, Hale 870, Howard 1088,
Janssonius 1154, Jones 1191, Kanehira 1206, 1209, Lecomte 1334, Nicoloff 1593, Pearson
and Brown 1679, Record 1780, 1783, 1787, 1800, 1801, 1818, 1825, 1843, 1851,
Record and Hess 1886, Record and Mell 1894, Stone 2203, Sudworth and Mell 2217,
Tang 2231, Tippo 2261, Webber 2377, Varossieau 2324.

129. MYROTHAMNACEAE
SUMMARY
(i) GENERAL
A
small family of xeromorphic, resinous shrubs, consisting of 2 species of
the single genus Myrothamnus from Africa and Madagascar. Important
anatomical characters include the occurrence of resin cells in the epidermis
of the leaf and of ranunculaceous stomata, whilst in the xylem of the axis
the small angular vessels mostly have scalariform perforation plates with
many bars.

(ii) WOOD
Vessels small, mostly solitary, perforation plates scalariform, members
moderately long. Parenchyma absent. Rays uniseriate and heterogeneous.
Fibres with bordered pits.
LEAF
Hairs absent. Cuticle fairly thick, but thinner above the resin cells of the
epidermis. Epidermis mostly composed of polygonal cells, but including
scattered, enlarged, resin cells. Stomata ranunculaceous. Mesophyll iso-
bilateral, consisting of tangentially elongated cells towards the centre, but
composed of palisade cells elsewhere. Vascular bundles of the veins em-
bedded in the mesophyll. Cluster crystals present.

Axis
YOUNG STEM
Xylem cruciform when young but later forming a continuous cylinder;
including, according to Solereder, small, quadrangular vessels with minute
bordered pits and perforation plates which are very oblique, with many bars,
or consist of small elliptical pits arranged in parallel rows thick- walled fibres
;

with inconspicuously bordered pits uniseriate rays. Pith composed of cells

;

elongated in the same direction as the axis.

WOOD 1

Vessels very small (mean diameter about 35 p)\ mostly solitary, but with
some groups; perforation plates scalariform with about 45 bars and very
oblique; intervascular pitting transitional and opposite; pits to ray cells
similar. Mean member length about 0-9 mm. Parenchyma absent. Rays

exclusively uniseriate, heterogeneous (Kribs's Type III). Fibres with bor-

dered pits.
1
Based entirely on the description given by Tippo (2261).
4594
Q q
 594 MYROTHAMNACEAE
TAXONOMIC NOTES
Myrothamnus was included in the Hamamelidaceae in the Bentham and
Hooker system. While it is still generally held that the genus is related to the
Hamamelidaceae, Hutchinson (1113) and Niedenzu and Engler (1599) treat
it as a distinct
family. The presence of resin cells in the leaf epidermis and
of ranunculaceous stomata are anatomical characters which favour the separa-
tion of the Myrothamnaceae from the Hamamelidaceae. Tippo (2261) has
pointed out that the wood structure in the 2 families is
very similar.

GENUS DESCRIBED
Myrothamnus.
LITERATURE
(i) On General Anatomy
Hutchinson 1113, Niedenzu and Engler 1599, Zemke 2505.

(ii) On Wood Structure

Record 1843, 1851, Thompson 2254, Tippo 2261.

130. BRUNIACEAE
(Fic. 131 on p. 598)

SUMMARY
Heath-like shrubs from South Africa. In correlation with the habit, the
leaf is often centric, but tends to be partly or wholly dorsiventral in some

species. The apex of the mature leaf, except in Audouinia, usually bears a
small projection composed of suberized cells. In species with strongly
adpressed leaves the palisade tissue is sometimes confined to the lower surface.
The distribution of the stomata, which are ranunculaceous but surrounded
by rather small epidermal cells, also varies somewhat according to the mor-
phology of the leaves and the extent to which they overlap or are adpressed
to the stem. The hairs are mostly long, slender and unicellular, with thick
smooth walls and narrow lumina. The narrow primary cortex of the stem
sometimes contains stone cells, whilst the xyiem is characterized by small
vessels having scalariform perforation plates with numerous bars, and fibres
with bordered pits. Crystals both solitary and clustered.

LEAF
Usually centric, but sometimes dorsiventral or intermediate between the
2 types. Hairs mostly long, slender, unicellular with a thick smooth wall and
narrow lumen; sometimes accompanied by shorter hairs in Audouinia. The
tip of the leaf, except in Audouinia, consists of a cap of cork cells replaced, as
the outermost ones become detached, by the activity of a central meristem.
Cuticle thick, especially on the outwardly directed abaxial surface; smooth
but longitudinally striate in Berzelia lanuginosa (L.) Brongn., and Nebelia
paleacea (Berg.) O. Kze. Cells of the epidermis large, especially above and
below the median vein, progressively smaller towards the margin; more or
less arched outwards sometimes papillose. Distribution of stomata varying
;

according to the external morphology of the leaf, several distinct types occur-
 BRUNIACEAE 595

ring within some of the genera; confined to the lower surface in Linconia
cuspidata (Thunb.) Swartz and Pseudobaeckea\ restricted to the upper side in
species with broad scaly leaves belonging to the genera Brunia, Lonchostoma^
Pseudobaeckea, and Raspalia; frequently arranged in 1-5 longitudinal rows
with the long axis parallel to the median vein, but exceptions occur. The
4-7 surrounding cells are often rather smaller than those of the remainder of
the epidermis. MesophylL Palisade tissue consisting of i, or occasionally 2,
layers; confined to the abaxial surface in species with adpressed leaves.
Sclerenchymatous cells recorded by Solereder in the mesophyll of Linconia
cuspidata and observed in Lonchostoma monostylis Sond. Water-storage
cells reported to occur at the apex of the leaf near the terminations of the
veins. Leaf often traversed by 3 veins, but 5 or even as many as 20 recorded
in certain species. Crystals usually present, except in Raspalia, in the

mesophyll or around the vascular bundles; more frequently clustered than

solitary. Mesophyll cells observed to have apparently tanniniferous contents
in Lonchostoma monostylis.

Axis
STEM (Fig.1310)
epidermis arched outwards. Cork thin- walled, arising imme-
Cells of the
diately below the epidermis. Primary cortex narrow; containing sclerosed
cells in speciesof Brunia Nebelia, Staavia, Tittmannia. Pericycle usually
y

containing a composite ring of sclerenchyma. Xylem, according to Niedenzu

and Harms (1600), characterized by small, mostly isolated vessels having
scalariform perforation plates with many bars (this was confirmed by direct
observation for Audouinia capitata (Thunb.) Brogn., but many of the vessels
in Lonchostoma monostylis Sond. were seen to be in tangential pairs, frequently

arranged to form irregular clusters) fibres with bordered pits scanty paren-
; ;

chyma; rays 1-3 cells wide with, according to Solereder, the distal ends
enlarged where traversing the phloem. Pith in Audouinia capitata and Lon-
chostoma monostylis seen to be composed of somewhat elongated, fairly thick-
walled, pitted cells. Resinous deposits observed in Audouinia capitata
situated in certain of the cells of the cork, phloem, medullary rays, and pith,
those in the phloem being elongated and resembling narrow sacs. Similar
deposits much less numerous or almost absent in Lonchostoma monostylis.

GENERA DESCRIBED
Audouinia,* Berzelia, Brunia, Linconia, Lonchostoma,* Nebelia, Pseudo-
baeckea, Raspilia, Staavia, Tittmannia.
*
Represented in the Kew slide collection.

LITERATURE
On General Anatomy
Niedenzu and Harms 1 600.
 (59*)

131. HALORAGACEAE
(FiG. 131 on p. 598; FIG. 132 on p. 600)

SUMMARY
Herbs, often very large, which grow in or near water in many parts of the
world. The plants are frequently covered with a variety of hairs, warts,
and emergences, some of the hairs being glandular. Some of the glandular
structures on the stem of Gunnera serve as channels for the ingress of colonies
of the blue-green alga Nostoc, which are normally to be found in the super-
ficial layers of the stem. The stomata are usually ranunculaceous, and are
said sometimes to be ephemeral. The mesophyll is dorsiventral or sub-
centric, but typical palisade cells are seldom differentiated. Clustered
crystals are common. The vascular system of the petiole of Gunnera con-
sists of separate steles. The primary cortex of the stem contains numerous

air-cavities, which are particularly well developed in aquatic species. The

vascular system of the stem exhibits varying degrees of complexity, cul-
minating in the polystelic network which occurs particularly in the upper
internodes of Gunnera. Each stele is surrounded by a well-defined endo-
dermis with Casparian thickenings. In the xylem the relatively narrow
vessels with simple perforations are embedded in ground tissue consisting of
fibres with thick walls, narrow lumina, and simple pits. Batham (151) has
shown that the cauline vascular structure of the relatively small species of
Gunnera which occur in New Zealand is similar to, although less complex
than that of, the large species from other countries. The vascular system of
the New Zealand species is said to consist of a tube with leaf trace gaps in the
sides. Toquote Batham's words: 'The "polystelic" state suggested by
when surveyed three dimensionally, into a
isolated sections resolves itself,
hollow monostele with foliar gaps and branches.'

LEAF
Dorsiventral or sub-centric. Hairs, (i) Unicellular in species of Gunnera
and Haloragis. (ii) Uniseriate in Haloragis^ Laurembergia, Loudonia, Meziella,
Myriophyllum, Proserpinaca. (iii) Glandular, sometimes flask-shaped, shaggy
types appear under the lens as small dots, in the leaf teeth, &c. composed of
;

polygonal cells containing highly refractive oily bodies in all genera except
Loudonia, Hemispherical warts (colleters), with the epidermal cells arranged
in the form of a fan as seen in longitudinal section, recorded in several species
of Gunnera (Fig. 132?). Other types of hairs and wart-like structures also
occur in the family. For further particulars of hairs and related structures in
Gunnera macrophylla Bl. see Skottsberg's (2121). article. Stomata usually
ranunculaceous, more numerous in species with aerial than in those with
submerged leaves; stated sometimes to be ephemeral; usually present on
both surfaces of the leaf; smaller and less numerous on the upper than on the
lower surface in terrestrial species of Haloragis and Loudonia. Mesophyll
frequently devoid of palisade cells, but these sometimes occur in Haloragis
and Loudonia. Petiole. Larger vascular strands of Gunnera consisting of
distinct steles, frequently composed of central xylem surrounded by an
 HALORAGACEAE 597
endodermis, but in certain species including a sclerenchymatous pith or
pseudopith. The pith, in the most complex types of all, is replaced by a
vascular system consisting of central strands of phloem enclosing some
xylem vessels, the whole being surrounded by 2 collenchymatous sheaths and
supported by 2 strands of fibres. Transverse sections through the distal end
of the petiole of Haloragis alata Jacq. (Fig. 131 F) exhibit an arc of widely
spaced vascular bundles, each surrounded by a distinct endodermis. Tannin
common. Crystals clustered; usually small, but large ones occurring in
Gunnera.

Axis
STEM (Fig. 131 E)
Glands resembling shaggy hairs, with only narrow canals between them,
but covered with a cap-shaped layer of epidermal cells (Fig. 132 G), occur
between the leaf bases in Gunnera macrophylla Bl. The glands secrete mucilage
containing tannin, which swells up and ruptures the cap of epidermal cells.
They sometimes serve as an entrance for colonies of the blue-green alga
Nostoc which become established in the outer part of the stem. An account
of a developmental study of the symbiotic relationship between Nostoc and
Gunnera has been published by Miehe (1516). Primary cortex containing
numerous those in aquatic being much larger than the ones in
air cavities,
Air cavities radially elongated and arranged in a ring in
terrestrial species.

Myriophyllum and Serpicula (Fig. 132 A). Cortex said to contain palisade
tissue and small strands of fibres in the sub-epidermal region of Loudonia\
outer part seen to consist of spongy assimilatory tissue in Haloragis alata.
Endodermis, especially in aquatic species, more or less well defined.
Isolated groups of sclerenchymatous cells recorded in the pericycle of a few
species of Haloragis, Loudonia, Proserpinaca. Vascular system exhibiting
various degrees of complexity in different genera and species, (a) Consisting
of an axile, fibro-vascular mass without true pith in Myriophyllum. A pseudo-
pith is formed in older stems of this genus by resorption of the central primary
vessels, (b) Structure more or less normal with a ring of cambium and a
continuous cylinder of secondary xylem traversed by narrow rays in terrestrial
species such as Haloragis alata (Fig. 131 E) and Loudonia aurea Lindl.
(c) With a large or small number of variously orientated, separate
steles in

Gunnera, particularly in the higher internodes. Each stele is surrounded by

a well-defined endodermis with Casparian thickenings on the radial walls,
followed on the inside by a pericycle of i or 2 cells broad, 2-6 isolated strands
of phloem, and finally the central xylem consisting of vessels embedded in
thin-walled parenchyma. For details concerning the relatively simple but
polystelic structure of the small species of Gunnera from New Zealand see
Batham (151). Xylem including vessels with narrow lumina, simple perfora-
tions, and, where in contact with the rays, bordered pits. Wood fibres com-
posed of elements with fairly thick walls, narrow lumina, and simple pits.
The above remarks concerning the xylem refer particularly to Haloragis, but
simple perforations in the vessels are stated to occur throughout the family.
Pith frequently absent, but fairly broad and consisting of somewhat spongy
parenchyma in Haloragis alata. Crystals, situated in hair-like cells, recorded
in the cortex of species of Haloragis, Laurembergia, Meziella, Myriophyllum^
 Fie. 131, COMBRETACEAE, A-D; HALQRAGACEAE, E-F; BRUNIACEAE, G;
HIPPUWDACEAE, H; RHIZOPHORACEAE, I-J
A, TerminaliachebulaRetz. Petiole x8. B, T. chebulo Retz. Stem Xi3. C, Calycopteris floribunda
Lam, Petiole x8. D, C. floribunda Lam, Stem x8. E, Haloragis data Jacq. Stem Xis. F H,alata
t

Jacq. Petiole xi 5. G, Lonchostoma monostylis Sond. Stem X 15. H, Hippuris vulgaris Linn. Stem
X 8. I, Bruguiera gymnorhiza Lam, Petiole x 8. J, B. gytnnorhiza Lam, Stem X 6.
 HALORAGACEAE 599
Proserpinaca, Serpicula. Solitary and clustered crystals observed in the pith
of Haloragis alata. Acicular crystals recorded in i species of Gunnera.

STOLON
Stolons of Gunnera exhibit 2 rings of xylem and phloem, the inner one
being inversely orientated. One or more steles present in different species.
For particulars of the vascular structure of the stolons of New Zealand species
of Gunnera see Batham (151).

ROOT
Roots of NewZealand species of Gunnera said by Batham (151) probably
to consist entirely of primary xylem and phloem.

ECONOMIC USES
The large species of Gunnera are frequently cultivated in marshy places in
big gardens on account of their striking foliage.

GENERA DESCRIBED
Gunnera, Haloragis,* Laurembergia, Loudonia, Meziella, Myriophyllum,
Proserpinaca, Serpicula.
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Batham 151, Miehe 1516, Skottsberg 2121.

132. CALLITRICHACEAE
Cosmopolitan, terrestrial or aquatic, annual or perennial herbs belonging
to the single genus Callitriche. The hairs of terrestrial species include glands
each having a foot and an 8-celled head covered with a vesicular cuticle.
Stellate hairs recorded in the section Eucallitriche. Stomata numerous on
both surfaces of the leaf and on the stem in terrestrial species absent from
;

the stem, but occurring sporadically on the leaves in submerged forms; con-
fined to the upper surface 01 floating leaves; sometimes ephemeral in sub-
merged species. Vascular system of the stem reduced to a weak, axile
bundle. Pith consisting of only 2 or 3 cells.

GENUS DESCRIBED
Callitriche.

LITERATURE
On General Anatomy
Pax and Hoffmann 1674.
 (6oo)

133. HIPPURIDACEAE
(Fic. 131 on p. 598; FIG. 132 on p. 600)

SUMMARY
A family represented by the aquatic herb Hippuris vulgaris Linn, which
occurs in many North Temperate regions,

LEAF
Leaves isobilateral to centric. Peltate hairs (Fig. 132 B-D) each with a
unicellular foot and multicellular head present. Stomata occur on both
surfaces. Outer part of the mesophyll towards both surfaces and at the

FIG. 132. HALORAGACEAE, A and E~G; HIPPURIDACEAE, B-D

A, Transverse section through a piece of the cortex of Serpicula repens, L. B-D, Peltate hairs of
Hippuris vulgaris L.; B in surface-view, C in section; D, fan-shaped peltate hair. E, Tip of a segment
of the leaf of Myriophyllum verticillatum L. F, Colleter from the leaf of Gunnera macrophylla Bl.
G, Longitudinal section through a stem-gland of Gunnera macrophylla. A and E by Solereder,
B-D after Rauter, F and G after Merker.

margins composed of a single layer of somewhat palisade-like cells, inter-

spersed by numerous air cavities, the latter being especially well developed
below the stomata. Palisade tissue not very clearly differentiated from the
spongy parenchyma at the centre of the mesophyll. Vascular bundles
poorly developed, embedded in the mesophyll, and surrounded by a single-
layered sheath of parenchymatous cells.

Axis (Fig. 131 H)

The axis consists of a rhizome bearing erect branches. The following
particulars refer to the latter.
Outer and inner tangential walls of the cells of the epidermis considerably
thickened, the radial ones less so. Primary cortex broad, the whole of it,
 HIPPURIDACEAE 601

except for 2 sub-epidermal layers of cells, permeated by large intercellular

spaces elongated in the same direction as the axis, the cavities being separated
from one another by multicellular plates only i cell wide. Intercellular spaces
also interrupted at the nodes by transverse plates of small-celled tissue
containing the leaf traces. Endodermis well defined for details of its develop-
;

ment see Barratt (142). Vascular system reduced to an axile strand of thin-
walled tissue, the outer part consisting of a narrow zone of phloem and the
inner part of a broader region of xylem. Vessels small in diameter, with
spiral or reticulate thickening. According to Solereder a pseudo-pith is
formed in old stems from the central primary vessels.

TAXONOMIC NOTES
Hippuris sometimes treated as a member of the Haloragaceae, but in the
is

Engler and Prantl system is given the status of a separate family. The
anatomical structure of the plant Js that of a hygrophyte, and provides little
information concerning the taxonomic affinities of the plant. The conspicuous
endodermis and the axile vascular system are not unlike those of the less
complex members of the Haloragaceae such as Myriophyllum.

GENUS DESCRIBED
Hippuris.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Barratt 142.

134. RHIZOPHORACEAE
(Fie. 131 on p. 598; Fio. 133 on p. 604; FIG, 134 on p. 606; FIG. 135 on p. 608)

SUMMARY
(i) GENERAL
Trees and shrubs, some of which constitute the chief component of the
mangrove swamps around tropical shores and estuaries. The mangrove
plants belonging to this family are characterized by stilt-roots, only the
lower parts of which are subterranean. The cortex of these stilt roots is
sponge-like in structure owing to the development of a complex intercellular
system. The leaf is usually dorsiventral, but tends to be centric in some
species. The hairs are mostly unicellular. The cuticle is often very thick.
The epidermis frequently consists of more than i layer, but a true hypo-
derm is also common, Stomata in the leaf, confined to the lower surface,
are frequently depressed and provided with a well-developed front cavity,
which is sometimes divided into 2 distinct parts. Subsidiary cells are not
usually differentiated from those of the remainder of the epidermis, although
the epidermal cells around the stomata are sometimes more oblong than their
neighbours. Sclerenchymatous idioblasts occur in the mesophyll of the
leaf, in the cortex and pith of the stem of Bruguiera and Rhtzophora, as well
as in the stilt roots. The cork in young stems generally arises superficially,
602 RHIZOPHORACEAE
and the xylem, in some genera, is traversed by fairly broad primary medullary
rays. Cells containing tannin are frequent in all tissues.

(ii) WOOD
GROUP Rhizophoreae (Bruguiera, Ceriops, Kandelia, and Rhizophord)
I.

Vessels moderately small, with fairly numerous and sometimes long

multiples and clusters; perforation plates exclusively scalariform with few
and thick bars; intervascular pitting scalariform; members of medium
length to moderately long. Parenchyma typically scanty paratracheal;
banded in Kandelia. Rays mostly 3-6 cells wide, often more than 2 mm.
high, uniseriate rays rare, homogeneous or heterogeneous, with the pro-
cumbent cells almost square. Fibres with small simple pits, of medium
length.

GROUP II. Gynotrocheae (Anisophyllea, Caraltia, Combretocarpus, Crosso-

stylis,and Gynotroches)
Vessels medium-sized to large and often very few perforation plates simple,
;

intervascular pitting typically alternate; pits to parenchyma commonly

elongated and unilaterally compound. Members of medium length to
moderately long. Parenchyma typically banded or aliform, the bands vary-
ing from narrow to broad and from regular and metatracheal to irregular and
confluent, often with some diffuse cells. Rays up to 10-15 ce ^ s wide and very
high, uniseriate rays numerous, markedly heterogeneous. Fibres with
bordered pits, moderately to very long.
GROUP III. Macarisieae (Anopyxis, Blepharistemma, Cassipourea, Macarisia,
and Sterigmapetalum)
Vessels small to large, often exclusively solitary; perforation plates simple
and scalariform (with fine bars), intervascular pitting opposite to alternate;
pits to parenchyma mostly elongated, tending to be horizontal to wood
parenchyma and irregular to ray cells; members moderately to very long.
Parenchyma typically predominantly paratracheal and limited to the abaxial
sides of the vessels, varying from aliform to confluent, with some diffuse in ;

predominantly apotracheal bands in some genera. Rays mostly 3-4 cells wide
and less than i mm. high, uniseriate rays numerous, markedly heterogeneous.
Fibres with large and distinctly bordered pits, moderately long.

LEAF
(Fig. 133 A)
Usually dorsiventral tending to be centric in species of Anisophyllea^
;

Ceriops, Kandelia. Hairs mostly unicellular with thick or thin walls; tufted
trichomes recorded in Macarisia. Glandular shaggy hairs, present on the
inside of the stipules in certain species, sometimes secrete a gum-like sub-
stance on the vegetative buds. Cork warts occur as small black spots on the
lower side of the leaf of species of Carallia, Rhizophora, Weihea. Similar
structures, described as glands and said to be formed by resorption of the
cells of the epidermis and hypoderm, recorded by Engler (633) in Poga okosa
Pierre. Cuticle often very thick; described by Arzt (45) as consisting of 2

chemically distinct layers in mangroves, the thicker outer layer staining yellow
and the inner an intense brown when treated with chlor-zinc-iodide. A
 RHIZOPHORA CEAE 603

single-layered epidermis on the upper surface recorded only in a few species

of Blepharistemma, Carallia, Weihea; locally 2-layered in several species of
Cassipourea. Hypoderm towards the upper surface i -layered in various
species of Bruguiera, Crossostylis, Gynotroches, Macarisia; i- or 2-layered in
Anapyxis\ 2- or more-layered in species of Ceriops, Poga oleosa, and Rhizo*
phora, the innermost layer in Poga oleasa including, according to Engler
(633), large, solitary, mucilage cells. According to Bowman (251) the number
of layers of hypoderm cells in Rhizophora mangle Linn, is controlled by the
concentration of salt in the water in which the plants are growing, high
salt concentration being associated with an increased number of hypodermal

layers. The leaves are also thicker when grown in normal than in diluted sea
water because the tanniniferous cells (see below) are larger. A 2-layered
epidermis accompanied by a layer of hypoderm is stated to occur in Dactylo-
petalum and Kandelia. A single-layered hypoderm towards the lower surface
also recorded by Mullan (1570) in Ceriops candolleana Arn. and in Poga
oleosaby Engler (633). Cells of the epidermis, hypoderm, and mesophyll
sometimes mucilaginous, e.g. in species of Blepharistemma, Carallia, Dactylo-
petalum, Gynotroches, Rhizophora. Stomata confined to the lower surface;
depressed and often provided with a front cavity, the latter sometimes divided
into 2 distinct parts surrounding cells sometimes more oblong than those
;

of the remainder of the epidermis in Blepharistemma, Bruguiera, Ceriops,

Kandelia. A mixture of ranunculaceous, cruciferous, and rubiaceous stomata
recorded by Sprague and Boodle (2173) in Anapyxis ealaensis Sprague.
Mesophyll. Palisade tissue consisting of 1-4 layers in different genera and
species. Spongy tissue usually with large intercellular spaces; very dense in
Weihea zeylanica Baill.; collenchymatous in Bruguiera gymnorhiza Lam.;
sometimes containing aqueous cells in Bruguiera and Kandelia. H-shaped
sclerenchymatous idioblasts (Fig. 133) occur in the palisade tissue, and
variously branched ones in the spongy mesophyll of Rhizophora. Solitary
sclerenchymatous cells also recorded by Engler (633) in the spongy mesophyll
of Poga oleosa. Vascular bundles of the veins mostly embedded in the
mesophyll; almost vertically transcurrent in a species of Macarisia; rarely
accompanied by well- developed sclerenchyma except in a species of Gyno-
troches; sclerenchyma above and below the smaller veins recorded by Sprague
and Boodle (2173) in Anapyxis surrounded by starch sheaths in Ceriops
;

candolleana Arn. according to Mullan (1570). Enlarged terminal tracheids

reported in Bruguiera, Ceriops, Kandelia. Petiole, in transverse sections
through the distal end, exhibiting: a crescent-shaped strand in Anapyxis
according to Sprague and Boodle (2173); structure similar in Bruguiera
gymnorhiza Lam. (Fig. 131 i); exhibiting a ring of bundles surrounding
additional medullary strands in Rhizophora mucronata Lam. similar but with a
;

single medullary bundle in Ceriops candolleana according to Mullan (1570).

Crystals mostly clustered in Bruguiera, Carallia (in the epidermis), Ceriops,
Gynotroches, Kandelia, Rhizophora', mostly solitary in Anapyxis (occasional in
the epidermis), Blepharistemma, Cassipourea, Macarisia, Weihea (particularly
in the epidermis). Secretory elements. Cells containing tannin recorded
in the mesophyll of Anapyxis, Ceriops (cells elongated), Rhizophora, and
probably present in other genera and species as well. Secretory canals with
brownish contents occur in the spongy mesophyll of Poga oleosa according
604 RHIZOPHORA CEAE
to Engler (633). Large globules of oil said to be present in the cells surround-
ing the stomata and in the palisade cells of Bruguiera.

Axis
YOUNG STEM and 133 B)
(Fig. 131 j
Epidermis of Bruguiera caryophylloides Blume composed of variously
shaped cells appearing conical in transverse sections; including vertically
divided cells in Ceriops. Stem of Bruguiera caryophylloides grooved when
sufficiently young. Cork arising superficially, usually in the hypodermis, in

FIG. 133- RHIZOPHORACEAE

A, Transverse section of the leaf of 'Rhizophora conjugate L.'. B, H-shaped idioblasts from the
cortex of the branch of Rhizophora mangle L. A, by Solereder, B, after Warming.

species of Anapyxis, Bruguiera, Carallia, Ceriops; Rhizophora spongy in Ceriops

;

and Rhizophora mangle Linn.; component cells provided with thickened

inner walls in Anapyxis. Primary cortex lacunar in species of Bruguiera,
Ceriops, and Rhizophora', outer part containing strongly lignified stone
cells in Bruguiera caryophylloides', including H-shaped sclerenchymatous
idioblasts in Rhizophora mangle (present in the pith as well in this genus) ;

occasional sclerotic cells recorded by Sprague and Boodle (2173) in Anapyxis.

Pericycle with a sub-continuous, composite ring of sclerenchyma in
Anapyxis', described as including very small fibres in Rhizophora mucronata
Lam. and isolated strands of fibres in Bruguiera caryophylloides and Ceriops
candolleana Arn. according to Mullan (1570). No pericyclic fibres observed
in young stems of Bruguiera gymnorhiza Lam. (Fig. 131 j). Xylem
traversed by rays 2-3 cells wide in Rhizophora mucronata and by rays
5-6 cells wide in Ceriops candolleana. Vessels with scalariform perforation
plates. Crystals of the same types as those in the leaf; crystalliferous cells
recorded in the pericycle of Ceriops. Secretory elements. Vertically
elongated secretory cells containing tannin and/or oil present in the cortex
and pith of Ceriops; similar elements probably occur in other genera as well.
 RHIZOPHORACEAE 605
BARK
Secretory cavities, originating from groups of cells which become
mucilaginous, recorded by Engler (633) in Poga oleosa Pierre. For anatomy
of the tanniniferous barks of Rhizophora, &c., see 'Economic Uses* on p. 611.

WOOD
The genera have been divided into 4 groups, as proposed by Marco (1439).
GROUP I. Bruguiera, Ceriops, Kandelia, and Rhizophora (Fig. I34A-C)
Vessels moderately small (50-100 mean tangential diameter), solitary
/x,

and in radial multiples often of more than 4 cells and up to 14, and in clusters;
mostly 5-35 per sq. mm.(Panshin (1649) gives 25-70 for Ceriops). Perfora-
tion plates scalariform, steeply inclined, and with rather thick and few bars.
Intervascular pitting scalariform, the pits narrow, crowded, and extending
completely across the member wall. Pits to ray cells differing from those to
the wood parenchyma; the former typically small and round to oval, 2 to
several such pits fitting a single large, vertical or obliquely elongated pit in
the ray cell wall; the ray pit very broad and shield-shaped in Ceriops and
often subtending more than i row of vertical vessel pits pit-pairs between
;

vessels and wood parenchyma typically oval or elongated in both walls, the
longer axes of the pits horizontal. Tyloses often present, sometimes abundant.
Mean member length 0-6-1*0 mm. Parenchyma paratracheal, scanty except
in Kandelia j and often consisting of only a few cells touching the vessels
(Fig. 1
34 A); more abundant in Kandelia, forming complete sheaths round
the vessels and in bands 2-4 cells wide linking up the vessels (Fig. 1340).
Strands of 4-8 cells. Rays mostly up to 3-6 cells wide, sometimes up to
10 cells in Bruguiera and Rhizophora (1439); seldom more than 3 mm. high
in Ceriops and Kandelia, often more than 3 mm. in Bruguiera and Rhizophora\
uniseriate rays typically scarce to almost absent, except in Ceriops in which
they are moderately abundant and composed of both upright and procumbent
cells; 6-10 rays per mm.; heterogeneous (Kribs's Type HA?) except in
Rhizophora, with 3 or 4 marginal rows of upright cells and up to 10 rows in
Ceriops, the inner 'procumbent' cells short radially and almost square; uni-
seriate rays similar. In Rhizophora the rays are homogeneous (Kribs's
Type II) and composed mainly of definitely procumbent cells and in Bruguiera
almost homogeneous. Cells commonly containing dark gummy contents and
single crystals containing silica in Kandelia candel Druce (794). Fibres with
;

small simple pits and thick walls, the walls often gelatinous (1439). Some
septate fibres observed in a specimen of Kandelia Rheedii W. et A. Mean
length i-i~i*7 mm.

GROUP II. Anisophyllea, Carallia, Combretocarpus, Crossostylis, and Gyno-

troches (Fig. 134 D-F)
Vessels medium-sized (100-200 p mean tangential diameter) in Crosso-
stylis and Gynotroches, large (more than 200 ft) in Anisophyllea, Carallia, and
Combretocarpus', solitary and in small multiples and clusters; 1-2 per sq. mm.
in the genera with large vessels, 6-20 per sq. mm. in Crossostylis and Gymno-
troches. Perforation plates simple; Marco (1439) notes some scalariform
plates in the first-formed secondary xylem of Gynotroches. Intervascular
6o6 RHIZOPHORACEAE
pitting scalariform in Gynotroches, alternate
and moderately small in the other
genera; pits to ray and wood parenchyma commonly elongated in various
directions and also round and unilaterally compound; pits in Anisophyllea
small and round only. Tyloses common, e.g. in Anisophyllea and Carallia\

FIG. 134, RHIZOPHORACEAE

A, Rhizophora conjugata Linn. B, Bruguiera parviflora Wight et Am. C, Kandelia rheedii Wight et
Am. D, Anisophyllea laurina R, Br. E, A. laurina R. -fir. F, Gynotroches axillaris Blume.

sometimes with gummy deposits, e.g. Carallia, and, according to Marco

(1439), often with chalky infiltrations completely occluding the lumina.
Mean member length 0*7-1 *o mm. Parenchyma typically abundant and
predominantly banded the bands varying from moderately regular apotracheal
;

forms in Anisophyllea (Fig. 1340) and Combretocarpus p.p. to irregular and

almost confluent (paratracheal) forms in some species of Combretocarpus and
in Camilla, in which the bands tend to form arcs between the large rays but
connect all the vessels and sometimes form wings from them; the bands vary
 RHIZOPHORACEAE 607
from several cells wide in Carattia, Combretocarpus, and Anisophyllea to i or 2
cells in Gynotroches (Fig. 134 F). Some
paratracheal parenchyma always
present and some isolated cells usually scattered among the fibres. Cells
usually with dark gummy contents, crystals present in the ordinary cells in
Camilla and in chambered cells on the edges of the bands and scattered
among the fibres in Carallia and Combretocarpus; silica present in Combreto-
1

carpus motleyi Hook. f. (794). Strands of 4-16, mostly 8 cells. Rays of 2

distinct sizes, the larger up to 10-15 ce U s wide; very high, commonly exceed-
ing 5 mm.; often showing evidence of dissection into smaller units; uniseriate
rays numerous and composed of upright cells; 4-10 rays per mm.; hetero-
geneous (Kribs's Types I and II A), with 2-10 marginal rows of high upright
cells and some sheath cells. Cells commonly with dark gummy contents and
with solitary crystals in Carallia. Fibres with small, flat, and sometimes
rather indistinct borders, e.g. in Anisophyllea ;
walls very thick and often
gelatinous. Mean length 1-8-2-6 mm.

GROUP III. Anopyxis, Blepharistemma, Cassipourea, Macarisia, Sterigma-

petalum (Fig. 135 A, B, and E)
Vessels mostly small to medium-sized, 100-200 p mean tangential diameter,
approaching 200 p, mean tangential diameter in Sterigmapetalum and some-
times exceeding it in Anopyxis\ exclusively solitary in Anopyxis, Cassipourea,
and Sterigmapetalum mostly 20-40 per sq. mm., sometimes more numerous
;

in Cassipourea, rather fewer in Sterigmapetalum, and only about 4 per sq. mm.
in some species of Anopyxis. Perforation plates typically both simple and
scalariform, the latter with delicate bars, the simple perforations often much
longer than wide and with parallel sides (1439). Intervascular pitting difficult
to observe in genera with solitary vessels, according to Marco (1439) alternate
to opposite and tending to scalariform in some genera; pits to ray cells and
wood parenchyma typically large and elongated, varying from mostly both
long and wide, e.g. in Anopyxis, to mostly round or slightly oval, as in some
species of Cassipourea, sometimes unilaterally compound but mostly approxi-
mately the same shape and size as the subtending pits in the parenchyma
walls; tending to be elongated horizontally in contact with wood parenchyma
cells and irregularly in contact with ray cells. Sclerotic tyloses and amorphous

deposits of gum present in some specimens of Cassipourea elliptica (1439) and

thin-walled tyloses present in some other species of Cassipourea. Mean
member length 1-0-1-4 mm. Parenchyma moderately abundant, typically
predominantly paratracheal and absent from the adaxial sides of the vessels ;
varying from aliform, e.g. in Anopyxis, to aliform and confluent, with some
diffuse, e.g. in some species of Cassipourea, or predominantly in uniseriate
apotracheal bands, e.g. in Blepharistemma. With chambered crystal cells in
some species of Anopyxis. Strands of 8-16 cells. Rays 2-5, mostly 3-4 cells
wide, widest in Anopyxis and some species of Cassipourea', typically less than
i mm.
high, but rather higher in most species of Cassipourea; uniseriate rays
numerous and composed of moderately to very high upright cells; mostly
5-12 rays per mm., sometimes slightly more numerous in Blepharistemma
and Cassipourea] heterogeneous (Kribs's Types I and II A); with 4 or more
marginal rows of upright cells, except in Macarisia, and with 10 or more rows
1
Described by Marco (1439) as crystailiferous fibres.
6o8 RHIZOPHORACEAE
in Anopyxis and Cassipourea p.p. ; cells sometimes containing crystals. Fibres

typically with large, distinctly bordered pits, tending to be more numerous on

the radial walls; borders less distinct in Blepharistemma and Macarisia\ walls
thick and often gelatinous; mean length i -6-2-2 mm.

V
D E
FIG, 135. RHIZOPHORACEAE
A, Cassipourea podantha Standley. B, C. podantha Standley. C, Pellacalyx saccardianus Scortech.
D, Poga oleosa Pierre. E, Anopyxis ealaensis Sprague.

GROUP IV. Pellacalyx and Poga (Fig. 135 c and D)

These 2 genera do not fit
satisfactorily into any of the other groups ;
their
chief characteristics are given below.
Pellacalyx. Vessels medium-sized, arranged in tangential groups and rows,
perforation plates simple and very occasionally delicately reticulate (1439),
intervascular pitting large and opposite, pits to parenchyma similar or
irregularly elongated, particularly to the ray cells silica reported by Gonggrijp
;

(794). Parenchyma enclosing the vessels and extending from them to form
bands between the large rays, in narrower apotracheal bands and sometimes
 RHIZOPHORA CEAE 609
scattered among the fibres. Rays up to 25 cells wideand very high; uni-
seriatesnumerous, composed of upright cells; heterogeneous. Fibres with
bordered pits.
Poga. Vessels large (more than 200 /A) and almost exclusively solitary,
fewer than i per sq. mm., perforations simple and nearly horizontal, inter-
vascular pitting alternate, tending locally to conspicuous coalescent apertures,
pits to parenchyma nearly always round but occasionally large, elongated and
almost simple, sometimes unilaterally compound mean member length about
;

8 mm. Parenchyma about the vessels, extending laterally from them in

narrow wings and in apotracheal bands between the large rays. Rays of 2
distinct sizes, the larger up to 20 cells wide and very high, uniseriate rays
numerous and low, heterogeneous. Fibres with a few distinctly bordered
pitswhich are almost entirely limited to the radial walls, mean length about
2*0 mm.
ROOT
The well-known 'breathing' roots ofmembers of the Rhizophoraceae
inhabiting mangrove swamps have been the object of numerous investigations.
The large intercellular spaces of the spongy cortex of these roots have for
long been thought to facilitate gaseous exchange in the special habitat in
which the plants grow. Troll (2282) has questioned this view, for he believes
that the form of the root system in Sonneratia (Family Lythraceae) and
Bruguiera is chiefly important because it enables a sequence of absorptive
roots to be produced in a substratum of which the level is liable to be raised
by the deposition of silt. Even Troll, however, subscribes to the previously
accepted view to the extent of pointing out that the spongy structure of the
cortical tissues may be of secondary importance in facilitating gaseous

exchange. The root structure for each genus is described separately below.
(i) BRUGUIERA
Cortex thick, containing abundant, large, intercellular spaces, arranged
radiately around the stele. According to Solereder, mechanical support is
provided for this otherwise thin-walled tissue by annular, lignified thickening
ridges to the cells which border on the intercellular spaces. In the light of
Bowman's (252) observations on the similar structures which occur in RhizQ-
phora (see next paragraph) it seems possible that they may represent cells
filled with mucilaginous sap. Intercellular spaces are larger in the sub-
terranean than in the aerial parts of the roots. Sclerenchymatous idioblasts
occur in the cortical parenchyma. Cork consisting wholly of suberized cells
below the ground, but of alternating layers of suberized and ordinary paren-
chymatous cells in the aerial portion.

(ii) RHIZOPHORA

Similar to Bruguiera but the primary cortex in the terrestrial part of the
root is composed of cells of 2 kinds (a) radially elongated cells connected
: .

with one another tangentially by short lateral arms; (b) rows of vertically
elongated cells which exhibit small, circular lumina in transverse sections.
Solereder, in common with other early authors, records the existence of
ridges of thickening which are said to give mechanical support to the radially
elongated cells in which they are alleged to occur, but according to the more
4594 R r
6io RHIZOPHORACEAE
recent work of Bowman (252) these 'Verdickungsleisten* are really cells filled
with mucilaginous sap. Mullan (1570) states that the cortex in the aerial part
of the root system of R. mucronata Lam. is much reduced and the lacunae
small, whilst the component cells are all alike and roundish to polygonal in
outline. Bowman (I.e.) also says that the cortex in the subterranean clusters
of roots, which lie below the mud but are attached to the ends of the stilt roots,
is composed of round cells interspersed with large intercellular spaces. Some
of these cells are arranged in short strands, but others radiate from a central
cell, the latter often being filled with starch and those of the radii with

mucilage. H-shaped, sclerenchymatous idioblasts project into the cortical

but are less numerous below
intercellular spaces in the aerial part of the root,
ground. Mullan records the existence of vertically elongated tubular
(I.e.)
cells, filled with tannin and
oil, situated at the junctions between the branched
cells in the terrestrial part of the root of R. mucronata.

(Hi) CERIOPS
Mullan (1570) describes the general structure of the root system of Ceriops
candolleana Arn. as similar to that of Rhizophora mucronata, except that, in
Ceriops, multiradiate idioblasts are absent from the cortex.

(iv) CARALLIA
Buscalioni (317) has described the morphology and anatomy of a fasciated
aerial root of Carallia integerrima DC.
For further details concerning the root system of mangroves belonging to
the Rhizophoraceae see Liebau (1368) and Emould (627).

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
Examination of the exomorphic and anatomical features of Anopyxis
ealaensis Sprague by Sprague and Boodle (2173) led to the conclusions that
this species has many features in common with the Rhizophoraceae-Legno-
tideae, the agreement with Macarisia pyramidata Thouars being particularly
close.

(ii) FROM WOOD STRUCTURE

According to Marco (1439), the wood structure does not fully conform with
any of the 3 taxonomic systems proposed by Bentham and Hooker, Ridley,
and Schimper. The tidal genera, Bruguiera, Ceriops, Kandelia> and Rhizo-
pkora, form a well-defined, homogeneous natural group, which is readily
separable from other members of this or any other family. The Legnotideae
(the inland genera), as proposed by Bentham and Hooker, includes woods of
very different types, which Marco has separated into 2 groups, which he
terms the Macarisieae and the Gynotrocheae. These groups, which correspond
to Groups II and III in the above description of the wood, he considers as
being as well-defined and natural aggregations as the Rhizophoreae. The
material examined by the author fully supports his treatment of the Rhizo-
phoreae, but provides less convincing evidence of the clear distinction between
the Gynotrocheae and Macarisieae.
Marco expresses some doubts as to the placing of Anopyxis and Sterigma-
 RHIZOPHORACEAE 6it

petalum in the family and only places them provisionally in the Macarisieae.
They appear to fit reasonably well in this group. He excludes Poga and
Pellacalyx and notes a superficial resemblance between the latter and
Embothrium of the Proteaceae. Both these genera, however, have many
characteristics in common with other genera in this family; indeed, nearly
all the details of their structure can be matched in some
species in the family,
though it is true that neither genus fits very well into any of the proposed
groups.
Panshin (1649), in a study of the woods of the Philippine mangrove
swamps, found no evidence of the anatomical structure of the wood being
influenced by the habitat.

ECONOMIC USES
Apart from the timbers derived from the family, the tanniniferous bark of
Bruguiera, Ceriops, Kandelia, and Rhizophora is one of the most important
economic products. This material is imported into European countries for
commercial use, although the tanning qualities are said to be inferior to those
of many other materials. The microscopy and chemical composition of man-
grove barks have been exhaustively described by Wenzel (2412), who gives an
extensive bibliography of other work on the same subject. According to
Bodenstab (211) commercial samples of Rhizophora bark consist of hard,
heavy pieces, 3-30 mm. thick, with little or no cork. The colour is red to
dark brown throughout, and the taste and scent weakly aromatic. Fracture
brittle and horny. Cork, mostly confined to young material, up to 2 mm.

thick, composed of thin- walled cells with brownish contents. A layer of

parenchyma 1-5 mm. thick beneath the cork includes fairly numerous stone
cells, the latter being solitary and in groups and provided with branched pits
in the concentrically zoned thickening of the cell walls. Groups of stone cells
towards the inside of the bark more elongated than those towards the exterior,
appearing to the naked eye as lines or bars. Cells containing solitary crystals
occur at the periphery of the groups of stone cells. Ground tissue consisting
of medullary rays, 5-6 cells wide, which contain abundant cluster crystals, and
of phloem strands which include occasional short thick fibres. All parenchy-
matous tissues contain tannin and a brownish-red colouring matter.
The family is not important as a source of timber, but the 'tidal* genera are
often in demand as poles and for making charcoal. Carallia lucida Roxb. is
used as a general construction timber in India and to some extent for
ornamental purposes, making use of its oak-like silver grain.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Anisophyllea, Anopyxis, Blepharistemma, Bruguiera,* Carallia, Cassi-
pourea, Ceriops, Crossostylis, Dactylopetalum, Gynotroches, Kandelia,
Macarisia, Poga, Rhizophora, Weihea.
* Kew slide
Represented in the collection .

(ii) FOR WOOD STRUCTURE

Anisophyllea, Anopyxis, Blepharistemma, Bruguiera, Carallia, Cassipourea,
612 RHIZOPHORACEAE
Ceriops, Combretocarpus, Crossostylis, Gynotroches, Kandelia, Macarisia,
Pellacalyx, Poga, Rhizophora, Sterigmapetalum.

LITERATURE
(i) On General Anatomy
Arzt 45, Bodenstab 211, Bowman 251, 252, Buscalioni 317, Emould 627, Engler 633,
Liebau 1368, Mullan 1570, Sprague and Boodle 2173, Troll 2282, Wenzel 2412.

(ii) On Wood Structure

et al. 164, Benoist 170, den Berger 179, 182, Besson 1 86, Burgerstein 310,
Becking
Cooper and Record 461, Dadswell and Record 533, Foxworthy 705, Gonggrijp 794,
Hopkinson 1083, Howard 1088, Janssomus 1154, Jentsch 1173, Kanehira 1204, 1206,
Kribs 1283, Lecomte 1334, Marco 1439, Memaud 1492, Panshin 1649, Pearson and
Brown 1679, Pfeiffer, J. Ph. 1713, Record 1783, 1851, Record and Hess 1886, Record
and Mell 1894, Riera 1937, Stone 2202, Tupper 2295.

135. COMBRETACEAE
(FiG. 131 on p. 598; FIG. 136 on p. 614; FIG. 137 on p. 616)

SUMMARY
(i) GENERAL
A mainly tropical family of trees, shrubs, and woody climbers. The leaf is
dorsiventral, or more
rarely centric, and provided with glandular and non-
glandular hairs of various kinds. Of these the simple unicellular and uni-
cellular 2-armed types are the most widespread. The leaf stomata are usually
confined to the lower surface, but occur on the upper side also in a few
species; they are ranunculaceous. The mesophyll, in some genera, includes
ramifying sclerenchymatous fibres which are generally connected with the
vascular bundles of the veins. Large stellate idioblasts containing cluster
crystalshave also been recorded in the mesophyll. These idioblasts are some-
times apparent as transparent dots if the leaf is not too thick. Cluster crystals
also occur in the parenchymatous tissues in other parts of the plant. The
vascular bundles of the veins are nearly always bicollateral, and accompanied
by a varying amount of sclerenchyma in different genera and species. Canal-
like secretory receptacles are present in association with the xylem of the
veins in a number of species of Terminalia. Cork in the young stem arises
superficially or in the pericycle in different genera and species. The secondary
phloem frequently characterized by islands of sclerenchyma, the latter
is

including chambered fibres filled with crystals. Both interxylary and intra-
xylary phloem are common in the family. Negatively geotropic roots, with
abundant intercellular spaces in the cortex, have been recorded in Laguncu-
laria and Lumnitzera.

(ii) WOOD
Vessels mostly medium-sized, either exclusively solitary or with numerous
multiples of 4 or more cells, perforations simple, intervascular pitting alternate,
pits to parenchyma similar to the intervascular pitting; vestured; members
of medium length. Parenchyma typically aliform to confluent with some
scattered cells, occasionally vasicentric and in terminal bands idioblasts con- ;

taining druses present in some genera. Rays most typically exclusively

 COMBRETACEAE 613
uniseriate or rarely biseriate, but up to 2-5 cells wide in several species,
particularly of Terminalia\ homogeneous or slightly heterogeneous; crystals,
either single or druses, commonly present. Fibres with small simple pits,
some fibres usually septate, of medium length. Vasicentric tracheids
present in i genus. Traumatic vertical intercellular canals occasionally

present. Included phloem of the 'foraminate* type present in 4 genera.

LEAF
Usually dorsiventral; rarely centric; middle of the mesophyll composed of
aqueous tissue surrounded by palisade in species of Laguncularia and
Lumnitzera.

Hairs
(a) Non-glandular, (i) Simple, unicellular, often somewhat bulbous at the
base and with a cellulose membrane more or less conically or convexly
arched outwards towards the filamentous distal end sometimes appear-
;

ing to be bicellular owing to this peculiar construction (Fig. 136 A-B).

This type is widely distributed, (ii) Unicellular, 2-armed in Cono-
*
carpus (Fig. 136 c); sometimes with an imperfectly developed second
arm in Ramatuela and Terminalia (Fig. 136 D).
(b) Glandular
I. Sessile or short-stalked, (i) Spherical, bladder-like, the material
secreted between the cuticle and the surface of the component cells
obscuring the internal structure of the gland in Calycopteris (Fig.
136 E), Combretum (pro parte), Guiera. (ii) Small glandular scales,
with little or no accumulation of secreted material in Combretum
(pro parte) (Fig. 136 F) and Thiloa.
II. With a more or less elongated stalk, (i) With a spherical or
ellipsoidal head in Cacoucia, Combretum (species at one time included
in the genus Poivrea), Quisqualis.

(c) Almost cap-shaped glands, situated at the bases of flask-shaped

sessile,
cavities, recorded on both leaf surfaces in Languncularia racemosa
Gaertn. (Fig. 136 H-I); and somewhat similar ones on the lower side
of the leaf of Conocarpus erectus Linn. Large, conspicuous glands also
occur on the petiole in^nogeissus, Conocarpus, Laguncularia, Terminalia.
Reddish glandular scales situated in small pits recorded by Hate (919)
on the lower surface of the leaf in Calycopteris floribunda Lamb. The
anatomy and development of the petiolar thorns of Quisqualis indica
Linn, have been described by Helm (946).

Epidermis papillose on the lower surface of 2 species of Combretum with

;

marginal pits to the cell walls in a number of species of Combretum and in

Ramatuela. Hypoderm said to occur locally below the upper epidermis in
Combretum ternatum Wall. Stomata ranunculaceous usually confined to the
;

lower surface, but present on the upper side as well in a few species of Com-
bretum, Conocarpus, Guiera, Laguncularia, Lumnitzera, Macropteranthes.
Hydathodes recorded in Laguncularia and Lumnitzera. Sclerenchymatous
fibres, branching off from the veins and extending irregularly throughout the
614 COMBRETACEAE
mesophyll, present in species of Anogeissus, Buchenavia, Bucida, Com-
bretum, Ramatuella, Thiloa. Vascular bundles of the larger veins bicollateral ;

enclosed by a ring or accompanied by an arc of sclerenchyma in species of

Anogeissus, Cacoucia, Combretum, Conocarpus, Guiera, Macropteranthes,
Quisqualis, Ramatuela, Terminalia, Thiloa\ accompanied by only a few fibres
in species of Calycopteris, Combretum, Macropteranthes, Terminalia', scleren-

FIG. 136. COMBRETACEAE

A, B, Ordinary combretaceou8 hairs. C, Two-armed hair of Conocarpus. D, One-armed hair of
Terminalia argentea Mart. E, Glandular hair of Calycopteris in section. F, External gland of Com-
bretum aculeatum Vent. G, External gland of Combretum decandrum t Roxb. H, I, Glandular leaf-pit
of Laguncularia racemose Gaertn. f, A, G, after Heiden; H, I, after Holtermann.

chyma absent from the veins in certain species of Combretum, Lumnitzera,

Terminalia. Vascular bundles of the smaller veins embedded in the mesophyll
in species of Anogeissus, Combretum, Conocarpus, Laguncularia, Lumnitzera,
Ramatuela, Terminalia, Thiloa, or vertically transcurrent by sclerenchyma
or thin- walled tissue in different species; accompanied by a large or small
amount of sclerenchyma in species of Anogeissus, Cacoucia, Combretum,
Guiera, Macropteranthes, Quisqualis, Ramatuela, Terminalia, Thiloa', scleren-
chyma absent from species of Anogeissus, Calycopteris, Combretum, Lagun-
cularia, Lumnitzera, Macropteranthes, Terminalia.
 COMBRETACEAE 615
Petiole. Transverse sections through the distal end, in the few species of
Calycopteris (Fig. 1310) and Combretum represented in the Kew slide collec-
tion, exhibiting a solitary crescent-shaped vascular strand, with outwardly
directed or incurved ends. Main vascular strand closed and dorsally flattened
in Terminalia chebula Retz. (Fig. 131 A). Canal-like secretory cavities,
provided with a definite epithelium and filled with mucilaginous contents,
situated above the xylem of the veins and extending into the mesophyll of a
number of species of Terminalia. Similar cavities also seen in the 'cortical 1

region of the petiole of Terminalia chebula. Secretory cells, with unidenti-

fied amorphous contents, observed in unlignified tissue in species of Caly-

copteris, Combretum, and Terminalia, and probably present in other genera

and species as well. Fatty bodies often present in the mesophyll. Crystals
almost exclusively clustered; occurring in the mesophyll and beside the
vascular bundles of the veins; numerous, large, solitary types noted in the
'cortical* region of the petiole in Terminalia chebula\ sometimes situated in

large, somewhat star-shaped idioblasts, the latter being visible as transparent

dots if the leaf is not too thick, in species of Anogeissus, Combretum, Cono-
carpus, Guiera, Laguncularia, Macropteranthes, Quisqualis, Terminalia, Thiloa.

Axis
YOUNG STEM (Fig. 131 B and D)
Cork arising superficially in certain species of Anogeissus, Calycopteris,
Conocarpus, Laguncularia, Lumnitzera, Macropteranthes, and Terminalia',
originating in the pericycle in other species of Anogeissus, Calycopteris,
Conocarpus, and Terminalia as well as in Cacoucia, Combretum, Guiera,
Quisqualis, Ramatuela, Thiloa', consisting mostly of cells with wide lumina
and thin walls, but sometimes including stone cells or cells thickened only on
one side. Cork cells radially elongated in Quisqualis. Phelloid cells stated to
be included in the cork of the same genus. Pericycle usually containing
isolated strands of fibres situated around a broad inner parenchymatous
portion. Secondary phloem frequently characterized by islands of scleren-
chymatous elements; including chambered fibres filled with small clustered
crystals and arranged in tangential bands as seen in transverse sections of
species of Anogeissus, Combretum, Conocarpus, Laguncularia, Lumnitzera,
Quisqualis, Terminalia, Thiloa. Phloem and xylem in most genera forming
a continuous cylinder traversed by narrow rays. Vessels with simple perfora-
tions. Strands of interxylary phloem, sometimes arranged in concentric
circles and composed of numerous clustered
soft tissue, often containing

crystals, recorded in species of Calycopteris, Combretum (particularly in

African species), Guiera, Thiloa. For further details see under Wood* on
p. 618. Intraxylary phloem, either in the form of a ring or in separate
strands at the periphery of the pith, recorded in species of Anogeissus,
Cacoucia, Calycopteris, Combretum, Guiera, Laguncularia (inconspicuous),
Lumnitzera (inconspicuous), Pteleopsis, Quisqualis, Ramatuela, Terminalia,
Thiloa. Mucilage cavities recorded in the intraxylary phloem of various
species of Terminalia, and in the interxylary phloem in species of Combretum.
Pith usually fairly small, mostly consisting of lignified, fairly thick- walled, or,
more rarely, of thin- walled cells containing groups of stone cells in Caly~
;

copteris and Terminalia sp. and vertically elongated elements filled with an
 FIG. 137. COMBRETACEAE
A, Guiera senegalensis Larn. B, Lumnitzera cocdnea Wight et Arn. C, Terminalia parviflora Presl.
D, T, ivorenris A. Chev. E, Combretum hartmannianum Schweinf.
 COMBRETACEAE 617
unidentified white substance in several species of Combretum. Secretory
elements. Mucilage canals recorded in the pith in a number of species of
Terminally cavities observed in the cortex and phloem in species of Caly-
copterisand Terminalia (see also under 'Intraxylary Phloem'). Secretory cells
with unidentified, amorphous contents seen in certain species of Combretum.
Crystals mostly clustered; solitary types less frequent; very small raphides
noted in the phloem of Calycopteris (see also under Thloem' and 'Interxylary
Phloem' above).

WOOD (Fig. 137)

Vessels mostly medium-sized (mean tang. diam. 100-200 /*), moderately
small (50-100 /t) in Anogeissus leiocarpus Guill. et Perr. and Lumnitzera, large
(about 280 fi) in Strephonema\ exclusively solitary in Combretum, Guiera,
Ramatuela (931 A) and Strephonema, with radial multiples of 4 to several
cells in Anogeissus and Lumnitzera (Fig. 1376); the beginning of the ring
marked by a zone without vessels in some species of Terminalia', mostly
5-20 per sq. mm., fewer than 5 per sq. mm. in Combretum p.p., Strephonema
and most species of Terminalia, about 20 per sq. mm. or slightly more in
some species of Anogeissus, Conocarpus, and Lumnitzera according to Record ;

(1851) ring-porous in some species of Combretum. Perforations simple.

Intervascular pitting alternate, medium-sized or occasionally rather small,
e.g. in Lumnitzera and Pteleopsis, sometimes with striations due to coalescent
apertures, e.g. in Buchanavia\ pits to parenchyma similar to the intervascular
pitting, pits to ray cells sometimes arranged in distinct horizontal rows, e.g. in
Combretum, Pits vestured (78). Tyloses abundant in Terminalia, deposits
of gum common and sometimes abundant, white chalky deposits noted by
Panshin (1649) in Lumnitzera. Mean member length 0-3-0-5 mm. Paren-
chyma predominantly paratracheal, typically abundant, aliform to confluent
and with some isolated cells scattered among the fibres (Fig. 137 D and E);
the paratracheal parenchyma varying from a few cells round the vessels, e.g.
in most species of Lumnitzera (Fig. 137 B), to broad, irregular confluent
bands, these bands more regular and possibly apotracheal in some species of
Terminalia, e.g. T. bialata Steudel with both aliform parenchyma and isolated
;

patches suggesting vestiges of continuous broad bands in Strephonema narrow ;

terminal bands sometimes present in Buchanavia, Bucida, and Terminalia.

1

Without crystals in most genera but chambered crystalliferous cells common

in Pteleopsis, large single crystals present in chambered or ordinary cells in
some species of Terminalia, and with large idioblasts containing druses in some
species of Combretum, Guiera, and Terminalia (e.g. T. catappa L.); small
druses also .present in the parenchyma surrounding the strands of included
phloem in Guiera commonly with gummy contents. Strands commonly of
\

6-8 cells. Solereder refers to the presence of unlignified parenchyma in

the twining plant Quisqualis indica Linn. Rays exclusively uniseriate or
with occasional biseriate portions in Anogeissus (most species), Buchenavia,
Combretum p.p., Conocarpus, Laguncularia, Lumnitzera, and Terminalia p.p.
(e.g.
T. arjuna Bedd., T. belerica Roxb., T, bialata Steudel, T. curtisii
1
Chowdhury (412-14) has shown that in Terminalia tomentosa W. et A. these bands are
the tissue
first formed in the growth ring and has suggested the term 'initial' in place of
'terminal'.
618 COMBRETACEAE
Ridl., T. exceha Liebm., T. mannii King (1679), T. myriocarpa Heurck
et Muell. (1679), T. obovata (R. et P.) Steud., T. oliveri Rands (1679), T.

paniculata Roth., T, pyrifolia Kurz., T. subspathulata King, T. superba Engl.

et Diels, T. teysmannii Koord. et Val. (i 154), and T. tomentosa Wight et Arn.),
commonly 2 wide in Combretum (a few species), Guiera, Pteleopsis, and
cells

Strephonema, 2-4 cells wide in Budda and many species of Terminally up to

5 cells wide in Quisqualis (2158); less than i mm. high; uniseriates moderately
numerous in woods with multiseriate rays and composed of procumbent cells ;

mostly 8-12 rays per mm., more numerous (12-18) in Anogeissus and some
species of Combretum, Guiera, Laguncularia, and Lumnitzera (1649), about
4-6 per mm. in a few species of Terminalia, e.g. T. ivorensis A. Chev. ;

homogeneous (Kribs's Types I and III) in Buchenavia, Bucida (apart from

crystalliferous cells), Combretum (a few species), Pteleopsis, and most species
of Terminalia; heterogeneous in the remainder but not markedly so, except in
Strephonema, with 4 or more marginal rows of square or upright cells and
uniseriate rays composed entirely of upright cells, and, according to Record
and Hess (1886), in Ramatuela; rays composed entirely of square or upright
cells in Terminalia p.p. Crystals common in Anogeissus, Bucida, Combretum

p.p., and Terminalia p.p. (particularly in species with uniseriate rays),

mostly as single crystals in square cells interspersed in rows between the
procumbent cells (Fig. 137 c), the crystalliferous cells often conspicuous in
tangential sections (particularly in Bucida) owing to their larger size or rounded
shape (Fig. 137 A); large idioblasts containing druses present in Guiera\
gummy deposits often present, conspicuous in Lumnitzera. Fibres typically
with small, often very inconspicuous, simple pits, more numerous on the
radial than on the tangential walls; pits with small borders in Lumnitzera and,
according to Holterman as quoted by Solereder, in Combretum decandrum
Roxb. A proportion of the fibres septate in some species of Anogeissus,
Combretum, Conocarpus, Guiera (2158), Pteleopsis, Strephonema, and Termi-
nalia, the septate fibres sometimes more numerous in the neighbourhood of
the parenchyma and, according to Solereder, sometimes containing crystals
in Combretum and Terminalia. Walls moderately to very thick. Mean length
0-8-1-7 mm. Vasicentric tracheids present in Strephonema. Intercellular
canals of the vertical traumatic type present in some specimens of Anogeissus,
Buchenavia, Bucida, and Terminalia. Growth rings. The seasonal develop-
ment of the wood of Terminalia has been investigated by Coster (481) and
Chowdhury (412-14). Chowdhury has shown that in T. tomentosa the
so-called 'terminal* band of parenchyma is in fact the first-formed tissue of
the growing season. Heiden (2158) refers to secretory spaces, with yellow
contents that give a mucilage reaction, in thick parts of branches of Terminalia
bellerica Roxb. and T. macroptera Mart. Included (interxylary) phloem
of the 'foraminate' type (c. I. foraminulatum) present in some species of
Calycopteris (2158), Combretum (Fig. 137 E), Guiera, and Thiloa (2158);
1

the strands of included phloem usually isolated, though tending to occur in

tangential rings, particularly in Thiloa (1679), and occasionally linked by
1
Solereder (2158) notes that included phloem occurs only in the African species of Com-
bretum and lists the following: C. erythrophyllum Sond., C. hartmannianum Schweinf., C.
kraussii Hochst., C. lepidotum Rich., C. nyikae Engl., C. rueppellianum Rich., C. salicifolium
E. Mey., C. schelei Engl., C. tenuispicatum Engl., C. tricacanthum Fres. To these may be
added C. multispicatum Engl. et Diels and C. guemzii Sond.
 COMBRETACEAE 619
parenchyma; radial strands occur occasionally in the rays in Combretum. The
strands contain no bast fibres; the parenchyma often contains numerous small
druses, e.g. in Guiera. For inter- and intraxylary phloem see also under
c 1

Young Stem .

ROOT
Negatively geotropic roots, with abundant intercellular spaces in the
cortex, recorded in Laguncularia racemosa Gaertn. and Lumnitzera racemosa
Wffld.

TAXONOMIC NOTES
The wood structure is very uniform throughout the family. Guiera and
Lumnitzera are slightly exceptional in having only scanty paratracheal paren-
chyma and the latter is further distinguished by its radial vessel multiples.
occurrence of included phloem in Combretum appears
It is of interest that the
to be limited to African species, not all of which, however, possess this
character.

ECONOMIC USES
Myrobalans are the and Indian Almonds
fruits of Terminalia chebula Retz.
the seeds of T. catappa Linn. Infusions of the leaves and young stem of
Combretum micranthum G. Don. are used by the natives in western tropical
Africa for fevers and internal complaints generally. Leaves and young twigs,
believed to have been derived from this species and submitted to Kew for
identification, showed the following features. Leaves mucronate, opposite,
somewhat glaucous on the upper surface, but bearing peltate, often yellow,
scales on the lower side. Mesophyll dorsiventral, containing particularly
large cluster crystals in special cavities. Petiole, in transverse sections through
the distal end, exhibiting a solitary, arc-shaped vascular strand with incurved
ends, the xylem vessels being numerous and arranged in radial rows. In the
young stem of the same species intraxylary phloem occurs on the inside of
the closed cylinder of xylem. Vessels small, solitary, or tending to be in radial
rows. The pericycle contains few fibres, and the broad phloem includes
numerous cluster crystals. Cork deep-seated in origin. Pith containing a few
stone cells.
timbers of some species of Terminalia are well known to commerce,
The
though perhaps not used in any very large quantities, e.g. Indian Laurel,
T. alata Roth., Indian Silver Grey Wood, T. bialata Steud., and 2 West
African species, Idigbo, 7". ivorensis A. Chev., and Afara or Limba, T. superba
Engl. et Diels. These and other species of Terminalia and of Anogeissus,
however, may be of considerable importance locally. Gamble (737) said of
T. tomentosa Wight et Arn., 'it is possible that there is no tree in the Indian
forests ... so important for the supply of the agricultural population'.
The wood of Anogeissus spp. is used in India for the axles, shafts, and
wheel-spokes of carts, and that of Bucida buceras Linn, is valued in the West
Indies as a durable construction timber.
620 COMBRETACEAE
GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Anogeissus, Buchenavia, Bucida, Cacoucia, Calycopteris,* Combretum,*
Conocarpus, Guiera, Laguncularia, Lumnitzera, Macropteranthes, Pteleopsis,
Quisqualis, Ramatuela, Terminalia,* Thiloa.
* Kew slide
Represented in the collection.

(ii)
FOR WOOD STRUCTURE
Anogeissus, Buchenavia, Bucida, (Calycopteris), Combretum, Conocarpus,
Guiera, Laguncularia, Lumnitzera, Pteleopsis, (Quisqualis), (Ramatuela),
Strephonema, Terminalia, (Thiloa).

LITERATURE
(i) On General Anatomy
Hate 919, Helm 946.

(ii) On Wood Structure

Bailey 73, 78, Bastos 147, Becking et al. 164, Benoist 170, den Berger 179, 182, Besson
186, British Honduras For. Department 274, Brown, F. B. H. 282, Burgerstein 310, 312,
Chalk et al. 364, Chalk and Chattaway 362, Chowdhury 412-14, Cooper and Record 461,
Coster 481, 482, Desch, H. E. 574, Foxworthy 705, Gamble 737, Hopkinson 1083,
Howard 1088, Janssonius 1154, Jentsch 1171, Jones 1191, Kanehira 1206, 1209,
Lecomte 1334, M&iiaud 1492, Panshin 1649, Pearson and Brown 1679, PfeirTer, H.
1712, Record 1780, 1783, 1787, 1801, 1818, 1825, 1834, 1843, 1851, Record and Hess
1886, Record and Mell 1894, Riera 1937, Stbne 2206, 2207, Torres 2269, Williams 2430.

136.
l^O. MYRTACEAE
K 1 A^HAr,
IVl I

(FiG. 138 on p. 622; FIG. 139 on p. 624; FIG. 140

14 on p. 626; FIG. 141 on p. 628)

SUMMARY
(i) GENERAL
Trees or shrubs from tropical and sub-tropical regions particularly com- ;

mon in Australia. A noteworthy feature is the presence of secretory cavities,

which are lined with epithelium when young, and almost invariably secrete
oily substances. According to Welch (2395) the epithelial layer is not con-
spicuous in the species of Eucalyptus which he examined. The same author
also concluded that the cavities are mainly schizolysigenous in origin. The
cavities are tobe found below the epidermis on either side of the leaf, and
when in this position appear as transparent dots. In some instances they are
enclosed in emergences. They also occur in the ground tissue of the young
stem, particularly the cortex, but have not been recorded in the root. The
mesophyll is often to centric, especially in those species
isobilateral
where the lamina of some or of the leaves normally assumes a vertical
all

position. The hairs are mostly unicellular, or sometimes apparently with

2 chambers, thus resembling those of the Combretaceae in a few genera they ;

tend to be or are 2-armed. The stomata are usually ranunculaceous except

in a few species, and occur on both surfaces of vertically placed leaves. The
cork is often stratified, and partly composed of phelloid cells* The secondary
phloem consists of alternating layers of fibrous and non-fibrous tissue except
in a few genera. The vascular bundles of the young stem, as well as those
 MYRTACEAE 621
of the petiole, are bicollateral in all investigated species. Intraxylary phloem
is believed to be universally present. Tannin is abundant in the tissues.
Both solitary and clustered crystals occur.

(ii) WOOD
Vessels most typically small, numerous, solitary and without definite
arrangement, but medium-sized to large, fewer, with numerous multiples and
a marked tendency to an oblique or radial pattern in some genera, particularly
Eucalyptus perforations simple, intervascular pitting alternate except in 2
i

genera, vestured, commonly minute and with pits to ray cells similar, but
sometimes larger and with oblong pits to ray cells; members of medium
length. Parenchyma typically diffuse or in uniseriate bands in the woods
with solitary vessels, predominantly paratracheal in woods with numerous
multiples, with some intermediate forms with both diffuse and paratracheal
types or with broader bands apparently intermediate between metatracheal
and confluent. Rays exclusively uniseriate or up to 2-6 (mostly 2-3) cells
wide; usually moderately heterogeneous, occasionally markedly so, some-
times homogeneous. Fibres typically with distinctly bordered pits moderately
;

short to moderately long, but usually of medium length. Vasicentric

tracheids commonly present. Intercellular canals of the vertical traumatic
type present in a few genera.

LEAF
Oftenisobilateral or centric, particularly in vertically placed leaves such as
those of Callistemon linearis DC. and in cylindrical leaves like those of
Calothamnus (Fig. 139 H), the centre of the leaf being occupied by aqueous
tissue in the last genus. Species of Eucalyptus and Eugenia possessing both
vertical and horizontal leaves sometimes exhibit isobilateral structure in the
former but are dorsiventral in the latter. According to Ohtani (1636)
mechanical tissue occurs in the leaf margin in Eucalyptus and Melaleuca and
to a lesser extent in Eugenia and Pimenta. Hairs usually simple, unicellular
or apparently 2-chambered, the structure in the last instance somewhat
recalling that of hairs in the Combretaceae sometimes 2-armed or with a
;

tendency towards that structure in Calyptranthes, Eugenia, Myrcia. Glandular

hairs absent. Cork warts, resembling lenticels, recorded in Eucalyptus and
Eugenia\ their mode of development in Eucalyptus globulus Lab. has been
described by Motte (1566). Epidermis coated with wax in some species of
Eucalyptus. Stomata usually occur on all parts of the surface of leaves with
isobilateral or centric structure; ranunculaceous in most instances, but
rubiaceous in Myrcia, and, according to Bandulska (132), in Rhodomyrtus.
The epidermal cells and stomata of Rhodomyrtus and Tristania have been
described in considerable detail by Bandulska (I.e.). Stomata of Darmnia
grandiflora Baker et Smith described by these authors (112) as having guard
cells shaped in transverse section like the arms of a pair of callipers. Hypo-
derm stated to occur below the upper epidermis in several species of Metro-
sideros and Psidium crystalliferous in P. guajava Linn, according to de Boer
;

(214). Mesophyll said to contain sclerenchymatous fibres in Jambosa sp.

and stone cells in certain species of Eugenia. Johnson (1185) has described
differences in the mesophyll structure of horizontal and vertical leaves of
622 MYRTACEAE
Eucalyptus globulus. Elongated water-storage tracheids with spiral thickening
recorded by Baker and Smith (i 12) at the boundary between the palisade and
loose spongy tissue in the centric leaf of Darwinia grandiflora Baker et Smith.
Petiole (Fig. 139 A-D and F). Transverse sections through the distal end in
the few species of Callistemon, Eucalyptus^ Eugenia, Feijoa, Myrtus, Pimenta,
and Syncarpia represented in the Kew slide collection exhibit a single,
principal, arc-shaped vascular strand, which is widely open, U-shaped, or with
somewhat incurved ends according to the species. Additional strands in the
wings observed in Eugenia caryophyllus (Sprengel) Sprague (syn. E. caryo-
phyllata Thunb.). Main vascular strand apparently bicollateral in all species

FIG. 138. MYRTACEAE, A-B; MELASTOMACEAE, C

A, B, Secretory cavity in an emergence of the leaf of Eucalyptus maculata var. citriodora Hook.
C, Stomatal pit of Mouriria eugeniaefoUa Spruce. A-B, after Lignier; C, after van Tieghem.

examined, but adaxial phloem not always very clearly defined; generally
surrounded by a continuous sheath of fibres varying in width in different
species. Solitary and clustered crystals common in all unlignified tissues.
Crystal-sand also recorded by de Boer (214) in the cortical region of the
petiole of Psidium guajava. Secretory elements. Cells containing tannin
abundant in all unlignified tissues. The family is characterized by secretory
with various kinds of, but generally oily, contents (said
cavities (see also 'Axis')
to be filled with mucilage in Tristania laurina R. Br.); situated below the
epidermis, usually on both sides of the leaf; also observed in the cortical
region of the petiole in the few species represented in the Kew slide collection ;
enclosed in emergences in a few species, e.g. in young leaves of Eucalyptus
maculata var. citriodora Hook. (Fig. 138 A-B). Similar nodules, but sur-
mounted by tufts of long hairs, recorded by Fourment and Melis (702) in
E. erythrocorys F. v. M. and E. preissiana Schau. and by shorter hairs in
'E. spatulataflore rubra Hook.'. Secretory cavities frequently appear as trans-

parent dots; probably occurring in all and definitely recorded in most genera.
Young secretory cavities usually lined with a distinct epithelium, but the
latter is soon obliterated by compression and suberization. The mode of
development of the secretory cavities of Eucalyptus globulus has been in-
vestigated by Fohn (689). Concerning the taxonomic value of secretory
cavities in Eucalyptus, Welch (2395) writes as follows: 'Although the distribu-
tion and number of glands is not of very great taxonomic value, yet certain . . .

variations do occur and without doubt hold good throughout the distribution
 MYRTACEAE 623
of the species. Again it is quite possible to recognize certain differences in
'
their arrangement which would permit of a rough classification into groups
Axis
YOUNG STEM (Fig. 139 E, G, i)

Stems, when sufficiently young, frequently provided with wings, e.g. in

Eucalyptus globulus Labill. (Fig. 139 E) and Pimenta officinalis Lindl. Position
of origin of the cork somewhat variable according to the species, arising super-
ficially in Eugenia caryophyllus (Spreng.) Sprague (syn. E. caryophyllata
Thunb.) and E. jambos Linn., but more deep-seated (usually pericyclic) in
certain other species of Eugenia as well as in species of Feijoa, Leptospermum,
Melaleuca (arising in the inner part of the cortex but externally to the peri-
cyclic fibres in M. leucadendron Linn.), Myrtus, Pimenta, Psidium (sometimes
in the centre of the cortex), represented in the Kew slide collection first-
;

formed phellogen sub-epidermal in Callistemon citrinus Skeels but a second

one observed to arise later in the pericycle of the same species. Cork usually
stratified, being composed of layers of radially elongated thin-walled cells
alternating with radially flattened cells sclerosed on their inner or outer
tangential walls. Spongy cork, composed of cubical cells with delicate walls,
occasionally to be found in Eucalyptus and Syzygium. Cortex containing stone
cells in species of Eucalyptus, Eugenia, and Metrosideros. Pericycle generally

including a sub-continuous ring of fibres. Secondary phloem usually con-

sisting of alternating layers of fibrous and soft tissue except sometimes in
Chamaelaucium, Darwinia, Myrtus, Verticordia where little or no scleren-
chyma has been recorded in this position. Stone cells, often very large,
present in the phloem of a few species of Eucalyptus and Syzygium. Phloem
and xylem generally in the form of continuous cylinders traversed by narrow
rays. Vessels with simple perforations. Bicollateral vascular bundles
recorded in Actinodium, Callistemon, Calycolpus, Calythrix, Chamaelaucium,
Darwinia, Eucalyptus, Eugenia, Leptospermum, Melaleuca, Metrosideros,
Myrcia, Myrrhinium, Myrtus, Psidium, Thryptomene, Tristania, Verticordia,
and probably occurring in other genera as well. Where the xylem is con-
tinuous, intraxylary phloem occurs throughout the family. Both solitary and
clustered crystals occur, often abundantly, in the unlignified tissues; solitary
types sometimes situated in chambered fibres in species of Callistemon,
Eucalyptus, Melaleuca', clustered forms present in similar fibres in species of
Eugenia, Myrtus, Syzygium, Secretory elements. Tanniniferous cells
abundant. Cavities containing various chemical substances, generally of an
oily nature, occur in the unlignified tissues, especially the cortex, of most of
the species examined; observed also in the wings, e.g. in Eucalyptus globulus
and Pimenta officinalis. Fourment and Melis (702) record the existence of
4 secretory canals in the pith of Eucalyptus maculata var. citriodora Hook, and
E. fidfolia F. v. M. but not in 29 other species of Eucalyptus, and nodules
containing secretory cavities, similar to those on the leaf, on the very young
1
stem of *". spafalata flore rubra Hook. According to Welch (2395) a trans-
.

parent elastic substance resembling rubber is secreted from papillose

epidermal cells on the young shoots of certain species of the Corymbosae

group of Eucalypts and also of certain Angophoras, where it is thought to
prevent excessive transpiration.
 62 4 MYRTACEAE
BARK
de Beuzeville and Welch (191) record the following facts concerning the
bark of Eucalyptus badjensis de Beuz. et Welch 'Oil glands very numerous,
:

situated in loose parenchymatous tissue formed by the irregular widening

of the medullary rays, chiefly arranged in groups visible to the naked eye.
The phellogen is developed at intervals in the secondary phloem, producing

J
FIG. 139. MYRTACEAE
A, Syncarpia laurifolia Tenore. Petiole X 14. B, Eugenia caryophyllus (Sprengel) Sprague (syn.
E. caryophyUata Thunb.). Petiole Xi8. C, Pimento, acris Wight. Petiole X 19. D, Eucalyptus obUqua
L'Herit. Petiole X 8. E, E. globulus Labill. Young stem x 8. F, Callistemon citrinus Skeels. Petiole
G
Xi 8* f Pimento acris Wight. Stem x 18. H, Cahthamnus laterah s Lindl. Lamina X44. I, Eucalyp-
:

tus obliqua L'Herit. Stem x ia.

c.e. Cutinized epidermis, p.t. Palisade tissue, s.c. Secretory cavity, s.m. Spongy mesophyll.
s.t. Stone cells, u.f. Unlignified fibres.

a succession of absciss phelloids in the periderm/ For anatomical details

concerning Eucalyptus bark used as a source of tannin see 'Economic Uses'
on p. 629.

WOOD (Figs. 140 and 141)

Vessels typically small (less than 100 p mean tangential diameter), very
small (less than 50 p) in at least some species or specimens of Amomis,

Eugenia, Feijoa, Krokia, Myrceugenia, Myrria, Myrrhinum, Myrtekmania,

Myrtella, Myrtus, Pimenta, Psidium, and Rhodamnia, medium-sized
(100-200 p) in some species of Calycolpus, Campomanesia, Eucalyptus,
 MYRTACEAE 625

Eugenia, Kjellbergiodendron, Leptospermum, Marlierea, Melaleuca, Metro-

sideros, Myrcia, Myrtus exaltata M. F. Bailey, Pleurocalyptus, Rhodamnia,
and Tristania, large (more than 200 p.) in some species of Angophora and
Eucalyptus typically exclusively solitary but with occasional to numerous
;

multiples of 2 or 3 cells in Calyptranthes p.p., Eucalyptus p.p., Eugenia, Gomi-

desia, Jambosa, Leptospermum, Myrtella, Myrtus p.p., and Syzygium, and com-
monly with multiples of 4 or more cells in Angophora (Fig. 140 H), Eucalyptus
p.p., and Eugenia p.p. the vessels usually without any definite pattern, but
;

forming loose oblique lines in Angophora, Calycorectes, Campomanesia,

Eucalyptus (Fig. 141 A and E), Kjellbergiodendron, Leptospermum, Marlierea,

Melaleuca, and Xanthostemon and with an occasional tendency to oblique
lines locally in some other genera. Vessels sometimes absent from zones
separating the growth rings, such zones sometimes very conspicuous, e.g. in
Eucalyptus gigantea Hook. often very numerous and crowded in woods with
;

small vessels, e.g. more than 100 per sq. mm. in some species of Eugenia,
Feijoa, Myrtus, and Psidium, but sometimes with small but relatively widely
spaced vessels; not more than 2-5 vessels per sq. mm. in some species of
Eucalyptus, Kjellbergiodendron, Metrosideros, and Rhodamnia, 5-20 per sq.
mm.- in some species of Calycolpus, Campomanesia, Eucalyptus, Gomidesia,
Jambosa, Melaleuca, Myrcia, Psidiopsis, Psidium, Syncarpia, Syzygium, and
Tristania, 40-100 per sq. mm. in some species of Amomis, Backhousia,
Calyptranthes, Myrceugenia, Myrcia, Myrrhinium, Myrtekmania, Myrtella,
Myrtus, Osbornia (1649), and Psidium', with spiral thickening in some species
of Eugenia and Myrceugenia and,- according to Janssonius (1154), in Eugenia
acuminatissima Kurz. Perforation plates simple, except in Myrceugenia and
Myrtus communis L., in which they are exclusively scalariform with about
15-25 fine bars. Scalariform perforation plates also observed in a young stem
of Eugenia sp. cultivated at Kew. Intervascular pitting typically alternate and
minute, small to moderately large in Angophora, Eucalyptus, Eugenia, Jambosa,
Kjellbergiodendron, Osbornia (1649), and Tristania, pits opposite or inter-
mediate in Myrceugenia and Myrtus communis L. vestured (78) pits to wood
; ;

parenchyma often rare owing to presence of vasicentric tracheids, pits to ray

cells usually similar to the intervascular pitting where this is small, often

large, oblong and simple in the above-mentioned genera with larger inter-
vascular pitting. Solid deposits of gum common, tyloses observed or recorded
in Angophora, Baeckia, Eucalyptus, Eugenia (525), Melaleuca, Rhodamnia,
Syncarpia, Tristania, and Xanthostemon. Mean member length 0*3-0*8 mm.
Parenchyma of 2 intergrading types, (a) predominantly apotracheal as
scattered cells or irregular uniseriate bands, in most of the genera with solitary
vessels (Fig. 140 K), and (b) predominantly paratracheal in the genera in which
vessel multiples are relatively common; entirely or predominantly apotra-
cheal in the following genera: Amomis, Backhousia, Baeckia, Callistemon,
Calycolpus, Campomanesia, Decaspermum, Eugenia (a few
Calycorectes,
species), Krokia, Leptospermum, Marlierea, Metrosideros, Myrceugenia, Myrcia,
Myrrhinium, Myrtekmania, Myrtus p.p., Pimenta, Psidiopsis, Psidium, and
Tristania', entirely or predominantly paratracheal in the following: scanty
paratracheal or vasicentric in Eucalyptus p.p., Gomidesia, and Xanthostemon,
aliform to confluent in Calyptranthes syzygium (L.) Sw., Eucalyptus p.p.,
Eugenia p.p. (Fig. 140 E), Kjellbergiodendron, Melaleuca p.p. (together with
4594 g s
 FIG. 140, MYRTACEAE
A, Jambosa jambos (L.) Millsp. B, Campomanesia crenata Berg, C, Calyptranthes syzygium Sw.
D, C. syzygium Sw, E, Eugenia urceolata King. F, Jambosa jambos (L.) Millsp. G, Decaspermum
paniculatum Kurz. H, Angophora intermedia DC. K, Myrtus communis Linn.
 MYRTACEAE 627
diffuse crystalliferous strands in addition) and Myrtella, the aliform
parenchyma sometimes limited to the abaxial sides of the vessels (Fig.
140 D); intermediates between the 2 main types are common, e.g. diffuse
and paratracheal parenchyma equally abundant in Angophora (Fig. 140 H)
(sometimes tending to form bands), Eucalyptus p.p., Melaleuca p.p.,
Osbornia (1649), Pleurocalyptus, Syncarpia, and Syzygium, with broader
bands of indeterminate type and often with long wings from the vessels in
Calyptranthes sericea Grisch., Eugenia p.p., Jambosa (Fig. 140 A), Myrtus
exaltata, Rhodamnia p.p., and Syzygium cordatum Hochst.; apparently
absent from Leptospermum javanicum Bl. (1154); terminal bands sometimes
present. Cells typically filled with a resinous or gummy substance that some-
times gives a tannin reaction; crystals comparatively rare, present in
chambered cells in some species of Angophora, Eucalyptus (e.g. the Blood-
wood Group), Eugenia, Feijoa, Kjellbergiodendron, Marliera, Myrceugenia,
Myrcia, Myrrhinium, and Psidium. Some of the crystalliferous cells of Myrcia
and Psidium are distinctly wider than those of the normal strand. Silica
grains have been reported by Janssonius (1154) in Eugenia sexangulata Koord.
et Valet, and E. densiflora Duthie. Strands usually of up to 8 cells. Rays

ocfcasionally with a suggestion of being of 2 distinct sizes typically up to 2-3

;

cells wide, exclusively uniseriate or with only occasional biseriate parts in

Eucalyptus p.p., Melaleuca, Pleurocalyptus, Tristania p.p., and Xanthostemon,

4-6 cells wide in some species of Baeckia, Calyptranthes, Campomanesia, Lepto-
spermum, Myrtus, and Rhodamnia, up to 8 cells wide in Gomidesia less than
;

i mm. high except in some of the species with more than 10 marginal rows of

upright cells uniseriates numerous, typically composed of square to upright

;

cells only, except in the woods with homogeneous rays and in Eucalyptus, and
often only i or 2 cells high; composed of mixed procumbent and upright to
square cells in some species of Eugenia, and wholly or mostly of procum-
bent cells in the other species of Eugenia and in Eucalyptus-, mostly 13-20
rays per mm., sometimes more than 20 per mm., e.g. in Calycorectes, Myrtek-
mania, Myrtus, and Rhodamnia, sometimes rather fewer than 13 per mm.,
for example in Angophora, Calycolpus, Calyptranthes, Eucalyptus, Eugenia,
Leptospermum, Psidium, Syncarpia, and Xanthostemon and about 3 per mm.
in Gomidesia\ typically heterogeneous (Kribs's Types II A and B) with 4-10

marginal rows of upright cells with more than 10 rows of upright cells (often
;

Kribs's Heterogeneous Type I), in some species of Jambosa, Myrceugenia,

Myrcia, Psidiopsis, and Rhodamnia', with only 1-3 rows in some species of
Calyptranthes, Campomanesia, Eucalyptus, Feijoa, Gomidesia, Krokia, Metro-
sideros, Myrtella, and Psidium', homogeneous (Kribs's Type I) in Angophora,
Eucalyptus p.p., e.g. E. maculata Hook., Myrcia p.p., and Tristania p.p.;
with sheath cells in Gomidesia. Cells typically filled with a dark gummy or
oily substance and without crystals; crystals abundant in Calyptranthes
syzygium (L.) Sw. silica has been reported (194, 794, 1126) in Eugenia,
;

Metrosideros, and Tristania. Fibres typically with bordered pits, the pits
varying from rather few to very numerous, usually equally distributed on
both the radial and the tangential walls, but sometimes more numerous on
the tangential walls; rarely more numerous on the radial walls; pits simple
or with indistinct borders in Eugenia, Gomidesia, and Jambosa. Accord-
ing to Gouwentak (799) both libriform fibres and fibre-tracheids occur in
628 MYRTACEAE
some species of Eucalyptus. Walls moderately to very thick. Often containing
gum, oil, or other matter, which sometimes produces the appearance of septa;
Gouwentak (799) notes the presence of tannin in the fibre-tracheids of
Eucalyptus rostrata Schl. Occasional septate fibres present in some species,
e.g. of Eugenia(ii$4)%ndMarlierea. Mean length 0*7-2*0 mm. Vasicentric
tracheids present in most of the genera, not observed or not clearly dis-
tinguishable from the fibre-tracheids in some species of Backhousia Baeckia, y

Decaspermum, Eucalyptus, Gomidesia, jfambosa, Krokia, Myrceugenia^ Myrtus,

Psidium, Syncarpia, Syzygium, and Xanthostemon. Intercellular canals of

FIG. 141. MYRTACEAE

A, Eucalyptus globulus Labill. B, E. resinifera Sm. C, E. regnans F. Muell. D, E, gigantea Det.
, E. marginata Sm.

the vertical traumatic type reported (1851) to occur in Angophora, Eucalyptus,

and Rhodamnia, those of Eucalyptus sometimes very large; their origin is
discussed by Jacobs (1136); radial schizogenous canals recorded by Janssonius
in the rays of Eugenia cuprea Koord. et Valet., and Leptospermum javanicum
Bl. Growth rings* The seasonal development of the wood in Eugenia and
Psidium has been investigated by Coster (481) and Chowdhury (414). The
nature of the ash has been used as an aid to distinguishing between species of
Eucalyptus (337, 338, 518, 522, 523). The effect of water-supply
on the struc-
ture of the wood and the rate of water conduction in Eucalyptus rostrata
Schlecht. has been investigated by Greiss (816-17).

ROOT
Musson and Carne (1576) have recorded the normal occurrence of adventi-
tious roots on the trunk of Melaleuca linariifolia Sm. Bird (199) describes the
cortex of the climbing roots of Metrosideros hypericifolia A. Cunn. as com-
posed of chlorophyllous cells when young-, but becoming lignified and serving
as mechanical tissue when older. Other features recorded in the same species
include the normal absence of root hairs except when their formation is
induced by treatment in a moist chamber; the conspicuous endodermis; the
vessels in the xylem of the central cylinder with small lumina and very thick
 MYRTACEAE 629
walls. The vessels in the absorbing roots are rather larger and the cortex
does not become converted to mechanical tissue.

TAXONOMIC NOTES
(i)FROM GENERAL ANATOMY
The Myrtaceae, as understood by Bentham and Hooker, included the
Lecythidaceae, now generally treated as a separate family. Comparison of the
anatomical features of the Lecythidaceae with those of the Myrtaceae sensu
shows that there are important differences between the two groups, and
stricto
favours their recognition as distinct families.

(ii)FROM WOOD STRUCTURE

There do not appear to be any consistent differences between the Myrteae
and the Leptospermoideae. Differences between species of the same genus
are sometimes unusually marked, e.g. between Calyptranthes sericea Griseb.
and C. syzygium (L.) Sw,, Myrcia spp., and Myrtus communis L. and M.
exaltata M. F. Bailey, the latter suggesting Eugenia. Gomidesia lindeniana
Berg, distinct in some respects. For comparison with Lecythidaceae see that
is

family. Dadswell and Ingle (531) state that the wood structure definitely
supports the revision of the genus Eugenia proposed by Merrill and Perry,
and that, anatomically, the species of the New World belonging to the genus
Eugenia are quite distinct from the majority of the Australian and New Guinea
species. They, however, consider that the two Australian species Eugenia
carissoides F. Muell. and E. macrophylla C. T. White et Francis are true

Eugenias, similar to those of the New World. On the other hand, these
authors have found little to support the reinstatement of the genera Acmena
and Cleistocalyx.

ECONOMIC USES
from the flowers and foliage of various members
Essential oils are distilled
of the family. The well-known Eucalyptus oil used in medicine is said to be
derived from Eucalyptus polybracteata R. T. Baker, E. dumosa A. Cunn., and
other species of Eucalyptus. Lemon-scented Eucalyptus oil is obtained from
E. maculata var. citriodora Hook., whilst the oil of E. dives Schau. is used in
the manufacture of thymol. Cajuput oil, also used in medicine, is derived
from Melaleuca leucadendron Linn, and other species of Melaleuca. Other
important products derived from the family include Cloves (Eugenia caryo-
phyllus (Sprengel) Sprague syn. E. caryophyllata Thunb.), Allspice (Pimenta
officinalis Lindl.); Guava Fruits (Psidium guajava Linn.), Rose Apple (Eugenia
jambos Linn.), and Jambolana Fruits (E, jambolana Lam.). The bark of
various species of Eucalyptus has been used as a source of tannin, and at one
time Mallet Bark (Eucalyptus occidentalis Endl.) was imported into European
countries from Western Australia for use in tanning. According to Wiesner
(2423) true Mallet Bark may be distinguished from that of other species of
Eucalyptus by the horny transverse fracture. Transverse surfaces of mature
specimens exhibit i or more layers of cavities filled with dark red, glistening
masses of Eucalyptus Kino. Commercial samples consist mostly of the inner
bark. The phloem parenchyma and rays which are 2-3 cells wide consist of
thin-walled, tanniniferous parenchyma. Small or large clusters of radially
 630 MYRTACEAE
arranged groups of fibres occur in the secondary phloem. Paired crystals of
calcium oxalate are characteristic of this as well as of other Eucalyptus barks.
Further particulars concerning mallet bark have been recorded by Bodenstab
(211). The many important timbers produced by this family almost all
belong to the sub-family Leptospermoideae and are mostly species of the
single genus Eucalyptus. Among the best known are Jarrah (Eucalyptus
marginata Sm.), Karri (E. diversicolor F. Muell.), and 'Tasmanian Oak* (E.
gigantea Hook, f., E. obliqua L'Herit., and E. regnans F. Muell. ). Many other
species are well known at least locally, e.g. Tallow wood E. microcorys F.
Muell. and the various 'Ashes', 'Boxes', Ironbarks, and Gums.
Two closely related species of Metrosideros, Kajoe Lara and Kajoe Nani
from the Netherlands East Indies, are reputed to be specially suited for marine
construction, the high silica content making them resistant to marine borers
(1126), and the same appears to be true of the Australian Turpentine, Syn-
carpia laurifolia Ten. The Brush Box, Tristania conferta R. Br., and Satinay,
Syncarpia billii Bailey, are important construction timbers in Australia and
species of Eugenia are used for building in Burma, India, and tropical and
sub-tropical America. Many of the other genera produce hard, tough timbers
that are used locally and the woods of the family as a whole are excellent for
fuel.

Many species are extensively grown as exotics in forestry plantations in

different parts of the world, e.g. Eucalyptus globulus Labill., E. maculata Hook,,
E. microcorys F. Muell., E. robusta Sm., E. rostrata Schlecht, and E. saligna Sm.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Actinodium, Angophora, Beaufortia, Callistemon,* Calothamnus, Caly-
colpus, Calycorectes, Calyptranthes, Calythrix,Campomanesia, Chamaelau-
cium, Cupheanthus, Darwinia, Decaspermum, Eremaea, Eucalyptus,*
Eugenia,* Feijoa,* Gaslondia, Jambosa, Kunzea, Leptospermum,* Lhotzkya,
Marlierea, Melaleuca,* Metrosideros, Myrcia, Myrtus,*Pileanthus, Pimenta,*
Psidiopsis, Psidium,* Regelia, Rhodamnea, Rhodomyrtus, Syncarpia,*
Syzygium, Thryptomene, Tristania, Verticordia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Amomis, Angophora, Backhousia, Baeckia, Callistemon, Calycolpus,
Calycorectes, Campomanesia, Decaspermum, Eucalyptus,
Calyptranthes,
Eugenia, Gomidesia, Jambosa, Kjellbergiodendron, Krokia, Leptospermum,
Marlierea, Melaleuca, Metrosideros, Myrceugenia, Myrcia, Myrrhinium,
Myrtekmania, Myrtella, Myrtus, (Osbornia), Pimenta, Pleurocalyptus,
Psidiopsis, Psidium, Rhodamnia, Syncarpia, Syzygium, Tristania, Xantho-
stemon.

LITERATURE
(i) On General Anatomy
Baker and Smith 107-11, Bandulska 132, de BeuzeviHe and Welch 191, Bird *99j Boden-
Fohn 689, Fourment and Mclis 702, Johnson 1185, Motte
stab 211, Boer 214, Brogli 277,
1566, Musson and Carne 1576, Ohtani 1636, Wallis and Santio 1535, Welch 2395, Wiesner
 MYRTACEAE 631
(ii) On Wood Structure
Anonymous 27, Bailey 73, 78, Baker 104, Beekman 167, Benoist 170, den Berger 179,
182, Besson 186, Betts, M. W. 188-9, Bianchi 194, Brown, F. B. H. 282, Burgerstein
310, 312, Campion 337, 338, Chowdhury 411, 414, Coster 481, Dadswell et al. 518, 522,
523, 525, 531, Diehl 582, Foxworthy 705, Garratt 744, Giordano 786, Gonggrijp 794,
Gouwentak 799, Greguss 2522, Greiss 816, 817, Howard 1088, van Iterson 1126, Jacobs
1136, Janssonius 1154, Jolly 1188, Jones 1191, Kanehira 1206, 1209, Kribs 1283,
Lecomte 1334* McNair, J. B. 1476, Milanez 1521, Panshin 1649, Pearson and Brown 1679,
Pereira 1687, Record 1780, 17835 1801, 1843, 1851, Record and Hess 1886, Record
and Mell 1894, Stone 2202, 2207, Tupper 2294, Webber 2377, Welch 2398, 2402, 2403,
2406, Williams 2430.

137. LECYTHIDACEAE
(FiG, 142 on p. 634; FIG. 143 on p. 638)

SUMMARY
(i) GENERAL
A
tropical family of trees and shrubs which differs from the Myrtaceae in
the absence of secretory cavities from the leaves and ground tissues of the
stem. Intraxylary phloem is also absent. The leaf is usually dorsiventral,
or, more rarely, centric. The stomata are usually confined to the lower
surface of the leaf, but sometimes occur on the upper side as well. They are
usually cruciferous. The structure of the petiole, which is frequently some-
what complex, provides useful diagnostic characters. The presence of cortical
vascular bundles is one of the most characteristic features of the axis.
These appear to be truly cauline, since they have been seen to be accompanied
by separate leaf trace bundles in certain species.

(ii) WOOD
Vessels very small tolarge, sometimes with pronounced radial multiples
or irregular clusters, perforations simple, intervascular pitting alternate and
minute to large, pits to parenchyma often large and simple, members of
medium length. Parenchyma typically in apotracheal bands, varying from
uniseriate and broken to wide and continuous, but predominantly aliform to
confluent in a few genera, long, chambered, crystalliferous strands charac-
teristic of the New World genera. Rays mostly 2-3 cells wide, but very
broad in some genera, distinctly heterogeneous to homogeneous. Fibres
with simple pits, of medium length to very long. Intercellular canals of
the vertical traumatic type present in some genera.

LEAF
Usually dorsiventral; centric structure recorded in certain species of
Asteranthos and Foetidia. Hairs simple, unicellular or uniseriate, occasionally
tending to be tufted. Glands present on the lower side at the base of the leaf
in Napoleona. Lower epidermis papillose in certain species of Couratari
and Lecythis. Stomata usually confined to the lower surface, but occurring
on the upper side also in a few species of Baningtonia, Foetidia, Gustavia;
generally cruciferous, but reported to be rubiaceous in Chytroma
idatimon
Miers. Ordinary stomata are accompanied by especially large ones in
Napoleona whitfieldii Decne. Hypoderm recorded below the upper epidermis
632 LECYTHIDACEAE
in a few species of Barringtonia and Foetidia. Mesophyll stated to contain
sclerenchymatous fibres in Asteranthos. Petiole. Transverse sections through
the distal end in the few species examined exhibit a main arc of widely spaced
collateral bundles, sometimes accompanied by accessory strands on the
adaxial side, e.g. in species of Bertholletia, Gustavia (Fig. 143 E), Lecythis, and
on the abaxial side as well in Gustavia. Solereder records the occurrence in
the petiole of a single arc of 5 bundles in the Napoleoneae, and 2 or 3 con-
centrically arranged arcs in the Barringtonieae and Lecythideae. A
petiole
of Napoleona imperialis Beauv. examined at Kew is shown in Fig. 143 H.
Crystals, according to Solereder, solitary in the Napoleoneae; mixed
solitary and clustered in the Lecythideae; star-shaped clusters in the
Barringtonieae. Secretory elements. Cells containing tannin usually
present. Secretory cavities absent.

Axis
YOUNG STEM (Fig. 143 J-K)
Cortical vascular bundles occur in Asteranthos, Barringtonia, Bertholletia
(Fig. 143 j), Carey'a, Couratari, Foetidia, Gustavia, Lecythis, Napoleona
(Fig. 143 K), Planchonia\ said to be inversely orientated in Barringtonia,
Foetidia, Gustavia, Planchonia; normally orientated in other genera. Smaller
cortical bundles tending to be centric. Cortical bundles of genera represented
in the Kew slide collection sheathed by thick-walled fibres accompanied, at
the outer periphery, by a single layer of cells containing solitary crystals,
except in Gustavia where cluster crystals occur in the corresponding position.
According to Solereder the number of cortical bundles in the Napoleoneae is
much smaller than in the remainder of the family. Cork arising superficially,
usually composed of flattened cells, sometimes stratified into layers with thin
cell walls alternating with others having the inner or outer tangential wall
thickened in species of Cariniana, Gustavia, Lecythopsis. Pericycle contain-
ing a somewhat interrupted ring of fibres in all of the few species represented
in the Kew slide collection, but the fibre strands become more widely spaced
as the stems grow older. Secondary phloem generally stratified into fibrous
and soft portions. Phloem and xylem usually in the form of continuous
cylinders traversed by narrow rays, but rays broader and provided with
enlarged, triangular, distal ends in Napoleona (Fig. 143 K). Vessels with
simple perforations; scalariform plates also recorded (see 'Wood'). Primary
phloem strands in the last genus also triangular with outwardly directed
apices. Intraxylary phloem absent. Secretory elements. Mucilage canals
recorded in species of Bertholletia, Couratari, Eschweilera, Lecythis, but
apparently not present throughout these genera. Cells with amorphous,
probably tanniniferous contents common in the unlignified tissues. Solitary
and clustered crystals common (see also under 'Cortical Bundles').

WOOD (Fig. 142)

Vessels mostly medium-sized (100-200 mean tangential diameter),
//,

moderately small (50-100 p) in Asteranthos p.p. (582), Careya p.p. (582),

Eschweilera^ Foetidia, Gustavia, and Napoleona, very small (25-50 /z) in
Grias, large (more than 200 /*) in Bertholletia, Cariniana, Couratari, and
Couroupita solitary and in small multiples, commonly with multiples of 4 or
;
 LECYTHIDACEAE 633
more Careya, Eschweilera p.p., and Lecythis p.p., with irregular clusters
cells in
in Allantoma, Careya, and Napoleona\ very variable in number, ranging from
i '5-5 per sq. mm. in Careya, Cariniana, Couroupita, Eschweilera^ and
Lecythis to 20-40 in Foetidia, Grias, and Gustavia, the latter sometimes with
more than 40 per sq. mm. (582); spiral thickening recorded by Janssonius
(1154) in Planchonia sundaica Bl. Perforation plates simple, except for occasional
scalariform or reticulate plates observed in Allantoma and one specimen of
Barringtonia speciosa L. and reported by Record and Hess (i 886) in Asteranthos
and Grias.Intervascular pitting alternate, except in Grias, in which it is
transitional between scalariform and opposite; varying in size from minute,
e.g. in Eschweilera, Foetidia, Grias, and Gustavia, to large, e.g. in Barring"
tonia, Combretodendron, Couroupita, and Petersianthus\ sometimes with occa-
sional pits elongated horizontally to about twice their usual width, e.g. in
Planchonia andamanica King and some species of Napoleona (582); pits to
ray and wood parenchyma typically including many large, elongated, slightly
bordered or simple pits, such pits tending to be circular in woods with minute
intervascular pitting, e.g. in Gustavia, particularly when adjoining ray cells;
absent from Chytroma. Tyloses observed in Allantoma, Asteranthos, Berthol-
letia,. Careya, Cariniana, Chydenanthus, Combretodendron, Couratari, and

Lecythis', sclerosed, according to Diehl (582), in some specimens of Aster-

anthos, Eschweilera, and Lecythis] deposits common in Foetidia
solid
mauritiana Lam. Mean member length 0-4-0*6 mm. Parenchyma typically
abundant and apotracheal banded with some paratracheal in numerous, ;

short, irregular (almost diffuse) lines in Asteranthos, Barringtonia (with vasi-

centric or aliform parenchyma in addition and sometimes predominant, see
Fig. 142 A) and Grias (1894) ; in continuous bands i, occasionally 2, cells wide
in Careya, Cariniana (Fig. 142 D), Couratari, Couroupita, Eschweilera p.p.,
Foetidia, Gustavia (Fig. 142 H), Lecythis p.p., and Napoleona, in broader
bands in Allantoma, Bertholletia, Eschweilera p.p., and Lecythis p.p.; the fine
bands sometimes very numerous, e.g. 25 or more per mm. in Gustavia
rhodantha Standl. and Napoleona parviflora Bak. f.; predominantly para-
tracheal, aliform to confluent and usually with some diffuse, in Chytroma,
Combretodendron (Fig. 142 E), Petersia (582), and Petersianthus', parenchyma
very scarce or absent from the only available specimen of Grias fendleri Seem.
Crystalliferdus strands are common in the New
World genera and, according
to Diehl (582), serve as a basis for separating these from the rest of the family,
with the exception of Foetidia. The crystalliferous cells often occupy the
complete length of the strand and are sometimes locally biseriate (tangential
section), e.g. in Cariniana the strands occur on the margins of the apotracheal
;

bands, except in Foetidia, in which they are scattered among the fibres. The
ordinary strands are most commonly of up to 6 or 8 cells. Rays very variable
in width, ranging from exclusively uniseriate in occasional specimens of
Eschweilera and Lecythis to up to 20 cells wide in Napoleona vogelii Hook.
et Planch.; mostly up to 2-3 cells wide; 4-10 cells wide in Barringtonia,
Bertholletia, Chydenanthus, Combretodendron, Couroupita, Gustavia p.p.,
and Petersianthus, more than 10 cells wide in Grias, Gustavia p.p., and
Napoleona', typically more than i, and often several, millimetres high,
but less than i mm. in Allantoma, Eschweilera, Foetidia^ Lecythis
p.p., Petersianthus, and Planchonia] uniseriate rays sometimes very
 634 LECYTHIDACEAE
scarce or absent, e.g. in Barringtonia p.p., Careya, Cariniana, Combreto-
dendron, Couroupita, Grias> Lecythis p.p., and Napoleona^ when present
usually composed of mixed procumbent and upright cells; mostly 5-12 rays
per mm., more numerous (up to 16 per mm.) in some species of Eschweilera,
Foetidia, and Planchonia, fewest (1-3 per mm.) in Grias and Napoleona\

FIG. 142. LECYTHIDACEAE

A, Barringtonia acutangula (L.) Gaertn. B, B. acutangula (L.) Gaertn. C, Cariniana estrellensis
(Raddi) Kuntze. D, C. estrellenris (Raddi) Kuntze. E, Combretodendron africanum Exell. F, Eschweilera
G
calyculata Pitticr. Napoleona vogelii Hook, ct Planch. H, Gustavia brasiliana DC.

commonly almost homogeneous, with i or 2 marginal rows of square cells

present in some of the rays, homogeneous (Kribs's Types I and II) in
Allantoma, Cariniana^ Couratari, Eschweilera, Lecythis, and Napoleona p.p.,
distinctly heterogeneous (Kribs's Type II, mostly A, occasionally B), in
Asteranthos, Barringtonia p.p., Careya, Combretodendron, Foetidia, Gustavia^
Petersianthus, and Planchonia\ the cells are nearly all almost square in
Asteranthos (1886); in Gustavia brasiliana DC. small groups of procumbent
cells alternate with square or upright cells sheath cells occasionally present
;
 LECYTHIDACEAE 635
in Some species of Barringtonia. Gummy
deposits common, often filling the
cells ;
some species of^Barringtonia, Cariniana,
crystals observed or reported in
Chydenanthus, Couroupita, Gustavia, and Napoleona, and, according to Diehl
(582), more common in the Old than in the New World genera. Inclusions
of present in the ray cells of Eschweilera longipes Miers (1126). Fibres
silica

typically with simple pits, but the pits distinctly bordered in Allantoma, and
with small, indistinct borders in a few genera, e.g. Couroupita and Peter$i-
anthus\ the pits more numerous on the radial than on the tangential walls.
Typically with thick to very thick walls, sometimes with a gelatinous layer,
e.g. in Barringtonia, Grias, Gustavia, and Petersia (582); walls moderately
thin in some species of Bertholletia, Cariniana, Couratari, Couroupita, and
Grias. Diehl (582) reports septate fibres in Planchonia andamanica King. Mean
length i -0-2-4 mm
Intercellular canals of the vertical traumatic type,
-

similar to the gum veins of Eucalyptus, reported by Record and Hess (1886)
in several specimens of Eschweilera, a few of Lecythis, and one of Cariniana,

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY

The absence of secretory cavities and intraxylary phloem, the presence of

cortical bundles in the axis, and the frequently complex vascular structure of
the petiole serve to differentiate the Lecythidaceae from the Myrtaceae.

(ii) FROM WOOD STRUCTURE

Diehl (582) has made a study of this family and comes to the following
conclusions :

'Excepting the occurrence of crystal strands, differences within the family,

i.e.between genera and groups of genera, are few, and, being of degree rather than
of kind, are at present of dubious diagnostic value.'
'The Lecythidaceae may be separated into two sections on the basis of the
presence or absence of crystal strands. These structures are a unique and distinctive
character and show a striking correlation with geographical distribution, being
absent in the Old World genera, except Foetidia, and constantly present in those of
the New World/
'The anatomical structure of the Lecythidaceae lends support to their segregation
1
from the Myrtaceae.
'On the basis of the anatomical structure, Asteranthos and Napoleona are properly
included with the Lecythidaceae/
'The Myrtaceae differ from the Lecythidaceae principally in having mostly
solitary pores, fibres with distinctly bordered pits, and wood parenchyma that is
paratracheal or diffuse instead of distinctly metatracheal.'

It should be pointed out, however, that the parenchyma distinction is not as

clear as Diehl implies. For example, in the author's opinion, the parenchyma
of Chytroma (not included by Diehl) should be classified as aliform and that
of Combretodendron and Petersianthus as predominantly paratracheal (aliform
to confluent); also the apotracheal parenchyma of Barringtonia p.p. and
Planchonia is as much diffuse (as opposed to metatracheal) as in some genera
of Myrtaceae the Myrtaceae also include some genera with predominantly
;

paratracheal parenchyma, e.g. Gomidesia, Kjellbergiodendron, and Melaleuca.

636 LECYTHIDACEAE
ECONOMIC USES
The
hard, lidded capsules of Lecythis are known as Monkey Pots and used
in South America as water vessels after removal of the seeds. The most
familiar product in European countries is the Brazil nut (Berthottetia exceha
H. B. et K.). Sapucaia nuts (Lecythis usitata Miers and L. ollaria Linn.) are
less well known. The Anchovy Pear is the fruit of Grids cauliflora Linn.
The Surinam timber Manbarklak, Eschweilera longipes Miers and E. sub-
glandulosa (Steud.) Miers, is remarkable for being more resistant to marine
borers even than Demarara Greenheart, owing to the presence of silica
particles (1126). It is reputed to be becoming of increasing importance for
marine construction, particularly in brackish water (1886).
The timber of Cariniana pyriformis Miers has been exported from Colombia
to Europe and the U.S.A. under the name Colombian Mahogany. Silica,
which is likewise present in this timber, is stated by Record and Hess (1886)
to cause excessive dulling of tools when the wood is being worked.
Some other timbers from this family are used locally for general utility
purposes and Record and Hess consider that the American members are
become important owing to their abundance, large dimensions, and
likely to
good form, and the wide range of properties among the different species.
Pearson and Brown (1679) include Barringtonia acutangula Gaertn. and
Careya arborea Roxb. as timber species in India.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Asteranthos, Barringtonia, Bertholletia,* Careya, Cariniana, Couratari,
Eschweilera, Foetidia, Gustavia,* Lecythis,* Lecythopsis, Napoleona,*
Petersia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

Allantoma, Asteranthos, Barringtonia, Bertholletia, Careya, Cariniana,
Chydenanthus, Chytroma, Combretodendron, Couratari, Couroupita,
Eschweilera, Foetidia, Grias, Gustavia, Lecythis, Napoleona, (Petersia),
Petersianthus, Planchonia.

LITERATURE
(i) On General Anatomy
Knuth 1257.
(ii) On Wood Structure
Benoist 170, den Berger 179, Cooper and Record 461, Dadswell and Record 533, Desch
574, Diehl 582, Dixon 592, Howard 1088, van Iterson 1126, 1127, Knuth 1257, Kribs
1283, Martin- Levigne 1450, Pearson and Brown 1679, Pfeiffer, J. Ph. 1713, Record 1802,
1836, 1843, 1851, Record and Hess 1886, Record and Mell 1894, Stone 2202, 2207,
Sudworth and Mell 2219, Tupper 2295, Williams 2430.
 (637)

138. MELASTOMACEAE
(FiG. 138 on p. 622; FIG. 143 on p. 638; FIG. 144 on p. 640; FIG. 145 on p. 646)

SUMMARY
(i) GENERAL
A
mainly tropical family of herbs, shrubs, and trees. Some species, e.g. of
Medinilla, are epiphytes. Various complex types of hairs, which may be
glandular or non-glandular, occur very widely and constitute a charac-
teristic feature. Simple hairs are relatively infrequent. A great many details

concerning their more

particular structure and
distribution, given by
Solereder, have not been repeated here, but they would be an aid to the
identification of genera and species if reinvestigated. The stomata are
usually ranunculaceous or cruciferous, but caryophyllaceous and, more rarely,
rubiaceous types have been recorded. Stomata are occasionally almost entirely
surrounded by a single cell, and sometimes confined to cavities of various
sizes. The
petiole, in transverse sections of all investigated species, exhibits
an arc composed of widely spaced vascular bundles which are frequently
accompanied by variously arranged accessory bundles. In the stem, the
outstanding features are the universal occurrence of intraxylary phloem,
and the common, but not universal, occurrence of cortical and medullary
bundles which may be present alone or together in different genera and
species. Interxylary phloem also occurs in a number of genera. Cork
arises superficially in most genera, or in the pericycle in others. It is occa-

sionally replaced by aerenchyma in submerged parts of a few aquatic

species. The endodermis is usually well defined, but the pericycle never
contains a composite and continuous ring of sclerenchyma, although a propor-
tion of sclerosed cells
is often present.

(ii) WOOD
Vessels usually small, occasionally medium-sized, small multiples and
clusters
common, perforations simple, intervascular pitting typically alternate,
commonly minute, but sometimes moderately large, vestured, pits to paren-
chyma either similar to the intervascular pitting or conspicuously elongated
and simple; members of medium length. Parenchyma typically para-
tracheal only and rather sparse, particularly in the Melastomoideae, but
varying from aliform to broad apotracheal bands in a few genera sometimes ;

in isolated groups among the fibres and characterized by conspicuous inter-

cellular spaces; paratracheal parenchyma always present. Rays of the
Melastomoideae typically uniseriate and composed almost entirely of square
or upright cells; 2-4 cells wide in a few species. Rays of the Astronioideae
and Memecyleae typically 2-5 cells wide, the larger rays composed mainly of
procumbent cells with 1-3 marginal rows of upright cells, the uniseriate rays
often very low; wholly uniseriate in a few species, composed mainly of
procumbent cells. Fibres of the Astronioideae and Memecyleae typically
with bordered pits and thick walls; those of the Melastomoideae with simple
pits, usually septate and with thin walls, commonly in distinct radial rows;
often with parenchyma-like patches or bands of thinner-walled cells with
 K
FIG. 143. MELASTQMACEAE, A-D, F-G, and I; LECYTHWACEAE, E, H, and J-K
A, Medinilla magnified Lindl. Petiole X4. B, Memecylon umbellatum Kostel. Petiole x 13. C,
Centradenia grandifolia Endl. Petiole x 13, D, Melastoma normale D. Don. Stem X 15. E, Gustavia
speciosaDC. Petiole x8. F, Memecylon umbellatum Kostel. Stem x 18. G, Tibouchina semidecandra
Cogn. Petiole X 19. H, Napoleona imperialis Beauv. Petiole x 8. I, Bredia hirsuta Blume. Petiole
x 15. ],BerthoIettia*xcelsaHumb.ttBonpl. Stem x 13. K, Napoleona imperialis Beauv. Stem X 17.
 MELASTOMACEAE 639

conspicuous intercellular spaces; very short to moderately long. Included

phloem of the 'foraminate* type present in 6 genera of the Astronioideae and
Memecyleae.
LEAF
Dorsiventral or centric; glabrous or hairy. Hairs (Fig. 144) consisting for
the most part of somewhat variable and complex types, such as those men-
tioned below, but these and other modifications also occur in many other
genera besides those mentioned by name. The more complex hairs are often
partly or wholly sclerenchymatous or even contain vascular bundles. The
relatively simple types described below under (i) and (ii) are comparatively
rare. Glandular hairs widely distributed.
A. Non-glandular, (i) Unicellular or resembling papillae in species of
Kibessia, Macairea, Marcetia, Miconia. (ii) Uni- or biseriate, with thick walls
and I-shaped lumina in Trembleya sp.; other uniseriate types recorded in
Sonerila and Veprecella. (iii) With multiseriate stalks and terminal cells

arranged like the hairs of a paint-brush in Tococa sp. (Fig. 144 B).
(iv) Candelabra hairs in Dichaetanthera, Dissochaeta, Dissotis, Kibessia,
Marumia, Medinilla (Fig. 144 D), Meriania, Omphalopus. (v) Stellate, with
short stalks in Miconia sp. (vi) Peltate in Astronia. (vii) Shaggy hairs as in
Fig. 144 E, i, and K.
B. Glandular, (i) Shaggy hairs, resembling some of the non-glandular
types described above, but with multicellular, glandular heads, e.g. in
Pyramia (Fig. 144 c). (ii) Small, with well-defined, multicellular, variously
shaped heads, (iii) Bladder-like glands, resembling those of the Labiatae, in
Blastus, Chaetostoma, Lavoisiera, Macairea, Microlicia, Trembleya.
According to de Cordemoy (468) the emergences on the trees and shrubs
belonging to the Osbeckieae consist of epidermal and cortical tissue, the
epidermis of the emergences being smooth in some but provided with conical
papillae in other species of Dichaetanthera. Cortical tissue of the emergences
often more or less lignified. Transitions between emergences and true hairs
also occur, e.g. in Tristania.
Cork warts recorded on the lower surface of the leaf in Pachyloma sp.
Epidermis sometimes consisting of gelatinous cells with wide lumina;
papillose on the upper surface in certain species of Aciotis, Allomorphia,
Bertolonia, Clidemia, Conostegia, Heterotrichum, Leandra, Marcetia, Medinilla,
Miconia, Monochaetum, Opisthocentra, Rhynchanthera, Sonerila] and on the
lower surface of species of Bellucia, Dichaetanthera, Dissochaeta, Henriettea,
Kibessia, Mecranium, Miconia, Pternandra, Tetrazygia; locally consisting of
2 or more layers on the upper surface of species of Comolia and Miconia',
wholly or locally palisade-like on the upper side in species of Calycogonium,
Clidemia, Leandra, Miconia, Ossaea, Tetrazygia\ and on the lower side in
species of Miconia. Hypoderm recorded in species of Anplectrum, Astronia,
Blakea, Calycogonium, Charianthus, Clidemia, Conostegia, Dichaetanthera,
Dissotis, Graffenrieda, Henriettea, Henriettella, Kendrickia (specially thick),
Loreya, Macairea (especially above the vascular bundles of the veins), Mar-
cetia, Marumia, Mecranium, Medinilla (occupying half the thickness of the
lamina in some species), Meriania, Miconia, Microlepis, Mouriria, Ochthocharis,
Pachyloma, Sonerila, Tetrazygia, Tibouchina.
 FIG. 144. MELASTOMACEAE
A, Clustered crystal of Centradenia floribunda Planch. B, Brush-like shaggy hair from the leaf of
Tococa sp. C, Shaggy hair of candelabra type with glandular head in Pyramid. D, Candelabra
hair of Medinilla sp. E, Shaggy hair on the leaf" of Tibouchina aemula Cogn. with root-like base to the
hair. F, Section through a bundle of sclerenchyma-fibres (hair-base), running beneath the epidermis
of the leaf, as a prolongation of the hair in Tibouchina chamaecistus Cogn. G, Section through the base
of a shaggy hair, adpressed to the leaf in the same plant. H, Base of a shaggy hair in surface-view in
the same species, i, Shaggy hair of Tibouchina decemcostata Cogn., sending out several bundles of
sclerenchyma-fibres at the base, which run subepidermally. K, Scale-like shaggy hair of Tibouchina
mathaei Cogn. A, C. and K after Pflaum, B and D after Lignier; the remainder by Solereder.
 MELASTOMACEAE 641
Stomata very variable in size, present on both surfaces or confined to the
lower side; surrounded by ordinary (often 3) epidermal cells in mature leaves
(ranunculaceous to cruciferous); often accompanied by caryophyllaceous
types in species of Bertolonia, Marcetia, Medinilla, Microlicia, Oxyspora,
Sonerila, and other closely related genera; rubiaceous in Memecylon, Miconia,
and related genera, Stomata sometimes with their pores parallel to one
another and to the median vein of the leaf in Dissotis, Microlicia, Osbeckia,
Pterolepis\ confined to variously shaped pits with wide or narrow openings in
most species of Mouriria (Fig. 138 c); arranged in groups on the inside, and
more rarely on the outside, of ant domatia in species of Myrmidone and
Tococa, as well as on the lower surface of the leaf in species of Calycogonium,
Leandra, Ossaea. MesophylL Palisade tissue consisting of 2 or 3 layers of
interlocking cells in species of Bellucia\ i -layered in some of the species of
Medinilla examined by de Cordemoy (472); including isolated cells with
reticulate thickening in species of Graffenrieda and Meriania and sclerosed
cells in species of Medinilla. Cells of the spongy mesophyll provided with
local thickenings, recalling those of certain Menispermaceae, in species of
Anplectrum, Graffenrieda, Medinilla, Melastoma, Meriania, Ochthocharis,
Pachyloma. Spiral tracheids recorded in the mesophyll of a few species of
Adotis, Bellucia, Henriettea, Sonerila, and sclerenchymatous cells of various
kinds in species of Gravesia (de Cordemoy 471), Henrietella, Huberia,
Lavoisiera, Leandra, Macairea, Medinilla, Memecylon, Miconia, Microlicia,
Mouriria, Ossaea, Plethiandra\ those of Memecylon and Mouriria stated by
Solereder to be of considerable specific diagnostic value. (The more detailed
statements on this subject recorded by Solereder need confirmation.) In this
connexion, reference must be made to the recent interesting study by Foster
(697) of the foliar sclereids in Mouriria. Foster observed that the range of
form exhibited by these elements in different species of Mouriria is quite
considerable, and he recognized 4 main types. The sclereids were found to
differ from those of many other genera in which they have been studied, in

being associated with the terminations of the veins, rather than lying in the
mesophyll independently of the vascular system. This raises the question of
whether they are homologous with vascular elements. From the taxonomic
standpoint, Foster concluded that the sclereids provide a good generic charac-
ter for Mouriria, but feels that their value in the identification of species
cannot, at present, be fully assessed. Nevertheless he lists species in which
each of his 4 types of sclereid occur, but issues a warning that *a number of
the entities used in the present survey may subsequently prove to have been
incorrectly determined*. Vascular bundles of the veins more often accom-
panied by collenchyma than by well-developed sclerenchyma; the latter,
when present, sometimes situated below the epidermis and not in direct con-
tact with the vascular bundles in species of Chaetostoma, Lavoisiera, Micro-
lepis, Microlicia. Vertically transcurrent veins recorded in Charianthus,
Clidemia, Henriettea', Leandra, Miconia, Tetrazygia, Tococa. Elongated rod
cells said to occur in the parenchymatous ground tissue of the veins in a
number of genera.
Petiole (Fig. 143 A-C, G, and i). Transverse sections through the distal end
in nearly all of the species examined exhibit an arc of widely spaced vascular
bundles, frequently accompanied by subsidiary strands, the latter often being
4594 T t
642 MELASTOMACEAE
very numerous and usually situated towards the adaxial side. Vascular struc-
ture quite complex in some species. The petiolar vascular structure would
be of considerable diagnostic value if better known. Ground tissue of the
petiole seen to be supported by scattered stone cells in species of Medinilla
and Miconia.
Crystals usually clustered (Fig. 144 A, F, and H), those in the mesophyll
sometimes especially large and appearing as transparent dots in the leaf.
Styloids occur, either alone or together with other types of crystal, in Astronia,
Beccarianthus, Belinda, Calycogonium, Centradenia, Henriettea, Henriettella,
Kibessia, Loreya, Memecylon, Mouriria, Myriaspora, Pternandra.
Developmental anatomy. The developmental anatomy of the leaf of
Heterotrichum macrodon Planch, and Clidemia hirta Don. has been described
by Weidt (2385).

Axis
YOUNG STEM (Fig. 143 D and F)

Frequently winged and provided with vascular bundles in the wings, e.g. in
Centradenia, Medinilla, Tibouchina. Hairs similar to those on the leaf also
occur on the stem. Cork arising superficially in Adelobotrys, Astronia,
Beccarianthus, Bertolonia, Blakea, Blastus, Boerlagea, Bredia, Calvoa,
Calycogonium, Carionia, Charianthus, Clidemia, Conostegia, Dalenia, Diplarpea,
Henriettea, Henriettella, Heterotrichum, Kendrickia, Loreya, Mecranium,
Medinilla (pro parte), Medinillopsis, Miconia, Microphysa, Myriaspora, Ossaea,
Oxyspora, Pachyanthus, Pachycentria, Platycentrum, Plethiandra, Salpinga,
Tetrassygia, Topobea; originating in the pericycle in Amphorocalyx, Anerin-
cleistus, Benevidesia, Calophysa, Calyptrella, Centronia, Kibessia, Meriania,
Microlicia, Mouriria, Ochthocharis, Octopleura, Osbeckia, Sagraea, Sakersia,
Tibouchina; originating in the inner part of the cortex in at least i species
of Melastoma (Fig. 143 D). Cork cells with i -sided thickenings on the inner
tangential walls in Astronia, Dichaetanthera, Henriettea, Medinilla (pro parte)
(de Cordemoy 472), Mouriria. Cork homogeneous in Dissotis; consisting of
alternating layers of flattened and tall cells respectively in Barbeyastrum,
Comolia, Heeria, Lavoisiera, Marcetia, Microlicia, Osbeckia, Pterolepis,
Tibouchina. Cork sometimes replaced by aerenchyma originating in the
pericycle of submerged parts of the stem and plder roots of marsh plants
included in Acisanthera and Rhynchanthera. Primary cortex containing
stone cells, sometimes arranged in groups, in species of Adelobotrys, Axinaea,
Barthea, Blakea, Blastus, Centronia, Graffenrieda, Medinilla (de Cordemoy
472), Meriania, Miconia, Myriaspora, Ossaea, Oxyspora, Pachyanthus, Pachy~
centria, Pogonanthera, Tetrazygia; stone cells sometimes arranged in a ring
in Conostegia, Dissochaeta, Mecranium, Memecylon, Meriania, Myriaspora,
Omphalopus. Fibres observed in the cortex of a species of Melastoma.
Endodermis usually well defined ; in many species composed of thin- walled
cells, but stated to include cells with U-shaped thickenings in species of
Anerinckistus, Calophysa, Dichaetanthera, Kibessia, Memecylon, and
Mouriria seen to possess casparian thickenings in Sonerila\ suberized in
i

Gravesia and Medinilla, according to de Cordemoy (472). Pericycle never

provided with a composite and continuous ring of sclerenchyma, but contain-
 MELASTOMACEAE 643

ing a variable proportion of fibres or thickened cells in species of Blakea,

Calyptrella> Centronix Creochiton, Henriettella, Kibessia, Loreya, Memecylon,
Meriania^ Plethiandra, Topobea.
Phloem and xylem, in the genera represented in the Kew slide collection,
in the form of continuous cylinders traversed by narrow rays. Vessels with
simple perforations. Pith very broad in many genera; consisting of lignified
or unlignified cells, sometimes including stone cells or scattered fibrous
cells.

Intraxylary phloem (Fig. 143 D and F) universally present in the very

considerable numberof investigated species, including many not mentioned
by name in the text; intraxylary phloem somewhat reduced in Gravesia
according to de Cordemoy (471). Interxylary phloem recorded in Kibessia,
Memecylon (Fig. 143 F) (in the root as well as in the stem), Mouriria, Pter-
nandra. (See also under 'Included Phloem* on p. 647.) Cortical and/or
medullary bundles, often with central xylem, but the latter sometimes very
much reduced or absent, occur in many but not all genera and species; their
number and distribution varying even within a single species. Cortical
bundles most frequent in species with rectangular stems; said to represent
the lateral strands of the leaf trace, or, more rarely, the midrib itself. The
medullary bundles stated to be continuous with the intraxylary phloem. The
distribution of medullary and cortical bundles is of some value for the diagnosis
of tribes and genera. (See table below and Solereder for further details.)

* Recorded in 3 or more Recorded only in i or 2 species in genera not

species.
marked with a star.
 644 MELASTOMACEAE

* Recorded in
3 or more species. Recorded in only i or 2 species in genera not
marked with a star.

Secretory cells, with unidentified, amorphous contents, fairly common

in unlignified tissues, especially in the cortex and phloem, but varying in
frequency in different species, de Cordemoy (468) refers to a system of
tanniniferous elements in the phloem of certain species of Dichaetanthera.
Crystals almost exclusively clustered in the species represented in the Kew
slide collection, solitary types seen only in Memecylon\ frequently abundant,

especially in the cortex or phloem. Cristarque cells (i.e. cells with cellulose
thickenings to the inner walls, each cell containing a solitary crystal), recorded
by de Cordemoy (470) in the exodermis of certain species belonging to the
Osbeckieae.

WOOD (Fig. 145)

Vessels mostly small (less than 100 mean tangential diameter), very small
/LC

(50-100 fi) in Brachyotum, Calycogonium, Henriettella, Leandra, Mecranium,

and Mouriria p.p., medium-sized (100-200 p) in some species of Astronidium,
Centronia, Conostegia, Dactylocladtis, Dissochaeta (2158), Henriettea, Huberia,
Meriania, Miconia y and Pternandra\ usually with numerous small radial
multiples and commonly with irregular clusters, though occasionally tending
to be mostly solitary insome species of Memecylon, e.g. M. edule Roxb. and
M .
subfurfuracewn Merr. multiples of 4 or more cells moderately common in
;

Graffenrieda, Huberia^ Miconia p.p., and Tococa, clusters most pronounced

in Brachyotum^ Henriettella, Heterotrichium> Mecranium, Melastoma, Miconia

p.p., Rhynchanthus, and Tibouchia\ with a tendency to a tangential pattern in

Memecylon p.p. and Rhynchanthera, and a radial or flame-like pattern in
Mouriria, e.g. in some specimens of M. parviflora Benth. and M. pseudo-
geminata Pitt. 4-80, mostly 7-25 per sq. mm., seldom giving the appearance
;

of being crowded even when numerous, less than 5 per sq. mm. in Conostegia,
Dactylocladus, and Meriania, 20-40 per sq. mm. int Aciotis, Astronia, Bellucia,
 MELASTOMACEAE 645

Brachyotum, Dissotis, Henriettella, Huberia, Medinilla, and Memecylon p.p.,

more than 40 per sq. mm. in Calycogonium, Clidemia, Leandra, Mecranium,
Memecylon p.p., Mouriria p.p., and Rhynchanthera p.p. Perforations exclusively
simple. Intervascular pitting typically alternate, but scalariform in Astronia,
Blastus (1206), and Medinilla, commonly very small to minute, e.g. in Bellucia,
Calycogonium, Clidemia, Henriettea, Henriettella, Heterotrichium, Leandra,
Mecranium, Memecylon, Miconia, Mouriria Ossaea, Pachyanthus, Tetrazygia,
,

and Tococa, moderately large in Dactylocladus, Huberia, Meriania, and

Tibouchina', occasionally with coalescent apertures, e.g. in Macairea and
Memecylon; pits to parenchyma simple and elongated, with their long axes
horizontal, oblique, or axial, in Astronia, Astronidium, Brachyotum, Caly-
cogonium, Calyptrella, Centronia, Conostegia, Dissotis, Graffenrieda, Henriettea,
Huberia, Loreya, Macairea, Mecranium, Medinilla, Melastoma, Meriania,
Pternandra (pits almost round), Rhynchanthera, Tibouchina, and Tococa\
similar to the intervascular pitting in the other genera, which include all those
of the Memecyleae. Usually without solid deposits; Williams (2430) refers
to calcium deposits in Miconia and Tococa; tyloses observed or reported in
Astronia, Huberia, Kibessia, Medinilla, Pternandra, Rhynchanthera, and Tococa,
those of Medinilla sclerosed. Mean member length 0-3-0*8 mm. Paren-
chyma in most of the genera paratracheal only and varying from a few cells

(scanty) to a complete sheath round the vessels (vasicentric) ; variable and

difficult to classify in the other genera, but some paratracheal always present;

aliform, with blunt or long uniseriate extensions from the vessels in the
Memecyleae, i.e. in Dactylocladus (Fig. 145 A), Memecylon p.p., and Mouriria
p.p.; in narrow, apparently apotracheal bands 1-2 cells wide in Clidemia^
Mouriria p.p., and Tococa\ in broader, regular to wavy bands in Calyptrella,
Graffenrieda, Kibessia, and Meriania (Fig. 145 i); in patches or discontinuous
bands, suggesting fragments of the broad bands described above and some-
times comparable in distribution to the groups of septate fibres or included
phloem described below, in Conostegia, Ossaea, Pachyanthus, and Tibouchina
(Fig. 145 G), often with conspicuous intercellular spaces; diffuse parenchyma
present in Kibessia, Memecylon, and Mouriria (Fig. 145 H) and, according to
Janssonius (1154), in i species of Melastoma', with apparently terminal bands
in some species of Dissotis and Mouriria. Seldom with gummy contents and
crystals not observed. Strands mostly of 4-6, occasionally 2 or 8, cells;
Janssonius ( 1 154) notes occasional fusiform cells in Kibessia azurea DC. Rays
of the Melastomoideae typically exclusively uniseriate or with only local
biseriations, and composed of upright and square cells, with very few or no
truly procumbent cells; occasionally with moderately numerous procumbent
cells (Kribs's Type Heterogeneous III), e.g. in Astronidium and Pachyanthus,
and sometimes composed almost entirely of procumbent cells, e.g. in Axinaea
(1886) and Conostegia. In Centronia, Macairea, and Medinilla some of the
rays are multiseriate (2-4 cells wide), but they are essentially similar to the
uniseriate type described above, being composed mainly of square or upright
cells and presenting in tangential section a loose mixture of cells of all sizes
and shapes (Fig. 145 D); multiseriate rays also present, according to Record
and Hess (1886) in Blakea and Topobea. In the Astronioideae and Memecy-
leae the rays are up to 2-5 cells wide in Kibessia^ Memecylon, Mouriria p.p.,
and Pternandra, and except in Mouriria, tend to be of 2 distinct sizes; the
646 MELASTOMACEAE
larger rays with a compact central portion of small procumbent cells and
1-3 marginal rows of square or upright cells, the uniseriate rays low and often
only i cell high; in Pternandra the central portion of the larger rays is largely
disintegrated in dry specimens, but appears to consist of tissue similar to that

/D

FIG. 145. MELASTOMACEAE

A, Dactylocladus stenostachys Oliv. B, D, stenostachys Oliv. C, Mouriria pseudogeminata Pittier.
D, Centronia excelsa Triana. E, Meriania urceolata Triana. F, Centronia excelsa Triana. G Tibouchina
lepidota Baill. H, Mouriria pseudogeminata Pittier. I, M.eriania urceolata Triana.

of the axial strands of included phloem; in Mouriria the rays, whether multi-
seriate or wholly uniseriate, as in M. parviflora Benth. and M. cyphocarpa
Standl., are similar to those described above for the Melastomoideae, as are
also the wholly uniseriate rays of Astronia and Astronidium\ in Dactylocladus
the rays, though uniseriate only, differ from those of the Melastomoideae,
except Axinaea and Conostegia, being composed entirely of procumbent cells
apart from an occasional marginal row of square cells. Usually with 7-25 rays
per mm.; species with only uniseriate rays mostly with 13-17 per mm., fewer
in Brachyotum and Meriania and more than 17 per mm. in some species of
 MELASTOMACEAE 647
Atiotis, Clidemia, and Miconia\ species with multiseriate rays mostly with
7-13 per mm., but up to 16 in some species, e.g. of Memecylon. Very com-
monly with gummy contents, which often fill the cells crystals not observed.
;

Fibres with distinctly bordered pits, usually equally numerous on both radial
and tangential walls, in most of the genera of the Astronioideae and Memecy-
leae, i.e. in Dactylocladus, Kibessia, Memecylon, Mouriria, and Pternandra,
and with thick walls except in Dactylocladus\ with simple pits, thin walls and
septa in Astronia and Astronidium. In the Melastomoideae: pits simple and
usually more numerous (often in more than i row) on the radial walls;
Solereder refers to narrow borders in Sonerila elegans Wight commonly septate
;

in Belinda, Blastus (1206), Calycogonium, Henriettea, Henriettella, Hetero-

trichium, Huberia, Leandra, Macairea, Mecranium, Medinilla, Melastoma,
Miconia, Ossaea, Pachyanthus, Tetrazygia, Tibouchina, and Tococa\ with
occasional fine septa in most of the other genera, but septa not observed or
very rare in Brachyotum, Calyptrella, Graffenrieda^ Meriania, Rhexia (2158),
and Rhynchanthera. Thinner-walled patches or bands of fibres, with con-
spicuous intercellular spaces, very characteristic the septate fibres sometimes
;

limited to these groups; Janssonius (1154) states that such septate fibres in
Astronia, Medinella, and Melastoma are shorter than the surrounding non-
septate fibres. In Tibouchina septate fibres occur in similar groups, which,
however, consist mainly of parenchyma cells; walls usually moderately thin
and often with a mucilaginous layer, thick in Meriania; commonly in very
distinct radial rows. Mean length 0-5-1-2 mm. Included (interxylary)
phloem of the 'foraminate' type, with isolated strands of included phloem
given off externally by the cambium (2158), present in some genera of the
Astronioideae and Memecyleae, i.e. in Kibessia, Lijndenia, Memecylon,
Mouriria (Fig. 145 H), Olisbea (1851), and Pternandra.

ANOMALOUS STRUCTURE
See last paragraph and Inter- and Intraxylary Phloem under 'Young Stem*.

ROOT
Tuberous roots, which occur in certain species of Medinilla, stated by de
Cordemoy (470, 472) to consist mainly of secondary phloem largely composed
of parenchyma.

TAXONOMIC NOTES
FROM WOOD STRUCTURE
The woods of the Astronioideae and Memecyleae are distinguishable from
those of the Melastomoideae, particularly on account of their fibre-tracheids,
included phloem, and multiseriate rays. Astronia and Astronidium, however,
are exceptions, being indistinguishable from the Melastomoideae. F. Mark-
graf (1442) considers these 2 genera to be separate and, though a direct
comparison of the woods has not been possible, from descriptions (376) there
appear to be some differences, e.g. in type of perforation.
Included phloem occurs only in the Astronioideae and Memecyleae, but
is not always present. The following genera have normal wood: Astronia,

Astronidium^ Axinandra, Beccarianthus, Dactylocladus, and Plethiandra,

Janssonius (1154) considers that Kibessia, Memecylon, and Mouriria should
 648 MELASTOMACEAE
be treated as a separate family between the Melastomaceae and the Myrtaceae.
Pternandra (not investigated by Janssonius) would fall into the same group
and possibly also Dactylocladus, though this differs in some respects.

ECONOMIC USES
No
products of great economic importance are derived from this family,
but a number of species are commonly cultivated for ornamental purposes.

GENERA DESCRIBED
(i) FOR GENERAL ANATOMY
Acanthella, Aciotis, Acisanthera, Adelobotrys, Allomorphia, Amphiblemma,
Amphorocalyx, Anaectocalyx, Anerincleistus, Anplectrum, Antherotoma,
Appendicularia, Arthrostema, Astronia, Axinaea, Axinandra, Barbeyastrum,
Beccarianthus, Behuria, Bellucia, Benevidesia, Bertolonia,* Bisglaziovia,
Blakea, Blastus, Boerlagea, Brachyotum, Bredia,* Brittenia, Bucquetia,
Calvoa, Calycogonium, Calyptrella, Cambessedesia, Carionia, Castratella,
Catocoryne, Centradenia,* Centronia, Chaetolepis, Chaetostoma, Charianthus,
Clidemia,* Comolia, Conostegia, Creochiton, Dalenia, Desmoscelis, Dicel-
landra, Dichaetanthera, Dinophora, Diolena, Dionychia, Diplarpea, Disso-
chaeta, Dissotis, Driessenia, Ernestia, Fritzschia, Graffenrieda, Gravesia,
Guyonia, Heeria, Henriettea, Henriettella, Heterotrichum, Huberia, Ken-
drickia, Kibessia, Lasiandra, Lavoisiera, Leandra, Lithobium, Loreya,
Macairea, Macrocentrum, Maieta, Mecranium, Medinilla,* Medinillopsis,
Melastoma,* Memecylon,* Meriania, Miconia,* Microlepis, Microlicia,
Microphysa, Monochaetum, Monolena, Mouriria, Myrmidone, Nepsera,
Ochthocharis, Omphalopus, Opisthocentra, Osbeckia, Ossaea, Otanthera,
Oxyrneris, Oxyspora, Pachyanthus, Pachycentria, Pachyloma, Phornotham-
nus, Phyllagathis, Platycentrum, Pleiochiton, Plethiandra, Pogonanthera,
Poteranthera, Pternandra, Pterocladon, Pterogastra, Pterolepis, Purpurella,
Pyramia, Rhexia, Rhodosepala, Rhynchanthera, Rousseauxia, Sagraea,
Sakersia, Salpinga, Sarcopyramis, Schwackaea, Siphanthera, Sonerila,*
Stenodon, Svitramia, Tetrazygia, Tibouchina,* Tococa, Topobea, Trem-
bleya, Triolena, Tristemma, Tulasnea.
* Kew
Represented in the slide collection.

(ii)
FOR WOOD STRUCTURE
Melastomatoideae: Aciotis, (Axinaea), Bellucia, (Blakea), (Blastus),
Brachyotum, Calycogonium, Calyptrella, Centronia, Clidemia, Conostegia,
(Dissochaeta), Dissotis, Graffenrieda, Henriettea, Henriettella, Hetero-
trichum, Huberia, Leandra, Macairea, Mecranium, Medinilla, Melastoma,
Menendezia, Meriania, Miconia, Ossaea, Pachyanthus, (Rhexia), Rhyn-
chanthera, (Sonerila), Tetrazygia, Tibouchina, Tococa, (Topobea). Astro-
nioideae: (Astronia), Astronidium, Kibessia, Pternandra. Memecyloideae:
Dactylocladus, (Lijndenia), Memecylon, Mouriria, (Olisbea).

LITERATURE
(i) On General Anatomy
de Cordemoy 468, 470, 471, 472, Foster 697, Ross 1959, Weidt 2385.
 MELASTOMACEAE 649
(ii) On Wood Structure
Bailey 78, Benoist 170, den Berger 182, Burgerstein 310, Chennery 2515, Chalk and
Chattaway 362, 376, Cooper and Record 461, Howard 1088, Janssonius 1154, Kanehira
1206, 1209, Kribs 1283, Markgraf 1442, Pfeiffer, H. 1712, Pfeiffer, J. Ph. 1713, Record
1843, 1851, Record and Hess 1886, Record and Mell 1894, Williams 2430.

139. LYTHRACEAE
(Fie. 146 on p. 650; FIG. 147 on p. 652; FIG. 148 on p. 658)

SUMMARY
(i) GENERAL
The most important feature common to the trees, shrubs, and herbs
belonging to this widely distributed family is the occurrence of intraxylary
phloem. Where the vascular system is in the form of bundles, e.g. in the
leaf, these are bicollateral. The hairs are mostly unicellular or bicellular,

although more complex types also occur. Mucilaginous cells are fairly
common in the leaf epidermis and occasionally elsewhere in the lamina. The
;

somewhat variable stomata are ranunculaceous in most species. Secretory

cavities are absent except from i species of Cuphea and in Heteropyxis, but
the last genus ought probably to be excluded from the Lythraceae. The cork
in the stem, which generally arises in the inner part of the pericycle, is often
somewhat loosely constructed, and may be replaced or accompanied .by
aerenchyma. Interxylary phloem recorded in the root of Lythrum salicaria
Linn.

(ii) WOOD
Vessels small to medium-sized, ring-porous in some species, perforations
simple, intervascular pitting alternate, small to medium-sized, vestured, pits
to parenchyma similar or large and simple; members of medium length to
moderately short. Parenchyma predominantly paratracheal, scanty or
vasicentric to aliform and confluent, sometimes with numerous septate
crystalliferous cells. Rays exclusively uniseriate or up to 2-3 cells wide,
heterogeneous to homogeneous. Fibres with simple pits and commonly
septate, thin-walled fibres containing starch or crystals sometimes occurring
with a parenchyma-like distribution ;
of medium length to moderately short.

LEAF
Usually dorsiventral, Adenaria, Ammannia, Crenea, Cuphea, Ginora,
e.g. in
Lafoensia, Lagerstroemia, Lawsonia, Nesaea, Physocalymma, Woodfordia; iso-
bilateral in Pemphis acidula Forst ; homogeneous in Peplis according to Gin

(785). Hairs (Fig. 146) include the following types, the first being
the most
widely distributed, (i) Unicellular or bicellular, elongated in some but short
and resembling papillae in other species of Adenaria, Cuphea, Decodon,
Diplusodon, Ginora, Lagerstroemia, Lythrum, Nesaea, Pemphis,
Grislea,
Peplis, Physocalymma, Pleurophora, Woodfordia. (ii) Simple, unicellular,

placed obliquely or almost parallel to the surface, e.g. in Pemphis, tending to

be 2-armed and provided with thickened silicified walls with warts in
Cuphea (Fig. 146 c). (iii) Tufted or branched and provided with uniseriate
 650 LYTHRACEAE
stalks in Decodon and Lagerstroemia. (iv) Glandular and non-glandular shaggy

types in Cuphea (Fig. 146 A~B). (v) Spherical glandular hairs, provided with
very short stalks, situated in pits and appearing as black dots on the lower
side of the leaf, in species of Adenaria, Grislea, Lagerstroemia, Pemphis,
Woodfordia (Fig. 146 D).

FIG. 146. LYTHRACEAE

A, B, Shaggy hair of Cuphea lanceolata Dryand. in Ait. Hort. Kew., with glandular base; the latter
cut transversely in B. C, Trichome of Cuphea appendiculata Benth. D, Section of an external gland
of Woodfordia florilfunda Salisb. A, B, after Martinet, C, D, by Solereder.

Epidermis containing mucilaginous cells in species of Adenaria, Ammannia,

Cuphea, Diplusodon, Ginora, Lafoensia, Lagerstroemia, Lawsonia, Lythrum,
Nesaea, Pemphis, Peplis, Woodfordia, and probably other genera as well.
Mucilaginous cells also occur in other parts of the lamina of Diplusodon,
Lagerstroemia, Nesaea, Pemphis, &c. Cells of the upper epidermis sometimes
horizontally divided in Ginara, Lagerstroemia, Pemphis occasionally papillose.
\

Hypoderm recorded in Ginora and Lagerstroemia. Stomata confined to the

lower surface or present on both sides, usually ranunculaceous pores approxi- ;

mately parallel to the main vein in species of Diplusodon, Heimia, Lythrum,

Nesaea, Pleurophora. Mesophyll of species with dorsiventral leaves con-
taining 1-3 layers of palisade cells.
Vascular bundles of the veins bicollateral. Smaller veins vertically trans-
 LYTHRACEAE 651
current in species of Decodon, Grislea, Lagerstroemia, Physocalymma\ those
of Pemphis acidula Forst. said by Kienholz (1236) to be devoid of scleren-
chyma and embedded in aqueous tissue. Storage and terminal tracheids
occur in the form of large pitted cells in species of Crenea, Diplusodon,
Lawsoriia. Petiole (Fig. 148 D). Transverse sections through the distal end
of the few species represented in the Kew slide collection exhibit a solitary,
slightly curved, crescent-shaped, bicollateral, vascular strand, sometimes
accompanied by small, circular, accessory bundles in the wings. Secretory
receptacles, appearing as transparent dots, recorded by Solereder in Cuphea
anagalloidea St. Hill, and in Heteropyxis. (The taxonomic position of Hetero-
pyxis is uncertain and it has been variously included in the Lythraceae,
Myrtaceae, and in a special family the Heteropyxidaceae.) Mucilage cells,
see 'Epidermis'. Crystals mostly clustered; abundant solitary crystals
recorded in association with the veins in Lagerstroemia. Small crystalline
rods, needles, or granules recorded in Adenaria, Decodon, Diplusodon, Heimia,
Lafoensia, Lawsonia, Lythrum, Nesaea, Physocalymma, Pleurophora, Wood-
fordia. Small clustered crystals situated in groups of special mesophyll cells
occur in Pemphis. Clustered or large solitary crystals stated to occur in
idioblasts and sometimes to appear as transparent dots in species of Diplusodon,
Galpinia, Lafoensia, Lagerstroemia, Nesaea. Sphaerocrystalline masses of
unknown chemical nature recorded in certain species of Decodon, Diplusodon,
Heimia, Lagerstroemia, Lythrum. Anatomical differences in the anatomy of
Cuphea balsamona Cham, et Schl. collected at different altitudes have been
described by Privault (1759). Bodmer (212), in the course of a very detailed
investigation of the developmental anatomy of Lythrum salicaria Linn., found
that submerged leaves are much thinner than those on the aerial part of the
stem, but the stomatal frequency was similar in both types of leaf.

Axis
STEM (Fig. 148 A)
Young stem commonly with 4 or 5 prominent angles, the latter sometimes
enlarged to form wing-like expansions. Cork generally arising in the inner
part of the pericycle in species of Ammannia, Crenea, Cuphea, Diplusodon,
Ginora, Heimia, Lagerstroemia, Lawsonia, Lythrum, Nesaea originating in the
;

primary cortex in species of Lafoensia (Fig. 148 A) and Pemphis. Cork cells
very small in species of Ammannia, Crenea, Lawsonia, Lythrum, Nesaea, Peplis.
Cork many-layered in species of Pemphis and Pleurophora partly composed
;

of phelloid cells in a number of genera; sometimes replaced or accompanied

by aerenchyma in Crenea, Cuphea, Decodon (forming a white spongy tissue
on the aerial stems of D. verticillatus Ell. according to Gin (785)), Heimia^
Lythrum, Nesaea, Peplis. Primary cortex described by Gin (785) as con-
taining a system of intercellular spaces in species of Ammannia, Cuphea^
Heimia, Lythrum, and 4 large air canals in Crenea repens G. F. W. and various
species of Peplis including P. portula Linn. Endodermis with casparian
thickenings recorded by the same author in species of Adenaria, Cuphea,
Lagerstroemia, and Peplis portula composed of cells with uniformly thickened
;

cellulose walls in other species of Lagerstroemia] cells thickened on the internal

and lateral walls in other species of Lagerstroemia, in Nesaea salicifolia H. B.
et K. and Physocalymma scaberrimum Pohl. Sclerenchyma poorly developed
 I
i

FIG. 147. LYTHRACEAE, A-F; OLINIACEAE, G and J; CRYPTERONIACEAE, H-I;

PUNICACEAE, K-M
A, Lagerstroemia flos-reginae Retz. B, L. flos-reginae Retz. C, L. ovalifola Teysm. D, Pemphis
acidula Forst. E, Lafoensia sp. F, Pemphis acidula Forst. G, Olinia usambarensis Gilg. H, Crypteronia
paniculata Blume. I, C. paniculata Blume. J, Olinia cymosa Thunb. K, Punica granatum Linn.
Crystalliferous fibre ( x 200). L, P. granatum Linn. M, P. granatum Linn.
 LYTHRACEAE 653
in or absentfrom the peri cycle, but isolated strands of fibres recorded in
Cuphea, Lafoensia, Lagerstroemia, Lythrum, or occasionally a sclerenchy-
matous ring in Ammannia, Diplusodon, Lafoensia, Lagerstroemia. Phloem
sometimes including concentrically arranged crystal cells. Secondary phloem
containing fibres in Lafoensia and Lagerstroemia. Sclerosed cells also noted in
the secondary phloem in a few species of Cuphea and Lafoensia by Gin (785).
Phloem and xylem in the form of continuous cylinders traversed by narrow
rays in of the species represented in the Kew slide collection. Vessels
all

with simple perforations. Xylem of Cuphea annulata Koehne said by Gin

(785) to appear, in transverse sections, as 2 equal arcs separated by unlignified
parenchyma.
Intraxylary phloem recorded or observed in Adenaria, Ammannia, Crenea,
Cuphea, Decodon, Diplusodon, Ginora, Heimia, Lafoensia (none clearly visible
in the specimen illustrated in Fig. 148 A), Lagerstroemia, Lawsonia, Lythrum,
Nesaea, Pemphis, Peplis, Physocalymma, Pleurophora, Rotala, Woodfordia, and
probably occurring throughout the family. Pith described by Solereder as
strongly sclerosed in Diplusodon and Lagerstroemia including large, sclerosed
;

cells in Adenaria sp. and Pemphis acidula Forst. Crystals mostly clustered
and frequently abundant, especially in the pith and cortex; solitary types
observed in Lagerstroemia (see also under 'Phloem' above). Secretory cells
with unidentified, amorphous contents, present in the unlignified tissues of
most of the genera examined at Kew.

WOOD (Fig. 147 A-F)

Vessels small(less than 100 //,
mean tangential diameter) to medium-sized
(100-200 fj)', solitary and in multiples of 2 or 3 cells; varying from about
5 per sq. mm. in some species of Lagerstroemia to about 20 per sq. mm. in
Pemphis ring-porous in most species of Lagerstroemia. Perforations simple.
;

Intervascular pitting alternate, small to moderately large; vestured (78); pits

to ray or wood parenchyma usually similar to the intervascular pitting, but
occasionally larger or simple, e.g. in Lagerstroemia. Sometimes with deposits
of gum and with tyloses in Lagerstroemia and Pemphis Pearson and Brown
;

(1679) note 2 types of tyloses in Lagerstroemia'. (a) thin- walled, large, and
cyst-like, and (b) small, sclerenchymatous, copiously pitted, and filled with
gum. Mean member length 0-25-0-5 mm. Parenchyma predominantly
paratracheal; abundant and aliform to confluent in Lagerstroemia (Fig. 147 A),
tending to be in broad bands in the outer part of the ring; scanty to vasi-
centric and diffuse in Pemphis', scanty paratracheal in most of the American
genera (1886), but reported to be abundant in Adenaria floribunda H. B. et K.
(2430). With conspicuous chambered crystals in Lagerstroemia, particularly
in cells touching the fibres, the parenchyma cells of normal height but sub-
divided by septa. Strands commonly of 4 cells. Rays exclusively uniseriate
in most species of Lagerstroemia (2-3 cells wide in most samples of L. flo$~
f

reginae Retz.) and in Lafoensia, 2 cells wide in Pemphis, i or 2, rarely 3 cells

wide* in the American genera (1886); rays rather low and with many very low
uniseriates only i or 2 cells high in all the material studied; about 7-17 rays
per mm.; homogeneous (Kribs's Type III) in Lagerstroemia and, according
to Record and Hess (1886), in Physocalymma; heterogeneous (Kribs's Types
II B and III) in the other genera. Commonly containing gummy deposits,
654 LYTHRACEAE
crystals not observed. References occur in the literature (1801, 2430) to the
occurrence of or mucilage cells in Physocalymma scaberrimum Pohl. but
oil
have since been shown to be incorrect (1830, 2430). Fibres with simple pits,
often septate, e.g. in Lafoensia and Lagerstroemia; walls usually rather thin,
but moderately thick in some species of Lagerstroemia, very thick and non-
septate in Pemphis. Record and Hess (1886) refer to thin- walled fibres with
large lumina in numerous distinct parenchyma-like bands in Ginora and
Physocalymma, the cells containing starch at first, but later crystalliferous,
and to similar starch-bearing, non-crystalliferous, indistinctly vasicentric
fibres in Lafoensia.Pearson and Brown (1679) record the occurrence in
Lagerstroemia of crystals in small locules separated by septa, particularly in
fibres contiguous with the rays. Mean length 0-8-1-3 mm.

ROOT
phloem, formed by secondary differentiation of the
Islands of interxylary
xylem parenchyma, recorded by Gin (785) in the root wood of Ly thrum
salicaria Linn.
TAXONOMIC NOTES
Sprague and Metcalfe (2175) concluded, partly on anatomical grounds,
that Rhynchocalyx should be regarded as a member of the Lythraceae.

ECONOMIC USES
Henna, which is used as a hair dye, consists of the powdered leaves of
Lawsonia alba Lam., sometimes mixed with other substances. The glabrous,
oval, somewhat acuminate leaves are 3-4 cm. long and 1-5-2-5 cm. broad.
The tabular epidermal cells are rectilinear or tend to be polygonal in surface
view. Some cells of the epidermis are larger than the remainder and mucilagi-
nous. Stomata numerous on both surfaces, surrounded by 3 or 4 ordinary
epidermal cells. Mesophyll dorsiventral, with 2 or more layers of palisade
tissue towards the upper surface, but locally isobilateral owing to a tendency
for the outer cells of the spongy mesophyll to develop as a second palisade.
A very detailed description of the history and uses of Henna is included in
Gin's (785) thesis. The same author also gives particulars of other members
of the family to which medicinal properties have been attributed.
The only genus producing timber of any importance is Lagerstroemia. Of
this there are several species in the East that produce very similar timbers,
which are suitable for high-class joinery work. The best known are those
furnishing the timber known as Pyinma, Lagerstroemia speciosa Pers. from
India and Burma and L. hypoleuca Kurz from the Andaman Islands.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Adenaria, Ammannia, Crenea, Cuphea,* Decodon, Diplusodon, Galpinia,
Ginora, Grislea, Heimia, Lafoensia,* Lagerstroemia,* Lawsonia, Lythrum,*
Nesaea,* Pemphis, Peplis, Physocalymma, Pleurophora, Woodfordia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Adenaria), (Ginora), (Grislea), Lafoensia, Lagerstroemia, Pemphis,
(Physocalymma).
 LYTHRACEAE 655
LITERATURE
(i) On General Anatomy
Bodmer 212, Gin 785, Kienholz 1236, Lourteig 1394, Privault 1759, Sprague and
Metcalfe 2175.

(ii) On Wood Structure

Beekman 167, den Berger 179, Besson 186, Collardet 447, Coster 481, Desch
Bailey 78,
574, Foxworthy 705, Gin 785, Howard 1088, Janssonius 1154, Jones 1191, Kanehira 1206,
1209, 1213, Lecomte 1334, Mniaud 1492, Pearson and Brown 1679, Record 1781, 1787,
1801, 1830, 1843, 1851, Record and Hess 1886, Williams 2430.

140. CRYPTERONIACEAE
(Fio. 147 on p. 652)

SUMMARY
(i) GENERAL
A
small family consisting of a few species of trees from India and the
Malay Archipelago, comprising the single genus Crypteronia. The general
anatomy has not received much recent attention, but one of the most signifi-
cant features is the occurrence of intraxylary phloem.

(ii) WOOD
Vessels medium-sized, perforations simple, intervascular pitting alternate
and small, vestured, pits to parenchyma similar, members of medium length.
Parenchyma predominantly apotracheal, scattered and in uniseriate lines,
also about the vessels. Rays up to 2-4 cells wide, markedly heterogeneous
with about 10 marginal rows, with conspicuous, canal-like intercellular spaces.
Fibres with bordered pits, of medium length.

LEAF
Solereder records the occurrence of a single, somewhat collenchymatous
layer of hypoderm, and the presence of cluster crystals.

Axis
STEM
The following features have been recorded by Solereder. Young stems
provided with wings, the latter consisting of epidermis and cortical tissue.
Secondary phloem conta ning elongated sclerenchymatous elements with
:

numerous pits. Xylem in the form of a continuous cylinder traversed by

narrow rays. Vessels with simple perforations. Pith including scattered
prosenchymatous elements, sometimes arranged in groups. Intraxylary
phloem present.

WOOD (Fig. 147 H-I)

Vessels medium-sized (100-200 p, mean tangential diameter); solitary and
in multiples of 2 or 3 cells; 5-8 per sq. mm. Perforations simple. Inter-
vascular pitting alternate, small, pits to ray and wood parenchyma similar to
the intervascular pitting; vestured (78). Mean length 0-35-0-7 mm. Paren-
chyma predominantly apotracheal, as scattered cells tending to form
irregular uniseriate lines and to be rather more abundant round the vessels.
656 CR YP TER O NIA CEA E
cells. Rays up to 2-4 cells wide less than i mm. high
Strands of 8 or more ; ;

numerous and composed entirely of upright cells about 20 rays

uniseriates ;

per mm.; markedly heterogeneous (Kribs's Type II A), with up to about 10

marginal rows of square to upright cells. Cells often filled with dark gummy
contents. Fibres with few and small, but distinctly bordered, pits, mostly
on the radial walls. Walls of medium thickness. Mean length about i -4 mm.
Intercellular canals. The ray cells are often rounded, with marked radial
intercellular spaces between the procumbent cells. Webber (2377) has pointed
out that there is no clear-cut distinction between such cavities and inter-
cellular canals, and the family is sometimes listed (1851) among those having
radial canals.

TAXONOMIC NOTES
The
presence of intraxylary phloem is interesting, since this feature occurs
also in the Lythraceae, in which family Crypteronia was included in the
Bentham and Hooker system.
ECONOMIC USES
One species, Crypteronia paniculata Bl. from Burma, is stated by Pearson
and Brown (1679) to be used for building and is referred to by Desch (574)
as a local substitute for Meranti.

GENUS DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Crypteronia.
LITERATURE
On Wood Structure
Bailey 78, Desch 574* Janssonius 1154, Lecomte 1334, Pearson and Brown 1679, Record
1843, 1851, Webber 2377.

141. OLINIACEAE
(FiG. 147 on p. 652)

SUMMARY
(i) GENERAL
A
small family consisting of a few species of shrubs or small trees comprised
in the single genus Olinia which occurs in St. Helena and South and East
Africa. The general anatomy has not received much attention in recent years,
but the presence of intraxylary phloem is a noteworthy character.

(ii) WOOD
Vessels small, in numerous small multiples, perforations simple, inter-
vascular pitting alternate, very small, vestured, pits to ray cells similar,
members of medium length. Parenchyma very sparse, scanty paratracheal.
Rays 2 cells wide, markedly heterogeneous with io or more marginal rows.
Fibres with simple pits, septate, of medium length.

Axis
STEM
The following features have been recorded by Solereder. Cork arising in
 OLINIACEAE 657
the sub-epidermis. Cortex containing slightly branched sclerenchymatous
idioblasts resembling short fibres. Secondary phloem including chambered
fibres containing solitary crystals. in the form of a continuous
Xylem
cylinder traversed by narrow rays. Vessels with simple perforations. Intra-
xylary phloem present.
WOOD (Fig. 147 G and j)
Vessels moderately small (50-100 fi mean tangential diameter); solitary
and in numerous radial multiples of 2 or 3 cells; with a slight tendency to
arrangement in loose oblique rows about 25 per sq. mm. Perforations simple.
;

Intervascular pitting very small to minute; vestured (78); pits to ray cells
similar. Mean member length about 0-5 mm. Parenchyma very sparse,
limited to an occasional cell touching a vessel; occasionally irregularly banded
at the ring boundary (2075). Solereder states that the wood of Olinia is note-

worthy on account of *unlignified groups of parenchymatous cells containing

solitary crystals and arranged concentrically'; such cells have not been ob-
served by the author. Rays up to 2 cells wide, the multiseriate parts short
and often more than i per ray; less than i mm. high; wholly uniseriate rays
relatively few; 15-20 rays per mm.; markedly heterogeneous with up to 10,
and occasionally more, marginal rows of square to upright cells (Kribs's
Type II A). Fibres with simple pits, rather more numerous on the radial
walls. Septate, but septa not common in Olinia usambarensis Gilg. Walls of
medium thickness to moderately thick. Mean length about 0-9-1-0 mm.

TAXONOMIC NOTES
The
presence of intraxylary phloem is noteworthy, since this feature also
occurs in the Lythraceae in which family Olinia was included in the Bentham
and Hooker system.
ECONOMIC USES
Olinia cymosa Thunb. produces a timber, known as Hard Pear, which is

used in South Africa for a variety of purposes (2075).

GENUS DESCRIBED
FOR WOOD STRUCTURE AND GENERAL ANATOMY
Olinia.

LITERATURE
On Wood Structure
Bailey 78, Record 1843, 1851, Scott 2075.

142. PUNICACEAE
(FiG. 147 on p. 652; FIG. 148 on p. 658)

SUMMARY
(i) GENERAL
A
small tropical and sub-tropical family of woody, sometimes spiny, shrubs
or small trees comprised in the single genus Punica. The widely cultivated
Pomegranate (P. granatum Linn.) is the only species which appears to have
4594 UU
 FIG. 148. LYTHRACEAE, A and D; PUNICACEAE, B-C; ONAGRACEAE, G-H and J;
LOASACEAE, E-F, I, and K
,
A, Lafoensia vandelliana Cham, et Schlect. Young stem X 33. B, Punica granatum Linn. Petiole
X 18. [Adaxial phloem not shown.] C, P. granatum Linn. Young stem X 18. [Stems of this species
frequently have only 4 wings.] D, Lagerstroemia indica Linn. Petiole X 18. E, Blumettbachia hieronymi
Urb. Stem x 8. F, B. hieronymi Urb. Petiole XI3. G, Oenothera mtssouriense Sims. Petiole x 8.
H, Fuchsia magellanica Lam. Stem x 13. I, Loasa vulcanica Andr^. Petiole X 13. J, Oenothera
missouriense Sims, Stem X 13. K, Loasa vulcanica Andrei Stem x8.
*
Intraxylary phloem omitted.
 PUNICACEAE 659
been investigated anatomically. One of the most interesting features is the
occurrence of intraxylary phloem.

(ii)
WOOD
Vessels very to moderately small, with some multiples of 6-8 cells, per-
forations simple, intervascular pitting alternate and very small, vestured, pits
to ray cells similar, members very short. Parenchyma absent. Rays uni-
seriate, composed entirely of square and upright cells, very numerous. Fibres
with simple pits, septate, crystalliferous, very short.

LEAF
Dorsiventral. Cells of the upper slightly and those of the
epidermis with
tower epidermis with very sinuous anticlinal walls. Stomata confined to
the lower surface mostly ranunculaceous. Mesophyll including i layer of
;

palisade cells. Idioblasts, containing very large solitary crystals, present along
the boundary between the palisade and spongy tissues. Vascular bundles of
the smaller veins embedded in the mesophyll, and each surrounded by a
sheath of rather large, parenchymatous cells. Midrib with an open, crescent-
shaped, bicollateral, vascular bundle. Transverse sections through the distal
end of the petiole (Fig. 148 B) exhibit a similar structure. Solitary and
cluster crystals present in the lamina and petiole (see also crystal-idioblasts
under 'Mesophyll' above).

Axis
YOUNG STEM (Fig. 148 c)
Provided with wings. Cork arising in the inner part of the pericycle.
Pericycle including a somewhat interrupted ring of fibres along the outer
periphery. Phloem, in 2- or 3-year-old stems, tending to be separated into
strands by the enlarged, triangular, distal ends of the medullary rays. Xylem
forming a closed cylinder, traversed by narrow rays. Vessels with simple
perforations. Intraxylary phloem present (Fig. 1480). Pith heterogeneous.
Cluster crystals abundant in the phloem, those in the secondary phloem
tending to be in concentrically arranged sacs. Secretory cells with unidenti-
fied, amorphous contents, observed in the cortex and pith.

WOOD (Fig. 147 K-M)

Vessels very to moderately small (mean tangential diameter about 50 /A);
mostly solitary and in multiples of 2 or 3 cells, but with some multiples of
7 or 8 cells or groups of 3 or 4 cells each almost contiguous radially; about
30 per sq. mm. Perforations simple. Intervascular pitting alternate and very
small, vestured (78); pits to ray cells similar. Mean member length about
0-23 mm. Parenchyma absent. Rays uniseriate, less than i mm. high,
composed entirely of square and upright cells, about 23 per mm. Fibres
with simple pits; septate, the septa commonly very numerous and dividing
the fibre into a number of chambers each containing i, or occasionally 2,
crystals (Fig. 147 K); the crystalliferous fibres thinner walled and in groups
that, in transverse sections, resemble irregular parenchyma bands. Mean
length about 0*65 mm.
660 PUN1C ACE AE
TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
Punica was included in the Lythraceae in the Bentham and Hooker system.
Its affinitywith that family is confirmed by the similarity of the anatomical
characters, especially the occurrence of intraxylary phloem.

ECONOMIC USES
Pomegranates are the fruits of Punica granatum Linn, The anatomical
structure of the dried rind of the fruit, which is used in medicine, has been
described in detail by Griffiths (822).

GENUS DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Punica.*
* in the Kew slide collection.
Represented

LITERATURE
(i) On General Anatomy
Griffiths 822.

(ii) On Wood Structure

Bailey 78, Greguss 2522, Record 1843, 1851.

143. SONNERATIACEAE
(FiG. 149 on p. 662)

SUMMARY
(i) GENERAL
Trees. Some of the species of Sonneratia inhabiting Mangrove Swarnps
from Africa to Australia are provided with vertical branches of the root
system projecting into the air or water above the mud. The occurrence of
intraxylary phloem in both Duabanga and Sonneratia is also noteworthy.

(ii) WOOD
Vessels medium-sized to moderately small, mostly in multiples of 2 or 3
cells,perforations simple, intervascular pitting alternate and moderately small
to large, vestured, pits to ray cells either similar or large and simple, members
of medium length. Parenchyma absent or vasicentric. Rays exclusively
uniseriate or up to 2 cells wide, homogeneous or heterogeneous. Fibres with
simple pits, septate in Sonneratia, of medium length to moderately short.

LEAF
Isobilateral in Sonneratia apetala Ham. Upper epidermis provided with
cuticular ridges, and the lower with papillae in Duabanga. Cork warts
recorded in certain species of Sonneratia. Stomata deeply sunken and present
on both surfaces in Sonneratia acida Linn. rubiaceous and equally numerous
;

on both surfaces in S. apetala Ham. Mesophyll including numerous large

mucilage cells, the central portion consisting of aqueous tissue containing
branched sclerenchymatous idioblasts.
 SONNERA TIA CEAE 661

Axis
STEM
Young stem of Duabanga and Sonneratia provided with collenchymatous
wings. Cork arising in the sub-epidermis in Sonneratia apetala Ham. Primary
cortex of Sonneratia mostly consisting of spongy parenchyma, outer part
containing branched sclerenchymatous idioblasts. Xylem in the form of a
continuous cylinder traversed by narrow rays. Vessels with simple perfora-
tions. Phloem including sclerenchyma. Pith supported by sclerenchymatous
elements, more conspicuous in Duabanga than in Sonneratia; composed of
pitted cells and distinctly lacunar in Sonneratia apetala according to Mullan
(1571). Intraxylary phloem present; that of Sonneratia apetala containing
a few fibres and crystal cells. Secretory cells occur in the phloem (including
the intraxylary phloem) of S. apetala, and, according to Mullan (1571),
crystals in vertical rows of special cells in the secondary phloem of the same
species.

WOOD (Fig. 149 A-D)

Vessels medium-sized (100-200 ft mean tangential diameter); solitary and
in numerous multiples of 2 or 3 cells, occasionally with a tendency to form
loose oblique or tangential lines; 2-10 per sq. mm. in Duabanga, usually about
20 per sq. mm. in Sonneratia. Perforations simple. Intervascular pitting
alternate, large in Duabanga, medium-sized to rather small in Sonneratia;
pits to ray cells often large and simple in Duabanga, mostly similar to the
intervascular pitting in Sonneratia, but with a few larger pits or with 2 pits
subtending i large ray pit; vestured (78). Tyloses often present, occasionally
sclerosed in Sonneratia (1649). Mean member length in Sonneratia 0-4-
0-5 mm. and in Duabanga 0-6-0-8 mm. Parenchyma absent from Sonne-
ratia paratracheal in Duabanga, as narrow to rather wide vasicentric sheaths
;

round the vessels (Fig. 149 A) and occasionally linking adjacent vessels. Rays
exclusively uniseriate, except for occasional biseriate parts, in Sonneratia\
more commonly 2 and occasionally 3 cells wide, in Duabanga, though pre-
dominantly uniseriate in some specimens; commonly about i mm. high in
the latter, shorter in Sonneratia', 10-18 per mm.; in Sonneratia composed
almost entirely of procumbent cells, apart from rows of crystalliferous cells
(Kribs's Homogeneous Type III); in Duabanga heterogeneous, with up to
about 10 marginal rows of square or upright cells, the uniseriates composed
of upright and procumbent cells. Cells of Sonneratia often filled with a dark,
gum-like substance and interspersed with rows of square cells, each cell con-
taining a solitary crystal. Fibres with simple pits; pits mostly on the radial
walls and rather few in Duabanga, but numerous in Sonneratia. Septate in
Sonneratia. H. P. Brown (1679) states that there is only i septum per fibre
in the Indian species; this is not so in other species. Walls thin in Duabanga,
moderately thick in Sonneratia. Mean length in Sonneratia 0-7-1-0 and in
Duabanga i'2~i'4 mm. Janssonius (1154) notes a distinct type of fibre round
the vessels in Sonneratia this is distinguished by being shorter, and having
;

rounded, untapered ends, thinner walls, and intercellular spaces; Brown

(1679), however, doubts if such a distinction can be drawn in the Indian
species; some markedly shorter fibres with blunt ends can be distinguished
in Duabanga sonneratioides Buch-Ham.
 662 SONNERA TIACEAE
ROOT
The following particulars concerning the structure of the vertical, negatively

FIG. i49 SONNERATIACEAE, A-D; ONAGRACEAE, E-I

;
A, Duabanga sonneratioides Buch-Ham. B, D. sonneratioides Buch-Ham. C, Sonneratia apetala
Ham. D, S. apetala Ham. E, Jus$iaea lattfolia Benth. F, J. latifolia Benth. G, Oenothera biennis
Lmn. H, Fuchsia nccartonii Hort. I, F. macrostemma Ruiz, et Pav.

geotropic portions of the root system ('pneumatophores') of Sonneratia

apetala Ham. have been recorded by Mullan (1571):
Cork consisting of a succession of lamellae each composed of
3 layers of
 SONNERA TIA CEAE 663
cells. Cells of the outer layer of cork rounded externally and not suberized ;
cells of the middle layer tabular and suberized ; those of the inner layer radially
elongated, rounded on the inside and suberized. Two layers of approximately
spherical cells, present between each succeeding cork lamella, facilitate separa-
tion at these points. Cortex, at the distal end of the 'pneumatophore', com-
posed of rounded cells interspersed with schizogenous, intercellular spaces;
also containing crystalliferous cells and branched sclerenchymatous idioblasts,
the latter less numerous away from the distal end. Cortex at the proximal end
of the root similar in structure to that of the subterranean root and supported
by curved, lignified spicules. Phloem including isolated strands of fibres
and vertical rows of crystalliferous cells. Xylem in the form of a broad
feebly lignified ring. Pith containing sclerenchymatous idioblasts at the
distal but not at the proximal end.
Earlier accounts of the structure of the 'pneumatophores' of Sonneratia by
Liebau (1368) and Emould (627) agree for the most part with that by Mullan.
Liebau failed to observe the S-shaped, spring-like cells previously described
by Westermaier from the cortex of the lower part of the 'pneumatophores*
and terrestrial roots. These are, however, probably identical with the 'curved,
lignified spicules* observed by Mullan. Liebau thought that the vascular
structure of the 'pneumatophore' resembles that of a stem rather than a root,
especially as the smallest vessels are those towards the centre of the axis,
whilst the phloem and xylem elements often occur along the same radii.
The spongy aerial roots of Sonneratia have, for many years, been regarded
as 'breathing' organs which serve for the intake of oxygen and liberation of
carbon dioxide, which would otherwise be difficult in roots growing in the
peculiar substratum in the muddy swamps of tropical estuaries. More
recently, however, Troll (2282) and Troll and Dragendorff (2283), whose
work was reviewed by Metcalfe (1494 A), have shown how the unusual roo'
system enables a succession of fibrous absorbing roots to be formed at pro-
gressively higher levels on the negatively geotropic roots. This occurs at
intervals as the level of the muddy substratum is raised, owing to the deposi-
tion of colloidal matter suspended in the water. Troll pointed out, however,
that the importance of the aerial roots in performing this function does not
exclude the possibility of their serving for gaseous exchange as well.
The terrestrial roots of the same species exhibit the following features.
Cortex lacunar; component cells mostly triradiate, and supported by thickening
ridges and by vertically elongated cells with thick, pitted, but unlignified walls.
Cork arising superficially.

TAXONOMIC NOTES
The wood Duabanga and Sonneratia differs considerably in
structure of
respect of the vessels, parenchyma, rays, and fibres. If these were the sole
taxonomic criteria it would appear doubtful if these genera should be included
in the same family. The general anatomy of Duabanga is not so well known
as that of Sonneratia, but here there are points in common between the 2
genera, notably the occurrence of intraxylary phloem. This character also
serves to connect the Sonneratiaceae with the Lythraceae in which they were
included in the Bentham and Hooker system.
664 SONNERATIACEAE
ECONOMIC USES
Both Duabanga and Sonnemtia furnish good packing-case timbers, which
are in demand in some localities in India.

GENERA DESCRIBED
FOR GENERAL ANATOMY AND WOOD STRUCTURE
Duabanga, Sonneratia.
LITERATURE
(i) On General Anatomy
Emould 627, Liebau 1368, Metcalfe *494 *494 A, Muilan 1571, Troll 2282, Troll and
Dragendorff 2283.

(ii) On Wood Structure

182, Foxworthy 705, Howard 1088, Janssonius 1154,
Bailey 78, den Berger 179,
Kanehira 1206, Panshin 1649, Pearson and Brown 1679, Record 1843, 1851.

144. ONAGRACEAE
(FiG. 148 on p. 658; FIG. 149 on p. 662; FIG. 150 on p. 666)

SUMMARY
(i) GENERAL
A temperate and sub-tropical family consisting mainly of herbs but includ-
ing some shrubs. The occurrence of intraxylary phloem in the axis provides
one of the most important diagnostic features, and is indicative of affinities
with the Lythraceae and other related families. Interxylary phloem has
also been recorded in a few genera. The hairs are nearly all simple and
include unicellular and uniseriate types. The stomata are surrounded by
3 or more subsidiary cells, sometimes resembling those of the Cruciferae.
The mesophyll is dorsiventral or centric, and the spongy and palisade
portions not always clearly differentiated. Raphides are common in most
genera. Sclerenchyma is generally poorly developed or absent from the veins.
There is a principal arc-shaped vascular strand in the petiole, sometimes
accompanied by lateral accessory strands. The cork, which often includes
phelloid cells, the latter sometimes in layers alternating with the suberized
cells, is usually deep-seated in origin. It is sometimes replaced by aeren-

chyma in species from marshy habitats, e.g. in Epilobium and Jussiaea.

(ii)
WOOD
Vessels small to moderately small, often with numerous .radial multiples
and clusters, perforations simple, intervascular pitting alternate, pits to
parenchyma simple and often elongated; members of medium length to
moderately short. Parenchyma paratracheal, scanty. Rays 2-9 cells wide,
commonly more than i mm. high, heterogeneous, composed almost entirely
of square or upright cells except in Oenothera. Fibres with simple pits,
usually septate, moderately short. Included phloem of the 'foraminate*
type occasionally present.
LEAF
Dorsiventral or centric, the spongy and palisade mesophyll not always
 ONAGRACEAE 665
distinct. Hairs nearly always simple, commonly with granular or verrucose
surfaces; unicellular of various shapes, often club-shaped, e.g. in species of
Clarkia and Oenothera*, short, uniseriate in species of Fuchsia, Jussiaea^ Lud-
wigia, long uniseriate inTrapa bispinosa Roxb. according to Sabnis (1977).
Leveille (1365) found the structure of the hairs to be valuable in classifying
species of Oenothera. Epidermis including mucilaginous cells in species of
Jussiaea and Trapa. Stomata surrounded by 3 or more subsidiary cells, the
latter sometimes consisting of 2 large and i small cells as in the Cruciferae;

occurring on both surfaces or confined to the lower side in most species, but
limited to the upper surface in Epilobium crassum Hook. f. and in
Trapa
bispinosa Roxb. according to Sabnis (1977). Sclerenchyma said to be generally
absent from the veins except in certain species of Fuchsia. Petiole (Fig. 148 G),
in transverse sections through the distal end of all investigated species, exhibit-
ing a single, usually flattened, arc-shaped, vascular strand, sometimes accom-
panied by lateral accessory strands. Betts (188) refers to canals surrounded
by epthelium occurring at intervals in the mesophyll of Epilobium pedunculare
Hook. f. and E. pubens A. Rich. Crystals. Raphides, usually situated in sacs
and sometimes embedded in or replaced by mucilage, recorded or observed
in Circaea, Clarkia, Epilobium, Eucharidium, Fuchsia, Gaura, Gayophytum,
Gongylocarpus, Hauya, Jussiaea, Lavauxia, Lopezia, Ludwigia, Megapterium,
Oenothera, Zauschneria. Raphides accompanied by clustered crystals in
Jussiaea and Ludwigia. Crystals said to be exclusively clustered in Trapa.
Styloids recorded in a species of Hauya. Oil cells recorded by Stein (2192)
in the epidermis of Ludwigia alternifolia Linn, and less regularly in species
of Circaea, Clarkia, Fuchsia, Gaura, Jussiaea, Oenothera, Trapa.

Axis
STEM (Fig. 148 H and j)
Cork arising in the pericycle of Clarkia as well as in all of the genera and
species represented in the Kew slide collection, and probably in others as
well; often including phelloid cells, the latter sometimes arranged in layers
alternating with the suberized cells. Cork often replaced by aerenchyma
(Fig. 150 A-C) in submerged portions of stems and also in the roots of species
inhabiting marshes, notably in Epilobium and Jussiaea. Primary cortex pro-
vided with well-developed intercellular spaces in species of Fuchsia,'Jussiaea,
Ludwigia, Oenothera. Endodermis in Epilobium pedunculare Hook. f. and
E. pubens A. Rich, described by Betts (188) as well defined and consisting of
a single layer of large cells with suberized walls. Pericycle usually containing
isolated strands or, less frequently, a continuous ring of fibres in species of
Circaea, Clarkia, Epilobium, Fuchsia (none seen in the specimen illustrated
in Fig. 148 H), Gaura,Gayophytum, Godetia, Gongylocarpus, Jussiaea, Kneiffia,
Lavauxia, Lopezia, Ludwigia, Megapterium, Oenothera, Raimannia, Zau-
schneria. Phloem reported as sometimes containing stone cells in Hauya and
fibres in Fuchsia. Xylem forming a continuous cylinder traversed by narrow
Kew slide collection. Vessels with
rays in all of the species represented in the
simple perforations; occasional multiperforate plates also recorded (see
'Wood'). Pith often becoming hollow; including large intercellular spaces in
Trapa. Intraxylary phloem, which is one of the most important diagnostic
features for the family, has been recorded in numerous species of Circaea,
 666 ONAGRACEAE
Clarkia, Epilobium, Eucharidium, Fuchsia, Gaura, Gayophytum, Godetia,
Gongylocarpus, Hauya, Jussiaea, Kneiffia, Lavauxia, Ludwigia, Megapterium,
Oenothera (Fig. 148 j), Raimannia, Trapa, Zauschneria and probably occurs
throughout the family. It adjoins the vascular bundles in some species, but
forms isolated strands in the pith of others. For interxylary phloem see

FIG. 150. ONAGRACEAE

A, Cross-section through an aerial root of Jussiaea repens Linn. B, C, Jussiaea peruviana L.: B,
cross-section; C, radial longitudinal section of the aerenchyma. A, after Goebel, B, C, after H.
Schenck.

1
under 'Anomalous Structure p. 667. Crystals occur chiefly in the form of
,

raphides, which are present in varying abundance in different genera and

species; and have been observed at Kew in species of Epilobium, Fuchsia,
Kneiffia, Lavauxia, Megapterium, Raimannia. Raphides sometimes embedded
in or replaced by mucilage. Oil cells recorded by Stein (2192) in the epi-
dermis of Ludwigia alternifolia Linn, and less regularly in species of Circaea,
Clarkia, Fuchsia, Gaura, Jussiaea, Oenothera, Trapa.

WOOD (Fig. 149 E-I)

Vessels very small to moderately small (25-100 /z mean tangential
diameter); solitary and in numerous radial multiples and irregular clusters of
several cells except in Jussiaea ; 7-25 per sq. mm. Perforations simple accord- ;

ing to Solereder, Grosse mentions the occurrence of a few multiperforate

 ONAGRACEAE 667

plates in the herbaceous stems of species of Codetta and Oenothera. Inter-

vascular pitting alternate, moderately small, vestured (1886); pits to ray
or wood parenchyma usually simple, similar in shape to the intervascular
pitting or elongated, sometimes almost scalariform. Ty loses sometimes
present. Mean length O'3-O'5 mm. Parenchyma sparse, limited to a few
cells round the vessels. Rays 2-3 in Jussiaea and Oenothera and
cells wide
occasional specimens of Fuchsia, iip to 9 cells wide in some specimens or
species of Fuchsia, e.g. F. riccartoni Hort., and with low uniseriate rays;
uniseriate rays high and numerous in the other genera; uniseriates typically
composed of upright cells only, but of upright and procumbent cells in
Oenothera; commonly more than i mm. high, except in Oenothera mostly
io~~i6 rays per mm., rather fewer in some species of Fuchsia; heterogeneous
in Oenothera] composed almost entirely of large square or upright cells in
Fuchsia and Jussiaea; with sheath cells, except in Oenothera. Gum and
crystals not observed, but numerous large crystals reported (1886) in Jussiaea
latifolia Benth. Fibres with very small simple pits, very numerous on the
radial walls in Fuchsia; septate in most species of Fuchsia and Jussiaea; walls
thin riiean length 07-0*8 mm. Included (interxylary) phloem of the
;

'foraminate' type, with strands of included phloem scattered in a normal

xylem, reported by Record (1851) in Epilobium and Oenothera; the strands
sometimes grouped in tangential rows in Oenothera (see also under 'Anomalous
Structure' below).
ROOT
Aerenchyma (Fig. 150 A-C) present in portions of the root system of
Jussiaea above ground-level. Lateral roots of Trapa each provided with one,
poorly developed strand of phloem and xylem.

ANOMALOUS STRUCTURE
Interxylary phloem present in the stem and/or root of various species of
Epilobium, Gaura, Lopezia, Oenothera (see also under 'Wood* above). Inter-
xylary periderm described by Moss (1564) as occurring in the perennating
organs of Epilobium angustifolium Linn., where it is formed 'each year over
the surface of the wood connected with the previous year's aerial shoots and
joins with external periderm in the basal region of each of these decadent
shoots'. The interxylary periderm is produced by a phellogen which arises
in a parenchymatous zone of the xylem. The ability of the perennating
organs to withstand adverse conditions is thought to be correlated with the
presence of interxylary periderm.

ECONOMIC USES
The fruits
of species of Trapa are known as Water Chestnuts, and the seeds
from them are ground into flour and used in various parts of the world
(e.g. China, India, Tropical Africa) to make bread. Species of Clarkia,
Fuchsia, Godetia, Oenothera, &c., are commonly cultivated for ornamental

purposes.
GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Chamaenerion, Circaea, Clarkia, Epilobium,* Eucharidium, Fuchsia,*
668 ONAGRACEAE
Gaura, Gayophytum, Godetia,* Gongylocarpus, Hauya, Isnardia, Jussiaea,
Kneiffia,*Lavauxia,* Lopezia, Ludwigia, Megapterium,* Oenothera,*
Oocarpon, Raimannia,* Trapa.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Epilobium), Fuchsia, (Godetia), Jussiaea, Oenothera,

LITERATURE
(i) On General Anatomy
Betts 188, Gates 750, LeveilM 1365, Moss 1564, Sabnis 1977, Stein 2192.

(ii) On Wood Structure

Record 1843, 1851, Record and Hess 1886, Williams 2430.

145. LOASACEAE
(Fic. 148 on p. 658; FIG. 151 on p. 670)

SUMMARY
A family occurring mainly and temperate parts of the New
in tropical
World. Members are mostly herbaceous or, more rarely, woody; provided
on both leaf and stem with a wide variety of characteristic hairs, some of
which possess powerful stinging properties, owing to the presence of an
irritant substance in the hairs themselves or the surrounding basal cells. The
irritant substance is readily liberated because the hairs are brittle. Cystoliths
are common in the hairs. The stem of herbaceous species contains a ring of
collateral vascular bundles and the woody species a cylinder of xylem,
whilst intermediate types also occur. Mechanical tissue in herbaceous species
is chiefly in the form of collenchyma in the outer part of the cortex, but it is

frequently interrupted by groups of chlorenchyma extending outwards to the

epidermis. In living material the chlorenchyma patches can often be seen
with the naked eye as dark-green spots or streaks on the stem surface.

LEAF
Centric. Hairs (Fig. 151) include the following and intermediate types.
A. Unicellular

(i) Simple, of variable length, with silicified or non-silicified walls,

(ii)Conical, pointed, strongly silicified, the surface beset with downwardly

directed spines, (iii) Strongly silicified and barbed, comparable with a many-
fluted anchor (Fig. 151 A~C), the stalk sometimes covered with curved spines
as well, (iv) Hooked or sickel-shaped, and provided with a multicellular
base, (v) Filiform, with local tuberous swellings, sharply attenuated and some-
times barbed at the apex, (vi) Elongated, silicified stinging hairs often with
slightly recurved, blunt, or capitate apices; filled with a yellowish irritant
substance, readily liberated owing to the brittleness of the hair and causing
serious injury if discharged into the skin of animals.
 LOASACEAE 669
B. Multicellular

(i) Glandular, with uniseriate stalks and uniseriate heads. Lime or silica
frequently present in structures resembling cystoliths (Fig. 151 D-G) situated
in the hairs themselves or cells at their bases in
species of Blumenbachia,
Caiophora, Eucnide, Gronovia, Loam, Mentzelia, Petalonyx', these bodies are
stated to suggest affinities between the Loasaceae and Cucurbitaceae. Stomata
ranunculaceous in Blumenbachia, Caiophora, Eucnide, Mentzelia,
Petalonyx',
confined to the lower surface in the dorsiventral leaves of Caiophora and
Eucmde, but on both sides in the isobilateral leaf of Pentalonyx thurberi
A. Gray. The mesophyll of this last species contains a free network of
tracheids, and lignified pitted cells above and below the midrib. Transverse
sections through the distal end of the
petiole exhibit a slightly crescent-
shaped vascular strand accompanied by smaller accessory bundles in the
wings in Blumenbachia hieronymi Urb. (Fig. 148 F); a dorsally flattened, some-
what interrupted cylindrical strand, accompanied by 2
medullary bundles
and others in the wings, observed in the corresponding position in Loasa
vulcanica Andre (Fig. 148 i).

Axis
STEM and K)
(Fig. 148 E
Outer part of the cortex of herbaceous species provided with a well-
developed but interrupted ring of chlorenchyma, extending outwards to the
epidermis. Cork examined only in Mentzelia, originating there on the inside
of the pericyclic fibres. Pericycle
containing yellow, vertically elongated
groups of thickened parenchymatous cells in Petalonyx a few scattered fibres ;

seen in Loasa vulcanica Andre. Herbaceous species

provided with a some-
what interrupted vascular ring (Fig. 148 E), but a cylinder of xylem occurs
in woody members of the
family, e.g. in species of Mentzelia and Petalonyx.
Intermediate types of vascular structure also occur, e.g. in Loasa vulcanica
(Fig. 148 K). The xylem includes fairly wide vessels with simple perforations;
wood fibres with fairly wide lumina and simple and/or bordered pits.

GENERA DESCRIBED
Blumenbachia,* Caiophora, Eucnide, Gronovia, Loasa,* Mentzelia, Peta-
lonyx.
* in the Kew slide collection.
Represented

LITERATURE
On General Anatomy
Gilg 776.

146. TURNERACEAE
(Fie. 152 on p. 672; FIG. 154 on p. 682)

SUMMARY
(i) GENERAL
A small, mainly tropical American family of herbs of which some tend to
be shrubby. A very detailed account of the anatomy of the family was pub-
 670 TURNERACEAE
lished by Berger (184), from whose thesis a large part of the present descrip-
tion has been taken. The diverse kinds of hairs are of considerable taxonomic
value, whilst the occasional occurrence of scalariform perforations plates in
the vessels in the neighbourhood of the primary wood of the axis is note-
worthy.

(ii) WOOD
Vessels very small to medium-sized, with numerous multiples, perforations
simple or simple and scalariform, intervascular pitting alternate and very

FIG. 151. LOASACEAE

A-C, Unicellular anchor-hairs. A, B, Eucnide lobat a Gray; C, Caiophora lateritia (Hook.) Klotzsch.
D, E, Petalonyx thurberi Gray; D, cystolith-hair; E, trichome with merely calcified, but strongly
thickened wall, without a cystolith, but with cystoliths in the subsidiary cells. F, G, Loasa cheUdoni-
folia Benth.; F, cystolith hair adpressed to the leaf, back-view; G, in side-view. By Solereder.

small. Parenchyma apotracheal, diffuse or reticulate. Rays up to 5 cells

wide, heterogeneous or composed entirely of square and upright cells. Fibres
with numerous indistinctly to distinctly bordered pits.

LEAF
Generally dorsiventral, with i or 2 layers of palisade tissue in different
species; isobilateral, with a single layer of palisade cells towards both surfaces
in Streptopetalum hildebrandti Urban, Turnera diffusa Willd. and the variety
aphrodisiaca Urban, T. hermanioides Cambess, and T. ulmifolia Linn. Hairs
include the following diverse types.

A. Non-glandular

(i) Simple, unicellular, usually with thick

sclerified walls and narrow lumina
in species of Erblichia, Mathurina, Piriqueta, Turnera, Wormskioldia. (ii)
Simple, uniseriate, with thin or thick walls in species of Hyalocalyx, Piriqueta,
 TURNERACEAE 671

Turnera; uniseriate hairs, sometimes covered with fine granulations, in species

of Hyahcalyx, Streptopetalum, Turnera. (iii) Various types of stellate hair
recorded, especially in Piriqueta (section Eupiriqueta), but also occurring in
a few species of Streptopetalum, Turnera, Wormskioldia. Variations in the
type of stellate hair are valuable for the identification of species.

B. Glandular

(i) Each having a short

multicellular stalk, bearing a very large, spherical,
terminal of Hyalocalyx and Turnera. (ii) Similar to (i) but
cell in species

pluricellular and somewhat resembling a mulberry fruit in Erblichia and

numerous species of Turnera. These glands appear very different in fresh
and dried material respectively. Similar glandular hairs occur in the Labiatae
and Chenopodiaceae. (iii) Clavate hairs with a uniseriate base and multi-
cellular head, recorded, especially on the lower surface of the leaf, in species
of Piriqueta and Turnera. (iv) Multicellular hairs each with a massive base,
but tapering to a biseriate to triseriate distal end, the terminal cell being
spherical, recorded in a few species of Piriqueta, Streptopetalum (Fig. 152 A),
Turnera, Wormskioldia. Glands each having a short peduncle, traversed
longitudinally By a vascular strand and provided with a head which is at first
concave and resembles a small Peziza fructification, but later becomes convex
at the onset of secretion, recorded at the base and margins of some of the
leaves in species of Mathurina, Piriqueta, Turnera (Fig. 152 B), Wormskioldia.
Disks of the glands composed of palisade cells containing a granular material
readily stained by Sudan III.
Epidermis sometimes containing mucilage, e.g. in Piriqueta, Strepto-
petalum, and certain species of Turnera. Cells of the upper usually with less
sinuous anticlinal walls than those of the lower epidermis. Stomata confined
to the lower surface in certain species of Mathurina, Piriqueta, Streptopetalum,
Turnera, Wormskioldia, but recorded on both surfaces in other species of
Piriqueta, Streptopetalum, Turnera, Wormskioldia; mostly rubiaceous, but
sometimes cruciferous and ranunculaceous all 3 types known to occur together
;

in a single leaf, e.g. in Turnera diffusa Willd. var. aphrodisiaca Urban. Meso-
phyll containing sclerenchymatous idioblasts in Turnera hilaireana Urban.
Petiole, in transverse sections through the middle portion, exhibiting, accord-
ing to Berger a solitary, more or less open, arc-shaped vascular strand
(I.e.),
in species of Turnera and Wormskioldia. A
slightly crescent-shaped abaxial
vascular strand accompanied by a smaller adaxial one, the xylem groups of
both being directed towards one another, observed at Kew in sections through
the distal end of the petiole of T. ulmifolia Linn. (Fig. 1540). An arc of
3 separate bundles, the median one much larger than the other 2, recorded in
species of Erblichia, Mathurina, Wormskioldia, and a closed circular strand in
Piriqueta. Very small, clustered crystals frequent in the spongy mesophyll
and, less often, in the palisade tissue. Secretory elements known to occur
only in the form of fairly frequent tanniniferous idioblasts.

Axis
STEM (Fig. 154 i)

Young stem usually provided with hairs of the same types as those
described for the leaf. Cork arising in the sub-epidermis in certain species
6 72 TURNERACEAE
of Turnera. Cortex varying considerably in width usually consisting mainly
;

or wholly of collenchyma. Isolated, rounded fibres recorded immediately

below the epidermis in Turnera hilaireana Urban. Pericycle including a
continuous ring of sclerenchyma, an interrupted ring of fibre strands, a circle
of solitary fibres, or entirely devoid of sclerenchyma according to the species.
Phloem always devoid of sclerenchyma present in the form of a continuous
;

cylinder, cylindrical strands, or scattered sieve tubes. Xylem nearly always

FIG. 152. TURNERACEAE, A-B; PASSIFLORACEAE, C~D

A, Glandular shaggy hair of Streptopetalum hildebrandtii Urb. B, Glands of Turnera annularis Urb.
C, Crystal-cell from the palisade-tissue of Adenia lobata (Jacq.) Engl. D, Glandular shaggy hair of
Passiflora clathrata Ma8t. in longitudinal section. A-C by Solereder, D after Harms.

forming a continuous, fairly thick cylinder traversed by narrow rays. Vessels

small (up to about 45 JUL in radial diameter in Turnera ulmifolia Linn.), usually
irregularly distributed but more rarely in radial rows; perforations mostly
simple apart from occasional scalariform plates with a few bars recorded in
the region of the primary xylem in Piriqueta and Turnera. Ground-mass of
the wood consisting of fibres with small pits, the latter described as distinctly
bordered in at least certain species. Wood parenchyma scanty. Rays usually
i or 2 cells wide, recorded by Berger as up to 5 cells wide in certain species of

Turnera. Wide rays have been recorded by Colozza (455) in Streptopetalum

and Wormskioldia. Leaf trace bundles sometimes traverse the cortex longitu-
dinally for a considerable distance and constitute cortical vascular bundles
(Fig. 154 1). Pith generally composed of cells with cellulose walls, but the outer
part thickened in some species, and islands of sclerosed pitted cells recorded
ip the centre of the pith in Turnera hilaireana. Pitted cells also noted in the pith
of Streptopetalum and other species of Turnera. Clustered crystals usually
plentiful in unlignified tissues, especially in the cortex and pith. Secretory
elements known to occur only in the form of thick-walled, tanniniferous
 TURNERACEAE 673
idioblasts, frequently arranged in longitudinal columns, the transverse walls
sometimes breaking down; recorded in the pith and, less frequently, in the
cortex and phloem of a few species of Piriqueta and Turnera, but not present
in numerous other species of Turnera.

WOOD 1

Erblichia
Vessels small to medium-sized, in multiples of 2-6 cells. Perforations
simple. Intervascular pitting alternate, very small Parenchyma apotracheal,
in fine reticulate lines. Rays up to 5 cells wide, markedly heterogeneous.
Fibres with numerous indistinctly bordered pits and thick walls.

Turnera
Vessels very small, solitary, and in short to long multiples; with spiral
thickening. Perforation plates simple and scalariform with numerous fine bars.
Intervascular pitting alternate and very small pits to ray cells often elongated
;

and scalariform. Parenchyma apotracheal, Rays up to 4 cells wide;

diffuse.

very numerous; composed almost entirely of square and upright cells. Fibres
with numerous, very small, distinctly bordered pits; walls thick.

ROOT
Cork poorly developed. Cortex composed of homogeneous parenchyma.
Endodermis not clearly differentiated. Pericycle sometimes including
fibres. Xylem forming a thick, compact cylinder composed of tracheids,
vessels with spiral thickening or scalariform pitting or with ovoid pits with
linear apertures. Crystals said to be absent. Berger noted little difference
in the structure of the secondary xylem in different genera and species.

TAXONOMIC NOTES
It is generally thought by systematists that the Turneraceae have close
affinities with the Passifloraceae, a view which is supported by the anatomical

similarity of the 2 families. Berger (184) has pointed out that Streptopetalum
and Wormskioldia are indistinguishable by means of their anatomical features.
Taylor (2237) states that a cursory examination of the wood gives an
indication that this family belongs to the Flacourtiaceae complex. It should,
however, be noted that the 2 most characteristic features of the Flacourtiaceae
septate fibres and parenchyma absent or paratracheal only are lacking
from the Turneraceae.
ECONOMIC USES
The dried leaves of Turnera diffusa Willd. var. aphrodisiaca Urban con-
stitute thedrug Damiana, to which aphrodisiac and mild purgative properties
have been attributed. Important microscopical characters of this material
include: the isobilateral mesophyll; the unicellular, thick- walled, warty
trichomes; the glandular hairs with short, unicellular stalks and few-celled
head sometimes containing reddish-brown material; the numerous, mostly
rubiaceous and cruciferous, but occasionally ranunculaceous stomata confined
to the lower surface; the petiole with an open arc-shaped vascular strand
1
Based entirely on the description by Record and Hess (1886).
4594 XX
674 TURNERACEAE
containing small, radially arranged vessels in the xylem, the strand being
supported by thick- walled fibres in the pericyclic region; the yellowish
material secreted in the midrib and some of the epidermal cells the mostly ;

clustered and occasional solitary crystals of calcium oxalate.

GENERA DESCRIBED
Erblichia, Hyalocalyx, Mathurina, Piriqueta, Streptopetalum, Turnera,*
Wormskioldia.
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Berger 184, Colozza 455, Gilg 775.

(ii) On Wood Structure

Record 1851, 1885, Record and Hess 1886, Taylor 2237.

147. PASSIFLORACEAE
(Fie. 152 on p. 672 ; FIG. 153 on p. 676; FIG. 154 on p. 682)

SUMMARY
(i)
GENERAL
A
tropical and sub-tropical family of very varied habit including many
climbers, but others are more woody and have the form of shrubs or small
trees. Adenia includes xerophytic species such as A. pechuelii (Engl.) Harms
which occur in the driest regions of Africa. In this and closely related species
the plant consists of a collection of short, fleshy, tuberous stems from which
erect or ascendant, sometimes thorny branches arise. Other species of Adenia
possess a single, partly or wholly subterranean, napiform tuber bearing some-
what woody climbing stems. A. venenata Forsk. (syn. Modecca abyssinica
Hochst) is a succulent species. The leaf is usually dorsiventral, but exhibits
isobilateral structure in some genera. Sessile or shortly stalked glands are
common in the petiole, and glandular spots less frequent on the lower side
of the lamina. The hairs are mostly unicellular or uniseritate, but glandular
shaggy types also occur in Passiflora. The leaf epidermis is sometimes
2-layered. The stomata,
usually confined to the lower surface, are ranun-
culaceous. Crystals, where present, are solitary rhombohedra or clustered.
Secretory elements include tanniniferous cells which are common in the
parenchymatous portion of both stem and leaf, whilst secretory cavities,
which also contain tannin, have been recorded in the leaf and axis of Adenia
(section Ophiocaulon),

(ii) WOOD
Vessels extremely small to large, few to numerous, sometimes in radial
multiples of 4 or more cells or with a radial or oblique pattern, perforations
usually simple, rarely with a few scalariform plates, intervascular pitting
alternate, minute to moderately large, with horizontal apertures, pits to
 PASSIFLORACEAE 675

parenchyma similar or larger and almost simple; members usually of medium

f

length to moderately long, but extremely long in Soyauxia 'fibriform vessel

,

members' characteristic of some genera. Parenchyma typically apotracheal,

diffuse to numerous uniseriate bands, occasionally with broader bands,
aliform or vasicentric. Rays up to 1-8 cells wide, high, usually composed
entirely of square or upright cells, but normally heterogeneous in a few species.
Fibres with simple or distinctly bordered pits, moderately to very long.
Soyauxia differs considerably from the other genera, particularly in having
solitary vessels, scalariform perforation plates, scalariform to opposite inter-
vascular pitting, less abundant and diffuse parenchyma, and a different type
of ray.

LEAF
Usually dorsiventral, but isobilateral in species of Adenia, Paropsia, Passi-
flora, Tryphostemma. Hairs unicellular or uniseriate in various species of
Passiflora, the unicellular ones hooked at the apex in certain species. Uni-
seriate hairs also recorded in Paropsia and Tacsonia, those of the last genus
sometimes very long or sufficiently interwoven to form a felt. Simple,
unicellular, sclerenchymatous, and tufted trichomes recorded in Abatia, the
tufted ones in A, verbascifolia H. B. et K. said to be formed by the union at
their bases of a number of unicellular hairs. Glandular shaggy hairs (Fig.

152 D), each usually with a spherical head on a multiseriate stalk of variable
length, the latter sometimes containing a vascular bundle, occur in a number
of species of Passiflora, especially in the section Dysomia. Sessile or short-
stalked glands recorded on the petioles of Adenia, Crossostemma, Deidamia,
Hollrungia, Paschanthus, Passiflora, Smeathmannia, Tacsonia, Tetrastylis and
glandular spots on the lower side in a few species of Adenia, as well as in
Passiflora, particularly in the sections Cieca, Decaloba, Murucuja, Pseudo-
murucuja, Psilanthus. Cuticle of Abatia said to exhibit markings like those
on etched glass. Cuticular projections to the epidermis common in species
of Passiflora, especially in the sections Cieca, Decaloba, Eumurucuja (this
applies to the stem also). Epidermis partly or wholly 2-layered, often with
crystals in the lower layer, in a few species of Abatia, Adenia, Hollrungia,
Smeathmannia', cells sometimes with mucilaginous inner membranes in
Barteria and Paropsia. Lower epidermis frequently papillose in Adenia and
Passiflora. Hypoderm recorded in a few species of Abatia. Stomata usually
confined to the lower surface; mostly ranunculaceous, but a proportion said
to be rubiaceous in Abatia. Central part of the mesophyll stated to contain
thick, pitted cells in i species of Mitostemma and sclerenchymatous idioblasts
in a few species of Passiflora. Vascular bundles of the veins usually embedded
in the mesophyll; not always accompanied by sclerenchyma. Petiole
examined only in i species of Passiflora (Fig. 1540), in which transverse
sections through the distal end exhibit a circle of individually distinct vascular
bundles accompanied by smaller strands in the wings. Secretory elements
include frequent cells and receptacles with tanniniferous contents, the
receptacles recorded particularly in species of Adenia belonging to the section
Ophiocaulon, where they sometimes appear as black dots. Crystals solitary
or clustered large solitary types said to occur in special sacs in certain species
;

of Adenia (Fig. 152 c) and Soyauxia (in the mesophyll).

676 PASSIFLORACEAE
Axis
STEM (Fig, 154 H)
Young stems sometimes winged, e.g. in species of Adenia and Passifiora.
Epidermis encrusted with a wax-like substance on young shoots of several

G
FIG. 153. PASSIFLORACEAE
A, Soyauxia grandifolia Gilg et Stapf. B, S. grandifolia Gilg et Stapf. C, Androsiphonia adenostegia
Stapf. D, A. adenostegiaStapf. E, Smeathmannia pubescent Sol. F ,
S. pubescens Sol. G, Crosso-
stemma laurifolium Planch. H, Dilkea johannerii Rodr.

species of Adenia. Warty excrescences arising in the hypoderm recorded by

Baccarini (60) on old stems of Adenia venenata Forsk. (syn. Modecca abyssinica
Hochst). Cork arising superficially, sometimes in the epidermis itself;
usually composed of thin- walled cells; spongy cork consisting of radially
elongated cells present in Passiflora suberosa Linn. Pericycle commonly
containing widely spaced strands of fibres, e.g. in species of Passiflora
(Fig. 154 H); but provided with a composite
and continuous ring of scleren-
 PASSIFLORA CEAE 677
chyma in Mitostemma glaziovii Mast, and relatively old branches of species
of Barteria, Paropsia, Paropsiopsis, Smeathmannia, Soyauxia. Walls of the
pericyclic fibres of certain species of Passiflora said to be stained violet, and
the middle lamella yellowish, when treated with chlor-zinc-iodide. Phloem
seen to include small groups of fibres in a species of Passiflora. Xylem in the
form of a more or less continuous cylinder, but frequently traversed by broad
primary rays ; that of the climbing species includes a proportion of vessels of
wide diameter (Fig. 154 H). Vessel perforation mostly simple, but, according
to Harms (894), scalariform plates are common in Barteria, Paropsia, Parop-
siopsis,Smeathmannia (see also Wood'). Secretory elements include tannin
sacs with wide lumina, often with thick walls and sometimes vertically
elongated, in the cortex and/or pith of a few species of Adenia, Hollrungia,
Passiflora. Crystals include solitary and clustered types.
Bailey (70) has pointed out that in an unswollen branch of Barteriafistulosa
Mast, the homogeneous pith consists of compact, thin-walled parenchyma.
When part of the stem of this species develops into an ant domatium, the
pith becomes heterogeneous and the reduced vascular system consists of
separate bundles. The cavity of the domatium is surrounded by a few thin-
walled cells containing an amber-coloured substance. The structure of the
ant domatia of B. dewerei v. Willd. et Dur. is similar.
Some of the climbing members of the family, e.g. species of Passiflora and
Tetrapathaea, possess tendrils believed to be morphologically equivalent to
inflorescence branches. The anatomical structure of these becomes modified
by the combined stimulus of contact and tension, whereby the development
of mechanical tissue is increased, particularly in those parts of the tendril
where it is most effective. Experiments have shown that the tendrils in
Passiflora do, in fact, possess a greater breaking strength in consequence of
these anatomical changes. Contact is said to be the main influence which
causes an increase in the number of xylem elements and in the thickness of
their walls, whereas tension causes the walls of the pith cells to become much
thicker. For details of tendril structure see Brush (298), Bird (199), Gangstad
(741). Stems of the succulent Adenia venenata Forsk. (syn. Modecca abyssinica
Hochst.) include abundant water-storage parenchyma formed, according to
Baccarini (60), partly by subdivision of the ground parenchyma, and partly
by cambial activity. The intervascular parenchyma becomes lignified in old
stems.

WOOD (Fig. 153)

Vessels very variable in size even in different species of the same genus,
small (less than 100 /z mean tangential diameter) in at least some species of
Androsiphonia, Paropsia, Passiflora, and Soyauxia, medium-sized (100-200 /z)
in some species of Crossostemma, Dilkea, Smeathmannia, and Tacsonia\
mean tangential diameter often more than 200 /z in climbing species, e.g.
Passiflora quadrangularis L., and with individual vessels up to 400-500 /z,
e.g. in P. vitifolia H. B. et K. (2466); solitary in Soyauxia, solitary
and in pairs
and threes in the other genera, with multiples or groups of 4 or more cells in
Androsiphonia, Paropsia p.p., Passiflora p.p., and Smeathmannia; forming a
radial pattern in Smeathmannia (Fig. 153 E) and in radial or oblique lines in
Passiflora p.p., e.g. P.gigantifolia Harms and P. emarginataVL. B. et K. tending
;
678 PASSIFLORACEAE
to be grouped tangentially in Tacsonia. Variable in number even within a
genus, e.g. 2-5 per sq. mm. in Paropsia vareciformis Mast, and 26 per mm. in
P. schliebenii Sleumer, less than 5 per mm. also in Passiflora p.p. and 20 or
more per mm. in Crossostemma and Soyauxia', tending to be ring-porous in
Passiflora p.p. (1851) and in Paropsia schliebenii. Perforations usually simple
only, but with a few scalariform plates in Androsiphonia and Mitostemma
1
glaziovUWLast. (2I58); perforation plates exclusively scalariform in Soyauxia.
Intervascular pitting typically alternate, minute, e.g. in some species of
Paropsia and Smeathmannia, to moderately large, e.g. in Dilkea, usually with
very distinct and sometimes exserted horizontal apertures that may coalesce,
e.g. in Crossostemma and Smeathmannia; scalariform to opposite in Soyauxia;
pits to wood and ray parenchyma usually similar to the intervascular pitting
but sometimes larger, rounded or oblong, or nearly simple in some species of
Dilkea, Mitostemma (2158), Passiflora, and Tacsonia, and scalariform in
Passiflora gigantifolia and Soyauxia. Contents rare, tyloses observed in occa-
sional specimens of Dilkea and Passiflora. Mean member length usually
o*5~i-o mm., but up to 2-0 mm. in Soyauxia grandifolia Gilg et Stapf.
Woodworth (2465, 2466) has recorded the occurrence of an unusual type of
cell, which he terms a'fibriform vessel member', in several species (all that
were examined) of Passiflora and in Smeathmannia pubescens Soland. and
Tacsonia mollissima H. B. et K. ; these cells are similar in length, shape, &c., to
the fibre-tracheids, being twice as long as the fusiform cambial initials, but
have simple perforations. Similar cells were observed by the author in
Paropsia vareciformis Mast., but none were observed in Androsiphonia or
Soyauxia. Chalk and Chattaway (358) note the occurrence of perforated ray
cells in Androsiphonia. Parenchyma moderately to very abundant, most
typically apotracheal, as scattered cells and numerous uniseriate bands
(Fig. 153 D and E); the bands very irregular and up to 4 cells wide in Paropsia
vareciformis\ less abundant and mainly as scattered cells in Soyauxia (Fig.
153 A); intermediate between broad metatracheal bands and aliform in Dilkea
(Fig. 153 H); vasicentric only in Crossostemma. Crystals present in the
ordinary cells in Paropsia and in chambered cells in Tacsonia. Strands often
of more than 8 cells. Rays exclusively uniseriate in Soyauxia and almost so
in Androsiphonia, up to 2-3 cells wide in Paropsia, Passiflora p.p., and
Smeathmannia, up to 5-8 cells in some species of Adenia (2158), Crossostemma,
Dilkea, Keremanthus (2158), Passiflora (2158), and Tacsonia, fewer and with
some procumbent cells in Crossostemma', multiseriate rays typically high, less
than i mm. only in Tacsonia uniseriate rays usually numerous, and composed
;

entirely of square to upright cells; often high; 8-18 rays per mm., least
numerous (8-10 per mm.) in Crossostemma and Tacsonia and most numerous
(15-18 per mm.) in Androsiphonia, Dilkea, and Passiflora p.p.; usually com-
posed almost entirely of square or upright cells, with square cells in the
centre, except in Soyauxia (Fig. 153 B), in which rows of square cells alternate
with upright cells (Kribs's Type Heterogeneous III); the central, multi-
seriate parts in Crossostemma laurifolia Planch., Paropsia vareciformis, and
Tacsonia mollissima H. B. et K. consist of procumbent cells, with uniseriate
margins of square or upright cells, the uniseriate margins commonly of 10 or
1
Harms (894) refers to the abundant occurrence of scalariform perforation plates in
Barteria, Paropsia, Paropsiopsis, and Smeathmannia.
 PASSIFLORACEAE 679
more cells, except in Tacsonia; consisting mainly of procumbent cells in
Crossostemma. In woods with few or no procumbent cells, particularly
Androsiphonia, Passiflora p.p., and Smeathmannia^ the cells, as seen in cross-
section, are barely distinguishable in size and shape from the wood parenchyma
cells, and Solereder notes that in Tryphostemma littorale Engl., and some

species of Adenia, the cells are often elongated more in the tangential than in
the radial direction. Crystals observed in the ordinary cells of Crossostemma,
Dilkea, Paropsia, Smeathmannia, and Tacsonia, often abundant. Fibres with
bordered pits, which are often large and distinct in Crossostemma, Dilkea,
Smeathmannia, Soyauxia, and Tacsonia, and also, according to Solereder, in
some species ofAdenia, Hollrungia, Passiflora, and Tryphostemma\ with simple
pits in some species of Androsiphonia, Paropsia, and Passiflora and, according
to Solereder, of Adenia and Keramanthus; walls moderately to very thick, the
latter particularly in the species with simple pits. Mean length 1*9-2*9 mm.,

longest in Soyauxia. Woodworth (2465) notes some libriform fibres in

Passiflora spp., a preponderance of libriform fibres, with very few fibre-
tracheids, in Smeathmannia pubescent Sol., and only a few fibre-tracheids in
Androsiphonia. Woodworth (2466) also refers to short septate fibre-tracheids
in Passiflora vitifolia H. B. et K.

TAXONOMIC NOTES
(i) FROM GENERAL ANATOMY
This family appears to have points in common with the Flacourtiaceae.
It is, in fact, difficult to decide to which of the 2 families some of the genera

belong. Harms
(894) has pointed out that Barteria, Paropsia, Paropsiopsis,
and Smeathmannia differ from most members of the family in having frequent
scalariform perforation plates to the vessels, as well as in the presence of
sclerenchymatous elements between the primary groups of fibres in the
pericycle.

(ii)
FROM WOOD STRUCTURE
Soyauxia has been included here, following Hutchinson. In some respects
it appears to have more in common with the Flacourtiaceae, cf. Idesia\ on the
other hand, it might be fitted into the Passifloraceae on the assumption that
it is a primitive type, of which there is ample evidence in the wood structure.

No 'fibriform vessel members' have been observed in Soyauxia, but its

ordinary vessel members are extremely long and rather narrow and are
actually longer than the 'fibriform vessel members' measured by Wood-
worth (2466).
Smeathmannia has been included here rather than in the Flacourtiaceae and
kept separate from Paropsia, though the woods of the 2 genera are very similar.
The occurrence of 'fibriform vessel members' in both genera supports their
affinity with Passiflora and Tacsonia, though there are many very obvious
differences between them; some of the latter may perhaps be related to
differences in habit.
Tacsonia has been kept separate from Passiflora. Woodworth (2466) found
macerated material of Tacsonia mollissima H. B. et K. to be strikingly similar to
that of Passiflora spp. On the other hand, in the limited material available
to the author, the 2 genera appeared to be distinct.
68o PASSIFLORACEAE
ECONOMIC USES
Species of Passiflora, known as Passion Flowers, are cultivated in Great
Britain under glass or in sheltered places out of doors. The fruits of certain
species of Passiflora are commonly eaten in tropical countries, especially
America and the West Indies. The best-known edible species are Grenadillas
(P. quadrangularis Linn, and P. macrocarpa Mast.), Belle Apple (P. laurifolia
Linn.), Sweet Cup or Pomme d'Or (P. maUformis Linn.).

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Abatia, Adenia, Barteria, Crossostemma, Deidamia, Hollrungia, Mito-
stemma, Paropsia, Paropsiopsis, Paschanthus, Passiflora,* Smeathmannia,
Soyauxia, Tacsonia, Tetrastylis,* Tryphostemma.
*
Represented in the Kew Slide Collection.

(ii)
FOR WOOD STRUCTURE
(Adenia), Androsiphonia, Crossostemma, Dilkea, (Hollrungia), (Kera-
manthus), (Mitostemma), Paropsia, Passiflora, Smeathmannia, Soyauxia,
Tacsonia.
LITERATURE
(i) On General Anatomy
Baccarini 60, Bailey 70, Bird 199, Bohmker 2x6, Brush 298, Gangstad 741, Harms 894.
(ii) On Wood Structure
Burgerstein 310, Chalk and Chattaway 358, Cooper and Record 461, Cozzo 494, Harms
894, Record 1843, 1851, Woodworth 2465, 2466.

148. ACHARIACEAE
SUMMARY
A
small South African family, consisting of the low half-shrub Acharia
tragioides Thunb., and the climbers Ceratiosicyos ecklonii Nees and Guihriea
capewis Bolus, the last species being a stemless herb with fleshy roots arising
from a rhizome. The anatomy has not been very fully investigated, and the
following account has been taken from Harms (895) and Solereder.

LEAF
Dorsiventral in Acharia and Ceratiosicyos, with about i layer of palisade
tissue. Hairs in Acharia simple, multicellular, with fairly thick walls.
Stomata in Acharia and Ceratiosicyos confined to the lower surface. Vascular
bundles of the veins not accompanied by sclerenchyma in either genus.

Axis
STEM
Young stemangular in Acharia, but a few layers of cork subsequently
ariseon the inside of the collenchyma in the angles, and in the sub-epidermis
between the angles. Pericycle in Acharia and Ceratiosicyos containing isolated
strands of fibres. Xylem generally including vessels with simple perforations,
but scalariform plates with a few bars recorded locally in Acharia, especially
 ACHARIACEAE 681

in the region of the primary xylem. Vessels of Acharia narrow, radially

arranged, with bordered pits, embedded in a ground-mass of thick-walled,
septate fibres with simple pits. Vessels of Ceratiosicyos mostly wide and
scattered, embedded in ground-tissue consisting of occasionally septate,
prosenchymatous, simple-pitted elements intermixed with parenchyma, the
latter being mostly situated around the vessels. Rays broad in Ceratiosicyos.

TAXONOMIC NOTES
The genera in this family were included under Passifloraceae in the Bentham
and Hooker system, but they were placed in a separate family by Harms (895)
owing to differences in floral structure.

GENERA DESCRIBED
Acharia, Ceratosicyos.

LITERATURE
On General Anatomy
Harms 895*

149. CARICACEAE
(FiG. 154 on p. 682)

SUMMARY
This tropical American family consists of the genera Carica^ Cylicomorpha,
Mocinna, and Jacaratia, members of which are shrubs or small trees with
somewhat fleshy trunks bearing terminal clusters of leaves. The Pawpaw
(Carica papaya Linn.) has received more attention from anatomists than any
of the other members of the family, and the following description refers to this
species, Chatterji (378) has described the anatomical changes which occur in
Carica papaya when affected by a physiological disease. See under 'Axis'
on p. 683.
LEAF
Dorsiventral, with well-developed intercellular spaces in the spongy tissue.
Long, club-shaped, glandular hairs with multicellular heads present on the
petiole and along the principal veins. Stomata confined to the lower surface;
ranunculaceous. Petiole (Fig. 154 A), in transverse sections through the
distal end, exhibiting a circle of numerous widely spaced collateral vascular
bundles surrounding a very large, parenchyrnatous pith. Other strands,
consisting wholly of phloem, also occur in the vascular ring interspersed
between the collateral bundles. Articulated laticiferous canals accompany
the vascular bundles of the veins and extend into the surrounding mesophyll,
All parenchyrnatous tissues stated to contain refractive grains of a substance
in the nature of an aldehyde. Clustered crystals of calcium oxalate fairly
abundant.
Axis
STEM (Fig. 154 E)
Glandular hairs, similar to those described for the present on the
leaf,

young internodes of the stem. Stem swollen at the base owing to dilation of
 CARICACEAE, A and E; BEGONIACEAE, B-C and
FIG. 154. F;
PASSIFLORACEAE, D and H; TURNERACEAE, G and I
A, Carica papaya Linn. Petiole x 5. B, Begonia manicata Gels. Stem X 5. C, B. echinosepala
Regel. Petiole xy, D, Passiflora racemosa Brot. Petiole x 13. E, Carica papaya Linn. Stem X5
F, Begonia manicata Gels. Petiole x6. G, Turnera ulmifolia Linn. Petiole Xi8. H, Passiflora
.

racemosa Brot. Stem XI3. I, Turnera ulmifolia 'Lirm. Stem Xi8.

 CARICACEAE 683
the primary cortex and pith. Primary cortex largely composed of collen-
chyma. Pericycle including massive, isolated strands of fibres, in the form
of caps to the well-developed, radially elongated groups of phloem. Secondary
phloem stratified into hard and soft portions. Conducting system consisting
of a circle of narrow, radially elongated vascular bundles, separated from
one another by broad, parenchymatous rays. Ground tissue of the xylem
composed of wedges of unlignified, parenchymatous tissue. Vessels mostly
about 200 p, in radial diameter, solitary, or, more frequently, in approximately
radial multiples of up to 5 or occasionally more members. Lateral walls of
the vessels with reticulate thickening and large, simple, gash-like pits where
in contact with the parenchyma, and almost circular intervascular pitting.
Perforations simple and horizontal. Wood fibres absent. A network of
articulated laticiferous canals, with contents which are stained yellow by
iodine, present in all parts of the ground tissue. Clustered crystals abundant
in the cortex.
Chatterji (378) has described an apparently physiological disease of Carica
papaya in which the leaves at first assume an unhealthy green colour, whilst
the margins become curled. Dark-green blisters also arise on the lamina.
Later on the lamina becomes dissected until, in extreme cases, only the mid-
rib remains. The leaves eventually turn brown and fall off, whilst the whole
plant becomes stunted. Degeneration begins in the laticiferous system, but
later the elements of the secondary phloem are affected. Hypertrophy
eventually sets in, not only amongst the laticiferous elements and phloem,
but also in the mesophyll.

ECONOMIC USES
The large, edible fruits of Carica papaya Linn, are much relished in tropical
countries, where the Pawpaw is Papain, a drug with
widely cultivated.
digestive properties, is obtained from the latex from the unripe fruits.

GENUS DESCRIBED
Carica.*
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Chatterji 378, Harms 896, Stephens 2194.
(ii) On Wood Structure
Howard 1088, Record 1783, 1843, 1851, Record and Hess 1886, Rowlee 1964, Williams
2430.

150. MALESHERBIACEAE
SUMMARY
Malesherbia consists of erect or procumbent herbs or half-shrubs which
occur in dry habitats in the Andes from Peru to Bolivia and in the Argentine.
The following account of the anatomy is taken largely from Harms (893).
LEAF
Isobilateral. Hairs of 2 types, (i) Moderately stiff unicellular trichomes.
 684 MALESHERBIA CEAE
(ii) Long, filiform, multiseriate, frequently glandular hairs, secreting a sub-
stance with an unpleasant smell.

Axis
STEM
Cork arising in the epidermis, sub-epidermis, or cortex in different species;
consisting of thin- walled cells. Cortex
said to include many layers of palisade
cells in M. fasciculata Don. Pericycle containing isolated strands of fibres,

Xylem including numerous, radially arranged vessels, mostly with simple,

circular or elliptical perforations, but scalariform plates with a few bars
recorded in the neighbourhood of the primary wood. Wood fibres consisting
of fairly short elements with very small, slit-shaped or elliptical pits. Rays
mostly 1-2, rarely 3, cells wide.

GENUS DESCRIBED
Malesherbia.

LITERATURE
On General Anatomy
Harms 893.

151. CUCURBITACEAE
(FiG. 155 on p. 686; FIG. 156 on p. 686; FIG. 157 on p. 688)

SUMMARY
Mostly herbs, often scandent or prostrate, but some species tending to be
woody. The family is noted for its rapid vegetative growth. Most members
of the family are tropical. Many species are provided with tendrils of which
those in Cucumis have been interpreted, according to Trinkgeld (2279), as
metamorphosed leaves, whilst those in other genera are homologous with
stems bearing leaves. Tendrils become coiled round cylindrical supports
with which they come into contact, but form anchoring pads when in contact
with flat surfaces. Debberman (553) has shown that in Gymnopelalum contact
between the anchoring pad and its supporting surface is maintained by hair-
like papillaewhich grow out from the pads and fit very exactly into crevices
and depressions in the surface. Debberman concluded, from less detailed
observations on other species, that this mode of attachment is general through-
out the family. Glandular hairs with multiseriate stalks are very charac-
teristic,but simple unicellular or qniseriate types as well as wart-like or spiny
trichomes also occur. Calcareous cystoliths and similar bodies occur, usually
at the bases of the hairs or in adjacent epidermal cells, more rarely in epidermal
cells away from the trichomes. They vary considerably in size and shape in
leaves of an individual species at different stages of development. Extra-
floral nectaries and glandular leaf-teeth are fairly frequent. The stomata,
which occur on both sides of the leaf or are confined to the lower surface, are
ranunculaceous. They are frequently raised above the general level of the
epidermis, especially in stems. The petiole, in transverse sections through
the distal end, exhibits a crescent or circle of vascular bundles of which the
 C UC URBITA CEAE 685

larger ones are bicollateral. In the axis the most noteworthy feature is the
predominantly bicollateral vascular bundles, separated from one another
by broad strips of ground tissue, and frequently arranged in 2 rings. Their
course in the axis has been described by Zimmerman (2508), Manteuffel
(1435), and other authors. The xylem in the old stems sometimes becomes
cleft through the development of secondary medullary rays. The sieve tubes,
which are usually large and conspicuous, occur scattered in the cortex as well
as in the phloem in a number of genera. There is a closed ring of scleren-
chyma in the outer part of the cortex of herbaceous species, and a continuous
ring of fibrous cells in the pericycle of young stems, but this may become
discontinuous when older. Anomalous structure is fairly frequent in thick
stems and roots.

LEAF
Usually dorsiventral, more rarely isobilateral. Hairs include the following
types, (i) Simple, unicellular or uniseriate, sometimes accompanied by sub-
sidiary cells at the base, (ii) Wart-like or spiny trichomes in species of Bryonia,
Cucumis, Cucurbita, Ecballium. (iii) Glandular hairs with uniseriate stalks of
variable length and spherical or disk-shaped heads in species of Abobra,
Benincasa, Citrullus, Corallocarpus, Cucumis, Cucurbita, Cyclanthera, Ecbal-
Hum, Fevillea, Gynostemma, Kedrostis, Lagenaria, Luffa, Melothria Momor* y

dica, Trichosanihes, Zanonia. (iv) Explosive hairs with uniseriate stalks of

5 or 6 cells and 2-celled heads in Curcurbita and Momordica, According to
Zimmerman (2508) the whole head of each hair is explosively cast off and
the contents extruded through the pore thus formed, (v) Water-containing
margins; those of Momordica spp. multi-
hairs, chiefly situated at the leaf
cellular, elongated; with pointed apices in species of Bryonopsis, Cocdnia,
Melothria, Raphanistrocarpus. (vi) A network of uniseriate hairs, covered
with waxy material, recorded in Telfairia. (vii) Hooked, calcified hairs in
Peponium. (Types iv to vii recorded by Zimmerman, 2508.) Leaf-teeth
sometimes secretory, e.g. in Cucumis. Extra-floral nectaries, occurring all
over the lower surface or confined to the bases of the leaves, recorded in
species of Abobra, Adenopus, Alsomitra, Bryonia, Cephalandra, Cucurbita>
Luffa, Momordica, Sphaerosicyos, Trichosanthes. Large,
Fevillea, Lagenaria,
black glands (i opposite pair near the distal end of the petiole, and others on
the lower surface of the lamina near the vein endings) noted at Kew in
Sphaerosicyos sphaericus (E. Mey.) Hook, f. The material secreted appeared to
be mucilaginous. Cells of the epidermis particularly large in Japanese
species of Actinostemma, Benincasa, Citrullus, Cucumis, Cucurbita, Gyno-
stemma, Melothria, Momordica, Schizopepon.
Stomata confined to the lower side or present on both surfaces ranuncu-
;

laceous frequently raised on papillose projections above the general level of

;

the epidermis in the same way as in the stem (see p. 687), although in fewer
species. Mesophyll containing silicified cells in Fevillea and Zanonia. Mid-
rib reported by Solereder to be provided with: (i) A
single vascular bundle in
species of Actinostemma and Schizopepon, (ii) A
large strand accompanied by
a smaller one in Melothria sp. (iii) A
large bundle and 2 small lateral ones in
Gynostemma sp. (iv) Alarge bundle with 2 smaller ones above it in species of
Benincasa, Cucumis, Lagenaria. (v) Four bundles arranged in the form of a
 B

FIG. 155. CVCURBITACEAE

Double cystoliths and groups of cystoliths in Momordicai A, B, Momordica sp.; C, Momordica
charantia L. After Penzig.

FIG. 156. CUCURBITACEAE

Deposits of calcium carbonate in the leaf of Hanburia mexicana Seem. A, Incrusted group of cells
in a transverse section of the leaf; B, in surface-view (before decalcification the central epidermal cells
;

lying above the cell-group are not figured) C, in surface-view after decalcification.
; By Solereder.
 CUCURBITACEAE 687
cross in Luffa, Momordica, Trichosanthes. (vi) A
ring of bundles in species of
Citrullus and Cucurbita. Petiole, in transverse sections through the distal
end, exhibiting an almost closed crescent or circle of bundles, the larger ones
bicollateral, in Bryonia (Fig. 157 D), Cucumis, Cucurbita, Cyclanthera (pro
parte), Ecbattium, Lagenaria, Luffa, Melothria, Sicyos. Bundles likewise
separate but arranged in a slightly more open crescent in species of Alsomitra
(Fig. I57F), Cyclanthera (pro parte) (Fig. 157 B), as well as in Echinocystis.
Chakravarty (348) has also drawn attention to the constancy in the number
and structure of the vascular bundles in the midribs and petioles of cucurbi-
taceous leaves, which he believes to be of generic diagnostic value.
Cystoliths (Figs. 155 A-c and 156 A-C), varying widely in size and shape
and frequently visible to the naked eye as white areas, commonly occur at the
bases of hairs and in neighbouring epidermal cells of numerous genera and
species. Less frequent in epidermal and adjacent mesophyll cells not immedi-
ately associated with the hairs. The form and size of cystoliths vary consider-
ably within a species at different stages in the development of the leaves;
those in species of Adenopus, Cucumis, Cyclantheropsis, Gerrardanthus,
Melothria described by Zimmerman (2508) as becoming disorganized and
disappearing from the leaves when sufficiently old. Crystals infrequent in or
absent from many genera and species recorded, chiefly as clusters, in a few
;

species of Gerrardanthus and Momordica (Zimmerman 2508) and solitary and

clustered crystals together with crystal-sand in the same genera (Chakravarty
348).

Axis
STEM (Fig. 157 A, c, E)and
Frequently angular, with collenchyma in the ribs. Stomata raised above
the general level of the epidermis, at the apices of small projections, in
species of Adenopus, Benincasa, Cucurbita, Luffa, Momordica, Peponium,
Physedra, Sechium, Sphaerosicyos. Stomata said by Zimmerman (2508) to be
at the same level as or only slightly raised in species of Coccinia, Cyclantherop-
sis, Gerrardanthus, Kedrostis, Melothria, Momordica, Raphanistrocarpus,
Telfairia, Trochomeria, Cortex often including a large proportion of
collenchyma, especially in the ribs, but with the collenchyma sometimes inter-
rupted by patches of assimilatory tissue extending to the epidermis. Outer part
of the cortex, containing a sinuous ring of sclerenchyma, the latter being con-
tinuous in young stems, hut becoming interrupted during secondary thicken-
ing, although the individual groups sometimes become more or less reunited
by secondary sclerenchymatous cells. Cortex and medullary rays in old
stems sometimes including groups of stone cells. Cork not observed in many
species, but, whenseen, arising at different levels, e.g. according to Zimmer-
man (2508) in the sub-epidermis in Kedrostis sp. ;
in the outer part of the cortex
in Cyclantheropsis sp. y on the inside of the sclerenchymatous ring in the cortex
in Melothria sp. Solereder records it as arising in the cortex in Zanonia
indica Linn, and near the outer periphery of the pericyclic sclerenchyma in
Trichosanthes. Pericycle containing a continuous ring of fibrous cells in very
young stems, but the ring becomes discontinuous when older. Vascular
bundles widely separated by broad strips of ground tissue; nearly always
bicollateral and frequently arranged in 2, more or less distinct, circles;
 688 CUCURBITACEAE
vascular bundles of the inner ring almost meeting at the centre of the stem in
some species; approximately constant in number and arrangement in an
individual species according to Ghosh (762) and Zimmerman (2508), provided
attention is confined to material of comparable age. Pellisier (1684) has
emphasized, however, that the number of bundles is greater in transverse

FIG. 157- CUCURBITACEAE

A, Cucurbita sp. Stem x 35. B, Cyclanthera explodes Naud. Petiole X55. C, Ecballium elaterium
A. Rich. Stem X 12. D, Bryonia dioica Jacq. Petiole X3O. E, Acanthosicyos horrida Welw. Stein
Xi 8. F, Alsomtra sarcophylla M. Roem. Petiole xa6.

sections of old compared with young internodes of Cucurbita. Vascular

bundles, situated outside the sclerenchymatous ring in the cortex, recorded
by Zimmerman in a few species of Actinostemma and Momordica.
Zimmerman recognized the 3 following arrangements of vascular bundles
in the young internodes. The differences are not constant throughout the
genera mentioned, (i) Five bundles in the outer and 5 in the inner circle in
Adenopus, Blastania, Bryonopsis, Coccinia^ Coraltocarpus, Cucumis, Kedrostis,
Luffa, Melothria, Momordica, Physedra, Raphanistrocarpus, Trochomeria.
(ii)
Internode with less than 10 bundles in a few species ofCucumis, Kedrostis,
Melothria^ Momordica. (iii) Internode with more than 10 bundles, i.e. with
2 circles of 5 bundles each, accompanied by accessory, frequently reduced
bundles in a few species of Coccinta, Cucurbita, Gerrardanthus, Lagenaria>
 CUCURBITACEAE 689
Peponium, Sphaerosicyos, Telfairia, Zimmerman also classified stems accord-
ing to the arrangement of the sieve tube tissue.
Further particulars concerning the course of the vascular bundles in Bryonia
have recently been recorded by Bouvrain (248, 249) and Fourcroy (700,
701).
Collateral strands, formed through loss of the inner phloem, are present
amongst the bicollateral bundles in numerous genera and species. Bundles
exclusively collateral in species of Akomitra, Anisosperma, Cyclantheropsis,
Gerrardanthus\ bicollateral structure also stated by Solereder to be obscure
in species of Fevillea, Gynostemma, Zanonia. Sieve tubes wide, with con-
spicuous transverse sieve plates; sometimes occurring in the cortex as well as
in the phloem in certain genera, e.g. in species of Cucumis, Cucurbita, Ecbal-
lium Lagenaria, Zanonia\ those of the cortex and phloem ^respectively being
y

united to one another in at least some species. Xylem and phloem often
become dissected by the formation of secondary rays produced by the
fascicular cambium, and the xylem by large groups of unlignified elements
also derived from cambial cells. Complete collateral bundles sometimes develop
in the secondary rays in species of Cyclantheropsis and Gerrardanthus. Inter-
xylary phloem (see 'Anomalous Structure* on p. 690) sometimes arises in
the unlignified tissues of the xylem, e.g. according to Zimmerman in old stems
of a few species of Adenopus, Luffa, Melothria, Momordica, Physedra, Sphaero-
sicyos. Interfascicular cambium in old stems of some genera also serves to
enlarge the primary medullary rays with additional unlignified tissue during
secondary thickening, but, in some instances, phloem and xylem are also
derived from it, e.g. according to Solereder in Lagenaria. Xylem, in very
young stems, including narrow vessels, but those formed subsequently have
conspicuously wide lumina, as is usual in rapidly growing scandent herbs and
lianes; perforations simple. Tyloses common, especially in old stems;
frequently becoming thick walled. Giant nuclei, up to 85 JJL in diameter,
recorded by Scott (2072) in the developing vessels of Echinocystis macrocarpus
Britton, and others up to 66 p in diameter in Cucurbita pepo L. Pith becoming
hollow in manyspecies, but persistent and lignified in others.
Bews (192) has described, in an undetermined species of Benincasa, how
the central cavity of the pith becomes filled with a mass of tissue originating
from a single cell which intrudes into the hollow centre of the stem. A
secondary cavity arises within the intruding tissue, and becomes lined with
thick-walled cells with abundant contents, while simple pluricellular and less
frequent glandular hairs, resembling those on the outside of the stem, project
into the secondary cavity from the surrounding cell layers.
Secretory elements apparently not recorded, and definitely absent from
species in the Kew slide collection. Cystoliths, see p. 687. Crystals infre-
quent in or absent from most genera and species; clusters recorded by
Zimmerman in the ground tissue, particularly of old stems, in species of
Gerrardanthus, Luffa, Momordica Physedra.
y

ROOT
Primary structure said by Holroyd (1081) to be tetrarch in 4 representative
seedlings.
4504 yy
690 CUCURBITACEAE
TUBEROUS ROOTS AND RHIZOMES
Tuberous roots and rhizomes exhibit a well-developed, amylifous, ground
tissue, with weakly developed conducting elements embedded in it, according
to Scott (2072) and Zimmerman (2508).
Root tubers of Coccinia engleri Gilg. described as follows by Zimmerman.
Covered externally by a brown layer of cork. Ground tissue consisting of
amylifous parenchyma, but cracks developing in it towards the centre of the
root. Xylem, as seen in transverse sections, composed of isolated strands
embedded in the ground tissue, most strands including a large solitary vessel.
Small phloem strands present on the outside of the cambium ring, and others
occur scattered in the ground tissue, particularly towards the inside of the
xylem strands.
Stem tubers of 'Melothria argyrea* somewhat similar in structure but

xylem consisting of scattered strands arranged along certain radii, the phloem
being situated chiefly along the same radii as the xylem.

ANOMALOUS STRUCTURE
The following anomalies have been recorded in roots, (i) Islands of soft
interxylary phloem in species of Cucurbita and Lagenaria. (ii) Bundles of
interxylary phloem, arising through the activity of interfascicular cambium,
in the unlignified tissue of the primary medullary rays of Thladiantha dubia
Bge. and, according to Zimmerman (2508), complete vascular bundles in the
corresponding position in Momordica sp. (iii) Concentric vascular bundles
formed in the wood of Bryonia dioica Jacq. by the activity of cambial tissue
surrounding individual groups of secondary vessels, (iv) Successive rings of
growth arising in the pericycle of Ecballium elaterium A. Rich, (v) Thick
roots of several members of the family provided with a pith containing groups
of intraxylary phloem, the latter sometimes becoming converted to inversely
orientated bundles. For formation of interxylary phloem and other anomalous
tissues in stems see p. 689.

PHYLOGENETIC NOTES
Worsdell (2470), after studying the course of the vascular bundles in the
Cucurbitaceae, both in stems and in Conservative* parts of the axis such as
the peduncle and node, drew conclusions concerning the phylogenetic origin
of bicollateral bundles in general and of those of the Cucurbitaceae in parti-
cular. His chief views are as follows.

(i)
'The vascular system of the Cucurbitaceae represents the vestige of a former
ancestral scattered system of bundles such as obtains in the Monocotyledons, of which
only two series of rings remain in perfect condition, the rest appearing in the form
of rudimentary external phloem strands (rarely bundles as well), "internal phloem"
l

strands, and medullary bundles or phloem strands.' (ii) ln the vegetative stem of
certain members of the order (family), and, as a rule, in the lower part only, the
internal phloem exists in the form of vascular bundles of which the xylem exists
"
entirely or for the most part, on the outer side.' (iii) 'The bicollateral" bundle of
the Cucurbitaceae is a compound structure consisting of the more or less intimate
association or attachment of two distinct vascular bundles, of which the innermost
has lost the xylem.'
 CUCURBITACEAE 691

Chakravarty (348) has attempted to arrange some of the genera within the
family in a phylogenetic sequence, assuming for this purpose that reduction
in the number of vascular bundles is an advanced character.

ECONOMIC USES
The members of this family are the well-known gourds.
fruits of certain
Familiar gourds include Cucumbers (Cucumis sativus Linn.), Melons (Cucumis
melo Linn.), Water-melons (Citrullus vulgaris Schrad.), Vegetable Marrows
(Cucurbita pepo Linn.), whilst many others are familiar in tropical countries.
Loofahs consist of the vascular systfcrn of the fruits of Luffa cylindrica Roem.
and other species of Luffa. For particulars of the development of this vas-
cular system see Sinnott and Bloch (2117). White Bryony or English Man-
drake, at one time used to allay coughing in pleurisy, is the root of Bryonia
dioica Jacq.

GENERA DESCRIBED
Abobra, Acanthosicyos,* Actinostemma, Adenopus, Alsomitra,f* Aniso-
sperma, Benincasa, Blastania, Bryonia,* Bryonopsis, Cephalandra, Citrullus,
Coccinia, Corallocarpus, Cucumis, Cucurbita,* Cyclanthera,* Cyclantheropsis,
Ecballium,* Echinocystis,* Fevillea, Gerrardanthus, Gymnopetalum, Gyno-
stemma, Hanburia, Kedrostis, Lagenaria, LufFa, Melothria, Momordica,
Peponium, Physedra, Raphanistrocarpus, Schizopepon, Sechium, Sicyos,*
Sphaerosicyos, Telfairia, Thladiantha, Trichosanthes, Trochomeria, Wil-
brandia, Zanonia.
t Alsomitra Roem = Neoalspmitra Hutch.
*
Represented in the Kew slide collection.

LITERATURE
On General Anatomy
Barkley 141, Bews 192, Bouvrain 248, 249, Chakravarty 348, Cogniaux 441, Crafts 496,
Debberman 553, Fourcroy 700, 701, Ghosh 762, Hagerup 865, Holroyd 1081, Hutchinson
1116, Manteuffel 1435, Pellissier 1683, 1684, Sabnis 1977, Scott 2072, Balwant Singh
2513, Sinnott and Bloch 2117, Trinkgeld 2279, Werner 2414, Worsdell 2470, Zimmerman
2508.

152. BEGONIACEAE
(Fie. 154 on p. 682; FIG. 158 on p. 692; FIG. 159 on p. 692)

SUMMARY
A mainly tropical family consisting chiefly of herbs which are often suc-
culent, but some species are shrubby or scandent, the xylem in the climbers
often being excentrically developed. Some species possess rhizomes while
others are stemless but provided with a basal tuber. The tissues have a well
developed capacity for producing adventitious roots, and some of the orna-
mental begonias are cultivated from leaf cuttings. The hairs include various
multicellular, non-capitate types as well as others with heads composed of a
few or of many cells. The stomata, confined to the lower surface, are generally
very characteristic in appearance owing to their being surrounded by 3-6
subsidiary cells often arranged in 2 rings. The stomata in some species are in
 A

FIG. 158. BEGONIACEAE

A Stomatal group of Begonia fuchsiaefolia (A. DC.) Warb. B, Branched idioblast from the leaf of
Scheidw. D, Double cystotyle of
Begonia arborescens Raddi. C, Double cystotyle of B. luxuriant
B. paleata A. DC. after treatment with alcohol. E, Double cystosphere of B. laetevirens
Van Houtte.
After Fellerer.

FIG. 159. BEGONIACEAE

B. * lan h
Trichomes of: A, Begonia hispida Schott. B, B.pawflora Poepp. et Endl. C,
;
;
f?*
D B smconeura Liebm.; E, B. sinuata Wail.; F, '-B. pretontenw Moore*; G, B. carohmaefoha%<:'
Reg.,
\

H, B. xtnthina Hook.; I, JB. rhisocarpa Fisch. After Fellerer,

 BEGONIACEAE 693
definite groups with relatively large areas of unperforated epidermis between
them in other species the stomata are solitary.
;
A
hypoderm of large water-
storage cells is frequently present towards one or both surfaces. The meso-
phyll sometimes contains sclerenchymatous idioblasts which may be crystalli-
ferous. The petiole is supplied by isolated vascular bundles, some of which
are in the medullary position in certain species. In the stem there is a ring
of widely spaced vascular bundles which are sometimes accompanied by
medullary strands, and more rarely by others in the cortex. In the xylem the
vessels exhibit a characteristic appearance, owing to their having scalariform
bordered pits where they are in contact with one another, whilst there are
both simple perforations and scalariform plates which often occur together
in the end walls. Several kinds of cystolith are common in the leaf and
occasionally in the stem. Crystals are both solitary and clustered, often in
the form of octohedra or quadratic prisms.

LEAF
Hairs (Fig. 159) include the following types.
A. Non-capitate
Always multicellular, including uniseriate, multiseriate, shaggy, and various
other types. Outer cells of the shaggy hairs sometimes with free mamilliform
tips, thus resembling cones of one of the Coniferae (Fig. 159 A). Shaggy
hairs in other instances consisting only of shortly spinous, star-like structures
(Fig. 159 B), or 2-armed. Transitions between shaggy hairs and emergences
also occur, these sometimes including sclerenchymatous mechanical cells.
Whip-like (Fig. 159 D), stellate (Fig. 159 E) and tufted hairs also recorded.
B. Capitate (Fig. 159 F, G, i)

variable; the few- or many-celled head being spherical, ellipsoidal,

Very
club- or hammer-shaped; stalk also varying in length and structure.
Multicellular, easily detached, shortly stalked pearl glands also occur, but
usually become detached from herbarium specimens whilst being dried and
mounted.
Epidermis on both surfaces usually consisting of large, thin-walled cells;
frequently papillose, especially on the upper surface; red cell sap common in
the lower epidermal cells. Cuticle thin, granular, striate or verrucose. One
or many layers of large-celled hypoderm occur beneath one or, more fre-
quently, both surfaces in many species; especially well developed and con-
stituting most of the thickness of the leaf in Begonia fuchsioides Hook.
Stomata not numerous, confined to the lower surface, each pair of guard
cells surrounded by 3-6 subsidiary cells, often arranged in 2 rings; solitary or
in definite groups (Fig. 159 A), the latter often recognizable with the naked
eye as white dots. Hydathodes occur sporadically on the upper surface near
the leaf margin. Mesophyll generally including i layer of palisade tissue;
more rarely consisting of ordinary parenchyma. Palisade cells in some species
short and conical, in others elongated and provided with concertina-like
foldings of the lateral walls. Assimilatory tissue restricted to a narrow band
in the innermost part of the lamina in Begonia fuchsioides. Air-filled spaces
between the epidermis and palisade cells give a silvery sheen to the leaves
 694 BEGONIACEAE
of certain species. Stone cells, idioblasts (Fig. 158 B), and similar mechanical
elements fairly frequent in the mesophyll or around the veins; palisade cells
sometimes locally replaced by stone cells. Ground tissue around the larger
veins often containing strongly developed collenchyma. Vascular bundles
sometimes accompanied by elongated parenchymatous cells. Petiole, in
transverse sections through the distal end, exhibiting isolated vascular bundles,
arranged in a circle in some species, e.g. jB. echinosepala Reg. (Fig. 154 c), but
others, e.g. B. manicata Cels. (Fig. I54F), including medullary strands as
well. Outer part of the petiole frequently collenchymatous. Crystals, both

solitary and clustered, often occurring in the form of single octohedra or

quadratic prisms. Several kinds of bodies, resembling cystoliths (Fig. 1 58 C-E),
occur in the palisade and spongy tissue of the mesophyll or occasionally in
the hypoderm or in the shaggy hairs of certain members of the family;
commonly giving rise to transparent dots. Some of these bodies resemble
those of Momordica (Cucurbitaceae). The following types are recorded by
Solereder. (i) Bodies devoid of crystalline infiltration, appearing as white
translucent structures in dry sections exposed to the air, but swelling up and
forming an apparently structureless mass in water, and subsequently con-
tracting and resembling decalcified double-cystoliths with concentric and
radial striations when treated with alcohol, (ii) Bodies exhibiting finely

granular structure and yellowish refraction of light in dry sections exposed to

the air; turning grey and exhibiting the granular structure and stratification
more clearly when treated with water; containing resinous infiltrations soluble
in alcohol, (iii) Brittle, strongly refractive, whitish, yellowish, or brownish
bodies appearing as gelatinous masses in sections mounted dry or in water, or,
in the living plant, as liquid, brightly shining, dull white or yellowish masses
of resinous secretion enclosed in special sacs. Similar types of cystolith-like
bodies often occur throughout closely related groups of species of Begonia,
thus serving as a test of affinity, and, to some extent, valuable for specific
diagnosis. For further details see Solereder.

Axis
STEM (Fig. 154 B)
Epidermis consisting of 1-4 layers. Cork arising in or immediately below
the epidermis. Outer part of the cortex collenchymatous inner part com-
;

posed of thin- walled tissue containing crystals and chlorophyll; sometimes

with red cell sap.Vascular bundles appearing, in transverse sections, as a
single ring, rarely accompanied by others in the cortex, or, more frequently,
in the pith; each bundle subtended on the outer side by elongated parenchy-
matous cells and by prosenchymatous elements with slit-shaped pits. Bundles
of the main ring frequently isolated but sometimes forming a more or less
closed ring. Tissue between the bundles, in most shrubby species, consisting
of elongated cells with lignified walls and slit-shaped pits; the corresponding
tissue in some of the root-climbers is composed of radially elongated or iso-
diametric, thin-walled cells, forming broad rays similar to those of the
Aristolochiaceae. Xylem including vessels in radial rows, and progressively
larger in diameter towards the exterior of the stem; surrounded by paren-
chyma. Vessels also provided with scalariform bordered pits where in con-
tactwith one another; simple, circular perforations, or scalariform plates with
 BEGONIACEAE 695

many bars present in the end walls, both types sometimes occurring together
in a single vessel.Tyloses observed in B. manicata Cels. Ground tissue of
the secondary wood composed chiefly of delicately septate prosenchymatous
elements with simple pits. Pith composed of large parenchymatous cells
with thin, pitted walls.
Pneumatothodes have been described by Vouk (2341) in the stems of
Begonia vitifolia Schott. where they resemble and replace typical lenticels.
The pneumatothodes are composed of (i) an epidermis of small, thin-walled
cells devoid of cuticle (ii) stomata with poorly developed or occluded aper-
;

tures; (iii) thin- walled photosynthetic tissue with a weakly developed inter-
cellular system which constitutes the main portion of the pneumatothode.

ROOT
The modeof origin of adventitious roots from the cambium in certain
species of Begonia has been described by Smith (2143).

TAXONOMIC NOTES
Irmscher (1122) has pointed out that the affinities of the family are not well
established. It is questionable whether the common occurrence of cystoliths
in the Begoniaceae and Cucurbitaceae is of any great taxonomic significance.

ECONOMIC USES
Numerous species of Begonia are cultivated for ornamental purposes.

GENUS DESCRIBED
Begonia.*
* Kew
Represented in the slide collection.

LITERATURE
On General Anatomy
Irmscher nil, Knagg 1248, Smith, A. I. 2143, Vouk 2341*

153. DATISCACEAE
(FiG. 1 60 on p. 696)

SUMMARY
(i) GENERAL
A
family which occurs in northern tropical and sub-tropical regions. The
species of Octomeles and Tetrameks are trees, the last genus being charac-
terized by large buttresses. Datisca, the remaining genus, consists of shrubs.
The anatomy of the family, apart from the wood structure, has not been very
fully investigated. Distinctive characters are lacking, but the sclerenchyma-
tous idioblasts which, according to Solereder, traverse the entire thickness
of the leaf of Octomeles sumatrana Miq. are noteworthy.

(ii)
WOOD
Vessels large, perforations simple, intervascular pitting alternate, pits to
parenchyma conspicuous, simple and irregularly elongated; members of
medium length. Parenchyma paratracheal, vasicentric to slightly aliform,
696 DATISCACEAE
storied. Rays up to 4-7 cells wide, with few uniseriates, 2-3 stories high,
heterogeneous. Fibres with small simple pits, very rarely septate, vaguely to
distinctly storied; of medium length.

LEAF
Dorsiventral, Shaggy hairs, each with a multicellular stalk of variable
length, and a spherical or ellipsoidal, multicellular, glandular head, recorded

FIG. 160. DATISCACEAE, A-B; CACTACEAE, C-D

A, Octomeles sumatrana Miq. B, O. sumatrana Miq. C, Carnegiea gigantea Britt. et Rose. D,
Lcmaireocereus hystrix Britt. et Rose.

in Datisca; scale hairs, each having a 4-seriate stalk bearing a i -layered,

circular shield with an entire margin, present in Octomeles. Stomata occur
on both surfaces in Octomeles, although more plentiful on the upper than on
the lower side; confined to the lower surface in Datisca\ ranunculaceous in
both genera. A i -layered
hypoderm recorded beneath the upper epidermis
in Octomeles. H-shaped extending from the upper to the lower
idioblasts,
epidermis, also occur in Octomeles. Small prismatic or needle-shaped
crystals recorded in Octomeles.

Axis
YOUNG STEM
The following details refer to Octomeles except where stated, Cork com-

posed of relatively thin- walled cells with wide lumina; arising superficially in
 DATISCACEAE 697
material examined at Kew. Branched sclerenchymatous idioblasts and groups
of stone cells occur in the pith and/or cortex. Pericycle containing a com-
posite and continuous ring of sclerenchyma. Isolated bands of fibres with
wide lumina recorded in the corresponding position in Datisca. Secondary
phloem containing groups of fibres except when very young. Xylem in
young stems exhibiting similar characters to those described below under
'Wood', but rays 1-3 cells wide. Vessels with simple perforations.

WOOD 60 A and B)
(Fig. 1

Vessels large (more than 200 p mean tangential diameter) or slightly

smaller; solitary and
in multiples of 2 or 3 cells, occasionally with a slight
tendency to an oblique pattern, e.g. in Octomeles sumatrana Miq, ; mostly
2-4 per sq. mm., though sometimes more numerous locally (up to about
9 per sq. mm.). Perforations simple. Intervascular pitting alternate, medium-
sized; pits to ray and wood parenchyma cells typically simple, though
Solereder mentions some bordered pitting in Octomeles, and with transitions
to simple pitting in Datisca; large and oblong, elongated horizontally,
obliquely or vertically. Tyloses sometimes present, but not abundant. Mean
member length 0-4-0-5 mm. Parenchyma paratracheal; vasicentric, forming
sheaths 1-2 cells thick round the vessels in Octomeles, rather more abundant
and slightly aliform in Tetrameles; scarce in Datisca (777). Strands usually
of 2-4 cells, occasionally up to 8 cells. Storied. Rays up to 4-7 cells wide;
both Brown (1679) and Janssonius (i 154) refer to the rays as being of 2 sizes;
i mm. or more high; uniseriates very few in Octomeles and Tetrameles small t

and numerous in Datisca (777); composed of square to upright cells; rays

usually 3-4 per mm. heterogeneous (Kribs's Type II B ?), with 1-3 marginal
;

rows of upright cells; Janssonius mentions sheath cells in Tetrameles nudifiora

R. Br. Contents inconspicuous. The multiseriate rays more than i story
high and the uniseriates too few to be storied. Fibres with very small simple
pits, mostly on the radial walls Janssonius refers to septate fibres in a single
;

specimen of Tetrameles nudiflora, and Gilg (777) states that the fibres are
occasionally septate in Octomeles. Walls thin to very thin. Distinctly storied
in Tetrameles and vaguely so in Octomeles. Tapering abruptly and arranged
in distinct radial rows. Mean length 0-9-1-4 mm.

ROOT
Tubercles on the roots of Datisca cannabina Linn, have been investigated
by Severini (2081).
TAXONOMIC NOTES
Although the of the family have been much disputed, it is, accord-
affinities

ing to Gilg (777), generally regarded as related to the Begoniaceae.

ECONOMIC USES
The light, perishable timbers of Octomeles and Tetrameles are used locally
for packing cases. Desch (574) notes that logs of Octomeles sumatrana Miq.
have been exported to Japan for paper pulp, but that the wood has elsewhere
been reported on as unsuitable for this purpose.
 698 DATISCACEAE
GENERA DESCRIBED
Datisca,* Octomeles, Tetrameles.
* Kew slide
Represented in the collection.

LITERATURE
(i) On General Anatomy
Gilg 777, Severini 2081*

(ii) On Wood Structure

den Berger 179, 1 82, Desch 574, Gilg 777, Howard 1088,'Janssonius 1154, Pearson and
Brown 1679, Record 1843, 1851.

154. CACTACEAE
(Fic. 160 on p. 696; FIG. 161 on p. 700; FIG. 162 on p. 700)

SUMMARY
(i) GENERAL
A highly specialized family consisting, for the most part, of stem-succulent
herbs which usually bear sharp, rigid spines. A few members of the family
are definitely shrubby, whilst herbaceous forms may attain a considerable
size. The thick succulent stems, in diiferent genera and species, may be

cylindrical, fluted, or composed of an aggregation of mammiform portions.

Even within a single genus such as Opuntia there are dwarf species, tall-
stemmed forms with a cylindrical woody axis, and thick-stemmed species with
a flattened axis (Reiche 1909). The leaves are usually reduced to scales, so
that photosynthesis takes place chiefly in the green stem, except in Pereskia
which bears typical fleshy leaves. The diverse morphology and anatomy of
the vegetative organs have been fully described by Gravis (806), Reiche (1909),
Haehnel (862), and Vaupel (2327). Most of the species inhabit desert regions
on the American continent, particularly in Mexico, although quite a number,
such as the notorious Prickly Pear (Opuntia), have become naturalized in
other parts of the world. Some species of Epiphyllum, Pereskia, and Rhipsalis
occur as epiphytes in moist woodland habitats, while at Kew even some of
the desert species have been successfully cultivated for several years in jars
of tap water by L. Portheim. The sharp spines, whose morphological nature
has been much disputed, vary in length and thickness, whilst in some genera
they are barbed (so-called glochidia). They are usually grouped in small,
approximately circular areas (areoles) on the surface of the stems. The
areoles frequently bear a dense covering of hairs, forming a velvet down at
the base of the thorns in some species, but in others the hairs are as long as or
even longer than the spines, in extreme cases forming a definite flossy coating
to the plant. Leinfellner (1345) comparatively recently confirmed the earlier
opinion of Goebel and others that, in most genera, the areoles represent short
shoots of which the leaves have become metamorphosed to thorns. Boke's
(218, 219) still more recent developmental studies on Opuntia cylindrica
(Lamarck) DC. also confirmed the opinion that 'the areole is best regarded as
an axillary bud, growing on a persistent leaf base, and that its appendages
should be regarded as the morphological equivalents of leaves. From this it
 CACTACEAE 699
1
also follows that spines and glochids can be homologized with bud scales.
In a species of Pereskia, Leinfellner noted that a short-lived adventitious root
arises from each areole, Stomata are more numerous than in typical Dicoty-
ledonous stems, and the orientation of the pore in relation to the long axis of
the plant is of taxonomic importance. The collenchymatous outer part of the
cortex resembles a gelatinous hypoderm of i to several layers, usually with
pit-like canals connecting the cells. The bulk of the stem is usually composed
of mucilaginous ground tissue in which a cylindrical network of vascular
bundles is embedded. In only a few species is the outer part of the ground
tissue differentiated as palisade. Increase in girth is effected by cell divisions
in the ground tissue as well as by cambial activity. Mucilage cells in most
and laticiferous canals in a few genera occur in the ground tissue. Clustered
crystals of calcium oxalate are often abundant, raphides also occur, but
According to Weingart (2386) and Kummer
solitary crystals are infrequent.
(1301) it sometimes
is possible to distinguish closely related species of
Cactaceae by microscopical differences in the epidermal cells, stomata and
parenchyma. Anomalous Structure recorded in a few species.

(ii) WOOD
Vessels small, except in Pereskia, solitary and in multiples and clusters,

perforations simple, intervascular pitting usually scalariform to opposite, pits

to parenchyma similar and often simple; members of medium length to
extremely short. Parenchyma paratracheal only, a few cells to complete
sheaths round the vessels. Rays mostly 6-10 cells wide and without uni-
seriates, often very high, composed almost entirely of square or upright cells.
Fibres with simple pits, commonly septate, moderately to extremely short.
Tracheids with an unusual type of spiral thickening present in some genera.
Large radial channels common.

LEAF
Very much reduced except in Rudimentary leaves of Opuntia
Pereskia.

provided with abortive stomata; numerous normal stomata present in

Pereskia.

Axis
STEM
Stems generally soiny and in most genera succulent and assuming a variety
*
of forms (see Summary* above); constituting the main part of the plant
body, and serving as the principal assimilatory organ. Spines circular,
flattened or angular in transverse section consisting of a central bundle of
;

thick-walled sclerenchymatous elements surrounded by others with thinner

but pitted walls, the whole thorn being covered externally by radial rows of
epidermal cells with wide lumina. Glochidia or barbed thorns (Fig. 161 D)
with free, imbricate, downwardly projecting points to the epidermal cells also
occur, chiefly in Opuntia. Hairs usually confined to the areoles (see *Sum-
mary'); scanty or short in many species, but in others much longer and more
numerous or even forming a definite floss sometimes unicellular, but mostly
;

uniseriate to multiseriate component cells short near the bases but larger
;

and broader towards the distal ends of the multicellular hairs, cells sometimes
 B

D
FIG. 161. CACTACEAE
Epidermis and Hypoderm: B, Cactus intortus Miller. C, Portion of
A, Cereus variabilis Pfeiffer.
the wood of Echinopsis eyrtesti (Turpin) Zucc. D, Thorn from the leaf-cushion of Opuntia ficus indica
(Linn.) Miller. A-C, after Schleiden, D, by Solereder.

FIG. 162. CACTACEAE

A, B Sphacrites of 'Phyttocactus sp.'; C-E, Proteid-bodies ; C, in Schlumbergera russelliana (Gard-
ner) Britton et Rose; D-E, in Zygocactustruncatus (Haworth) Schumann. A, B after Kohl, C-E after
Molisch.
 CACTACEAE 70!

pitted or superficially striate. Extra-floral nectaries, secreting a sugary

fluid, recorded in species of Cereus, Echinocactus, Epiphyllum, Neomammittaria,
Opuntia, Rhipsalis; those of Epiphyllum (Phyllocactus) stated by Weingart
(2387) to originate as abnormal stomata. For mode of secretion see Lloyd and
Ridgway (1384). Cuticle frequently very thick; sometimes ridged. Surface
of the stem often coated with wax, which is easily detachable, and described
by Vaupel (2327) as sometimes present in sufficient quantities to give a
greyish or white colour to young stems, e.g. in Cephalocereus leucocephalus
(Poselger) Britton et Rose (syn. Cereus houlletii (Lem.) Berg.), and Lemaireo-
cereus pruinoms Otto (syn. C.pruinosus (Pfeiff.) S. Dyck). Stomata numerous
compared with those in a normal herbaceous stem, but less frequent than in
many typical mesophytic leaves; often situated in depressions; with or with-
out subsidiary cells parallel to the pore. Stomatal pores orientated trans-
versely to the long axis of the shoot in species of Cereus, Echinopsis, Pfeiffera,
Rhipsalis\ parallel to the long axis in species of Epiphyllum, Opuntia, Rhipsalis',
irregularly orientated in species of EC hinocactus, Epiphyllum, Neomammillaria*
Stomata in certain members of the family become closed by the development
of thylloid cells, with or without thickenings, which according to Bukvic (307)
arise from the subsidiary cells and/or those of the mesophyll. Long, tubular,
respiratory cavities beneath the stomata sometimes perforate the sub-
epidermal collenchyma. A collenchymatous, mucilaginous, pseudo-hypo-
derm (Fig. 161 A-B) of i to several layers of cells recorded in certain genera
including Cactus, Cereus, Echinocactus, Echinopsis, Opuntia (Nommeusen
1604); sometimes consisting of 6-7 layers in Echinocactus or 12-14 in Cereus ;

the component cells interconnected by branched or unbranched, pit-like

canals. Hypoderm not well developed in Ariocarpus, Epiphyllum, and
Pereskia. Cork arising in the epidermis, hypodermis, or ground tissue; some-
times forming unsightly patches on the surface of the plants, especially,
according to Bukvic (307), when grown in unfavourable conditions. For
further particulars of cork development, including the formation of in-
tumescences in Epiphyllum (Phyllocactus) see Nommeusen (1604), Heinricher
(942), Wolff (2450), and, for wound cork in Opuntia, Coutant (487). Ground
tissue composed chiefly of spherical cells, usually interspersed with well-
developed intercellular spaces except near the epidermis and in .the primary
rays including thick- walled cells in many species of Rhipsalis. Outer part of
;

the ground tissue constituting the chief assimilatory region, but only rarely
differentiated as prlisade, e.g. according to Solereder in Carnegiea giganiea
(Engelm.) Britton et Rose (syn. Cereus giganteus Engelm.). Inner part of the
ground tissue composed of cells with copious, mucilaginous contents.
The cells of the ground tissue remain meristematic for a long time, and,
by dividing, contribute largely to the increase in girth of the axis. This
persistent meristematic power also enables the Cactaceae to be readily pro-
pagated from cuttings or by grafting on other members of the same family.
The vascular network, in transverse sections of most members of the
Cactaceae, appears as a principal ring of separate vascular bundles, accom-
panied, in some species, by additional strands in the cortex, and, less frequently,
the pith. Vessels with simple perforations. Medullary bundles recorded in
Trichocereus candicans (Gillies) Britton et Rose as well as in species ofEchmo-
cactus, Echinopsis, Neomammillaria, but not in N. mammillans (L.) Britton et
702 CACTACEAE
Rose and N. glochidiata (Mart.) Britton et Rose. Cortical bundles, according
to Solereder, arranged in 2 rows in the angles in Rhipsalis crispata (Haworth)
Pfeiffer, R. pachyptera Pfeiffer, and R. rhombea (Salm-Dyck) Pfeiffer; in all
of the angles and sometimes between them in R. micrantha (H.B. et K.) DC.,
R paradoxa Salm-Dyck, R. pentaptera PfeiflFer; situated chiefly in the angles
t

in Lepismium cruciforme (Vellozo) Miquel and Pfetffera ianthothele (Monville)

Weber. Cortical bundles arranged in small, usually circular groups in Hatiora
salicornioides (Haw.) Britton et Rose and in the following species of Rhipsalis:
R. cassutha Gaertn., R. cereuscula Haw., R. cribrata (Lem.) Rumpler, R.
floccosa Salm-Dyck., R. grandiflora Haw., JR. mesembryanthemoides Haw., R.
teres (Vellozo) Steudel. Mechanical support for the main cauline ring of
bundles is provided by thickened elements at the outer periphery of the
phloem in the large but not in the small species. Bundles of the main vascular
ring remain separate for a long time before becoming connected to form a
continuous cylinder following the formation of interfascicular cambium.
Xylem constituting a large proportion of the axis in Pereskia, Laticiferous
canals with white, fairly dense, fluid contents, which sometimes harden and
turn yellow, present in Coryphantha macromeris (Engelm.) Lemaire, Leuchten-
bergia principis Hook., and many species of Neomammillaria. Contents of the
canals consisting of a mixture of resin and rubber-like material, soluble in
chloroform and ether and stained bright red by tincture of alkannin. Canals
isolated when present in the inner part of the plant, but more numerous and
anastomosing in the cortical parenchyma, extending outwards to the inner
boundary of the sub-epidermal collenchyma. Crystals of calcium oxalate
very numerous, usually clustered, the clusters in Opuntia being stellate or
irregular. Raphides recorded in Malacocarpus ottonia Lehmann and in
Opuntia spp., and sphaerocrystals (Fig. i62A-B) locally in Coryphantha
octacantha DC., Neomammillaria wildii (Dietrich) Britton et Rose, 'Phyllo-
cactus\ Selenicereus hamatus (Scheidw.) Britton et Rose, S. pteranthus (Link
et Otto) Britton et Rose. Sphaerocrystalline masses of unidentified material

reported to occur in specimens preserved in alcohol Crystalliferous cells

scattered in many species, but concentrated in a sub-epidermal layer in others,
both arrangements occurring in different species of a single genus in some
instances. sometimes accompany the Crystalliferous cells.
Mucilage cells
For concerning the deposition of mucilage see Stewart (2200).
details
Cortical mucilage cavities also occur in species of Opuntia. According to
Reiche (1911) mucilage, extruded from damaged areoles and other wounds
in Opuntia tomentosa Salm-Dyck, is at first a yellowish-brown fluid but
becomes hard and brittle.
The following arrangements of Crystalliferous and mucilaginous cells
are recorded for certain species of the genera mentioned.

I. Scattered Crystalliferous cells, unaccompanied by mucilage cells in

Ariocarpus, Coryphantha^ Dolichothele, Echinocereus, Echinopsis, Neo-
mammillaria> Pereskia^ Pfeiffera.
II. Crystalliferous cells arranged in a definite sub-epidermal layer, not
accompanied by mucilage cells, in Astrophytum, Echinocactus, Neo-
mammillaria, Opuntia^ Pelecyphora, Pereskia.
III. Scattered mucilaginous and Crystalliferous cells occur together in
 CACTACEAE 703

Cereus, Echinocereus, Echinopsis, Epiphyllum, Gymnocalycium, Lepis-

mium, Malacocarpus, Pereskia, Pfeiffera, Rhipsalis.
IV. A sub-epidermal layer of mixed mucilaginous and crystalliferous cells
in Coryphantha octacantha (DC.) Britton et Rose and Opuntia
subulata (Muhl.) Engelm.

Curiously shaped 'proteid bodies* (Fig. 162 O-E) recorded by Solereder

in the epidermis and adjacent parenchyma of Schlumbergera and Zygocactus.
Saponin said to occur in Cereus.

ETIOLATED SHOOTS
Etiolated shoots of Opuntia have been described by Brown (290) as differing
from the normal in the following respects. Cuticle absent. Surface becoming
covered with small papillae each provided with an apical stoma. Normal
stomata less numerous, Hypoderm of pitted cells, palisade tissue, and sub-
epidermal crystalliferous layer not developed. Air cavities reduced. Etiolated
shoots, when placed in normal light, lose the stomata at the apices of the
papillae by decortication.

WOOD (Figs. 1 60 c and D and 161 c)

Vessels mostly very small (25-50 /z mean tangential diameter), rather
larger in Carnegiea, and medium-sized (100-200 /u,) to large in Pereskia\
solitary, in small multiples and irregular groups or clusters; with a tendency
to a tangential arrangement in Cereus (1894), mostly 10-30 per sq. mm.,
fewer in Pereskia', sometimes semi-ring-porous in Cereus. Perforations
typically simple, but with a few irregularly reticulate plates in Harrisia.
Intervascular pitting typically scalariform to opposite, the pits with wide
apertures and narrow borders; pits round and intermediate between opposite
and alternate, or with no very obvious arrangement in Cereus pits to paren-
;

chyma similar in shapeand distribution and simple. Contents not observed.

Mean length o-i-o*5 mm. Parenchyma typically paratracheal and rather
sparse; vasicentric or only partly enclosing the vessels, sometimes tending to
link together groups of vessels in Carnegiea, Cereus, and Pereskia, and occa-
sionally forming broad bands in the latter; diffuse parenchyma reported (533)
as sometimes present. Strands usually of 2-4 cells. Rays mostly up to 6-10
cells wide, sometimes more than 10 cells wide in Carnegiea and Pereskia', often
several millimetres high, but with various degrees of dissection into shorter
units; uniseriate rays absent; mostly 2-3 rays per mm.; typically composed
almost entirely of square to slightly upright cells, with marginal rows of high
upright cells; with some procumbent cells in Neoabbottia; sheath cells often
present. With occasional druses in Cereus, Dendrocereus, Leptocereus, and
Pereskia. Strands of small tracheid-like vessels observed in a few rays in
i specimen of Leptocereus. Fibres with simple pits, more numerous on the

radial than on the tangential walls. Septate in Cereus, Harrisia, Lemaireocereus,

Leptocereus, Neoabbottia, Nopalea, and Pereskia. Walls moderately thin to
thick. Mean length O'4-o9 mm. Tracheids. According to Schleiden, as
quoted by Solereder, the wood of most of the Opuntias and of Cactus,
Echinocactus, and Neomammillaria is characterized by elementary organs of
quite a special nature (Fig. 161 c); these are broad cells with very thin walls,
704 CACTACEAE
in shape midway between that of a barrel and that of a spindle; they are
provided with locular or spiral thickening ridges, which are inserted on the
thin wall by their narrow edge, and project far into the cell lumen. Generally
these tracheids are also distinguished from the actual vessels by lack of
perforations, but in Echinocactus and Mammillaria rows of such cells occur
connected by perforations. In Opuntia these cells are absent from some
species, particularly those with flat oval shoots of the type of Opuntia ficus
indica (L.) Miller; in some species they occur sparingly in the articulations
between the segments, quite apart from the vascular bundles, in the rays or
'at the margins of the
pith in others they are an integral part of the bundles.
;

They are found in larger quantities in Echinocactus and Melocactus, where,

except in the lowest internodes, they form the most important part of the
xylem. In certain species of Neomammillaria all elements other than the
tracheae of the primary xylem disappear and these cells form the bulk of
the xylem. Solereder also quotes Preston as stating that in Cereus fendleri
Engelm. similar cells occur in broad zones alternating with narrow zones of

spiral vessels. Intercellular canals. Radial channels often present, giving

a pitted appearance to the tangential surface of the wood (1886).

ANOMALOUS STRUCTURE
Anomalous secondary thickening recorded in Rhipsalis by Milan ez (1523)
and Echinofossulocactus multicostatus (Hildm.) Britton et Rose (syn. Echino-
cactus multicostatus Hildm.) by Haehnel (862).

ROOT
Roots generally narrow and woody compared with the stem; tap roots
sometimes well developed, e.g. the fleshy roots in certain species of Neo-
mammillaria. Tuberous roots present in Peniocereus greggii (Engelm.) Britton
et Rose (syn. Cereus greggii Engelm.) and species with thin stems, e.g. Wilcoxia

poselgeri (Lemaire) Britton et Rose (syn. Echinocactus tuberosus Rump.) the

rat's tail cactus. String-like roots a yard long recorded in Opuntia. Swollen
roots covered with cork and filled with mucilaginous internal tissue. Xylem
and phloem strands in the lateral roots of Echinofossulocactus multicostatus
(Hildm.) Britton et Rose (syn. Echinocactus multicostatus Hildm.) opposite
and not alternate as in most roots. Roots in the same species covered with
cork but devoid of mucilage cells. Anchoring roots of epiphytic species
originate near the central vascular cylinder of the aerial axis, and grow out
through the cortical ground tissue. Anchoring roots of Selenicereus hamatus
(Schweidw.) Britton et Rose (syn. Cereus hamatus Scheidw.) octarch, but
those of certain other epiphytic species ribbon-shaped and polyarch. AH of
the above information is recorded by Haehnel (862). The following particu-
lars recorded for Neomammillaria by Hemenway and Breazeale (948). Hairs

present when young but disappearing when older; cork occurring on old
roots; xylem including spiral thickening and thin cellulose walls. Contractile
roots of Echinocactus said by Nommeusen (1604) to become split by enlarged
groups of wound cork. Laticiferous canals present in the roots of those
species in which they have been recorded in the stem (see 'Stem' on p. 702).
Secondary thickening in the root of Rhipsalis has been described by Milanez
 CACTACEAE 705
TAXONOMIC NOTES
not easy to establish the affinities of such a specialized group as the
It is
Cactaceae. Chorinsky (408) noted that the structure and mode of origin of
the emergences in Pereskia and Rhipsalis resemble those of the emergences of
Anacampseros (Portulacaceae), and claimed this as additional proof of the
existence of affinities between the Cactaceae and Portulacaceae. Wettstein
(2416) has also drawn attention to the floral and anatomical similarities between
the Cactaceae and other families included in the Centrospermae. Britton and
Rose's monograph (275) has become the standard work on the general
taxonomy of the Cactaceae, and in the present account the nomenclature of
those authors has been followed as closely as possible.

ECONOMIC USES
The and succulent stems, especially of Opuntia, have been used as
fruits
fodder in dry countries. The dead bodies of certain scale insects which infest
species of Opuntia and Nopalea yield the red dye cochineal. The Prickly
Pear (Opuntia and Nopalea spp.) has become a very serious weed in certain
parts of the world to which it has been introduced, notably in parts of
Australia (see Shirley and Lambert (2090)). The thorns of Prickly Pear and
possibly of other Cactaceae are used as gramophone needles. An extract of
Selenicereus grandiflorus (L.) Britton et Rose (syn. Cereus grandiflorus Mill.)
has been used as a heart stimulant, but its therapeutic value is not well estab-
lished. Rouhier (1961) has described the use of certain members of the family
by Mexican Indians for ritual purposes, and also states that some Cactaceae
influence the nervous system through the action of the alkaloids which they
contain. The distribution of alkaloids in a species of Echinocactus has been
investigated by Steiner-Bernier (2193). Many members of the family are
commonly cultivated on account of their unusual and frequently grotesque
appearance.
The wood, according to Record and Hess (1886), is sometimes considerably
used locally owing to the scarcity of other timber. The Card6n, Cereus or
Cephalocereus^ in parts of northern Venezuela supplies attractive, easily
worked timber for furniture and general construction.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Ariocarpus, Astrophytum, Carnegiea, Cephalocereus, Cereus, Coryphantha,
Dolichothele, Echinocactus, Echinocereus, Echinofossulocactus, Echinopsis,
Epiphyllum, Gymnocalycium, Hatiora, Lemaireocereus, Lepismium, Leuch-
tenbergia, Malacocarpus, Neomammillaria, Nopalea, Opuntia, Pelecyphora,
Peniocereus, Pereskia, Pfeiffera, Rhipsalis, Schlumbergera, Selenicereus,
Trichocereus, Wilcoxia.

(ii)
FOR WOOD STRUCTURE
(Cactus), Carnegiea, Cephalocereus, Cereus, Dendrocereus, (Echinocactus),
Harrisia, Lemaireocereus, Leptocereus, Neoabbottia, Neomammillaria,
Nopalea, (Opuntia), Pereskia.
4594 ZZ
7o6 CACTACEAE
LITERATURE
(i) On
General Anatomy
Bedclian 1 66, Boke 2lS, *!9t Britton and Rose 275, Brown, J. G. 290, Bukvic 307,
Chorinsky 408, Coutant 487, Dauman 545, Gravis 806, Haehnel 862, Heinricher 942,
Hemenway and Breazeale 948, Jeffrey and Cole 1167, Rummer 1301, Leinfellner 1345,
Lloyd and Ridgway 1384, Markgraf I44*> *44* Milanez 1523, Nommeusen 1604,
Pfeiffer, H. 1707, Reiche 1909, 1911, Rouhier 1961, Scaramella 2026, Shirley and Lambert
2089, 2090, Steiner-Bernier 2193, Stewart 220O, Vaupel 2327, Weingart 2386, 2387,
Wettstein 24169 Wolf 2450.

(ii) On Wood Structure

Burgerstein 312, Cozzo 494* Dadswell and Record 533, Kanehira 1209, Milanez 1523,
Record 1781, 1787, 1843, 1851, Record and Hess 1886, Record and Mell 1894,
Tupper 2295.

155. FICOIDACEAE
(Fic. 163 on p. 708)

SUMMARY
Mostly succulent herbs, but a few genera tend to be shrubby. The family
occurs chiefly, but not exclusively, in desert regions in the tropics and sub-
tropics, reaching its maximum development in South Africa and Western
Australia. The leaves usually constitute the most fleshy part of the plant,
and, in very specialized members of the family such as Lithops and Lapidaria,
the aerial part of the plant consists of opposite fleshy leaves between which the
flowers arise. These remarkable members of the family strongly resemble the
desert stones amongst which they grow. The family also includes species
with reduced leaves, the main part of the plant then consisting of assimilatory
branches, the habit resembling that of species of Cytisus. Others are more like
Salicornia. The epidermis of both leaf and stem generally includes large,
bladder-like cells, which suggest the name *ice-plants' which is sometimes
applied to members of the family. The stomata are generally ranunculaceous,
but, in a few instances, rubiaceous. The mesophyll of the leaves is most
frequently centric, but there are numerous exceptions. It usually consists
wholly of palisade cells, but there is frequently a core of aqueous tissue in
which the vascular system is embedded. The aqueous tissue extends to the
epidermis at the apex of leaves of the type found in Lithops which has caused
these organs to be termed 'window* leaves (Fensterblatter). The leaves some-
times exhibit a characteristically chalky appearance due to the deposition of
small crystals in the epidermis. The most noteworthy features of the axis
are the common occurrence of anomalous secondary thickening in both
stem and root, the frequent presence of a reticulum of leaf trace bundles
in the cortex, and the infrequence of rays in the xylem. Secretory elements
are known only in the form of large tannin sacs in a few species of Mesembry-
anthemum* Crystals solitary, in clusters, or in the form of raphides.

LEAF
Nearly always centric; with or without a central core of aqueous tissue, but
relatively flat leaves isobilateral, e.g. in Sesuvium porttdacastrum Linn. Iso-
1
For the sense in which the name MesembryatUhtmum is used in this book see 'Taxonomic
Notes' on p. 711.
 FICOIDACEAE 707
bilateral leaves also recorded by Sabnis (1977) in species ofLimewn, Mollugo,
Trianthema (see also
Orygia, and dorsiventral ones in Gisekia, Mollugo,
'Mesophyll* on p. 709). Surface sometimes coated with wax. Hairs simple,
unicellular in certain species of Mesembryanthemum; unicellular, 2-armed in
species of Aizoon, Galenia, Plinthus;
each having a uniseriate stalk bearing
a unicellular, stellate head in species of GKnus (Fig. 163 c). Leaf tips of
species of Trichodiadema provided
with cork cushions bearing a cluster of
hairs or papillae according to Oberstein (1626). Glandular hairs with a i- or
2-celled stalk and a large globular head, situated in small depressions on both
surfaces of the leaf recorded by Pilger (1720) in Glischrothamnus ulei Pilger.
A dense coveringof stellate hairs also recorded by Sabnis (1977) in Mollugo
hirta Thunb. Epidermis composed of large, bladder-like, water-storage
cells intercalated between much smaller ones, in species ofAizoon (Fig. 163 B),
Sesuvium (enlarged
Cryophytum (Fig. 163 A), Galenia, Mesembryanthemum,
cells not seen by Mullan (1571) in plants of 5. portulacastrum Linn, growing
on the sea-shore), Tetragonia, Trianthema\
the large cells sometimes provided
with hair-like or papillose extensions, e.g. in species of Mesembryanthemum and
Tetragonia. Epidermis, in a few species
of Hereroa examined by Zemke
the latter being protected externally
(2505), consisting wholly of small cells,
of Mesem-
by a thin layer of cuticle. Reule (1925), who examined 81 species
bryanthemum (sensu lato) collected in South Africa and cultivated in Germany,
found the structure of the epidermis and stomata to provide features of con-
siderable taxonomic interest. He recognized 7 more or less distinct types of
epidermis connected by transitional forms.
TYPE I (Normal Type). Epidermal
cells isodiametric, polygonal, outer wall at most slightly sinuous when viewed
in transverse sections, and, at most, projecting only slightly into the .sub-
I observed in species of
epidermal layer. Thickness of cuticle 3-17 p. Type
Cono-
Bergeranthus, Carruanthus, Chasmatophyllum, Cheiridopsis, Conophyllum,
phytum (few species only), Faticaria, Fenestraria, Gibbaeum, Glottiphyllum,
Nananthus, Rhombophyllum, Semnanthe. TYPE II (Conophytum
Pleisopilos,
Type). Epidermal covered externally by a very thick, flat, continuous
cells

layer of cuticle, the latter extending over the whole surface except where
interrupted by the stomata. In general similar to Type I but with much
thicker cuticle. Type II observed in species of Argyroderma, Conophytum
(10 spp.), Ruschia. TYPE III (Lithops Type). Epidermal
cells protected
a thick of the latter tending to form rounded, blunt
externally by layer cuticle,
cells. Cuticle more definitely papillose
papillae over the individual epidermal
in some genera than in others. Type III observed in species of Conophytum
(i sp.), Corpuscularia, Lapidaria, Lithops (6 spp.),
Rimaria. TYPE IV (Kegel-
zellentypus). Similar to Type III, but outer walls of the epidermal cells
provided with more pointed cuticular papillae. Type IV observed in certain

species of Cheiridopsis. TYPE V(Cheiridopsis Pilansii Type). Cuticle

deposited in the form of hair-like structures on the outer walls of the epidermal
cells. Type V observed in certain species of Cheiridopsis and Gibbaeum.
TYPE VI (Riesenzellentypus). Some of the epidermal cells very much en-
larged compared with the remainder. Type VI observed
in species of

Deksperma and Trichodiadema. TYPE VII. A

proportion of the epidermal
cells united in groupswhich are collectively much larger than the remaining
epidermal cells. Type VII observed in Qdontophorus and Psammotropha*
4504 Z252
7o8 FICOIDACEAE
Reule also found the stomata to vary in structure and the extent to which they
are sunk in the leaf tissue, species resembling one another in epidermal struc-
ture also exhibiting similar stomata. The types of epidermal structure
enumerated above tend to correspond to the genera, but it will be noted that
there are discrepancies. Thus in some of the genera, e.g. Conophytum and
Lithops, the epidermal structure was frequently of a uniform type in most but
not all of the species available for examination. Other genera included species

FIG. 163. FICOIDACEAE

A, Transverse section through the margin of the leaf of Cryophytum crystallinum (L.) N. E. Br.
B, Transverse section through the leaf of Aizoon canariense L. C, Stellate hair of 'Glinus cambcssedesii
Fenzl.' A, B after Volkens, C by Solereder.

each showing a different type of epidermal structure, e.g. in Ruschia. Finally

Reule noted some correspondence between epidermal structure and dryness
of habitat, but the ecological significance of this could not be fully assessed
without further physiological experiments.
Reule's results could be only partly confirmed by Oztig (1648), who found
the structure of the epidermal cells to vary in plants of a single species from
different localities or even in different parts of a single leaf. According to
Oztig the cuticle of the leaf epidermis is nearly always very thin, and seldom
striated* In extreme xerophytic forms such as species of Conophytum and

Lithops the epidermal cells are strongly thickened in a peculiar way, cutinized
lamellae of cellulose being present below the cuticle itself. The cutinized
layers sometimes extend to the lateral walls, especially in Lithops, and appear
wedge-, top-, rod-, or spindle-shaped in transverse sections. In some species
of Conophytum and Lithops they extend inwards from the outside wall, form-
ing a lens or peg in the centre of the cell. Unlike Reule, Oztig was unable to
observe complete cutinization of the outer wall of the cell. Stomata some-
what variable; orientated transversely to the longitudinal axis in some but not
all species of Mesembryanthemum\ said to be ranunculaceous in species of

Adenogramma, Gisekia, Limeum, Mesembryanthemum, Polpoda, Psammotropha,

 FICOIDACEAE 709

Tetragonia, but rubiaceous in other species of Mesembryanthemum. Meso-

phyll. Palisade tissue in Trianthema portulacastrum Linn, said by Kienholz
of the mesophyll, especially around the
(1236) to be confined to the centre
veins. Sabnis (1977) also records the occurrence of green bundle sheaths in
certain species of Mollugo and Trianthema. Palisade tissue in Trianthema
triquetra Rottl. et Willd. said by Sabnis to be confined to arcs of tissue
towards the lower surface, the remainder of the mesophyll in this species
consisting of aqueous tissue. Pax and Hoffmann (1675) cite Brenner as having
recorded the occurrence, in Lampranthus curvifolius (Haw.) N. E. Br., of a
circle of small bundles on the inside of 3-5 layers of palisade tissue, the main
vascular strand being embedded in the central core of water-storage tissue.
The small outer bundles in the same species are continuous with 2 lateral
1
strands given off from the central bundle near the base of the leaf. 'Window
leaves (Fensterblatter), in which aqueous tissue replaces the chlorenchyma
at the distal ends, recorded by Brown, Tischer, and Karsten (293) in all

species of Fenestraria and Lithops, in certain species of Conophytum and in

the monotypic genera Frithia and Imitaria.
The physiological significance of 'window' leaves has been investigated by
Huber (1104), Kean (1226), and Schmucker (2042). Contrary to previously
accepted views, Schmucker demonstrated that, in spite of the absence of
stomata, the 'window* tissue is the chief transpiratory surface of the leaf, loss
of water occurring via the cuticle, and reaching a maximum in damp air and
at an average temperature. Furthermore, sinking the plant deeply in a
sandy substratum accelerates transpiration. The intensity of the light reaching 1
the inner surface of the cylinder of assimilatory tissue through the 'window
is sometimes reduced, e.g. in Lithops pseudotruncatella (Berg.) N. E. Br. by

deposits of calcium oxalate on the surface, and, in Hypertelis, Orygia, &c., by

a waxy coating.
Large tannin sacs occur below the epidermis in a few species of Mesem-
bryanthemum according to Oberstein (1627). The translucent, mucilaginous
tanniniferous cells sometimes serve as lens cells according to Kean (1226).
Lens cells in the epidermis also described by Summers (2221). Small
crystals in the epidermal cells of Mesembryanthemum sometimes give the
leaves a white, chalky appearance. Outer walls of the epidermal cells in
Mesembryanthemum more or less encrasted with calcium oxalate crystals
according to Oztig (1648). Encrustation most strongly developed and indivi-
dual crystals especially large in extreme xeromorphic species. Large cluster
crystals embedded in the outer walls of the epidermal cells in a few species,
e.g. of Lapidaria and Lithops. Amount of crystalline deposit said to vary
according to the intensity of light to which the leaf is exposed. Crystals rarely
present in the inner and anticlinal as well as in the outer walls of the epidermal
cells. Solitary crystals recorded by Solereder and other authors in Adeno-

gramma, Coelanthium, Cypselea, Potpoda; clusters in Glinus, Hypertelis,

Limeum, Mollugo, Orygia, Pharnaceum, Sesuvium, Trianthema; raphides in
Gisekia, Mesembryanthemum, and Mollugo crystal-sand in Hypertelis acida
;

(Hook.) K. Mull. Cubical crystals of calcium phosphate and spherical

crystals of calcium malate recorded by Kean (1225) in spirit material of
Mesembryanthemum. Oxalic and malic acids said to occur in Mesembry-
anthemum, and malic, citric, and phosphoric acids in Cryophytum crystallinum
7 io FICOIDACEAE
(L.)N. E. Br, Reddish-brown colour of the leaves of certain species of
and Lithops found by Oztig (1648) to be due to orange-coloured
Aloinopsts
chromoplasts in the epidermal and sub-epidermal cells as well as to
anthocyan pigments mostly in the sub-epidermal cells. Contrary to earlier
statements in the literature, Oztig found no reddish-brown cystoliths. Colour
in Conophytum mainly due to anthocyan alone.

Axis
STEM
Epidermis in Mesembryanthemum including bladder-like aqueous cells
similar to those of the leaf. Cork
arising in the inner part of the cortex in the
few species of Mesembryanthemum in which its formation has been examined.
Outer part of the primary cortex sometimes differentiated as assimilatory
tissue. (2505) refers to an endodermis in 'Mesembryanthemum salt-
Zemke
which becomes suberized at an early stage, a phellogen subsequently
cornioides*

arising on the inside of the layer of endodermal cells. Pericycle either

devoid of sclerenchyma, e.g. in Trianthema pentandra Linn., or including
stone species of Limeum, Mollttgo, Orygia.
cells, e.g. in A
reticulum of cortical
leaf trace bundles also frequent in Mesembryanthemum. Xylem including
vessels with bordered pits and simple perforations embedded in ground tissue,
the latter usually consisting of prosenchymatous elements with simple pits,
but including a varying proportion of parenchyma. Delicate spiral thickening
of the vessel wall recorded in Galenia. Medullary rays generally absent but
present in Gisekia, Macarthuria (relatively broad), Orygia (narrow). Crystals
include (i) raphides, sometimes in suberized sacs, in the ground tissue of
Mesembryanthemum', (ii) styloids in the phloem of Aizoon sp. and in the pith
of Tetragonia spicata Linn, (iii) clusters in the cortex of Limeum, Mollugo,
;

Sesuvium portulacastrum Linn., and Tetragonia expansa Murr. See also under
'Leaf* on p. 706.

ROOT
Cortex lacunar in Sesuvium portulacastrum Linn, according to Mullan
(1571). For normal and anomalous secondary thickening see 'Anomalous
Structure* below.

ANOMALOUS STRUCTURE
Anomalous common in both stem and root especially in woody
structure
species, caused by the development of secondary bundles from successive
rings or arcs of meristem in the phloem or pericycle. (A) Stems. Two main
types of anomaly recognized, (i) Numerous bundles arranged in more or less
distinct concentric rings, embedded in prosenchymatous ground tissue in
many species of Mesembryanthemum as well as in a few of Acrosanthes, Galenia,
Macarthuria, Sesuvium, Trianthema. (ii) Alternating, more or less complete
rings of xylem and phloem in species of Aizoon, Glinus, Limeum, Orygia,
t

Pharnaceum, Sesuvium, Tetragonia, Trianthema. Transitions between (i) and

(ii) also occur.
Anomalous thickening in a special zone above the root
described by Schmid (2035) in Lithops pseudotruncatella (Berg.) N. E. Br.
(Mesembryanthemum pseudotruncatellum Berg.). Secondary thickening said to
be normal in the herbaceous genera Adenogramma, Gisekia, Limeum, Psammo-
 F1COIDACEAE 711

tropha, and in the woody genus Polpoda. (B) Roots. Anomalous thickening
recorded in species of Mesembryanthemum as well as in Aizoon hispanicum
Linn,, Mollugo radiata Ruiz, et Pav., Sesuvium port ulacastrum Linn., Tetra-
gonia expansa Murr. Normal thickening stated to occur in Pharnaceum
incanum Linn, and Psammotropha quadrangularis (L.) Fenzl.

ECONOMIC USES
No
important economic products are obtained from this family, but many
of its members
are cultivated on account of their unusual morphology or for
their beautiful flowers. A few species have been used as vegetables, for
example, according to Zwicky (2512) by Hottentots in the South African
Karoo.
TAXONOMIC NOTES
Certain members of the Ficoidaceae resemble the Centrospermae suffi-
ciently to suggest that their affinities lie in this direction. This is, to some
extent, confirmed by similarities in the anatomical structure, particularly by
the widespread occurrence of anomalous secondary thickening. In this con-
nexion Pax and Hoffmann (1675) particularly emphasize the resemblances
between the Ficoidaceae and Phytolaccaceae, but also point out that con-
siderable differences exist between the Ficoidaceae and Cactaceae.
Considerable difficulties have arisen in recent years concerning the
taxonomy of Mesembryanthemum, which has been split into numerous genera
by N. E. Brown (291), Schwantes (2057), and L. Bolus (222). Owing to the
lack of general agreement between those authors there has been much con-
fusion. Pax and Hoffmann (1675) in the main follow N. E, Brown, but also
retain the genus Mesembryanthemum in a wide sense. K. von Poellnitz (Repert.
Nov. Regn. Veg. 32, 1933, and Monatschr. Kakteenkunde, 1933) has attempted
to clear up the existing confusion. Owing to the difficulty of correlating the
old species names under Mesembryanthemum with their modern synonyms,
it has been necessary, in the present book, to follow Pax and Hoffman in

retaining the name Mesembryanthemum in a wide sense. This has been done
when citing information recorded in the literature. Whenever possible, how-
ever, the names given to the species by N. E. Brown have been used, as these
are generally accepted by British and German investigators. A complete
anatomical revision of the family could usefully be undertaken as soon as the
taxonomy of the family is more stable.

GENERA DESCRIBED
Acrosanthes, Adenogramma, Adenostemma, Aizoon, Aloinopsis, Argyro-
derma, Bergeranthus, Carruanthus, Chasmatophyllum, Cheiridopsis,
Coelanthium, Conophyllum, Conophytum, Corpuscularia, Cypselea, Delo-
sperma, Faucaria, Fenestraria, Frithia, Galenia, Gibbaeum, Gisekia, Glinus,
Glischrothamnus, Glottiphyllum, Hypertelis, Imitaria, Lampranthus,
Lapidaria, Limeum, Lithops, Macarthuria, Mesembryanthemum, Mollugo,
Nananthus, Odontophorus, Orygia, Pleisopilos, Plinthus, Polpoda, Psam-
motropha, Rhombophyllum, Rimaria, Ruschia, Semnanthe, Sesuvium,
Tetragonia, Trianthema, Trichodiadema.
 712 FICOIDACEAE
LITERATURE
On General Anatomy
Bolus 223, Brown, N. 291, Brown, Tischer, and Karsten 293, Huber 1104, Kean
.

1224, 1225, 1226, Kearney 1228, Kienholz 1236, Marloth 1444, Mullan 1571, Oberstein
1626, 1627, Oztig 1648, Pax and Hoffmann 1675, Pilger 1720, Reule 1925, Sabnis 1977,
Schmid 2035, Schmucker 2042, Schwantes 2057, Summers 2221, Zemke 2505,
Zwicky 2512.

156. UMBELLIFERAE
(FiG, 164 on p. 712; FIG. 165 on p. 714; FIG. 166 on p. 716; FIG. 167 on p. 718)

SUMMARY
(i) GENERAL
The family ismainly herbaceous, but includes a few species which tend to
be woody. It occurs chiefly in temperate regions and on tropical mountains,

FIG. 164. UMBELLIFERAE

Trichomea of: A, B, Xanthosia pilota Rudge; C, Botolesia tropaeoK/olia Gill, et Hook. -By Solercdcr.

but the various species grow in very different habitats. Some of the xerophytes
show anatomical specializations ('Ecological Anatomy', p. 720). In spite of these
ecological specializations, the basic structure is remarkably uniform through-
out the family. The stems are often ribbed, whilst the centre is occupied by
a pith which often becomes hollow apart from the septa at the nodes. The
ribs on the stems usually consist of collenchyma, or, more rarely, of scleren-
chyma. The leaf is usually dorsiventral, except in species which show
ecological specializations. The hairs, which are nearly always non-glandular,
include unicellular, dendroid, and stellate types. The stomata are sometimes
accompanied by variously orientated subsidiary cells, but others are ranun-
 VMBELLIFERAE 713
culaceous. Secretory canals, which contain a mixture of oils, resin, and
mucilage, are a particularly characteristic feature. They occur in the primary
cortex, pericycle, pith, and sometimes the secondary phloem of the axis, but
extend into the petiole and leaf lamina and also into the root. The petiole is
usually provided with an arc or ring of vascular bundles, which sometimes
surround medullary strands. In the stem there is always a ring of vascular
bundles, which may be accompanied by medullary or, more rarely, cortical
strands. The bundles of the main ring are sometimes embedded in scleren-
chyma. Anomalous secondary thickening sometimes occurs in the stem
and more frequently in the root. The anomalies include the development of
numerous concentric bundles with central xylem; the formation of an extra-
fascicular cambial ring which produces xylem on the inside and phloem on
the outside. A fissured xylem mass has been recorded in Azorella. Crystals
are infrequent and often absent, but clusters and needles have been observed
in a few species.

(ii) WOOD
Vessels very small to medium-sized, often in clusters and sometimes with
a tangential pattern; perforations usually simple, but occasional scalariform
plates sometimes present; intervascular pitting mostly alternate to opposite,
sometimes almost scalariform, pits to parenchyma similar, sometimes simple;
members of medium length to extremely short. Parenchyma paratracheal
(scanty to vasicentric) and sometimes terminal. Rays usually up to 4-5 cells
wide, sometimes more, heterogeneous to almost homogeneous. Fibres with
simple pits, very to extremely short. Intercellular canals present in the rays
of some genera.

LEAF
Usually dorsiventral, but centric in leaves having narrow or terete segments,
e.g. in Bupleurum or Foeniculwn vulgare Mill. isobilateral in Daucus carota
;

Linn. Hairs (Fig. 164) include the following types, (i) Simple, unicellular,
(ii) Unicellular,
bladder-like, (iii) Dendroid (Fig. 164 A-B), each having a
few short basal cells and tiers of thick- walled ray cells with narrow lumina.
(iv) Stellate (Fig. 164 c), with biseriate
or multiseriate pedestals, (v) Small
glandular hairs with 2- to 4-celled heads recorded on the lower side of the
leaf of Pimpinella saxifraga Linn. Glandular hairs also reported from the

peduncle of Astrantia, and on the umbels of Heracleum. Upper epidermis

papillose in species of Angelica, Anisotome, Bupleurum, Carum, Cicuta,
Conium, Conundrum, &c.; consisting of specially large cells in Hermas sp.
Epidermal cells surrounding the hair bases frequently forming cushions.
Stomata occurring on both surfaces or confined to the lower side, differences
in this respect being mainly of specific diagnostic value; arranged parallel to
one another in species of Eryngium with a monocotyledonous habit. Rubia-
ceous stomata recorded in species of Eryngium and Hydrocotyle. Stomata
ranunculaceous in species of Ammi, Bowlesia, Caucalis, Conium, Conundrum,
Laserpitium, Mulinum, Peucedanum, Prangos, Sanicula, Seseti, and other
genera. Hypoderm recorded in species of Aciphylla, Bupleurum, Eryn-
gium> &c. (see also 'Ecological Anatomy' on p. 720).
Vascular bundles of the veins nearly always accompanied above and
 FIG. 165. UMBELLIFERAE
A, Eryngium campestre Linn. Petiole X 7. B, Foeniculum vulgare Mil. Petiole X 8. C, Centum
maculatum Linn. Petiole xy. D, Carum carvt Linn. Petiole x 18. E, Apium graveolens Linn.
Petiole XQ. F, Oenanthe silaefolia Bieb. Petiole xia. G, Trinia glauca (L.) Dum. Petiole X 18.
H, Meum athamanticum Jacq. Petiole X 15. I, Archangelica officinalix Hoffm. Petiole X 7. J, Aegopo-
dium podagraria Linn. Petiole X 8. K, Crithmum maritimum Linn. Petiole X 13. L, Seseli libanotis
(L.) Koch. Petiole X 33.
The small unshaded circles represent secretory canals.
 UMBELLIFERAE 715
below or on one side only by collenchyma or parenchyma, the arrangement
of which possesses diagnostic value. Sclerenchyma seldom present in the leaf,
but recorded in association with the vascular bundles in Eryngiwn campestre
Linn, and, according to Meyer (1509), between the marginal bundles and the
actual margin in Sanicula europaea Linn.
Petiole, in transverse sections through the distal end of material examined
at Kew, exhibiting a considerable range of structure (Fig. 165 A-L and
1 66 A, c, D, E, F, and H). Vascular bundles widely spaced and usually numer-

ous, arranged in crescents, circles, or irregularly distributed in different genera

and species. Variously shaped crescents of bundles noted in species of
Aegopodium j), Apium (Fig. 165 E), Bupleurum (Fig.
(Fig. 165 166 D), Carum
(Fig. 165 D), Chaerophyllum (Fig. 166 H), Coriandrum (Fig. 166 F), Crithmum
(Fig. 165 K), Daucus, Echinophora (Fig. 166 E), Eryngium (Fig. 165 A), Fal-
caria, Meum (Fig. 165 H), Pimpinella, Sanicula (Fig. 166 c), Seseli(Fig. 165 L),
but additional medullary strands present within the crescent in Chaerophyllum
(Fig. 1 66 H), Daucus, and Eryngium. Complete circles of separate bundles
observed in species of Archangelica (double circle) (Fig. 165 i), Conmm
(Fig. 1650), Foeniculum (Fig. 1653), Ligusticum (Fig. i66A), Oenanthe
(Fig. 165 F), Peucedanum (double circle), and Trinia (Fig. 165 G). Numerous,
scattered vascular strands present in Ferula communis Linn, and Oenanthe
crocata Linn. Petiolar medullary bundles described by Solereder as initially
collateral, or appearing to be bicollateral owing to the fusion of the xylem
portions of 2 contiguous bundles. Centric medullary bundles sometimes
formed from collateral strands by the development of xylem externally to the
phloem. Sub-epidermal strands of collenchyma frequently occur opposite
the peripheral vascular bundles of the petiole. Secretory canals, more
fully described under 'Axis', present in the pith, pericycle, and cortex of the
petiole, and in the phloem as well in certain species; in some species extend-
ing into the lamina beside the vascular bundles. These canals probably occur
in all members of the family. Their arrangement in relation to the vascular
bundles of the veins is said to be of specific diagnostic value. Secretory
cavities also occur occasionally, e.g. in the mesophyll of Aegopodium.
Crystals often absent, but, where present, generally clustered; recorded by
Meyer (1509) in Astrantia, Eryngium, and Sanicula. Small, solitary or
clustered crystals, of undetermined chemical nature, recorded, also by Meyer,
in a few species of Anethum, Angelica (crystal-sand), Apium, Foeniculum,
Heracleum (crystal-sand), Levisticum, Petroselinum, Pimpinella. Plumose
masses of an unidentified material, probably hesperidin, also stated to occur
in species of Aethusa, Bupleurum, Chaerophyllum, Conium, Coriandrum,
Cuminum, Oenanthe, Sium, Trinia. Spherical aggregations of yellowish
acicular crystals also noted at Kew in the epidermis and vessels of the petiole
of Crithmum maritimum Linn. Tannin noted by Meyer (1509) in a few species
of Anthriscus, Chaerophyllum, and Siler. For examples of special leaf struc-
ture see under 'Ecological Anatomy* on p. 720.

Axis
STEM (Fig. 166 B, G, i, j, K)
Stems frequently with ribs, the latter consisting largely of collenchyma or,
more rarely, of sclerenchyma; sometimes with palisade chlorenchyma between
 Fio. 166. UMBELLIFERAE
A, Ligusticum scoticum Linn. Petiole x B, Foeniculum vulgare Mill. Stem X 28. C, Sanicula
8.
gregaria Bickneil. Petiole xiz. D, Bupleurwn fruticosum Linn. Leaf base x xi. E, Echinophora
anatolica Boiss. et Heldr. Petiole X 12. F, Coriandrum sativum Linn. Petiole x 19. G, Apium
graveolens Linn. Stem X 8. H, Chaerophyllum nodosum Crantz. Petiole x 12, I, Daucus hispidissimut
Sennen. Stem X 19. J, Smymium olusatrum Linn. Stem X 8. K, Crithmum maritimum Linn*
Stem X 15, showing intercellular cavities in the cortex. The small unshaded circles represent
secretory canals.
 UMBELLIFERAE 717
the ribs. A ring of collenchyma present beneath the epidermis in other species.
Fine structure of the collenchyma of Heracleum sphondylium Linn, described
in detail by Majumdar and Preston (1428). Epidermis sometimes sclerosed
(see 'Ecological Anatomy' on p. 720). Apart from the chlorenchyma and collen-
chyma already mentioned in the primary cortex, the inner part of this region
consists of thin-walled compact tissue with secretory canals embedded in it.
Cork described by Solereder as originating in the sub-epidermis in Bupleurum
fruticosum Linn., Heteromorpha arborescens Cham, et Schlechtd., and in species
of Hydrocotyle and Trachymene; pericyclic in Mulinum spinosum Pers. Sub-
epidermal cork recorded by Lemesle (1346) in Hydrocotyle arbuscula
Schlechtd., Peucedanum capense Sond., 'P. ferulaceum Eckl. et Zeyh/,
'Trachymene ericoides Sieber.', and *T. linearis Spreng.'. Cork described by
the same author as arising in the inner part of the cortex in Eryngium carli-
noides Boiss., Hydrocotyle solandra Linn., Mulinum spinosum, and Pituranthos
spp. Pericycle sometimes including strands of fibres, but sclerenchyma in
this region seldom well developed. Vascular bundles consisting mainly of

primary tissue, and usually arranged in a ring, but accompanied by medullary

or, less frequently, by cortical strands in some species. The bundles of the
principal ring vary in size and distribution, and exhibit the following alterna-
tive arrangements, (i) Isolated bundles separated from one another by ground
tissue, (ii) The xylem portions of the bundles united to one another by inter-
fascicular, mechanical elements, (iii) Bundles wholly embedded in scleren-
chyma. (ii) and (iii) occur particularly in xerophytic species. Vessels mostly
with simple perforations; scalariform plates occasional (see 'Wood' on p. 719).
Medullary bundles recorded in species of Apium, Cachrys, Cenolophium,
Crithmum, Eryngium, Ferula, Laserpitium, Magydaris, Oenanthe, Peucedanum,
Pimpinella, Silaus, Sium, and Thapsia; their number varying considerably in
different species; irregularly distributed or arranged in a ring; sometimes
inversely orientated. Cortical bundles recorded in species of Eryngium
with a monocotyledonous habit, as well as in Siler trilobum Crantz. and
Mulinum. Pith. Central portion, in many species, becoming disorganized
except for persistent septa at the nodes. Whole pith more durable and some-
times sclerified in species of Coriandrum, Eryngium, Ferula, Sison, Sium, &c.
Secretory canals, containing a clear, or turbid, milky-white or yellow
mixture of oils, resin, and mucilage when fresh, present in the inner part of
the primary cortex, as well as in the pericycle, and sometimes the secondary
phloem of most if not of all members of the family. Secretory canals also
occur at the periphery of or scattered in the pith of most members of the
family, but none recorded in this tissue in a few species of Bupleurum, Hydro-
cotyle, Xanthosia-, canals inthe phloem occur particularly in woody species
such as Bupleurum fruticosum and Heteromorpha arborescens. Canals in the
aerial part of Ferula communis Linn, described by Perrot and Morel (1697) as

consisting of those which are solitary except at the nodes and ramify in the
parenchyma of the cortex and pith, and those accompanying and forking in
the same way as the vascular bundles. Primary secretory canals in species of
Archangelica, Imperatoria (Peucedanum), and Levisticum examined by Elias
(625) said to arise in the pericycle, the later ones originating in tissue derived
from the cambium. Layer of cells surrounding the cavity of the canals in the
same genera described as producing mucilage, with fine cellulose threads
7 z8 UMBELLIFERAE
embedded in it. No resiniferous layer detected in these genera. Contents of
the secretory canals of Cicuta maculata Linn, poisonous. According to
Sifton (2097) the less poisonous nature of the aerial than of the underground
parts of this plant is correlated with the smaller number of canals in the part
of the plant above ground. Secretory canals in the cortex of Smyrnium

B
FIG. 167. UMBELLIFERAE
A, Steganotaenia araliacea Hochst. B, Heteromorpha arborescent Chain, et Schlect. C, H,
arborescent Cham, et Schlect. D, Steganotaenia araliacea Hochst. E, Eryngium inaccessum
Skottsberg.
i.e. Intercellular canal.

olusatrum Linn, seen at Kew to be in pairs, one being situated just outside
and the other just inside each patch of collenchyma. For details concerning
the development and mode of secretion of the canals in certain members of
the family see Pirschle's (1726) paper. Crystals usually infrequent or lacking,
mostly clustered when present. Clustered crystals noted at Kew in 2 species
of Eryngium and acicular types in Critkmum and Ferula, Developmental
anatomy. Details concerning the developmental anatomy of various mem-
 UMBELLIFERAE 719
bers of the family have been recorded by Brandscheit (256), Chaerophyllum;
Majumdar (1424, 1425, 1427), Heracleum; Esau (651, 654), Apium.
WOOD (Fig. 167)
Vessels very small to medium-sized (mean tangential diameter ranging
from 40 to 125 ft); often in clusters and with a tangential or oblique pattern
(Fig. 167 c) in Heteromorpha arborescens (Thbg.) Cham, et Schlect. and
Pituranthos (1493); number per sq. mm. varying from about 5 in Peucedanum
and Steganotaenia to 20-25 in Eryngium and Heteromorpha. Perforations
typically simple, but occasional scalariform plates with few bars reported by
Solereder to accompany the simple perforations in Aethusa, Bupleurum,
Carum, Hydrocotyle, Oenanthe, Peucedanum, and Xanthosia. Intervascular
1

pitting usually alternate to opposite, the mouths of the pits horizontal and
sometimes coalescing; often locally transitional between opposite and scalari-
form, particularly in Heteromorpha, in which the pitting is sometimes almost
scalariform; pits to ray and wood parenchyma similar, sometimes simple.
Mean length 0-25-0-45 mm. Parenchyma paratracheal, scanty to vasicentric,
forming sheaths usually only i cell wide round the vessels; terminal bands
observed in Peucedanum and Steganotaenia. Strands varying between 1-4
cells, e.g. in Eryngium and Heteromorpha, and 4-6, occasionally 8, cells, e.g. in
Peucedanum and Steganotaenia. Rays usually up to 4 or 5 cells wide, up to
8 cells in some species of Eryngium seldom more than i mm. high, except in
;

some species of Eryngium; uniseriates typically few and composed of both

procumbent and square to upright cells, more numerous in Eryngium
bupleuroides Hook, et Arn. rays mostly between 2 and 8 per mm., more
;

numerous where uniseriates are common; usually slightly heterogeneous

(Kribs's Type II B), with i or 2 marginal rows of square to upright cells;
almost homogeneous in Steganotaenia, and composed almost entirely of
square to upright cells in Eryngium inaccessum Skottsberg. Fibres with
numerous, small, simple pits in the radial walls and few in the tangential
walls; the pits larger and equally numerous on the tangential walls in Eryn*
gium inaccessum; walls rather thin; mean length 0-3-0-75 mm. Majumdar
(1422) has described the elongation of the fibres in Heracleum by 'intrusive*
growth. Intercellular canals observed in the rays of Eryngium, Peucedanum,
and Steganotaenia. Messeri (1493) refers to lysigenous secretory pockets
('tasche') in the rays of Pituranthos.

ANOMALOUS STRUCTURE
The development of an extrafascicular cambial ring, producing phloem on
the outside and xylem on the inside, recorded in the stem of a few species
of Eryngium. Transverse sections of the mature roots of certain species of
Angelica, Anthriscus, Apium, Carum, Chaerophyllum, Cicuta, Daucus, Magy-
daris,Oenanthe, Sium exhibit numerous centric bundles, with central xylem,
arranged in 2 or 3 concentric rings. The mode of development of this
anomaly has been examined particularly in Oenanthe crocata Linn. In the
young root of this species there are, around the pith, 5-14 groups of vessels,
and between them a corresponding number of phloem strands. Each group
1
Multiperforate plates have also been recorded by Majumdar (1421) in Heracleum sphondy-
lium Linn., where they appear to be confined to the protoxylem of the leaves and the meta-
xylem in the nodal region of the stem.
720 UMBELLIFERAE
of vessels becomes surrounded by cambial tissue, which forms a few new
vessels and prosenchymatous elements on the inside; on the outside a con-
siderable amount of parenchymatous tissue is produced. In this way the
so-called centric bundles arise. A second ring of similar bundles then
originates on the inside, and, by a repetition of this process, the structure of
the mature root is attained. Secondary meristematic rings forming inversely
orientated bundles recorded in Myrrhis odorata (Linn.) Scop. Xylem mass
becoming fissured in Azorella selago Hook. f.

ROOT
Primary structure diarch in Pimpinella according to Michlin (1512).
Endodermis with casparian thickenings recorded in the same genus. Secre-
tory canals, similar to those of the stem and leaf, also occur in the root;
those in the primary tissues said by Solereder to be situated immediately
within the epidermis, opposite both the phloem and xylem groups of the
vascular strand. Secretory canals in the primary tissues of Pimpinella
described by Michlin ( 1 5 1 2) as pericyclic. Canals also present in the secondary
phloem of older roots of most members of the family; occurring in the xylem
as well in species of Myrrhis and Opopanax. Perrot and Morel (1697) have
recorded the following information concerning the root and rootstock of
Ferula communis Linn. In the lower part of the root secretory canals are
present in the pericycle immediately within the cork, and other canals in the
secondary phloem. Transverse sections at higher levels in the root show more
abundant secondary phloem and more numerous canals. Owing to pressure
against the outer corky layer as growth in thickness proceeds, the primary and
outer part of the secondary phloem is crushed and the medullary rays become
sinuous. The canals in the crushed tissue likewise become compressed, but
the pericyclic canals, protected by the cork, persist. In the upper part of the
rootstock, where a pith is differentiated, the xylem parenchyma and rays
become mucilaginous, and the groups of vessels can be seen in the partly
disorganized tissue. At a still higher level, transverse sections show the
xylem as part of a ring of vascular bundles separated by wide medullary rays.
The centre of the rootstock in this region is occupied by a cavity, surrounded
by an internal layer of cork. The secretory canals are solitary in the lower but
anastomose in the upper part of the root. Lunan (1401) refers to convoluted,
radiating strands of amyliferous tissue in the adult root of Ligusticum scoticum
Linn, due to unequal strains set up between the rigid xylem and the confining
cork tissue. The structure of the contractile roots of various members of
the Umbelliferae in relation to their physiological function has been described
by Berckemeyer (175). The root structure of carrot seedlings has been
described by Havis (922), and the developmental anatomy of the carrot
by Esau (654).

ECOLOGICAL ANATOMY
Several authors have drawn attention to the structural plasticity of certain
members of the Umbelliferae. Funk (733), for example, has pointed out
that the distribution of collenchyma and other mechanical elements in
umbelliferous stems is liable to vary in response to environmental factors, to
undetermined causes, or at different stages in the development of the plant.
 UMBELLIFERAE 721
The arrangement of the mechanical tissue is, therefore, of restricted diagnostic
value, Mokeeva (1546) found that the stem structure of certain species of
Muretia and Scaligeria varies so much in plants growing under different
ecological conditions that the genera cannot be distinguished by the micro-
scopical features of the stem. All of the stem tissues in each individual species
are also very plastic. Lunan (1401) has demonstrated that the uppermost leaf
of Ligusticum scoticum Linn, is more xerophytic than the remainder,
Friedel and Yen (719) made an anatomical comparison between Eryngtum
campestre Linn, and Drypis spmosa Linn, (family Caryophyllaceae). These
2 xerophytes, although wholly unrelated in the taxonomic sense, show a
superficial resemblance to one another. Friedel and Yen showed, however,
that both species are anatomically similar to the family to which each belongs,
thus demonstrating that the hereditary anatomical characters are not pro-
foundly modified by environment.
Lemesle (1346) made a comprehensive study of the stem structure of
xerophytic members of the family from different parts of the world. The
following information is taken from the summary of his long, detailed article.
The epidermis is provided with a thick cuticle, or, in certain species from
the Mediterranean region, steppe regions, Chile, and Australia, the whole
epidermal layer becomes completely sclerosed. The collenchyma of the
primary cortex is sometimes strengthened or wholly replaced by fibres, whilst,
in certain species of Eryngium from the Mediterranean region,, the fibres
form a continuous cylinder. In other species, e.g. Chamarea capensis (Thunb.)
Eckl. et Zeyh. (syn. Carum capense-Sond.) and some Australian members of
the family, although the cortical collenchyma is reduced or absent, there are
sub-epidermal fibres which are either widely spaced or form an almost con-
tinuous ring. In other species the ground tissue of the cortex tends to become
sclerified, this feature being particularly well developed in Aptumpanul (Gay)
Reiche var. araucarwn (Phil.) Reiche (syn. Ligusticum pansil Bert.) from Chile.
A sclerified cortex was not found in species from Australia, but is common in
those from steppe regions. The cork is sometimes especially well developed,
and constitutes a cylinder of mechanical tissue. This occurs in species of
Eryngium, Hydrocotyle, Mulinum, Peucedanum, Pituranthos, and 'Trachymene*
from the southern hemisphere, particularly South Africa and Australia, in
some of which the cork arises in the sub-epidermis, but in others from the
inner part of the cortex (see 'Stem' on p. 717). Certain members of the family
from steppe regions have the cortex reduced to 2 or 3 layers of small cells.
Lemesle, after pointing out that the pericycle nearly always includes arcs of
fibres, that the xylem and phloem, particularly in species from South Africa
and Australia, are in the form of continuous cylinders or have fibrous inter-
fascicular tissue, that the pith is often persistent, reduced in size, and sclerified,
ends by pointing out that the secretory canals, especially those in the pericycle,
are often more numerous and of larger diameter in xerophytes than in species
growing where the climate is less severe.
Chodat and Vischer (402), who examined the anatomy of various Umbelliferae
from Paraguay from the ecological standpoint, found an interesting range of
structure in different species of Eryngium especially in those with a habit
y

recalling that of the monocotyledons. In this genus there are species with
xerophytic and others with hygrophytic features, whilst in a third category
722 UMBELLIFERAE
there a mixture of characters of both kinds. Thus in E. paniculatum Cav.
is
there an almost continuous zone of hypodermal sclerenchyma a xerophytic
is

feature combined with large intercellular spaces in the mesophyll as is

characteristic of hygrophytes. Xerophytic features are especially well
developed in E. hassleri Wolff, and related species with grass-like leaves.
Here the leaf area is reduced, palisade tissues are developed below all parts of
the surface, and hypodermal fibres are numerous. E. sanguisorba Cham, et
Schlechtd., with leaves like those of Luzula or Plantago^ also exhibits xero-
phytic characters, but of a rather different kind. Here there is no hypoderm,
but the cuticle is very thick; there are no lacunae in the mesophyll, which is
supported by strands of fibres radiating outwards from the aqueous tissue
around the vascular bundles. Hygrophilous forms include E. floribundum
Cham, et Schlechtd. var. serrioides Urb., belonging to the taxonomic group
areata. Here the cuticle is thin, the hypoderm is not sclerified, and the
palisade tissue is in direct contact with the upper epidermis. In pandani-
".

folium Cham, et Schlechtd., another hygrophyte, sclerosed hypoderm occurs

beneath each epidermis, there is assimilating tissue towards each surface,
whilst the mesophyll includes large intercellular spaces.
Chodat and Vischer's work provides an interesting addition to Solereder's
earlier statement that species of Eryngium with a monocotyledonous habit are

provided with longitudinal air canals, interrupted at intervals by transverse

diaphragms, whilst the vascular bundles, mostly in pairs of which the upper
member is lie longitudinally in the tissue between the
inversely orientated,
air canals. Solereder also records that in some species of Eryngium, broad
strips of palisade tissue, alternating with narrower strips of sclerenchyma,
occur below the epidermis or hypoderm towards both surfaces of the leaf,
Solereder also notes that Foeniculum vulgare Linn, has a leaf which, in
transverse sections, is circular in outline, with an envelope of palisade tissue
beneath the epidermis, locally interrupted by bundles of collenchyma. The
centre of the leaf in this species is occupied by a ring of vascular bundles on
the same radii as the collenchyma.
For further details concerning the species of Eryngium with a monocoty-
ledonous habit see Wolff's (2453) summary of Moebius's earlier work.
The ecological anatomy of 'Anisotome aromaticum Hook, f.* has been
described by Betts (188).

TAXONOMIC NOTES
Hoar (980) found the anatomical resemblances between the Araliaceae and
Umbelliferae to be closer than those between either of these families and the
Cornaceae. He therefore proposed that the Cornaceae should not be included
in the same cohort as the other 2 families. The fundamental similarity between
the Araliaceae and Umbelliferae has also been confirmed at Kew by Dr. C. L.
Hare. Amongst the more important resemblances he noted are the following.
The principal mechanical tissue in young stems of both families is in the form
of peripheral collenchyma, pericyclic sclerenchyma being scanty or absent.
Thin-walled, aqueous tissue is abundant in the stems of both families.
Although secretory cells were not observed in either family, similarly dis-
tributed secretory canals occur in both. Except in Dizygotheca no solitary
crystals were observed in either family. Medullary vascular bundles occur in
 UMBELLIFERAE 723
several genera of both families. Wherever these occur they are often collateral
and inversely orientated, or more or less centric with central xylem. The
range of vascular structure in the petioles of the 2 families is also very much
alike,types with numerous scattered bundles or with the vascular strands
i to several rings being the most frequent. Differences noted
arranged in
between the 2 families are mainly those correlated with the more woody
character of the Araliaceae a compared with the dominant herbaceous habit
in the Umbelliferae.
l

Majumdar (1421) attaches no phylogenetic

significance to the multiperforate
perforation plates which he observed in the primitive, spirally thickened vessel
members in the protoxylem of leaves and in the more specialized, pitted vessel
members which occur in the metaxylem of the nodal regions of stem of
Heracleum sphondylium Linn.

ECONOMIC USES
Certain members of the Umbelliferae are eaten as vegetables, others are the
source of gum-resins used in perfumery and medicine, and a third group
yield other products of medicinal value. Serious cases of poisoning have been
caused by a few species.
Familiar vegetables, derived from cultivated varieties of species belonging
to this family, include the carrot (Daucus carota Linn.), the parsnip (Pastinaca
sativa Linn.), celery (Apium graveolens Linn.), and parsley (Petroselinum
crispum (Mill.) Nym.). Angelica, prepared from the stems of Archangelica
officinalis Hoffm., is used in confectionery. The anatomy of the vegetative
organs of the parsnip has been described in some detail by Warning (2363),
9
who showed that the mature 'root consists of the true primary root and the
hypocotyl. Most of it is built up of amyliferous phloem parenchyma, the
central part being occupied by xylem, and, in the region of the hypocotyl, a
pith. The primary structure is diarch, although apparently triarch roots are
known. The oil ducts in the pericycle and primary phloem are relatively
ephemeral. The floral axis, initiated during the first year, develops rapidly
during the second. Hayward (927) has also described the anatomy of Apium
graveolens. According to Lambeth (1315) the medullary bundles of Apium
graveolens are cauline in nature, they may arise and terminate at any level in
the stem, and have no connexion with the leaf traces. The occurrence of
roots in the internal cavities in a celery 'tuber' has been described by Fourcroy
(699). Aseedling carrot with an abnormal cotyledon was investigated by
Parrot (1656), and 'the results of other work on the seedling anatomy of the
carrot published by Fourcroy (698) and Havis (922). For developmental
anatomy of the carrot see Esau's (654) article.
Gum-resins derived from this family include galbanum (Ferula galbamflua
Boiss. et Buhse and other species of Ferula); asafoetida (Ferula foetida Regel
and other species of Ferula); ammoniacum (Dorema ammoniacum D. Don).
All of these are obtained from Iran and adjacent regions. Moroccan ammo-
niacum is said to be derived from Ferula communis Linn. var. nodiflora Linn.,
and Cyrenian ammoniacum from F. marmarica Aschers et Taub.
Caraway seeds and oil are obtained from Canon carvi Linn., which is
cultivated in many parts of Europe; the fruits of Peucedanum graveolens
Benth. yield an oil used in the preparation of dill- water; oil from the fruits
734 UMBELLIFERAE
Mill.) is employed for flavouring and
and leaves of fennel (Foeniculum vulgare
in medicine, whilst aniseed (the seeds of Pimpinella anisum Linn.) is also
valued in medicine. The oil from Coriandrum sativum Linn, is used for
flavouring and in veterinary medicine.
Sumbul, which consists of pieces of the dried rhizome and rootstock of
Ferula sumbul Hook. f. and probably of F. suaveolens Aitch. et HemsL,
obtained from Turkestan, was at one time valued in medicine, but is not now
much employed. The commercial article consists of pieces about 3-6 cm.
long and wide. The brown to black outer surface is transversely wrinkled,
whilst fibrous leaf- trace bundles project from the scars of fallen leaves. Trans-
verse surfaces exhibit, internally to the thin layers of cork and bark, a ring of
yellowish vascular bundles, and additional medullary strands scattered
irregularly in the parenchymatous ground tissues in which resinous material
is also deposited.
Numerous other Umbelliferous plants have been used in folk medicine, but
are now unimportant. The root anatomy of 28 such species has been described
by Liermann (1370). According to Michlin (1512) the roots of Pimpinella
saxifraga Linn, can be distinguished from those of other members of the
Umbelliferae with which they might be confused by the fact that the cells of
the medullary rays are not radially elongated.
Members of the family with poisonous properties include Conium maculatum
Linn., Oenanthe crocata Linn., &c.

GENERA DESCRIBED
(i)
FOR GENERAL ANATOMY
Aciphylla, Aegopodium,* Aethusa, Ammi, Anethum, Angelica,* Aniso-
tome, Anthriscus, Apium,* Archangelica,* Astrantia, Azorella, Bowlesia,
Bupleurum,* Cachrys, Carum,* Caucalis, Cenolophium, Chaerophyllum,*
*
Cicuta, Conium,* Coriandrum,* Crithmum,* Cuminum, Daucus ; Echino-
phora,* Eryngium,* Falcaria,* Ferula,* Foeniculum,* Heracleum, Hermas,
Heteromorpha, Hydrocotyle, Laserpitium, Levisticum* Ligusticum,* Magy-
daris, Meum,* Mulinum, Muretia, Myrrhis,* Oenanthe,* Opopanax, Petro-
selinum, Peucedanum,* Pimpinella,* Pituranthos, Prangos, Sanicula,*
Scaligeria, Seseli,* Silaus, Siler, Sison, Sium, Smyrnium,* Thapsia, 'Trachy-
rnene', Trinia,* Xanthosia.
* Kew
Represented in the slide collection.

(ii) FOR WOOD STRUCTURE

(Aethusa), (Bupleurum), (Carum), Eryngium, Heteromorpha, (Hydro-
coytle), (Oenanthe), Peucedanum, (Pituranthos), Steganotaenia, (Xanthosia).
LITERATURE
(i) On General Anatomy
Berckemeyer 175, Betts 188, Brandscheit 256, Buchet 304, Chodat and Vischer 402,
Elias 625, Esau 651, 654, Fourcroy 698, 699, Friedel and Yen 719, Funk 733, Gue>m 837,
Ha vis 922, Hayward 92 7> Hector 929, Hoar 980, Lambeth 1315, Lemesle 1346, Liermann
1370, Lunan 1401, Majumdar 1421, 1424, 1425, 1427, Majumdar and Preston 14289
Meyer 1509, Michlin 1512, Mokeeva 1546, Parrot 1656, Perrot and Morel 1697, Pirschle
1726, Schwarz 2058, Sifton 2097, Tunman 2292, Tunnel 2299, Warning 2363, Wolff
2452, 2453, 2454.
(ii) On Wood Structure
Hoar 980, Majumdar 1422, Messeri 1493* Record 1843, 1851.