GENERA OF THE POTTIACEAE:
MOSSES OF HARSH ENVIRONMENTS
GENERA OF THE POTTIACEAE:
MOSSES OF HARSH ENVIRONMENTS
By Richard H. Zander
Illustrated by Patricia M. Eckel
BULLETIN OF THE BUFFALO SOCIETY OF NATURAL SCIENCES
VOL. 32
BUFFALO, N.Y.
1993
The Buffalo Society of Natural Sciences
Buffalo Museum of Science
1020 Humboldt Parkway
Buffalo, New York 14211-1293 USA
Bulletin of the Buffalo Society of Natural Sciences
Volume 32
Published December 1993
(Volume 33 was published in 1988.)
Cover illustration: see page 128.
Library of Congress Cataloging-in-Publication Data
Zander, Richard Henry. 1941Genera of the Pottiaceae: mosses of harsh environments I by
Richard H. Zander; illustrated by Patricia M. Eckel.
p. em. -(Bulletin of the Buffalo Society of Natural
Sciences. ISSN 0096-4131; v. 32)
Includes bibliographical references and index.
ISBN 0-944032-51-6 (hardcover)
1. Pottiaceae-Classification. 2. Pottiaceae-ldentification.
I. Title. II. Series.
OK539.P8Z36 1993
588'.2-dc20 93-37139 CIP
ISSN 0096-4131
ISBN 0-944032-51-6
Copyright © 1993 by The Buffalo Society of Natural Sciences
All Rights Reserved
The Bulletin of the Buffalo Society of Natural Sciences first appeared in 1873. Please contact the Publications Sales
Division, Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, NY 14211 USA, for back issues and price lists.
v
TABLE OF CONTENTS
Preface........................................................................................... vi
Introduction .................................................................................... !
Previous Work ......................................................................... 1
Taxonomic Characters ............................................................. 3
Technique ................................................................................ 8
Practical ldentification ........................................................... lO
Most Recent Suprageneric Classification .............................. 11
Evolutionary Relationships .......................................................... 12
Phylogenetic Analysis ........................................................... 16
Cladograms ............................................................................. 33
Data Set.. ................................................................................ 50
Taxonomic Section ...................................................................... 52
Pottiaceae ............................................................................... 52
Key to Suprageneric Taxa ............................................... 53
Key to Genera .................................................................. 54
Timmielloideae ...................................................................... 68
1. Timmie/la .............................................................. 68
Erythrophyllopsoideae ........................................................... 71
2. Erythrophyllopsis .................................................. 71
3. Erythrophyllastrum ............................................... 72
Gertrudielloideae ................................................................... 74
4. Gertrudiella ........................................................... 74
Chionolomoideae ................................................................... 76
5. Chionoloma ........................................................... 76
6. Pseudosymblepharis .............................................. 79
7. Pachyneuropsis ..................................................... 80
Trichostomoideae .................................................................. 82
8. Calymperastrum .................................................... 83
9. Eucladium ............................................................. 84
10. Trichostomum ...................................................... 86
11. Tuerckheimia ....................................................... 94
12. Streptocalypta ..................................................... 94
13. Pleurochaete ....................................................... 96
14. Calyptopogon ...................................................... 98
15. Tortella .............................................................. 100
Merceyoideae ....................................................................... l05
Tetracoscinodontieae ..................................................... l05
16. Tetracoscinodon ................................................ l05
Bryoerythrophylleae ...................................................... 107
17. Dialytrichia ....................................................... 107
18. Rhexophy/lum .................................................... llO
19. Mironia .............................................................. ll2
20. Bryoerythrophy/lum .......................................... 112
21. Pseudocrossidium ............................................. 116
Leptodontieae ................................................................ 120
22. Triquetrella ....................................................... 120
23. Reimersia .......................................................... 123
24. Hymenostylium .................................................. l23
25. Trachyodontium ................................................ 127
26. Streptotrichum ................................................... l30
27. Leptodontiella ................................................... 130
28. Leptodontium .................................................... 132
Barbuleae ....................................................................... l36
29. Anoectangium .................................................... l36
30. Gyroweisia ........................................................ 138
31. Bellibarbula ...................................................... 140
32. Streptopogon .................................................. 142
33. Barbula .......................................................... 145
34. Gymnostomum ................................................ 15l
35. Scopelophila ................................................... 153
36. Gymnostomiella ............................................. 156
37. Didymodon ..................................................... 157
Pottioideae ........................................................................ 163
Hyophileae ................................................................. 163
38. Hypodontium .................................................. l 64
39. Hymenostyliella .............................................. l66
40. Molendoa ....................................................... 167
41 . Hyophila ......................................................... l70
42. Plaubelia ........................................................ 174
43. Teniolophora .................................................. l76
44. Weissia ........................................................... 176
45. Weissiodicranum ............................................ l84
46. Quaesticula .................................................... 186
47. Ganguleea ...................................................... 186
48. Weisiopsis ...................................................... 189
49. Luisierella ...................................................... 193
50. Stegonia .......................................................... l 95
51. Crossidium ..................................................... 196
52. Pterygoneurum ............................................... l 99
53. Globuline/la ................................................... 202
54. Aloina ............................................................. 203
55. Aloinella ......................................................... 206
Pottieae ....................................................................... 208
56. Leptobarbula .................................................. 208
57. Tetrapterum .................................................... 209
58. Trachycarpidium ............................................ 2 11
59. Aschisma ........................................................ 213
60. Bryoceuthospora ............................................ 216
61. Uleobryum ...................................................... 2l8
62. Tortula ............................................................ 2 l 8
63. Willia .............................................................. 233
64. Saitoe/la ......................................................... 236
65. Microbryum .................................................... 237
66. Crumia ........................................................... 240
67. H ennediella .................................................... 242
68. Dolotortula ..................................................... 248
69. Phascopsis ...................................................... 249
70. Stonea ............................................................. 252
71. Acaulon .......................................................... 253
72. Sarconeurum ................................................. 255
73. Chenia ............................................................ 255
74. Syntrichia ....................................................... 258
75. Hilpertia ......................................................... 270
76. Sagenotortula ................................................. 273
Excluded Taxa ........................................................................ 274
Updated List of Genera, Species
and Infraspecific Taxa ...................................................... 279
Bibliography ........................................................................... 348
Glossary .................................................................................. 372
Index ....................................................................................... 373
Corrigendum ........................................................................... 378
vi
PREFACE
This study is a taxonomic treatment of the moss family Pottiaceae at the genus level, and is summarized in the Table of Contents. It was begun in 1984 but involves more than two decades of specialization on the part of the author. The genera are
viewed in the traditional sense as groups of species sharing either a unique combination of several character states (or a
majority of such states) or standing out in sharing (or having, in the case of the several monotypic genera) one or more
unusual features. Subfamilies and tribes, however, are distinguished with the aid of cladistic analysis, answering the question:
"What would be the best suprageneric classification assuming the family as presently conceived was monophyletic and its
genera as now circumscribed were each monophyletic?" Certain generic concepts have been considerably emended to
increase the chances that future study of the individual genera will find them at least in large part monophyletic.
As critics have noted, the original Hennigian cladistic method of evolutionary analysis is overly simplistic in that evolution as we understand it is incompletely modeled, e.g. there are no provisions for dealing with introgression, convergence,
fossils or anagenetic change. Thus, no matter how excellent the data and "rigorous" the analysis, the simple model produces a
simplified and surely at least partially incorrect evolutionary hypothesis. On the other hand, the present popularity of phylogenetic cladistics among taxonomists reflects a general apprehension that, for all its faults, it remains an excellent method of
analyzing large taxonomic data sets, and it is possible to modify the method to some extent when we actually have information about the aspects unmodeled. It succeeds in using large data sets to parsimoniously group taxa, however simplistically,
on the basis of shared, recently evolved traits, a valuable first step in evaluating evolutionary relationships. That the largescale results are commonly similar to evolutionary relationships suggested by past systematic study, as is the case in this
work, is further support for an assumption that the higher resolution possible in cladistic studies also reflects valid hypotheses
of relationships. The details of standard cladistics-generated evolutionary hypotheses, however, should be examined with
other methods that reflect better theory.
In practice, the present suprageneric classification is based totally on a cladistic analysis. That convergence in sporophyte
features is a reality in many genera of the Pottiaceae is obvious to the specialist, and is well illustrated in this text. In view of
the above reservations, I have had no qualms about differentially weighting the characters used in the analysis against possible convergence through reduction. Also, when discussing genera, I point out those relationships between certain taxa that
seem plausible, probable and due to convergence, even if such relationships are contrary to the results of the cladistic analysis. Nor do I spurn the past work of non-cladists. In pursuit of a phylogenetic classification, a revisionist can with traditional
methods build on what is already known, or start afresh with a new, much heralded but immature analytic technique, or, as
here, in the face of a taxonomic group impossible to comprehend in only one lifetime, necessarily attempt both.
Considerable new nomenclature is presented in this treatment: four new subfamilies, four new tribes, two new genera,
two new subgenera, and two new sections, and one new variety. There are also 37 new names for taxa and 317 new combinations. The total number of recognized taxa in the Pottiaceae now includes seven subfamilies, six tribes, 76 genera, 1457 species, 31 subspecies, 536 varieties, 339 formae and seven subformae.
Part of the cost of research and publication has been underwritten by the National Science Foundation, Grant BSR8314843. I thank H. Bischler and R. A. Pursell for help in obtaining specimens from the Bizot herbarium at PC, William D.
Reese for comments on portions of the text draft, and J. Cargill, R. Halling, W. Margadant and G. Zijlstra for correspondence
regarding nomenclature and associated problems. J.-P. Frahm and P. J. Lightowlers made valuable comments on slide mounting media. W. R. Buck, R. Magill and W. Reese gave helpful observations on various taxonomic and nomenclatural topics.
D. Wagner discussed terminology for directions of helical twist in plant parts. W. Gall, R. Vineyard, E. DeLuna, B. Mishler
and B. Tan provided valuable comments on the phylogenetic analysis, but the execution, results and interpretation are
entirely my own. C. Delgadillo, D. Griffin, B. Murray, D. Norris, P. Sollman, and B. Tan tried out the keys and provided
many worthwhile observations. M. R. Crosby commented on details of the myriad nomenclatural changes. Correspondence
and discussion with more than a· few bryologists over the years helped firm the taxonomy developed here, and I thank these
colleagues. Many workers have openly shared with me their opinions on aspects of taxonomy of the Pottiaceae; I have, to the
best of my recollection, given their ideas, when reflected here, proper attribution. I also appreciate the diligence and courtesy
extended to me by the curators at the institutions from which I borrowed specimens; this work could not have been done
without their cooperation and patience. I greatly appreciate the help of Allen Press, especially G. Dresser and K. Blair, in
expediting the printing of this volume from our camera-ready copy. The Buffalo Museum of Science and especially its Director, Ernst Both, provided significant research support and an atmosphere conducive to uninterrupted, long-term study.
This book is dedicated to Patricia M. Eckel, colleague and spouse, who provided unflagging support and sensitive suggestions throughout the long evolution of this work. Her fine illustrations furnish an essential reinforcement for the generic concepts proposed here and round out the descriptions in an elegant manner.
INTRODUCTION
The taxonomy of the Pottiaceae is considered by many to be less than easy and largely the province of adepts. This is due to intrinsic problems such as polymorphy, taxonomic importance of anatomical characters, reduced size of the plants, obscure areolation,
and sterility of many specimens, and to extrinsic factors such as lack of regional identification manuals (especially for tropical
areas), few revisions, many poorly conceived "geographic" species, and the heterogeneity and large size of some genera.
The latest intensive treatment of the family at the generic level
was published by Brotherus in 1924, being a series of short
descriptions with keys to genera and lists of species (some species
lists presented with keys). Of the 85 genera recognized in the
family (of 795 in the mosses fide Crosby & Magill 1981) just previous to the present study, Brotherus described only 60 and illustrated only 41. His keys generally require a specimen that has a
sporophyte. Many new genera and generic synonymies have been
published since 1924 in various journals that may be hard to
obtain by botanists without easy access to the large libraries of
developed countries. A major problem in identification of mosses
of tropical and other non-Western areas is the lack of a modern
key to genera that reflects recent (past 50 years) work on taxonomy. The twenty-five accepted genera of Pottiaceae not found in
Brotherus' (1924-25) work must be tracked down through a large
and scattered literature.
This study is an overview of the family providing a taxonomic
introduction and means of identification for the genera of the
Pottiaceae. The family is the largest in the Bryopsida in number
of genera, 76 being recognized here, and consists of an agglomeration of many small, fairly well-defined genera and a few
large, poorly defmed genera much in need of revision. Because
the author believes that important morphological transformation
series may cut across presently recognized generic boundaries,
no concerted attempt was made at extensive revision (that is,
reapportionment of groups of species) at the generic level
except in the Phascum-Pottia-Desmatodon-Tortula complex
and in the Weissia-Trichostomum relationship. Instead, a key to
recognized genera, a cladistic evaluation, comprehensive
descriptions, discussion of problems, a bibliography, detailed
illustrations, and a list of the correct names of the taxa are provided to stimulate further study, floristic, taxonomic and phylogenetic. This is not a "type revision"; the descriptions are based
on the study of a selection of specimens of representative species, names of which are listed with citations of herbaria consulted. The discussions are the result of the author's 25 years of
specialization in the group and of new observations made during this study.
PREVIOUS WORK
The more than 1400 species of the Pottiaceae are characteristic of variable or harsh environments, and may form a conspicuous portion of the vegetation of ruderal, arid land, alpine or arctic areas. They exhibit a great variety of apparent morphological, physiological and genecological adaptations to their particular environments, mostly poorly or not at all understood.
The Pottiaceae is here reviewed with certain modifications at the
generic level reflecting modern recognition that morphological
reduction series are characteristic of many genera of the Pottiaceae, cf discussions below of Barbula, Bryoerythrophyllum,
Didymodon, Hennediella, Tortula, Trichostomum and Weissia,
among others.
Several common species of Pottiaceae in England were
described and illustrated by J. Dillenius (1741) in his early bryological manual. J. Hedwig (1801) described and illustrated many
pottiaceous species from regions worldwide in his "Species
Muscorum," from which moss nomenclature (except that of
Sphagnum) begins. The genera of Hedwig, however, were usually
taxonomically heterogeneous.
The name Pottiaceae was first used by Bruch et al. (1843) in
the "Bryologia Europaea" (1836-1855). The authors included
only three genera, while other genera now placed in the Pottiaceae
were recognized in segregate families (Phascaceae, Weissiaceae
or Trichostomaceae), these based largely on sporophytic characters. Most other authors in the 19th Century followed the practice
of dividing the Pottiaceae into smaller families. Mitten (1859)
recognized only one family, however, the Trichostomaceae,
which he later called the Tortulaceae (Mitten 1869). Brotherus
(1902-09) also recognized only one family, the Pottiaceae, with
46 genera in four subfamilies. Later, Brotherus (1924-25), in a
treatment for Engler and Prantl 's "Die Natiirlichen
Pflanzenfamilien," recognized 71 genera in five subfamilies in
the Pottiaceae. Brotherus' treatment remained the standard
compiliative work on the family to this date. Recent authors
generally follow the single family concept of the Pottiaceae, but
Cinclidotus is variously treated in the Pottiaceae or in its own
family, the Cinclidotaceae. Saito (1975a) gave a detailed discussion of the historical development of family and suprageneric
taxonomic concepts for the Pottiaceae.
Three papers published since Brotherus' ( 1924-25) treatment contributed significantly to generic and suprageneric classification.
Hilpert's (1933) "Studien zur Systematik der Trichostomaceen" was a treatment at the generic level of the
"Trichostomaceae," which was composed of what have come to
be recognized (sensu Saito l975a) as the tribes Trichostomeae,
Barbuleae and Pleuroweisieae (= Hyophileae here) of the Pottiaceae. Hilpert recognized three subfamilies and three tribes in
the Trichostomaceae while the rest of the genera that Brotherus
had recognized in the Pottiaceae sensu lato were placed by Hilpert in the Pottiaceae and Cinclidotaceae, and not studied. Hilpert discussed at length morphology and taxonomic importance
of characters of the areolation, costa, curvature of the leaf margins, differentiation of areolation at the leaf margins, papillae
2
GENERA OF THE POTTIACEAE
and mamillae, propagula, perichaetial leaves, annulus and peristome. He pointed out important differences between the
Trichostomaceae and the Pottiaceae in such morphological characters as leaf shape, areolation, costal anatomy and paraphysis
shape. The morphology of each genus was discussed in relationship to related genera. Many species names were transferred from
one genus to another. He described as new the genera Semibarbula, Prionidium and Macroglossum. He suggested that Sarconeurum was probably near to or a synonym of Tortula, that Leptodontiopsis belonged in the family Orthotrichaceae, that Chrysoblastella and Rhamphidium (see Excluded Taxa below) belong in
the Ditrichaceae, and that W eisiopsis belonged in the Pottiaceae
sensu stricto (=Pottioideae sensu Saito 1975a). The chief value of
Hilpert's (1933) work is that it provided the only discussion of the
many tropical genera published since Brotherus' (1924-25) treatment, and presented modern charts showing purported supraspecific phylogenetic relationships.
Chen's (1941) "Studien tiber die ostasiatischen Arten der
Pottiaceae" is a landmark treatment of the Pottiaceae. Primarily a
floristic work concerning only those species in eastern Asia, this
study included detailed descriptions and extraordinary illustrations of species in 32 genera, discussions of morphological characters at the genus level, floristic relationships of eastern Asian
species, and a lengthy evaluation of intra-familial phylogeny with
a detailed chart of presumed relationships.
Chen recognized six subfamilies, the Cinclidotoideae, Pottioideae, Trichostomoideae, Eucladioideae, Leptodontioideae and
Barbuloideae. These were distinguished by such characters as
capsule and costal anatomy, leaf shape and margin recurvature,
length of the operculum relative to that of the theca, differentiation of the leaf base and morphology of the laminal papillae.
Characters that distinguished the genera were variations and combinations of these plus such characters as plant size, shape of leaf
apex, length of the costa relative to that of the leaf, etc. Chen
made many new combinations at the species level, and described
a few new species and the genera Reimersia and Bellibarbula. He
provided Bryoerythrophyllum as a nomen novum for the illegitimate homonym Erythrophyllum (Lindb. in Braithw.) Loeske, and
was the first worker to recognize Bryoerythrophyllum as a rather
large genus. Of the generic complexes most juggled about in identification manuals today, Chen lumped Hymenostomum and
Astomum under Weissia, Didymodon under Barbula, and
Hymenostylium under Gymnostomum, but recognized as separate
Tortula and Syntrichia (sensu Chen, not Zander).
Saito's (1975a) "Monograph of Japanese Pottiaceae," gave
extensive discussion of the generic and suprageneric classification
of the Japanese taxa, and presented a nicely explained classification system based in part on several new characters and thorough
morphological and anatomical study. The most conservative characters in the family were considered to be curvature of the leaf
margins, shape of the area of differentiated basal leaf cells, the
occurrence of gemmae, and characters of the sporophyte. Saito
found that variations in the peristome (reduction series and
"complication" series) within certain generic complexes were correlated with variation in other characters of the sporophyte, such
trends varying in parallel. He made new statuses of names at the
intrageneric level and described a new subgenus (Barbula subg.
Odontophylla Saito). He merged Astomum and Hymenostomum
with Weissia, Hymenostylium with Gymnostomum, and Syntrichia
(sensu Chen) p.p. with Tortula (in an earlier paper, Saito 1973a),
but kept as separate Barbula, Bryoerythrophyllum and Didymo- ·
don. Saito, like Chen (1941), provided excellent illustrations of
all species including such important details as papillae morphology and transverse sections of the leaves. Saito's major contributions to supraspecific classification are the well-characterized
and well-justified subfamilies, tribes, genera, subgenera and
sections of Japanese Pottiaceae, often based in part on new
anatomical and morphological characters.
Ninety genera were recognized in the Pottiaceae by Crosby
and Magill (1981) in their "Dictionary of Mosses." Since the
publication of this list, revisionary and floristic work has
resulted in the synonymizing of some generic names with others
and in the description of new genera, reducing the total to 85 at
the beginning of the present study in 1985. There are, however,
only 76 recognized genera at its completion, reflecting considerable synonymy and removal of genera to other families.
My recent publications on the family, too, have been generally limited to one floristic region, the New World or parts of it.
In this study, however, several genera were deemed insubstantial and synonymized with other genera with previously published names. For example, Trichostomopsis (Zander 1978e)
and Husnotiella (Zander 1981c) were made synonyms of Didymodon, and Barnesia a synonym of Streptocalypta. Leptodontiella was erected as a segregate of Leptodontium (Zander &
Hegewald 1976). I agreed for the most part with Saito (1975a)
in distinctions between Barbula and Didymodon (Zander 1978e,
1979f, 198lc). I used color reactions of leaf laminae to various
acidic and basic reagents (Zander 1980a) as characters valuable
in distinguishing Barbula and Bryoerythrophyllum, showing
that twisted peristomes are not restricted to the former genus. I
used the phenological feature of date of sporophyte maturation
to characterize some genera of the Pottiaceae and to place certain species in their proper genus (Zander 1979d). My taxonomic methods and concepts are more extensively summarized
elsewhere (Zander 198lc, 1982e, 1982g, 1985b).
I have specialized in the taxonomy of the Pottiaceae since
1967. My larger papers include treatments of Leptodontium
(1972), Tuerckheimia (1978f); and Streptocalypta (1983a) in
the New World; the tribe Pleuroweisieae in Middle America
(1977c); Barbula and Pseudocrossidium (1979f) and Didymodon (1978e) in North America north of Mexico; Bryoerythrophyllum and Morinia [= Mironia] in the New World (1978g);
Barbu/a, Pseudocrossidium and Bryoerythrophyllum p.p. in
Mexico (1981a); and Didymodon in Mexico and California
(1981c). Treatments of 33 of the genera of Mexican Pottiaceae
for a moss flora of Mexico (A. Sharp et al. 1993), and of the
Trichostomoideae for a moss flora of the North America,n Arctic and Greenland (G. Mogensen, ed.) have been completed.
Significant recent taxonomic studies in the Pottiaceae by
other authors include treatments of Aloina, Aloinella and Crossidium by Delgadillo (1975a and in Sharp et al. 1993); of Weissia subg. A.stomum in Africa by Crundwell and Nyholm (1972a,
1974); of Tortula sect. Rurales by Kramer (1978, 1980, 1988);
of Globulinella by Magill (1977a); of Pottia in Great Britain by
Chamberlain in A. Smith (1978); of Hymenostyliella and
Trichostomopsis by Robinson (1970, 197la); of Tortula by
Mishler (in Sharp et al. 1993) of Triquetrella by Casas de Puig
et al. (1993); of Phascum by Guerra et al. (1991); and of Weissia by Stoneburner (1985)-see also the section on Practical
Identification below and the Bibliography of works on the Potti-
IN1RODUCTION
aceae. A relatively few identification manuals, such as those of
Gangulee (1969-80) for eastern India, Crum and Anderson (1981)
for eastern North America, Catcheside (1980) for South Australia,
Magill (1981) for South Africa, Norris and Koponen (1989) for
Papua New Guinea, Eddy (1991) for Malesia (Malaysia and the
East Indies), and the multiple author treatment edited by Sharp et
al. (1993) for Mexico, provide exacting taxonomic treatments and
considerable new information on the Pottiaceae.
Recent developments in taxonomic techniques have resulted
in the use of new taxonomic characters in the Pottiaceae. Most of
these are simply the results of more detailed observation of morphology and anatomy. Some examples may be mentioned. Saito
(1974a, 1975a) asserted that the subfamily Trichostomoideae may
be distinguished from other subfamilies of the Pottiaceae by the
five, rather than four, amphitheciallayers of the capsule. Insertion
patterns of rhizoids are distinctive features distinguishing the
Pottiaceae from certain other families with similar gametophytes
according to Norris (in Norris & Zander, 1981). Saito (1975a)
made great use of the morphology of axillary hairs in distinguishing genera of the Pottiaceae. He did not consider the hymenium of
Hymenostomum to be an organ distinct from the columella, and,
for this reason, recognized Hymenostomum only at subgeneric
rank under Weissia. The scanning electron microscope was used
in studies of spore and lamina ornamentation for taxonomic evaluations of some genera of Pottiaceae by Lewinsky (1974), Saito
and Hirohama (1974a,b), Zander (1972), Zander and Vitt (1979),
and others.
The present study reflects an emphasis on leaf characteristics
in defining higher categories begun by C. Miiller with his "Syn-
3
opsis Muscorum" (1849) and continued by Braithwaite (1887)
and Mitten (1869) in their works. Most recent authors, including
myself, have emphasized at least some of a number of unobtrusive anatomical or morphological features as being of considerable importance at the generic and specific levels, e.g. presence
or absence of a stem central strand, or of a stem hyaloderrnis, or
of a costal epidermis or hydroid strand (Begleiter cells); the
morphology of the laminal papillae; characters of the lamina!
cell walls and areolation patterns; position and size of propagula, and so forth. These relatively new characters, however,
have been used in revisionary or floristic studies of only a small
proportion of the species and, until the present work, of the genera of the family. A large number of species remain unstudied
or uncritically examined since their original description during
the main exploratory phase of bryological floristic development
in the 1800's and early 1900's.
The genera studied are illustrated with one or more species,
these selected to demonstrate the range of variation in morphological and anatomical features considered of taxonomic importance. Usually pictured are at minimum a habit, leaf, leaf apex,
leaf basal cells, sections of stem and leaves; also commonly
shown are upper lamina! papillae, propagula, peristome details,
operculum, and calyptra. Since size as a taxonomic character is
usually given as a range, magnifications are not provided for the
illustrations; however, the reader may assume that for all but
very small and very large taxa, figures of comparable features
are enlarged to a similar extent. Actual measurements for plants
and their morphological features should be obtained from the
descriptions.
TAXONOMIC CHARACTERS
Comments on specific morphological and anatomical features of taxonomic importance are given in the following paragraphs, while
evaluations of phylogenetic polarizations of characters are related in the section on cladistics. A number of treatments describe the
morphology of the mosses in detail; those of Flowers (1973a) and of Saito (1975a) include much up-to-date discussion and illustration of the morphology of the Pottiaceae.
Color
Emphasis is placed on natural coloration (see discussion of
MArtensson & Nilsson 1975) as a character ~nly for those taxa in
which color correlates with other, taxonomically important characters, e.g. Erythrophyl/opsis (red leaves) or Scopelophila (blackish leaves). Even such genera with characteristically red or black
leaves can be mimicked by placing a related but less colorful
genus in KOH solution to develop leaves of a red coloration or in
ferric chloride solution to create blackish leaves (cf Field 1977).
Thus, substrates that are highly alkaline or are associated with
iron deposits may produce false color characters in certain taxa,
just as characteristic coloration of other taxa may be suppressed in
acid or iron-deficient habitats. As a rule, highly colored plant
parts are yellow in acid solution and red in alkali, but this often
varies per genus. A basic chemistry seems to permeate collections
growing on calcareous rock, judging, for instance, from the strong
bubbling of gas commonly produced from capsules of calciphiles
placed in dilute HCl solution. One of the long-used cha:acters of
Barbula convoluta is the yellow seta. This seems to be a stable
character, as the color becomes merely light orange in dilute KOH
solution, quite different from the deep red evoked in, say, the
setae of B. unguiculata. The peristomes of both species, however, show the same reactions to dilute HCl and KOH solutions,
being light yellow in the first and deep red in the second.
Changes in color can be expected, too, for observations of peristomes in lactophenol gel or Hoyer's solution mounting media
(both acidic) or the highly alkaline KOH solution that might be
used to help remove the operculum. For accuracy, color of plant
parts should be used as a taxonomic character only when such
parts are immersed in the acid or basic media standard for such
observations.
The present taxonomic treatment emphasizes color
responses of leaf lamina! walls to two-percent KOH solution.
These color reactions are usually either brick red or a bright yellow, occasionally orange, seldom negative (colorless, no
change, or with the color of chlorophyll). Apparently (R. Mues,
pers. comm.), the yellow and red reactions are characteristic of
phenolic compounds-those compounds with only one or no
hydroxyl groups giving a yellow color in KOH, while those
with two or more closely associated hydroxyls will yield red.
All other descriptions of coloration are given here as the natural
state, and the general taxonomic value of these is not empha-
,-
4
GENERA OF THE POTITACEAE
sized. Often, the yellowish green of chlorophyll masks somewhat
the reaction of the cell walls to KOH (e.g. in the case of Chenia
leptophylla, which at first glance appears light yellowish green in
KOH under the dissecting microscope); when in doubt, the taxonomist should look at the lamina under high magnification to determine the actual color of the cell walls (which are actually bright
purplish red in C. leptophylla).
The color reaction of the laminae to KOH solution is apparently unaffected by at least some chemical treatments involved in
collection of mosses. Plants of Bryoerythrophyllum ferruginascens were boiled in 38% formaldehyde solution and also in 95%
ethyl alcohol and, after rinsing, the laminae retained their ability
to switch from red to yellow in dilute hydrochloric acid and back
to red in two-percent KOH.
Stems
The stem of the Pottiaceae when highly developed (e.g. Timmie/la
spp.) has four morphologically distinctive layers, the central
hydroid strand (hereafter usually termed simply the central strand)
of thin-walled, often partially collapsed cells; the central cylinder
of parenchyma; the cortex, often differentiated as a sclerodermis
of stereid or substereid cells; and the hyalodermis of enlarged
epidermal cells, usually thin-walled and often collapsed in mature
parts of the stem (e.g. Pl. l, f. 7). There is much variation in the
presence or absence of these features, and in the degree of expression, but these are often taxonomically important and examining
stem sections must become standard practice.
Central strand. Hilpert (1933) found that the central strand was
sometimes absent in species in which it was otherwise usually
present, Trichostomum tenuirostre being a case in point. Saito
(1975a) indicated that Didymodon asperifolius always lacked the
central strand, but Zander (1978e) questioned this. In Hymenostylium recurvirostrum, one variety occasionally has a central strand,
but it is otherwise absent (Pl. 32, f. 1) in the typical variety. In
spite of some variation, however, presence or absence of the central strand is generally a good character at the generic level in the
Pottiaceae, being, for instance, always absent in Leptodontium
(Pl. 36, f. 2-3; 37, f. 10) and Scopelophila (Pl. 47, f. 2). Occasionally, the stem is hollow (Pl. 3, f. 2), in which case a central strand
was probably present (section near the stem apex to check this).
"Satellite" strands visible in sections in the cortex of the stem
occur in Alaine/la and Calymperastrum, and are connected with
the leaf strand, but not with the central strand of the stem.
Central cylinder. The central cylinder of cells is usually of
medium-sized, thin-walled parenchymatous cells, but may occasionally be anatomically distinctive. Its cells may be uncommonly
large, as in species of Barbula (Pl. 43, f. 2), or thick-walled, as in
many species of Trichostomum (Pl. 11, f. 7), Tarte/la (Pl. 18, f.
16) and related taxa.
Cortex. The cortex of the stem of Pottiaceae usually consists of
cells with smaller lumens than those of the central cylinder. It is
sometimes differentiated as sclerodermis, which is well developed
in many genera of Pottiaceae and consists of small, thick-walled,
longitudinally fusiform cells, the "stereids." In most cases, the
presence of a sclerodermis is easily determined. In Barbula, especially, the sclerodermis is usually strongly differentiated from the
enlarged parenchyma of the central cylinder, but occasionally, the
outer cortex consists of "substereid" (Pl. 31, f. 2; 39, f. 3) cells
difficult to interpret.
Hyalodermis. This consists of usually one layer of enlarged,
thin-walled cells on the surface of the stem, commonly in combination with a sclerodermis or at least an outer cortex of smalllumened cells. Like the central strand, this character is variable
in expression in some taxa, both as to presence and degree of
differentiation, yet it is often of value taxonomically. In Leptodontium (Pl. 36-37), the hyalodermis is characteristic of certain
sections of the genus, but not of others (Zander 1972). A collapse of the thin outer walls of the hyalodermis in mature stem
parts of many but not all taxa may give a "fluted" appearance to
the stem section.
Axillary hairs. The filaments (Pl. 44, f. 21; 59, f. 15-16) borne
in the axils of the leaves of Pottiaceae are generally from 2-15
cells in length, consisting of hyaline, uniseriate, cylindrical cells
or rarely these bulging and thus producing a moniliform filament (cf Molendoa and Gymnostomiella). The basal 1-2 cells
are either similar to the rest or are more thick-walled and
brownish or yellowish. The terminal cell is apparently never terminally pored as in the axillary mucilage hairs of species of certain other families (e.g. Brachymitrion jamesonii Tayl. of the
Splachnaceae), but occasionally may be similarly swollen (e.g.
Globulinel/a globifera). Ligrone (1986) has described the
ultrastructure and cytochemistry of the mucilage-secreting
axillary hairs of Timmie/la.
Leaves
The cauline leaves of the Pottiaceae are typically lanceolate to
spathulate. Although the basal cells are differentiated in most
genera, leaves with more or less sheathing bases are less common, and are characteristic of the Leptodontium (Pl. 36-37)
group, Pleurochaete (Pl. 16), and certain other genera-these
taxa usually of relatively large physical size. Such leaves may
be squarrose-recurved, reflexed at the top of the basal sheath
(Pl. 3-4) or rarely wasp-waisted as in Timmie/la (Pl. 1). Leaf
shape may be variable in a characteristic fashion such that,
within the same species, there appears to be a standard differential development of character states. In species with much variation in leaf shape, leaf width commonly varies far less than leaf
length, perhaps because stem width also varies proportionately
less than leaf length. Thus, in some one species or genus,
shorter leaves are ovate, and are characteristically stiff and erect
in stance when dry, and longer leaves are generally scarcely
wider to sometimes twice as wide, but may be three to four
times as long as are the short leaves, and then mostly lanceolate
in shape, more easily twisting or curling when dry simply
because they have a high length to width ratio. So what may be
perceived as a number of characters (including leaf length and
stance), correlated with plant size may all be based on a single,
simple developmental feature found in many species with variable leaf shapes. Leaves that twist, flex or curl or become tubulose when dry may in that way lessen heat stress. In vascular
plants, research on Spartina pectinata Link (Gramineae) by S.
Heckathorn and E. DeLucia (Anon. 1990b) demonstrated that
leaf rolling reduced surface exposure to sunlight, lowering leaf
temperature by five degrees Cent.; J. Lebkuecher and W.
Eickmeier (Anon. 1990b) found that leaf curling in Selaginella
,·
INTRODUCTION
lepidophyl/a (Hook. & Grev.) Spring. (Selaginellaceae) reduced
high radiation damage that occurred during desiccation. See also
discussion of surface wax by Proctor (1979b).
There are at least six distinct trends in the family towards
elaboration of specialized photosynthetic leaf tissue:
l. Corrugated or rasp-like leaves, such as are developed in
Anoectangium aestivum variants (see Zander 1976).
2. Costal lamellae on the ventral laminal surface, as in Pterygoneurum (Pl. 72, f. 9) or species of Acaulon (Pl. 102, f. 12,
24-25), or the dorsal surface as reported in species of Hymenostyliella.
3. Ventral costal filaments, as in Aloina (Pl. 75, f. 8, 14), Aloinella, Crossidium and Pseudocrossidium.
4. Bulging ventral costal surface, as in the ventral surface of
certain species of Tortula sect. Tortula, especially T. atrovirens
(Pl. 85, f. 8) and T. revolvens (Pl. 89, f. 4), with the epidermal
cells generally higher than wide in transverse section.
5. Protected tissue inside leaf margin revolutions consisting of
swollen, thin-walled and coarsely hollow-papillose tissue in species of the genus Pseudocrossidium (Pl. 27, f. 13, 17), and in Hilpertia (Pl. 112, f. 4-5) forming a densely chlorophyllose cylinder
along each lateral margin.
6. Ventrally bulging and dorsally nearly flat upper laminal
cells that may enhance photosynthesis in periods of dim light and
dew in arid habitats by focusing light on the chloroplasts. Papillae
may obscure the asymmetric bulge of such cells, and sections are
often necessary to demonstrate these. Leaves with this distinctive
areolation are found, for example, in genera of the Hyophila complex (Hyophileae sensu this treatment; Pl. 54, f. 7, 16) and in Gertrudiel/a and Timmiella. Upper laminal cells that are strongly
bulging on both sides, as in some species of Syntrichia (Pl. 105, f.
7), may be an equivalent adaptation to dim light. Delgadillo
(1984) listed several taxa characteristic of the Yucatan Peninsula
of Mexico with a similar phenotype including ventrally bulging
upper laminal cells.
Costa
Wyatt (1985) has reviewed the complex and various terminology
that has been applied to moss costal anatomy.
Genera that consistently lack a ventral epidermal layer are
Aschisma (Pl. 81, f. 7-8, 17), Gymnostomiella, Leptodontiella,
Leptodontium (Pl. 36, f. 8), Reimersia, Streptotrichum, Trachyodontiutn, and Triquetrella. This is also characteristic of Hymenostylium (Pl. 31, f. 7, 16) except in one variety of H.
recurvirostrum (Pl. 32, f. 6).
The costal hydroid strand (the Begleiter cells of many earlier
authors, and hereafter referred to in context simply as the hydroid
strand) when present is found between the guide cells and the dorsal stereid band, or rarely also between the guide cells and the
ventral stereid band (e.g. Calymperastrum, Pl. 9, f. 9; Barbula
riograndensis, Pl. 45, f. 12). The hydroid strand is usually single,
but in some taxa (e.g. Pseudocrossidium, Pl. 27, f. 8, 13, Timmie/la, Erythrophyllopsis), there may be two or more distinct strands
all dorsal to the guide cells. In very young leaves, the several individual, thin-walled cells of the strand may be seen in section (Pl.
9, f. 9; 69, f. 10-11), but in mature leaves, the strand is usually
represented by an angular, three-armed or sometimes star-shaped
(with concave sides) opening between the central guide cells and
the stereid band (Pl. 3, f. 5; 23, f. 9). Taxa that consistently lack a
--,
5
stem central strand apparently never have a leaf hydroid strand.
In taxa with both stem and leaf strands, there is no evidence of a
hydroid "leaf trace" connecting the leaf and the stem strands. In
some taxa with a stem central strand, the leaf strand may be
seen (in stem sections) penetrating the outer cortex for a short
distance (Pl. 77, f. 2).
The location of the hydroid strand in the center of the single
stereid band of Hennediella stanfordensis (Pl. 98, f. 11), Phaseapsis rubicunda (Pl. 100, f. 8-10), Acaulon robustum and other
taxa, and the central location of the single stereid band in the
costa of Scopelophila ligulata (Pl. 47, f. 6) and Streptopogon
spp., may both be explained as the guide cells being absent in
essentially double stereid-banded species, but correlation of
other characters indicates that this is probably not the case,
except possibly the latter in Scopelophila. Also, developmental
series studied by Saito (l975a, p. 388) indicate that the ventral
stereid band of Barbuleae is elaborated from the same tissue
that forms the parenchymatous layer between the guide cells
and ventral epidermis in Pottieae. Additionally, certain taxa
with variable development of the ventral stereid band (e.g. species of Streptocalypta, Pseudocrossidium and Tortula) show
what is apparently a thickening of the walls of pre-existing cells
ventral to the guide cells, giving the appearance of what seems
to be a second stereid band.
The shape of the dorsal stereid band is considered important
here in taxonomically distinguishing between advanced genera
of the Pottieae. It must be noted, generally, that the dorsal
stereid band in species with much reduced leaves and apparently narrowed costa will be reduced from a crescent-shaped
state to a rounded state (e.g. Gyroweisia). Taxa with rounded
dorsal stereid bands in the upper part of the leaf may show
crescent-shaped bands in the lower part of the leaf (e.g. Leptobarbula, Pl. 78, f. 9-10, and species of Tortula). This character
is, as are many in the Pottiaceae, of greatest value with plants of
comparatively large stature and, in any case, sections should be
taken at midleaf, most definitely above a sheathing leaf base.
Upper Laminal Cells
Cell wall thickness may have a significant effect on survival in
some species. Gers{!n ,(l987) found that thickened cell walls in
mosses prevent feeding by mites with short mouth parts.
For consistency, measurements of laminal cells in this paper
are of the width (perpendicular to the length of the leaf) with
indication of length to width (in that order) ratio following. Norris and Koponen (1989) indicated that laminal cells arranged in
rows was characteristic of the Pottiaceae, and could be used to
distinguish such genera as Streptopogon from, for instance,
those of the Splachnaceae.
Trigones. Cell wall thickenings where three walls are contiguous (the so-called "comers" of cells, Pl. 52, f. 4) are characteristic of species of several taxa, including Calyptopogon, Hymenostylium, Hymenostyliella, Leptodontium, and Reimersia, correlated with hygric habitat. Leptodontium, Hymenostylium and
Reimersia are apparently closely related (see phylogenetic
analysis).
Papillae and mamillae. The laminal cells of most taxa may be
described as papillose, but this is an overly general term for a
rather complex superficial ornamentation. Ignoring papillae
6
GENERA OF THE POTTIACEAE
sensu stricto for the moment, examination of transverse sections
of laminae demonstrates that species generally have characteristically superficially bulging or flat or slightly convex superficial
cell walls. This character has been little used taxonomically (the
exceptions are treatments of Tortula by Kramer 1980 and Mishler
1986a) frrstly because it is masked by papillae when they are
present and, secondly, because ,secondary thickening of the cell
walls can modify the degree of convexity. Some taxa (e.g. species
of Tortula) that are quite mamillose, with deep grooves around
each !aminal cell such that the cells meet only in a narrow band
around their periphery, are commonly not described as being
intensely bulging because of obscuring high papillae.
Taxa with ventrally mamillose and dorsally flat or weakly
convex laminal cells are distinctive in appearance (Pl. 1, f. 2; 52,
f. 6; 55, f. ll; 60, f. 5; 64, f. 8-9; 65, f. 8), and the lens-like (commonly non-papillose) nature of the ventral bulge may play some
part in focusing light during photosynthesis. Recently, R.
Donahue (Anon. 1990a) used fiber-optic light detectors, SEM
techniques, and a gas-exchange system to trace light paths within
leaves of Thermopsis montana Nutt. (Leguminosae). Shade plants
had cells with significantly more convex surfaces than sun plants,
capturing diffuse light at greater angles and focusing it on a particularly sensitive photosynthetic region of the cell. In the same
report (Anon. 1990a), G. Martin used finely detailed molds of leaf
surfaces to make replicas in agarose gel. He found that shade
leaves can intensify captured light up to 26 times better than sun
leaves, while in intense light potentially damaging radiation may
be focused on a protective, absorbing layer. As for papillae,
Simon (1987) has pointed out that the papillae of Tortula ruralis
increase its leaf surface area by a factor of 30 to 40. Proctor
(1979a) discussed papillae as efficient capillary systems in the
Pottiaceae.
How these observations might apply to the majority of species
of Pottiaceae, which are well known to survive in harsh environments, would be worthwhile to investigate. The character of ventrally convex an<J dorsally flattened upper !aminal cells is often
recognized as contributing to distinctiveness at the generic level
(e.g. Gertrudiella, Luisierella, and other genera and species).
Weissia condensa, however, exhibits variation in degree of
ventral mamillosity, this additionally obscured by its papillae.
Weissia breutelii, W. veviridis and other species, likewise, have
ventrally mamillose leaf cells, yet are not otherwise distinguished
as a group from other members of the genus. Hyophila is a genus
with ventrally mamillose and dorsally flat cells characteristic of
some species, but not others. There is a strong possibility that
eperistomate taxa of Trichostomum are masquerading as Hyophila
species, and, especially because of the large size of the genus
Trichostomum and its evident heterogeneity, Hyophila will probably be significantly reduced in size by some future monographer.
The papillae are often taxonomically important features at the
infrageneric level, but, like cell wall convexity, their appearance
may be modified by secondary thickening of the cell walls. Hollow papillae, e.g. those characteristic of species of Leptodontium
sect. Verecunda, may be "filled in" with secondary wall thickening in some specimens. Likewise, papillae that are characteristically bifid may, in some thick-walled specimens of the same species, appear to be fused into massive "multiplex" papillae (Pl. 38,
f. 13).
Hagen (1929, p. 14) described three distinctive types of
laminal papillae in the Pottiaceae: (1) large, rounded or platelike,
scattered over the lumens, (2) cylindrical and centered over the
lumens or otherwise occurring in groups, and (3) ring or
horseshoe-shaped papillae. In the literature, many species are
commonly described as having "cee-shaped" papillae. In most
cases, these are actually hollow, simple papillae (like blisters).
Because they are arranged on a convex cell wall surface, they
are viewed from above at an angle, and are seen in optical section because of the shallow depth of field at high magnification.
The optical section of the walls of these blisters takes the form
of the letter "c" or "o". Andrews (1945) correctly called "ringshaped or horseshoe-shaped" papillae an "optical illusion," urging more careful focusing on the part of the microscopist.
Simple papillae in the Pottiaceae exclusive of Leptodontioideae occur usually only singly or in mixed ones and twos over
each upper cell lumen (Pl. 10, f. 8); plants morphologically similar to Pottiaceae and with many simple papillae over each
lumen may be looked for in the Orthotrichaceae or Rhachitheciaceae. Bifid papillae (Pl. 22, f. 8; 107, f. 17) are common in the
Pottiaceae, occurring usually two or more over each upper
!aminal cell lumen. There may be a high saddle of tissue
between the two salients, and in this case the pairs form bifid
papillae that are genuinely cee-shaped in both morphology and
gross appearance, i.e. the cee shape is not an optical section of a
thin hemisphere. Some species, e.g. Leptodontium flexifolium,
may have optically cee-shaped papillae in thin-walled plants
and bifid papillae in thick-walled plants, so there may be a
developmental relationship between the two types. Scanning
electron micrographs of the papillae of the Pottiaceae have been
published by Mishler (1985b, 1986a, 1987a), Proctor (1979a),
Robinson (1974) and Zander (1972), among others.
Lamina/ thickness. The laminae of some taxa are typically
multistratose (Pl. 1, f.2; 3, f. 5), yet this character is variable at
the specific level in others. Some genera may have some species
with bistratose laminae (Pl. 108, f. 15) and some species with
unistratose laminae, but most genera have unistratose laminae.
Considerable variation is evident in the genus Didymodon
(Zander 1982a), especially the species related to D. rigidulus
and D. vinealis. Sayre (1952) briefly described similar variation
in Grimmia (Grimmiaceae). When laminal cells are bistratose in
patches medially, each of the cells is usually only half as thick
through (measured from ventral surface of the leaf to the dorsal)
as the cells of the unistratose portion, giving the appearance of a
single lamina! cell with an interior periclinal wall (Pl. 4, f. 8; 13,
f. 15). When leaf margins are bistratose, however, the cells are
generally each as thick through as the medial cells, inflating the
border whether the marginal cell walls are thickened (Pl. 17, f.
8; 33, f. 8) or similar to the medial cells in thickness (Pl. 8, f. 8;
9, f. 9; 23 , f. 9).
Basal Laminal Cells
The basal !aminal cells are sometimes called hyalocysts (cf
Edwards 1980a) when much enlarged and lacking chlorophyll.
They are usually also epapillose or weakly papillose and sharply
differentiated from the smaller, green, papillose upper cells.
Hyalocysts are characteristic of Calymperaceae, but are also
found occasionally in the Pottiaceae in various species of such
genera as Leptodontium, Bryoerythrophyllum and Tortula. In
most species of Pottiaceae, however, the basal laminal cells
more or less gradually intergrade in size and degree of
INTRODUCTION
papillosity with the upper cells. The basal cells may be wide or
little wider than the upper cells, and are generally smooth and
elongated. Well-differentiated basal cells are commonly bordered
along the leaf margins by narrow, rectangular cells that may serve
to inhibit breakage of the leaf, the basal cells apparently being
weak structurally. Occasionally a distinct area of unusually thickened rectangular cells is discernible in the leaf just distal to the
area of hyaline basal cells, as in species of Trichostomum subg.
Oxystegus.
Edwards (1980a) noted that resorption pores are found in the
hyalocysts of all three major traditional groups of the Pottiales, in
the leucobryaceous leaf, and in Sphagnum. He reviewed past references to such pores by other authors, and was surprised that
they have not been mentioned more commonly. He found, in
Calymperaceae of west tropical Africa, variation in number per
cell, position and shape, such characters being of taxonomic value
at the family and infrageneric levels. Zander and Cleef (1982)
described such pores in Kingiobryum Robins., now recognized in
the Dicranaceae but which may belong to the Pottiaceae.
Propagula
Propagula in the Pottiaceae are generally borne on stalks arising
from the axils of cauline leaves (e.g. some species of Barbula (Pl.
43, f. 20; 44, f. 10), Didymodon, and Hyophila), but species with
propagula borne on long rhizoids buried in the soil are also common (e.g. species of Barbu/a, Chenia and Tortula). Less frequently, propagula may be born on the tips or margins of leaves
(e.g. Streptopogon spp., Pl. 41, f. 12-14; 42, f. 17), on the ventral
surface of the costa (e.g. Didymodon spp., Syntrichia rigescens
(Pl. 110, f. 19), Trichostomum tenuirostre), and on a naked branch
(e.g. Leptodontium stoloniferum, Pl. 37, f. 7). My report of propagula in a cup at the apex of a leaf (Leptodontium stellaticuspis,
Zander 1972) is incorrect; the cup apparently produces only rhizoid initials. Rhizoids are also commonly produced at unmodified
leaf tips of Leptodontium viticulosoides and Didymodon
occidentalis. Long-elliptical propagula of species of Streptopogon
have longitudinal internal walls (Pl. 41, f. 16); similarly shaped
propagula of species of Calymperaceae lack longitudinal internal
walls.
Unicellular propagula are rather rare in the Pottiaceae, to date
having been found in Bryoerythrophyllum inaequa/ifolium (Pl. 24,
f. 8; Zander 1968), Didymodon revolutus (Zander 198lc), and
Didymodon perobtusus (Zander 1978e). The three species are
curiously similar in their small stature and broadly obtuse, elliptical to ovate leaves, among other characters, and may be products
of convergent evolution. The fine nature of the diaspores is a possible adaptation for non-sexual long-distance dispersal or local
saturation of a large and homogeneous habitat.
Whitehouse ( 1980) discussed both the literature and provided
his own observations on gemmae produced directly on the
protonema of many species of Pottiaceae. To date, such protonemal propagula are known for species of the pottiaceous genera
Barbula, Desmatodon, Didymodon, Eucladium, Gyroweisia and
Tortu/a, and may well be a common occurrence in hygric habitats.
Sexuality
Terminology for sexuality in the mosses is discussed at length by
Lewis (1961), Zander (1984) and Wyatt (1985), among many others. Wyatt (1985) proposed a terminology for bryophytes reflect-
7
ing the great variation in sexual condition in this group. Unfortunately, Wyatt indicated that, although distinguishing between
haploid and diploid sexuality was important, this could ascertained by "context." This is surely insufficient,. being a source
of confusion when discussing sexuality in general. Allen and
Magill (1987) have published a good response to Wyatt's article, emphasizing the need for a separate terminology for bryophytes. The Wyatt system is recommended here as it nicely parallels the terminology used for angiosperms but with the use of
"-oicous" endings (to indicate gametophyte sexuality) (see
Zander 1984 for a review of the history of "-oecious" and
"-oicous" endings in bryophyte terminology).
Perichaetia
The perichaetialleaves of the Pottiaceae may be similar to those
of the remainder of the plant, or differentiated, and sheathing
the inner perichaetium (and ultimately the basal portion of the
seta) to various degrees. The basal cells of differentiated perichaetialleaves of Pottiaceae taxa are generally elongate and rectangular with a small percentage of cells having tapering, blunt
ends. They are seldom prosenchymatous (i.e., with tapering,
pointed proximal and distal ends). They may have thickened,
porose walls (e.g. Trachyodontium) or thin walls (e.g. Bryoerythrophyllum spp.).
Perigonia
Perigonia (Pl. 1, f. 8; 15, f. 18; 16, f. 9, 16; 18, f. 9; 38, f. 9) are
generally bud-shaped in the Pottiaceae, occasionally flattened.
Some species have swollen, nearly spherical perigonia with
tightly appressed bracts, while some others have perigonia that
are small and loosely foliate, thus often difficult to find on gross
examination. The perigonia occur terminally or laterally on
entirely perigoniate plants, or laterally in the axils of the leaves
of sporophyte-bearing, monoicous plants, in which case they are
sometimes flattened.
Theca
The thickness of exothecial cell walls in the descriptions of the
genera presented here refers to that of the anticlinal walls. The
superficial walls may be, and usually are, much thicker than the
anticlinal walls (Pl. 59, f. 11; 79, f. 10). The stomates (Pl. 93, f.
8) are usually rather transparent compared to the other exothecial cells, but this is obvious only in split-open capsules.
Stomates in the Pottiaceae are confined to the base of the capsule above or on the short neck, and their numbers are generally
correlated with capsule size. They are seldom entirely absent,
and variation among the genera is discussed by Paton (1957).
The exothecial walls of Byroceuthospora, Eucladium, Trachycarpidium (Pl. 80, f. 9), Uleobryum (Pl. 89, f. 9) and species of
Weissia are bulging, to a greater or Jesser degree, while capsules
of Gangu/eea (Pl. 65, f. 10-11) and species of Weisiopsis (Pl.
67, f. 20) are distinctly plicate, or, in the case of Tetrapterum
(Pl. 79, f. 10), are weakly winged.
Peristome
The peristomes of most taxa are either straight or nearly so, or
are twisted to various degrees counterclockwise (as viewed
from above with direction of twist coming upward); peristomes
are seldom twisted clockwise. The counterclockwise direction
of torsion is known as sinistrorse, and may also be described as
8
GENERA OF THE POTTIACEAE
'•
the appearance of the threads of a standard, right-handed screw
(right-handed because it is screwed in with an over-the-top right
twist, not because the threads twist to the right), Of all genera
studied, only Timmie/la has, in some species, a peristome distinctly twisted clockwise; the peristome of Leptodontiella is either
straight or very weakly twisted clockwise, Most genera have
spiculose peristome teeth, while a few (e.g. Leptodontium) are
characterized by smooth or striate peristome teeth'=' A basal membrane (Pl. 84, f. 12-13; 97, f. 16; 106, f. 9), may be differentiated
in various genera.
Operculum
The operculum is usually rostrate and straight or weakly curved
or inclined. When immature, rostrate opercula are often merely
long-conic, e,g. as in Trichostomum brachydontium.
Calyptra
The calyptrae of Pottiaceae species are usually cucullate (Pl. 51,
f. 11), but are sometimes mitrate (Pl. 42, f. 8) or conic-mitrate
(Pl. 17, f. 17; 41, f. 9), Species of Microbryum, Hypodontium
and Streptopogon may have papillose calyptrae.
Spores
A list of published electron micrographs of spores of Pottiaceae
was given by Ireland (1987). In the present treatment the main
character used taxonomically is relative size of the spores.
Anisospory in the Pottiaceae is discussed in the treatment of
Leptodontium.
TECHNIQUE
Study of the Pottiaceae entails techniques common to the study of most mosses with a few modifications that address the need to
examine features of internal anatomy, use of color reactions to KOH solution, or to make permanent microscopic mounts of delicate
tissue. Zander (l979g, 1980a, 1983b) and others (e.g. Lightowlers 1981; Long 1982b; Frahm 1981, 1990a) have detailed aspects of
this, but a review is appropriate here.
Packets are folded from 16-pound substance 100 percent rag or
buffered paper to 10 em in height and 14 em in width using an
easily constructed device consisting of a square board to which is
affixed a low ledge below with a metal template the bottom of
which is screwed to the ledge, held parallel and about 3 mm
above the board. The template is slightly less than 10 by 14 em in
dimension. Packets can be folded quickly and accurately about
this flange, Subpackets for fragments are made from thin rag or
buffered paper (we use a rag tracing paper) and may be cut to various sizes and folded into square or triangular packets. Glue for
labels is made from polyvinyl alcohol dissolved in water and used
as refill for commercially available "roll-on" glues. Commercially
available polyvinyl acetate glue is also adequate and perhaps less
liable to degradation through crosslinking. (A planchet of
polyvinyl acetate glue was dried in 1970 and to date retains its
original elasticity after about 20 years exposed to fluorescent
lighting. Evidently any plasticizer included in the formulation is
not volatile or degraded in this time frame.) Water-based glues
are, of course, subject to weakening in very humid environments.
In annotating specimens, care must be taken to use inks that
do not fade with time. This is also true with making slide labels,
which may fade surprisingly quickly when exposed to sunlight.
India ink is, of course, ideal. Attaching annotation labels to packets may be done with glue or with plastic or plastic-coated paper
clips (to avoid rust marks). It is helpful to keep some insectmounting pins available for attaching annotation labels to specimens borrowed from herbaria that require such pinning.
Certain standard and some modified microdissecting tools are
used in the study and processing of specimens. Fine watchmaker's forceps may be further sharpened with a file. Dissecting
probes are re-constructed with sewing needles of medium size and
little flexibility inserted into a standard wooden or composition
probe handle to replace the original comparatively coarse and
bendable needle. Single-edge razor blades for sectioning should
be discarded after five to ten uses since they dull fast. One holds a
leaf or stem crosswise with a stiff dissecting needle, then slices
the material with a razor blade held longitudinally against the
far side of the needle, meanwhile rolling the needle slowly
towards oneself to gradually expose uncut portions of the material. Practice (and a relatively fresh blade) makes this technique
quite effective, even for very small leaves. Remember to scrape
off sections (especially stem sections) adhering to the razor
blade with a dissecting needle after cutting. The usual pair of
compound and dissecting microscopes are needed, but using an
additional illuminator with the dissecting microscope for fme
dissections rather than just a single lamp will prove surprisingly
advantageous for observation of fine features.
Round cover slips of 18-mm size and the thickest weight
(number 2) are used for routine examinations in water or KOH
solution because of ease of handling and cleaning for reuse.
Square 18-mm cover slips of medium thickness (number 1) are
used for making permanent mounts because they are less expensive and can be luted (i.e,, sealed to prevent evaporation) more
easily, To avoid annoying spills, a box of cover slips may be
glued to the microscope base or to the bottom of a low, flat tray.
Stains are not standardly used, although they might be
worthwhile in examining the pores in the basal lamina! cells of
some taxa, notably species of the genus Tortu/a, Color reactions
to alkali provide, however, important data, and a two percent
potassium hydroxide solution is kept at hand in a plastic
squeeze bottle. Potassium hydroxide solution helps hydrate dry
plants (cf G. Smith 1971, p, 2), Hagen (1929, p. 14) suggested
the use of a 10 percent KOH solution soak for the leaves of
Pottiaceae species to enlarge papillae and make them more easily visible; this effect has not been evident to me, at least at the
two percent concentration recommended. Glass containers
should not be used because KOH reacts with the walls to form a
precipitate. Dilute hydrochloric acid is a valuable reagent to
help determine, from observation of gas bubbles, whether or not
a collection was made from a calcareous habitat. It will also
help clean plants of limy incrustations.
Pohlstoffe (Wagner 1981; Christy 1987) may be used to
\
INTRODUCTION
hydrate specimens quickly for examination. The formula for the
stock solution is one part di-octyl sodium sulfosuccinate, 24 parts
methanol and 75 parts water. A few drops of the stock solution
are added to a dropping bottle of distilled water to make a fme
wetting agent. Mounts on microscope slides for examination at
high magnification should, however, always be made in one or
two drops of two percent potassium hydroxide solution. This
brings out characteristic color reactions in leaves and other plant
parts. This also helps remove the operculum (Lauridsen 1972) to
reveal the peristome. Heating the slide with a butane cigarette
lighter (if the flame does not touch the glass slide, there is no
soot) aids in loosening the operculum, and placing a cover slip on
the preparation before heating lessens evaporation. If the peristome persists in being broken off at the base when removal of the
peristome is attempted, allow an intact capsule to soak in a mixture of KOH and Pohlstoffe 15 to 20 minutes or longer. Since
Pohlstoffe will eventually precipitate from basic solutions, one or
two drops of a concentrated (4 gin 20 cc water) stock solution of
sodium N-lauroylsarcosine (trade name "Gardol") may be added
to the bottle of KOH solution as an effective surfactant, although
small amounts of any commercial detergent may substitute.
Taking time to make a good microscopic preparation will save
time later during identification. My standard method is to place
one or two drops of two-percent KOH solution on a microscope
slide, and add a stem of the plant to the slide. The material is
cleaned of debris (at this time one can search for rhizoid-borne
propagula) and the !aminal color reaction to the KOH solution is
noted. If the material is to be transferred to an acidic mountant
like lactophenol gel (see below), a drop or two of dilute hydrochloric acid solution is now added. The best preparation consists
of one slide with free leaves that have been removed from the
stem, especially near the apex to reveal the axillary hairs, and sections of the stem and leaf, and a second slide with the capsule
showing the peristome, spores and perichaetial leaves. Taking
care to flip some of the leaves so that the ventral side is uppermost will save time later. When making permanent slides, two
cover slips may be mounted on each slide with a slide label on the
left for consistency.
Permanent slides may be made with Hoyer's solution (Anderson 1954) or polyvinyl lactophenol (Frahm 1990a), these being
very convenient mountants that need not be luted, especially if
sufficient glycerine is used in formulating Hoyer's solution to
counteract dry air in the storage area.
Many species of Pottiaceae, however, have large, delicate
!aminal cells that collapse in Hoyer's solution or polyvinyl
lactophenol. Lactophenol gel (Zander 1983b), which does not collapse the cells of most species; may be used instead for sensitive
species or as a standard mountant for all species. It has a high
index of refraction. The formula for lactophenol gel is:
30 cc lactic acid(= 2-hydroxypropanoic acid)
15 g phenol, crystal(= carbolic acid)
15 cc distilled water
6 g methyl cellulose, powder(= cellulose methyl ether, of
viscosity 25 cP in 2% solution or lowest viscosity
available)
35 cc ethylene glycol(= 1,2-ethanediol).
Mix the phenol with the lactic acid, dissolving it with gentle heat.
Add the water and stir. Heat to just boiling (use a fume hood).
9
Add the methyl cellulose powder and stir vigorously into the
hot solution to dissolve (reheat if necessary). Add the ethylene
glycol last. Pour into a glass cylinder and let stand to allow bubbles and undissolved material to rise. After a day or two,
remove any floating particles and pour the clear liquid into a
storage bottle. A small bottle with an applicator wand built into
the lid or a plastic squeeze bottle with a fairly wide aperture (4
mm) is used to place a drop or two on a slide. Specimens
incrusted with carbonate deposits must first be soaked in a drop
or two of dilute hydrochloric acid to prevent bubble formation
in the lactophenol gel.
Specimens soaked in KOH should also be neutralized with a
drop or two of dilute HCI before mounting in lactophenol gel. A
precipitate may appear when overly moist plants are placed in
lactophenol gel but this is redissolved on stirrjng. Plants may be
manipulated, and leaves and stems sectioned, while in the gel.
The slides prepared as noted above, after placement of the
cover slip, are adequate as semi-permanent mounts for one or
two months. Permanent mounts may be made by sealing the
cover slip to the slide. So as to further solidify the gel and help
prevent migration of the plant material, water portion of the
mounting fluid may be allowed to evaporate somewhat by leaving the freshly mounted slide exposed in a dry place, such as a
fume hood, overnight, but sealing the slide immediately upon
preparation is generally best. To seal the slide, clean cover slips
and slides must be used to insure a complete bond. Clear fmgernail polish, although commonly used as a lutant, is generally
inadequate since pyroxylin (nitrocellulose) does not adhere well
to glass.
A good lutant is poly (ethyl methacrylate) with butyl benzyl
phthalate as plasticizer (as Krystalon™, Harleco, Gibbstown,
NJ 08027 USA), an artificial balsam with good long-term
adherence to glass. Another, more easily obtainable sealant with
excellent adhesion to glass, and good flexibility when dry (but
somewhat reactive, so do not store slides in strong light) is the
commercial liquid "polyurethane" gloss finish for wooden
floors, of various formulations but based mainly on
diisocyanates (such as tolylene diisocyanate). Isocyanates
adhere to glass extremely well apparently because they react
with an always present thin film of water strongly adsorbed to
glass (Skeist 1962). Keep the polyurethane container well
sealed and, if necessary, add a few drops of artist's drying retardant to the commercial product to slow the gelling during polymerization caused by oxidation of linseed oil, a common additive. Any lutant will stay liquid longer and be more easy to use
if kept in a "balsam bottle" or, better, in a small, dispo~able
applicator bottle. Keep the bottle more than half full to help
exclude air. Also acceptable as glass sealants include various
commercially available silicone rubber glues or caulks, and
"hot-melt glue" applied with a glue gun, but these are difficult
to work with and leave a ridge that may obstruct the high-dry
objective lens. Apply lutants liberally to ensure a good seal;
most seals can be scraped off in patches if a morphological
detail is obscured.
Disposable serviettes or wipers are important to have on
hand to clean tools and slides of reagents. A small fan is valuable at times to disperse any acrid fumes.
Sexing specimens is often required for identification, but
this can be as difficult as sectioning. Poor lighting makes this
more difficult, and one should not be satisfied with only a single
10
GENERA OF THE POTTIACEAE
standard illuminator for the dissecting microscope. Dry plants
may show bulbiform perigonia better than moist ones, but plants
should be moistened before removal or manipulation of leaves.
Sexuality cannot be determined accurately without location of
antheridia (in perigonia, in antheridiate buds or naked on the
stem, and preferably as observed on several plants in the collection to weigh degree of variation). Perichaetiate collections
entirely lacking antheridia are often assumed, with some degree
of confidence, to be dioicous, but note Steere's (1940) discussion
of dichogamy in Syntrichia princeps (as Tortula princeps).
Perigoniate plants are best looked for near the periphery of a
clump of sporangiate plants. Most genera show a distinct apical
thickening of the plant apex in perigoniate plants, but this is not
always the case. Autoicous buds can be searched for by carefully
stripping leaves of archegoniate plants. Paroicy and synoicy is
determined by attentive dissection of perichaetia with fine forceps
under high magnification under the dissecting microscope with
bright illumination. Antheridiate buds or short perigoniate plants
located on soil at the very base of archegoniate plants are a sign of
possible rhizautoicy, but this cannot be defmitely determined
without dissection of the rhizoid system or cultivation of single
spores.
Measurements are made with a transparent plastic metric ruler
or a more accurate "Minitool" 5-mm ruler marked in 0.1 mm
increments. An ocular micrometer for microscopic use is of
course essential.
Specialists in the taxonomy of particular groups are often
asked by other bryologists to render identifications of unusual
specimens or other puzzling collections. The accepted way to
have this done is to send a "duplicate for identification," which
is a labeled portion of a collection that the specialist can retain
for his/her herbarium. A name for each specimen sent is then
returned to the sender by the specialist. If duplicates cannot be
made, each specimen to be returned should be clearly marked
"unicate." The cover letter should point out which specimens
may be retained by the specialist and which returned. Generally,
a quick response may be had if only one to three specimens are
sent at a time; a package of several specimens may be placed
aside by a specialist pending availability of a larger block of
time for work on them. Identifications of large sets may not be
completed for many months, and, for these, preliminary inquiries of the specialist should be made. Botanists preparing grant
proposals that necessarily involve more than a little identification work by others should consider inclusion of support
through subcontracting.
PRACTICAL IDENTIFICATION
Recommended identification guides for Pottiaceae include Brotherus' (1924-25) world treatment, and bryological manuals for particular floristic regions (see Lane 1978b and Schofield 1985, p. 412 ff). Two family monographs stand out: the treatment for China
by Chen (1941) and that for Japan by Saito (1975a).
Among generic monographs, revisions or keys, sometimes only
regional, that are valuable for identification are those of Bell
(1974) for Willia; Delgadillo (1975a) for Aloina, Aloinella and
Crossidium; Crundwell and Nyholm (1972a, 1974) for Weissia
and (1962) Tarte/la; Kramer (1978, 1980, 1988) for Tortula sect.
Rurales (here treated as (Syntrichia); Lightowlers (1986a,b,c) for
Tortula s. lat.; Magill (1977a) for Globulinella; Robinson (1970,
1971a) for Trichostomopsis (here treated as Didymodon sect.
Asteriscium) and Hymenostyliella; Saito (1973a) for Pottioideae;
Salmon (1903) for Streptopogon; Sloover (1977) for Gymnostomiella; Stoneburner (1985) for Weissia; Warnstorf (1916) for
Pottia (here split between various genera of Pottioideae); Zander
for Pleuroweisieae (the species here placed in other sections,
1977c), Barbula, Bryoerythrophyllum and Pseudocrossidium
(1979f, 1981a), Didymodon (1981c), Leptodontium (1972), Neohyophila (= Plaubelia, 1983c), Scopelophila (1967), Streptocalypta (1983a), and Tuerckheimia (1978f).
Floristic treatments with especially interesting or useful treatments of the Pottiaceae or portions thereof, in addition to being
thorough and well illustrated, are those of Agnew &
Vondia~ek (1975) for Iraq, Catcheside (1980) for South Australia, Crum and Anderson (1981) for eastern North America, Dixon
(1913-1929) and Sainsbury (1955) for New Zealand, Noguchi
(1988) for Japan, Eddy (1991) for Malesia (Malaysia and the East
Indies), Flowers (1973a) for the North American West, Sharp et
al. (1993) for Mexico, Gangulee (1969-80) for eastern India,
Magill (1981) for South Africa, Norris and Koponen (1989) for
Papua New Guinea, Nyholm (1989) for Finland and Scandinavia, Savicz-Ljubitzkaja and Smirnova (1970) for northern Asia,
A. Smith (1978a) for the British Isles, and Steere (1938a,b,
1939a) for North America. There are, of course, a number of
other fme treatments of smaller areas, or which examine only
relatively common species, or which are not or are little illustrated, but the above are those estimated to prove most valuable
for practical identification.
Many taxa of moss families other than the Pottiaceae
approach the Pottiaceae in gametophytic features and may be
confounded with them when sterile. It will be to the advantage
of bryological taxonomists, both beginners and adepts, to keep
in mind the following list of Pottiaceae "look alikes": Andreaeaceae: Andreaea; Andreaeobryaceae: Andreaeobryum;
Aulacomniaceae: Aulacomnium; Bartramiaceae: Breutelia;
Diphysciaceae: Diphyscium; Cinclidotaceae: Cinclidotus;
Dicranaceae: Aongstroemiopsis, Cnestrum, Dichodontium,
Dicranoweisia, Holomitrium, Kingiobryum, Oreoweisia,
Pseudohyophila, Rhabdoweisia, Wilsoniella; Ditrichaceae:
Astomiopsis, Ceratodon, Cheilothela, Rhamphidium; Encalyptaceae: Bryobartramia, Bryobrittonia; Grimmiaceae: Jaffueliobryum, Tridontium; Orthotrichaceae: Amphidium, Kleioweisiopsis, Leptodontiopsis, Uleastrum, Zygodon; Ptychomitriaceae:
Campylostylium, Ptychomitrium; Rhachitheciaceae: Rhachithecium, Tisserantiella; Serpotortellaceae: Serpotortella;
Splachnaceae: Brachymitrion; Splachnobryaceae: Splachnobryum; Timmiaceae: Timmia; Viridivelleraceae: Viridivellus.
INTRODUCTION
11
MOST RECENT SUPRAGENERIC CLASSIFICATION
The latest arrangement of genera in suprageneric categories for the Pottiaceae at the world level, which may serve for purposes of
comparison, is that of Walther (1983):
Trichostomoideae
Trichostomeae: Timmiella, Pseudotimmiella, Trichostomum,
Rhamphidium, Hymenostyliella, Weissia (including
Hymenostomum and Astomum), Chionoloma, Phasconica,
Aschisma, Trachycarpidium, Kleioweisiopsis.
Tortelleae: Tortella, Pleurochaete, Oxystegus, Pseudosymblepharis, Stephanodictyon, Serpotortella, Barnesia, Streptocalyptra.
Barbuleae: Barbula (including Didymodon and Trichostomopsis), Bryoerythrophyllum, Streblotrichum, Semibarbula,
Husnotiella, Pseudocrossidium, Erythrophyllopsis, Morinia [= Mironia], Gertrudiella, Sebillea, Prionidium, Bellibarbula, Geheebia, Hydrogonium, Leptobarbula, Tetrapterum.
Hyophileae: Hyophila, Gymnostomiella, Teniolophora, Uleobryum.
Eucladieae: Eucladium, Gyroweisia, Gymnostomum, Hymenostomum, Tridontium, Reimersia.
Pleuroweisieae: Anoectangium, Molendoa, Pleuroweisia.
Pottioideae
Pottieae: Tortula (including Syntrichia), Desmatodon, Stegonia, Neohyophila (= Plaubelia), Crumia, Globulinella,
Uleopsis, Willia, Sarconeurum, Phascopsis, Aloina, Crossidium, Pterygoneurum, Aloinella, Ulea (= Uleopsis),
Streptopogon, Calyptopogon, Tisserantiella, Pottia, Acaulon, Phascum, Byroceuthospora, Hyophilopsis.
Scopelophileae: Scopelophila, Weisiopsis.
Cinclidotoideae
Cinclidoteae: Dialytrichia, Cinclidotus, Pachyneuropsis.
Leptodontioideae
Leptodontieae: Leptodontium, Tuerckheimia, Triquetrella,
Rhexophyllum, Luisierella, Leptodontiella, Streptotrichum.
Walther's compiliation is clearly a synthesis of the major elements of Brotherus' (1924-25), Chen's (1941) and Saito's
(1975a) treatments, and several recent papers of various authors
in addition to his own ideas (see also discussion of Cladograms
17-19 under Phylogenetic Analysis). In the main, it serves well as
a summary of modem thought for comparison with the taxonomic concepts and nomenclature recognized in the present
study. Although there is little basis for comparison since
Walther's arrangement is not based on rigorous analysis of all
characters, the present study supports certain of the groupings
above (see Table of Contents for an overview) but proposes
many new relationships.
Study with electron (TEM) and light microscopy of the
spore wall ornamentation of several genera of European Pottiaceae led Rejment-Grochowska (1978) to divide the Pottiaceae
into informal "sections" grouping the following taxa: Cinclidotus; Anoectangium; Eucladium, Gymnostomum, Gyroweisia,
Hymenostylium, Hymenostomum, Weissia, Trichostomum, Tortel/a; Barbu/a, Bryoerythrophyllum; Pottia, Acaulon, Phascum,
Pterygoneurum, Desmatodon; Tortula; Aloina. Saito and
Hirohama's (1974a) similar study (but using a scanning rather
than a transmission electron microscope) of 19 species of 14
genera of Japanese Pottiaceae (one collection examined for each
species) found only one apparent difference at the suprageneric
level: the Trichostomoideae differed from the other taxa studied
by the presence of "multistalked verrucae" on most but not all
of its species studied. They pointed out that major differences
between spore ornamentation may exist within the same genus
(e.g. between Trichostomum crispulum and T. p/atyphyllum),
while spores of different genera may be quite alike.
A major new character (Zander 1980a, 1989) that greatly
affects the classification of the taxa recognized here is the upper
lamina! color reaction to two percent potassium hydroxide solution. The genera are for the most part well and consistently
characterized intragenerically by KOH !aminal color reaction,
and suprageneric classification (see below) is in part based on
this character. The species of Tortula, when split into KOH yellow (Tortula s. str.) and KOH red (largely Syntrichia and Hennediella) sets, are more similar to each other than to groups outside the Pottioideae, and share such apparently advanced characters as the broad leaf shape, lack of a ventral stereid band
even in well-developed plants, and rather large upper !aminal
cells.
EVOLUTIONARY RELATIONSHIPS
Taxonomically important character states are generally not morphologically complex in the Pottiaceae, e.g. leaf margin recurvature,
presence or absence of a stem central strand or a hyalodermis, red color reaction to KOH, etc., see also discussion of Rohrer (1985,
p. 230), for a similar problem with characters in the Hylocomiaceae. Character states are apparently easily re-evolved (or perhaps
de-suppressed). This contributes much homoplasy or taxonomic "noise." For many of these character states, functions, if any, are
not clear, but their appearance may be pleiotropic to complex physiological functions.
The occasional presence of a twisted peristome in genus Ditrichum (Ditrichaceae; cf discussion by Robinson 1968) is difficult
to interpret as indicative of either primitive or advanced status for this character in respect to the base of the Pottiaceae tree. There
is, apparently, evidence for reticulate evolution in at least some mosses (Wyatt et al. 1992). On the other hand, the Pottiaceae has
many morphological and anatomical characters, and an analysis based on these many characters ought to provide a testable evolutionary hypothesis or at least point to an acceptable classification.
The Pottiales
Fleischer (1920) defmed the Pottiales in the commonly accepted
modem sense as: small, mostly turf-forming, upright plants;
leaves radially arranged on the stem, always with a costa, seldom
much-elongate or setaceous, of one layer of cells; upper !aminal
cells parenchymatous; sporophyte acrocarpous, seldom
cladocarpous (i.e. borne on a short side branch); capsule mostly
upright and smooth, commonly cleistocarpous; peristome single
or absent, of 16 teeth, the dorsal layer thicker than the ventral,
usually papillose, sometimes with a basal membrane. Families
included were Calymperaceae, Syrrhopodontaceae, Encalyptaceae, and Pottiaceae. Robinson ( 1971 b) on the other hand placed
the genera of the Pottiales in a much larger Dicranales, a judgment the present study tends to support.
Shaw et al. (1987, 1989) have indicated that Buxbaumia
(Buxbaumiaceae) and Diphyscium have a haplolepideous peristome. The relationship of Bu.xbaumia to Diphyscium (Diphysciaceae) is clear in its capsule shape and peristome (cf Taylor 1962,
p. 205; Crum & Anderson 1981). Diphyscium is similar to families of the Pottiales (see also Shaw et al. 1987, p. 68) in its
gametophyte morphology, without unique characters (the
bistratose !aminal cells are staggered, overlapping each other as in
Timmiella). The basal membrane of the pleated peristome does
not have the tesselations of that of the high basal-membraned species of Pottieae, but the ornamentation is similarly branchingspiculose (note that Shaw et al. 1989 showed evidence that the
haplolepideous peristome is homologous to the cilia of Bryumtype peristomes, not the endostome segments). There seems to be
no problem with the inclusion of both the Buxbaumiaceae and the
Diphysciaceae (including Diphyscium, Theriotia and
Muscoflorschuetzia) in the Pottiales along with Calymperaceae
and Encalyptaceae (the latter includes Bryobartramia, but see
cogent remarks on the non-haplolepideous character of the peristome of Encalypta by Edwards 1979, p. 342). The leaves of Buxbaumia aphylla Hedw. are deep red in KOH. The three species of
Diphyscium seen in this study had laminae reddish orange in
KOH, although the cell walls of the laminae of young leaves were
often uncolored or merely vaguely pinkish. The cladograms of
Mishler (l986b) and Mishler and Churchill (1984) show the
Buxbaurniales as the closest sister group for the Bryales.
Another family worthy of classification near the Pottiales (but
see Vitt 1984) is the Timrniaceae, of which the single genus, Timmia, has an inner peristome strikingly like the high basalmembraned, fenestrate peristome typical of many Tortula (s. lat.)
species, except that the inner peristome of Timmia is more
coarsely granulose and does not show the 2:3 primary peristomial
layer to inner peristomiallayer ratio patte111 characteristic of the
Pottiaceae and the Haplolepideae in general as discussed by
Edwards (1979). The gametophyte has many features otherwise
characteristic of the Pottiaceae: ligulate leaves with ventrally
marnillose and dorsally flat cell walls, sheathing leaf base;
upper !aminal cells isodiametric and bulging ventrally but only
weakly convex dorsally; and the costal transverse section with
two stereid bands, hydroid strand, guide cells, and ventral and
dorsal epidermises well differentiated. In addition, the !aminal
KOH reaction is uncolored in young leaves (laminae clear green
in toto, the walls are uncolored) with reddish orange blotches in
patches, especial!y on the leaf base (T. megapolitana Hedw.),
but pink in mature leaves. Akiyama and Nishimura (1993, p.
191), on the other hand, felt that the Timmiaceae was only
remotely related to the Bryineae on the basis of branch development analysis.
Encalypta species examined (E. ciliata Hedw., E. mutica
Hagen, E. procera Bruch, E. rhaptocarpa Schwaegr., E.
spathulata C. Mtill., E. streptocarpa Hedw. and E. vulgaris
Hedw.) in the course of this study have KOH color reactions
characteristically bright yellow (becoming red-orange in older
leaves) in the upper lamina and deep red in the basal cells, and
species of Encalyptaceae have inner peristomes with high basal
membranes (Horton 1982). Edwards (1979) indicated, however,
that Encalypta, because it did not have the characteristic 2:3
pattern of peristomial development, was probably not a member
of the Haplolepideae, while Shaw et al. (1987, p. 68) indicated a
close phylogenetic relationship between the Encalyptaceae and
Buxbaumiaceae based on their own work and that of others. In
any case, a high basal membrane is probably not an advanced
character in the Pottiaceae, since four acrocarpous families with
papillose or ventrally mamillose, isodiametric upper !aminal
cells (Buxbaumiaceae, Diphysciaceae, Encalyptaceae and Timrniaceae) have high basal-membraned inner peristomes. Of
these families, all have either reddish orange or deep red
laminal KOH color reactions (in Encalypta restricted to the
basal cells), which indicates that a yellow KOH reaction is an
advanced character.
Edwards' (1979) indication that the Ptychomitriaceae (two
stereid bands, yellow in KOH) belong with the Haplolepideae is
agreeable on general morphological grounds: none of the major
gametophyte features would be out of place in the Pottiales,
while the sporophyte is also similar to thos'! of the Pottiales,
excepting only the large, plicate, rnitrate calyptra of Ptychomitrium.
Species of Calymperaceae examined reacted to KOH with a
EVOLUTIONARY RELATIONSHIPS
yellow color. Edwards (1980a) characterized the Calymperaceae
as having hyalocysts abruptly differentiated from the green upper
laminal cells and having resorption pores, teniolae (a strong,
mostly intramargina1leaf border) present, and propagula borne on
the lamina. Only the presence of resorption pores is constant,
according to Edwards. All the characters noted by Edwards
(1980a) are present in the Pottiaceae. 1n combination, however,
these characters are rather different than those of the Pottiaceae.
Reese (1987b) listed several advanced characters of the
Calymperaceae (Pottiales), including tropical environment,
corticolous habitat, well-defined cancellinae (basal "windows" of
hyaline, inflated !aminal cells) with superficial pores, reduced or
absent peristomes, broad and sheathing leaf bases, and propagula
usually present. The differences between Calymperaceae and
Pottiaceae are discussed at length by Reese ( l987b) and Reese
and Zander (1988). Hypodontium (q.v.) is here included in the
Pottiaceae although previously long accepted as Calymperaceae
(see also discussion of Calymperastrum); Reese (1987b) felt that
Hypodontium is intermediate between Calymperaceae and Pottiaceae.
Regarding evidence of gametophytic characters, certain families with somewhat similar gametophytes and haplolepideous
peristomes (Calymperaceae, Encalyptaceae, Dicranaceae, Ditrichaceae and Seligeriaceae) are shown by Kawai (1968) to have both
one and two stereid bands, and costal morphology with clear differentiation into dorsal and ventral epidermises and medial guide
cells. Other families gametophytically similar to the Pottiaceae
(Timmiaceae, Ptychomitriaceae) have two stereid bands. Other,
less closely related families (Mniaceae, Polytrichaceae) have one
or two stereid bands. The Orthotrichaceae, which often have the
general appearance of Pottiaceae, have mostly a single stereid or
substereid band and poor costal differentiation into cellular layers.
The Bartramiaceae, on the other hand, have a single band and
good cellular differentiation, as have certain genera of Grimmiaceae. Pleurocarpous families appear to have internally fairly
undifferentiated, thin costae. Given the often rather high degree of
costal differentiation of closely related families and of such relatively primitive families as Polytrichaceae (see cladogram of
Mishler 1986b), one might expect that the Pottiaceae and related
families were derived from ancestors in which two stereid bands
and possibly a high basal membrane, too, had already been
evolved. Additional evidence against convergence is that certain
pottiaceous taxa normally having a single stereid band (species of
Pseudocrossidium, Tortula s. lat., Streptocalypta) may develop a
few stereid cells between the ventral epidermis and the guide
cells, an exact positioning more indicative of partial
desuppression (cf Basile & Basile 1984) of a ventral stereid band
than of de novo elaboration of one. Thus, the line of costal evolution in Pottiaceae probably began with taxa with two stereid
bands and progressed through loss of the ventral band to those
with a single strong stereid band. The fact that some taxa that usually have two stereid bands may have only one band in occasional
collections of small stature is probably not apropos here, since
genera with only one band may be quite large in stature (e.g.
Crumia, Hennediella, Syntrichia and Tortula) in some species and
have their own reduction series involving sporophyte characters
and stem length but without much variation in costal morphology.
Reduced sporophytes are correlated with small gametophyte stature (but cf Willia).
Cladogram 1, discussed at length below, of both the genera of
13
the Pottiaceae and haplolepideous and diplolepideous genera
selected for having gametophytes similar to those of the Pottiaceae, with scored character states restricted to those found in
the Pottiaceae and using Polytrichum (Polytrichaceae, which
has many of the anatomical and morphological characters of the
Pottiaceae) as outgroup, shows most non-pottiaceous genera
derived from lines of the Pottiaceae. This may not show the
relationships of non-pottiaceous genera to each other, since nonpottiaceous characters are not used, but it at least indicates
likely relationships based on shared apomorphies of these genera with the Pottiaceae. The distance in steps of non-pottiaceous
genera from the root of the tree indicates the degree of weighting of particular pottiaceous or non-pottiaceous (and not used)
characters that would be necessary to bring these genera down
to the base of the tree; it also indicates that certain genera (e.g.
Ceratodon and Encalypta) high in the tree have a large number
of advanced characters that can only doubtfully be accommodated by weighting. It will be important to study details of the
peristomes of all taxa if sufficient additional characters are to befound to justify viewing the Pottiaceae as presently constituted
as monophyletic.
Intrafamilial relationships of the genera
It will be apparent from the discussions presented here that
there are probably far more genera worthy of recognition than
are presently accepted; however, much more work will be necessary to adequately identify and describe these additional genera. In the absence of a massive cladistic evaluation at the species level, there are two obvious places to look for unrecognized
genera; first, as transformation series within large genera, such
as Trichostomum (q.v.), and, second, as rather different species
brought together in small "wastebasket" genera, such as Gyroweisia. If clusters of closely related species were recognized as
genera or sections, there would be at least double the number of
supraspecific taxa presently accepted. On the other hand, segregation of such clusters would leave an overabundance of species
not assignable to any segregate and remaining in the original
genera by default.
"Good" characters may be variable. Many species in the
Barbuleae vary in production of a second stereid band (usually
in well-developed plants or larger leaves). Thus, it is the potential for development of a second stereid band that provides this
traditionally major character. A complication is the occasional
production of a second stereid band in some species otherwise
clearly belonging to the Pottioideae. One of the aims of this
study was to examine the possibility that separating genera at
the suprageneric level on the basis of presence or absence of a
second stereid band may mask relationships of closely related
species losing a second stereid band in a reduction series. Certainly small plants of Didymodon and Gymnostomum, among
other genera, have only one stereid band. It was found, however, that there were few (e.g. Calyptopogon and Streptopogon)
clear-cut overall morphological similarities between species traditionally distinguished by numbers of costal stereid bands that
might be inferred as being due to a close phylogenetic relationship marred only by lack of a second stereid band, i.e. the number of stereid bands is a valuable character.
A "separate evolution of gametophytic and sporophytic
characters" (cf discussion by Rohrer 1988) is often invoked to
explain incongruent character assemblages. Although sporo-
14
GENERA OF THE POTIIACEAE
phyte characters have much utility at the suprafamiliallevel in the
mosses, only a few such characters in the Pottiaceae were found
important at the generic level (cf treatments of Aschisma, Trachycarpidium and related genera) in the Pottiaceae. Many character
states once widespread among ancestors of present-day mosses
may now be suppressed or lost. In the Pottiaceae, few uncommon
characters, if any, are found in one genus and not also in another,
apparently distantly related, genus. Striking convergent evolution
is apparent in a large number of cases. This may even include the
one most obvious "unique" character in the family, the twisted
peristome of many of the genera. Although Mishler (l985b, p.
389) could identify no apomorphies for the Pottiaceae, the twisted
peristome may be considered a synapomorphy for the family if
genera of apparently reduced and simplified morphology, including shortening (and subsequent straightening) or loss of the peristome are considered derived, or if such genera are considered
relicts of larger lineages with members having twisted peristomes
now extinct. A twisted peristome, in any case, is present in widely
divergent pottiaceous genera and not in others, and it is present in
two species of Ditrichum (Ditrichaceae), namely D. tortipes
(Mitt.) Par. and D. ambiguum Best (cf Crum & Anderson 1981,
Grout 1927 and Robinson 1968). These two species of Ditrichaceae (discussed further under treatment of Barbula) have a distinctly twisted peristome much like that of the Pottiaceae, and the
laminae of these species have yellow KOH reactions. These two
species or the genus Ditrichum itself may actually belong near
Barbula sect. Hydrogonium. One might note, in this respect, that
Distichium (Ditrichaceae) has a pottiaceous peristome, albeit
mostly untwisted.
Apparent convergences may be genetically complex: one
might cite the production of a propaguliferous leaf apex in Leptodontium proliferum, Tortella fragilis, and Syntrichia angustifolia
with associated elongation of the upper marginal laminal cells,
and the fleshy capsules of Tridontium (here placed with the
Scoulerioideae of the Grimmiaceae) and Dialytrichia; but more
usually such convergences involve apparently simple traits, such
as basal margins of lamina serrulate in Eucladium verticillatum,
Molendoa hornschuchiana and occasional collections of Gymnostomum aeruginosum; the lack of a ventral stereid band in Calyptopogon, Pseudocrossidium, Streptopogon and Tortula; systylious
capsules in Hymenostylium and species of Hennediella (taxa previously recognized under Desmatodon).
Although certain families of mosses (e.g. Archidiaceae,
Ephemeraceae) are rightly characterized by relatively small
sporophytes with short setae and cleistocarpous capsules, the
assumption that these characteristics are conservative in the Pottiaceae has proven to be unacceptable to certain modem authors,
e.g. those dealing with the Weissia-Hymenostomum-Astomum
complex. In the Pottieae, Pterygoneurum is an example of a genus
with so distinctive a gametophytic character (the ventral costal
lamellae) that species with sporophytes with peristomes and spe"
cies without peristomes have long been acceptable within that
genus. Elsewhere in the Pottieae, species with eperistomate capsules have occasionally been thrown by previous authors into genera (e.g. Tortula) that are generally peristomate but with gametophytes that are fairly heterogeneous, and the complexity of these
groups of disparate species has discouraged further work by traditional methods. Such reduction is evident in other acrocarpous
families; for instance, Fife (1985), in his treatment of the genera
of the Funariaceae, stated that "The dominant evolutionary trend
in the Funariaceae has been the progressive simplification of the
sporophyte." He associated large stature, small spores and
"elaborate morphological structures for the regulation of spore
release" in patchy environments through wind dispersal, being
most suitable for opportunistic species, and at the other end of
the spectrum small stature, large spores and lack of peristomes
or even opercula is associated with growth in habitats widely
and continuously or at least periodically available, often characterized by disturbance or flooding. A parallel is evident in the
Pottiaceae, although the gametophytes of the Pottiaceae are not
nearly as uniform in morphology as those of the Funariaceae.
This last supports my apprehension of several independent
reduction series. On the other hand, Mishler and Churchill
(1987) emphasized that "postulating reduction series ... depends
on the use of outgroup comparison, not appeal to an a priori
evolutionary scenario ... ," a caveat addressed below in the
phylogenetic analysis.
Longton (1988) reviewed the literature on life history strategies in desert-dwelling bryophytes and noted that "most bryophytes are poikilohydric: they have little control over rates of
water uptake or loss, but in compensation the gametophytes of
many species can tolerate severe and prolonged cytoplasmic
desiccation." He found several trends, among them: " ... towards
reduced gametophyte life-span, early and prolific production of
spores and/or asexual propagula, and monecism in sexually
reproducing species: shifts from K- tor-selection and from tolerance to evasion of environmental stress" and " ... towards
reduced duration of habitat availability but with habitats tending
to recur predictably within a community. Parallel trends towards
increase in spore size and loss of feature[s] promoting spore dispersal in regularly sexually-reproducing species." The apparent
morphological reduction series seen in some genera of Pottiaceae (notably Hennediella, Tortula and Weissia) is possibly
associated with increased fitness in the environments of arid
lands.
Many nomenclatural novelties are presented here, most
being just transfers for long-unstudied exotic species necessary
to reflect modem generic constructs, but many others involve
recognition of probable evolutionary tracks often signalled by
unique character states or character state combinations.
A beginning at a new synthesis based on recognition of
apparent transformation series that may cut across traditional
generic limits is indicated in the discussion of Trichostomum,
Weissia and related genera, and the Phascum-PottiaDesmatodon-Tortula complex. An alternative view emphasizing
the taxonomic utility of peristome morphology in the Pottiaceae
is presented at length by Hagen (1929, p. 8-13), probably as a
reaction to Dixon's (1924) extensive generic lumping in the
family. Reduction of the sporophyte in Pottiac,eae (and other
acrocarpous families colonizing dry areas) may be assessed as
an evolutionary adaptation correlated with two enviJ;onmental
influences: (l) capsules may be exerted to their detriment above
the boundary layer of stagnant humid air for considerable periods of time (sporophytes generally take one year to mature, cf
discussion by Zander 1979d), and (2) fitness of mosses with
massive diaspores, such as large spores and cleistocarpous capsules, or with diaspores developed close to the substrate
("atelochory" in the terminology of van der Pijl 1972, and
"precinctiveness" of Carlquist 1966, 1974) may be improved in
patchy environments (cf discussion by Zander 1979f).
EVOLUTIONARY RELATIONSHIPS
Phylogenetic evaluations: discussion
Phylogenetic opinions of past authors were apparently based on
one or more of a combinations of "marker characters," being
unique or rare shared characters, of presumed great phylogenetic
weight, to distinguish genera and higher taxa. The mass of nonweighted characters were apparently assumed to be considerably
less important and prone to abundant convergence and parallelism. One must evaluate, however, the assumption that morphological convergence between taxa in a particular (especially stressful)
envirorunent may involve the evolution quite naturally of a combination of many characters the existence of which in clearly
unrelated taxa in the same envirorunent must otherwise be imagined as improbably fortuitous. (Thus the particular low weighting
of 22 reduction-related characters in the cladistic analysis below.)
Taxonomic evaluation of intrageneric relationships is here
mostly based on general phenetic similarity; when transformation
series are clear, however, the generic limits as given presumably
reflect probable intrageneric phylogenetic relationships, i.e.
grouping those taxa most readily derived from common ancestors
through evolution of traits that are unique or rare in the immediate
group. In the case of what appear to be extant little-different
descendants of ancestral species, the decision whether to keep
closely related species representing a branching series together in
one genus or assign them (perhaps arbitrarily) to their respective,
different phyletic lines is here usually made in favor of the latter.
Thus, Pseudocrossidium crinitum is quite closely related to the
genus Barbula (differing mainly in the shape of the section of its
dorsal stereid band and in the sheathing perichaetial leaves), yet
because it may be construed as a stem species for the Pseudocrossidium transformation series, it is placed with that genus.
It may also prove fruitful to examine the Barbuleae (as was
begun by Saito in his lucid treatment of the distinguishing characteristics of Barbula and Didymodon) and identify transformation
series that begin in genera with well developed peristomes and
end in those with reduced or absent peristomes. This may result in
some splitting of the well-peristomed genera and possible disappearance of some eperistomate genera, e.g. Gymnostomum and
Gyroweisia. A typical transformation series is described in a synopsis of Pseudocrossidium (Zander 1979f) in which a south to
north transformation series from the Andes to the Arctic showed
that P. revolutum and P. hornschuchianum belonged in the genus
even though lacking many of the more salient characters (but possessed of the distinctive costal section anatomy).
Unfortunately, in this light, revisions of single genera of the
Pottiaceae will commonly not reveal complete transformation
series. If such series are found to be worthwhile as contributing to
the clarification of generic limits, future revisionists must resign
themselves to careful examination of several genera at once to
distinguish the full range of potential series. At present, the genera of the Pottiaceae are in some cases neither "natural" nor
instructive of possible phylogenetic relationships, and, in fact,
many present generic definitions seem to cut across natural lines.
Exceptions are probably those genera with sporophytes that have
unusual (rare or autapomorphic) features or modifications (mainly
of the peristome) different than what may be interpreted as mere
reduction. These include Aschisma (exothecial cells in palisadelike rows and stomates absent), Leptodontium (and other taxa
with unvaryingly smooth teeth), Steptotrichum (elaboration of
internal lateral cell walls interior to the teeth), Leptodontiella and
Trachyodontium (teeth cleft into several rami), Uleobryum (and
15
other genera with pustulate exothecial cells), and Weisiopsis (16
spaced teeth).
In the absence of a species-based cladistic study, genera are
here characterized either by an observed decided lack of species
of transitional morphology between two groups (a phenetic gap)
or by some obvious division of evolutionary directions (nonrigorous evaluation of salient advanced characters). It is, of
course, most satisfactory when both obtain, such as is the case
with some of the smaller, isolated genera like Tetrapterum.
Transitional species between apparent major evolutionary
thrusts, however, are disconcertingly common. It is clear that
cladistic evaluations are capable of generating more rigorous
(based on defined methodologies) and more reasonable classifications than an "omnispection" technique grouping taxa with
general overall resemblance. This study attempts a classification
based on cladistically derived lineages of genera as these are
presently delimited.
An emphasis on using only presumed derived characters
(Hennig 1966) as a basis for phylogenetic analysis has been
extensively rationalized recently, and has been used in studies
of the mosses (e.g. Bremer, 1981; Cao & Vitt 1986; Churchill
1981, 1985; Frahm 1991b; Koponen 1968, 1973, 1982; Mishler
1985b,c, l986a,b; Mishler & Churchill 1984, 1985b; Rohrer
1985, 1988; Waters et al. 1992, imd others). On the other hand,
cladistics has become a new orthodoxy (see also Heywood 1983
and Robinson 1986a) that, in sweeping clean, has discounted
the value of past classifications; even taxonomy is eliminated
from one recent definition our field: " ... phylogenetics and
biogeography, together referred to as systematics" (Erwin
1991)! Judging from the good match of the three traditional Pottiaceae subfamilies (Trichostomoideae, Barbuloideae and Pottioideae) with the results of the present study, formal parsimony
is reflected to a significant extent in past attempts at the creation
of a simplest classification that takes into account the available
data with emphasis on shared characters or character state combinations, even if the effort does not involve a large data set,
base relationships only on derived characters, and use an exact
algorithm. It is expected that the generic assigrunents of infrageneric taxa will be supported to a considerable extent by future
species-level cladistic studies, except in genera pointed in the
taxonomic treatments as clearly heterogeneous (e.g. Barbula,
Didymodon, Hyophila, and Trichostomum).
Plausible phylogenetic relationships are satisfactorily inferable within identified transformation series, but the direction of
evolution is sometimes far from obvious. A series might start in
the middle, at either end, or in two or more places. Assuming,
however, that similar complexes of many apomorphic characters are more likely to be derived from shared ancestors than
evolved anew for each set of individuals supports an interpretation of most transformations as reduction series. This does not
imply that reduction is a process that occurs more easily than
elaboration, but that an explanation of a transformation series as
reduction requires the assumption of fewer parallel evolutionary
events than does elaboration.
There are two ways to view "splitting" and "lumping." The
first is to see these as two different ways of deviating from good
taxonomy. The second, and better approach, is to see these as
analytic and synthetic methods of dealing with information,
splitting thus being appropriate at exploratory stages and lumping at revisionary stages.
16
GENERA OF THE POTTIACEAE
Many of my publications attribute rather wide variation to
certain species and species complexes (e.g. Didymodon fa/lax
complex, Didymodon vinea/is complex, Hymenostylium
recurvirostrum complex, Molendoa sendtneriana s. lat.). For
example, see charts of variation in Hymenostylium (Zander 1977c,
Zander & Eckel1982), which demonstrate independently varying
character states and are surely evidence of considerable genetic
variation. It should not be surprising that such intraspecific
phenetically distinctive swarms exist and can be expected to
exemplify complex intraspecific genetics inasmuch as genetic
expression in mosses is expected to be essentially genotypic in the
gametophytic generation. Additionally, the considerable variation
within species or species complexes confounds easy identification. Often, characteristics that may not always be present, but are
nevertheless distinctive, may be helpful. For instance, peculiar to
the Didymodon vinea/is group is the usual absence of the quadrate
ventral costal cells at the extreme leaf apex, resulting in a short,
boat-shaped groove bottomed by epapillose elongate cells.
Hymenostylium species commonly have longitudinally elongate
mediallaminal cells, but this is not always the case.
Species are here seen as inferential populational units composed of individuals sufficiently similar to form a probable biologically predictable unit, and when this is not clear (such as
when species are known from single gatherings or when taxonomy is doubtful), a species is put forward as representing a presumed "basic taxonomic unit." Genera are more or less clearcut apparent phyletic series of species or groups of such
phyletic series, or simply groups of similar species when no
evolutionary series are evident. Unusual characters are viewed
here with some diffidence: the lens-like central thickening of
the exothecial cells of Byroceuthospora, U/eobryum and
Trachycarpidium is a clearly advanced feature, yet this is not a
complicated character and may well have been developed
through evolutionary convergence. If these three genera were
seen, however, as a single suprageneric tax.on, the quite different morphology of the gametophytes would require that they be
considered remnants of a once larger, complex group with taxa
of intermediate morphology now extinct. Since these are aridland genera with austral affinities, such a view is an acceptable
possibility and the cladistic analysis below supports it.
PHYLOGENETIC ANALYSIS
The object of the study was to obtain a most-parsimonious (minimized homoplasy) hypothesis of evolutionary relationships of the
genera of the Pottiaceae as presently conceived, and to use this as a basis for a suprageneric classification. This evaluation is complex and a precis, as overview, follows .
SUMMARY OF THE ANALYSIS
Timmia (Timmiaceae, diplolepideous) and Polytrichum
(Polytrichaceae, nematodontous) have gametophytes that are quite
similar to certain genera of Pottiaceae, but are considered distant
from the Pottiaceae and from each other because of their much
different peristomes, which they share with other genera not in the
Pottiaceae. Because these two genera are both distant and share
many characteristics with the Pottiaceae, they are necessarily
expected to appear at the base of any cladogram including both
them and the Pottiaceae. The Polytrichales is shown to be more
primitive than the Bryales in cladistic evaluations by Mishler
(1986b) and Mishler and Churchill (1984) although the
Buxbaumiales is the nearest sister group (but which is not used
here as an outgroup because of a lack of characters that is probably due to reduction).
Of the various haplolepideous genera in families evaluated as
potential outgroups (Cladograms 1-6), Ptychomitrium
(Ptychomitriaceae) was the outgroup that best kept Timmia and
Polytrichum low in the tree when ten non-pottiaceous genera were
added to the Pottiaceae data set. Analysis was then made with
only a single non-pottiaceous genus as outgroup. With
Ptychomitrium as outgroup (Cladogram 10), Timmie/la was found
to be the most primitive pottiaceous genus, and this genus was
also at the base of the tree with Polytrichum (Cladogram 7) and
Timmia (Cladogram 9) as outgroups. Timmie/la (Pottiaceae) was
then used (Cladograms 13-16) as functional outgroup to avoid the
possibility that Ptychomitrium was less distant from the Pottiaceae than Aloina (cf Cladograms 1, 4 and 8) or Gertrudiella, and
to avoid problems introduced by reduction in Ptychomitrium.
Cladograms 1-4 and 7-10 support the use of Timmie/la as
functional outgroup in Cladograms 13-16. They likewise show
that at least some genera of Pottiaceae retain characteristics of
the more primitive mosses Polytrichum and Timmia. Also, putative sister group genera (Bryobartramia, Ceratodon, Diphyscium, Ditrichum, Encalypta, Grimmia, Syrrhopodon) in the Pottiales, Dicranales, and Grimmiales are mostly too modified in
morphology to serve as outgroups in that too many characters
are lacking or fixed in states deemed here to be the result of
evolutionary reduction. These cladograms demonstrate that
many genera are found in similar or even identical lineages in
cladograms generated with different outgroups, implying evolutionary development of quite distinctive synapomorphies and
supporting more firmly the present hypothesis of relationship.
Cladograms 11 and 12 are from data sets restricted to terminal taxa of two major branches of Cladograms 9 (outgroup Timmia) and 10 (outgroup Ptychomitrium) that are identical
between cladograms. These consensus trees are based on a
known number of equally parsimonious trees and generally support details of branching patterns in the other cladograms, for
which only a portion of all trees could be kept in computer
memory and thus contribute to the consensus tree.
The cladogram that presents the hypothesis used here for
suprageneric classification is the single tree shown in Cladogram 16. Character state changes for this tree are detailed in
Cladogram 15. This single tree was chosen from more than
1250 equally most-parsimonious trees as summarized in the
strict consensus tree in Cladogram 14. The major subclades of
Cladogram 16 are identified as the subfamilies and tribes recognized in this treatment.
EVOLUTIONARY RELATIONSHIPS
INTRODUCTION:
Outgroup Selection
The argument for past emphasis on sporophyte characters in distinguishing genera of the Pottiaceae is (1) such characters work
well in distinguishing taxa in other groups, and (2) taxa sharing
similar sporophytes in the Pottiaceae also share similar gametophytes. The present study of the genera of the Pottiaceae has
shown, however, that although the general morphology of the
well developed capsule and its peristome is similar and typically
pottiaceous (with notable exceptions, e.g. Leptodontiella, Streptotrichum, Trachycarpidium and relatives), there is considerable
variation in expression of individual sporophyte features, i.e. in
degree rather than in kind. Also, detailed morphological analyses
demonstrate that this variation is often considerable among taxa
with gametophytes that share many morphological and anatomical
features not previously evaluated across the family (e.g. the genera Hennediella, Tortula and Weissia as emended here). Arguments for special weighting of sporophytic characters (cf Crosby
1974, Dixon 1932, Miller 1979) in the Musci generally are not
supported at the generic level in the Pottiaceae.
The rationale used here for selecting an outgroup with shared
presumed primitive characters is that haplolepideous sister groups
to the Pottiaceae (e.g. Calymperaceae, Ditrichaceae, Encalyptaceae, Grimmiaceae) are evolutionarily much reduced and simplified in both gametophytic and sporophytic morphology. This is
true when such groups are compared to the range of morphotypes
seen in the Pottiaceae. It is necessary to examine the other
haplolepideous and even diplolepideous and nematodontous taxa
for genera retaining characters of the ancestral morphology. The
"generalized" ancestor is not necessarily a gametophytically relatively characterless taxon (e.g. Grimmia), but may be characterrich.
Ptychomitrium (Ptychomitriaceae, haplolepideous) was found
to be the closest sister group to the Pottiaceae that was not so
morphologically reduced as to have important plesiomorphic
character states (also present in more distant ancestors Timmia
and Polytrichum as will be demonstrated below) much modified
or completely eliminated. This genus is considered the primary
outgroup, and was used to establish a functional outgroup among
the pottiaceous genera; cladograms were generated with additional alternate non-pottiaceous outgroups in an attempt to evaluate character state polarizations more globally.
Although its peristome is nematodontous, Polytrichum is considered here an acceptable outgroup because it shares many characters with the Pottiaceae that are not associated with apparent
reduction. Polytrichum is evidently a moss with very primitive
characters, sharing with ferns, for instance, a leaf trace connecting
the leaf hydroid strand and the stem central strand, and it has been
demonstrated cladistically to be of a lineage more primitive than
the Bryales (discussion of Mishler & Churchill 1984). Polytrichum is also distant from the Pottiaceae because of ( 1) the several
unique characters of its peristome, which presuppose a considerable lineage involved in their development, and (2) the number of
the species in its genus (and family), representing multiple evolutionary events. The similarity of the gametophytes of several genera of the Pottiaceae with Polytrichum indicates that the Pottiaceae, unlike other families of the Pottiales, retains members with
primitive morphologies. Characters of Polytrichum that are, on
the other hand, typical of reduced members of the Pottiaceae
include epapillose upper !aminal cells, lamellae present on the
17
ventral surface of the costa, and annulus vesiculose.
The heterolepideous genus Timmia (Timmiaceae, diplolepideous) is also a major source of information on plesiomorphic
features of the Pottiaceae. This genus of the monotypic family
Timrniaceae is extraordinarily similar in gametophytic characters to Timmie/la, Gertrudiella and other large and presumably
non-reduced genera of the Pottiaceae. Like Polytrichum, Timmia has a wealth of characters, all of which are found in the
Pottiaceae except the following: inclined capsule position,
stomates occurring in several rows in the lower half of the capsule, exothecial cells with sinuous walls, operculum shortmammillate, outer peristome is present, and inner peristome,
although similar to that of the Pottiaceae in being filamentous,
with 64 segments, and the Primary Peristomial Layer to Inner
Peristomial Layer cell ratio (cf Edwards 1979, Shaw et al.
1989) is apparently that of the Diplolepideae rather than the
Haplolepideae. Timmia shares the diplolepideous type of peristome with other families of non-pottiaceous taxa, and would
not be expected to appear high in the Pottiaceae tree. Vitt
(1984) treats Timmia as quite a derived taxon in the Bryales.
Timmia, Timmie/la and Gertrudiella are also quite like
Polytrichum in range and development of gametophytic characters; Polytrichum is possibly farther removed from the Pottiaceae than Timmia because of its nematodontous peristome, but
it must be recognized that the sharing of numerous
gametophytic characters generally not found in similar combination elsewhere in the mosses is evidence of a close phylogenetic relationship between these haplolepideous, diplolepideous and nematodontous genera. The weighting of the three
peristome types depends (or should depend) on the number of
characters involved in each and is not pursued to any extent in
this work. Consideration should be given to the possibility that
haplolepideous families may have evolved more than once, and
that past weighting of certain characters of the peristome (e.g.
presence or absence of the outer peristome) should be eliminated. Shaw et al. (1989) found that although peristome developmental data unite haplolepideous mosses, there is as yet no
information as to whether or not this is a synapomorphic condition or not.
Timmia also has characters associated with reduced members of the Pottiaceae, including stem sclerodermis absent,
leaves tubulose above, costa elliptical in transverse section,
sometimes monoicous, and spores rather large. It is here considered that the gametophytes of Polytrichum, Ptychomitrium and
Timmia represent the character states of a non-reduced ancestor
of the Pottiaceae better than do other Pottiales. The possibility
that the Pottiaceae may be been derived from a reduced immediate ancestor (e.g. other Pottiales) is less probable because the
character state combinations of the many Pottiaceous taxa of
large stature, which share so many characters with Polytrichum
and Timmia, must then have been derived independently. The
nematodontous peristome of Polytrichum and its relatives in the
Polytrichaceae may well have been derived from arthrodontous
and secondarily eperistomate ancestors with gametophytes similar to that of Timmia.
Again, outgroups for the phylogenetic analysis were not
selected from only apparently close sister groups, e.g. other Pottiales such as Calymperaceae or Encalyptaceae, or from the
Grimmiaceae (see Churchill 1981), because of the potential
masking effects of extensive reduction in these groups, at least
18
GENERA OF THE POTTIACEAE
compared to taxa of the Pottiaceae. This is clearly demonstrated
in cladograms including genera of non-pottiaceous families
(Cladograms 1-4) in which some non-pottiaceous genera appear,
not at the bottom of the tree, but at or near the ends of branches,
among what are here considered advanced pottiaceous taxa.
Cladograms 5 and 6, with Encalypta and Grimmia respectively as
outgroups, place Timmia and Polytrichum together at the end of
an apparently highly evolved subclade comprising the hereaccepted (Cladograms 14-16) basal stem of the Pottiaceae. This
would require re-evolution of a large number of character states
that in combination phenocopy the gametophytes of two unrelated
non-pottiaceous taxa.
The essential characters of many non-pottiaceous sister genera
are those of reduction, and are therefore likely to result in convergence. If this is the case, the primitive members of such groups
are unknown, and may well not be as similar to the stem genera of
Pottiaceae as are Ptychomitrium, Polytrichum and Timmia, which
appear near the base of the trees.
Thus, if Polytrichum and Timmia are both distant from the
Pottiaceae and from each other because (l) of the number of
unique characters in their peristomes, (2) the number of species
(major evolutionary events) in their genera, (3) the fact that other
genera share the non-pottiaceous characteristics, and (4) they
share more traits with the Pottiaceae than other mosses, then they
should appear at the base of the Pottiaceae cladogram. This
assumes that the Pottiaceae is monophyletic, of which the twisted
peristome found in various subclades is evidence. Of the eight
non-pottiaceous genera (other than Polytrichum and Timmia) that
were used as outgroups, only Ptychomitrium forced Polytrichum
and Timmia low in the tree. The other seven non-pottiaceous genera are considered, therefore, too modified (probably by morphological reduction and simplification) to act as outgroups in calculating a hypothetical phylogenetic tree.
Other Comments
Although two species of Ditrichum (Ditrichaceae), D. tortipes
(Mitt.) Par. and D. ambiguum Best, have the twisted peristomes
otherwise unique to the Pottiaceae, the gametophytes of that
genus are much reduced, and a case might be made for deriving
the genus from ancestors of Barbula sect. Hydrogonium. The
cladograms, however, do not support for this (but see Cladogram
4).
Encalypta (Encalyptaceae), is variously placed in the
Haplolepideae or Diplolepideae (see Edwards 1979 and Vitt
1984). Encalypta is apparently a closer sister group than Timmia
because of the quasi-haplolepideous peristome, but there is considerable modification of characters, including monoicy, loss
(compared to Timmia) of stem central strand (in most species),
gain of yellow KOH reaction of upper !aminal cells, loss of costal
ventral stereid band, and elaboration of additional layers of costal
guide cells. Bryobartramia (Encalyptaceae, see treatment of
excluded taxa) is a taxon probably reduced from Encalypta-like
ancestors (note that this study will suggest that monotypic genera
of reduced morphology are probably relicts of once larger and
more complex genera now all but extinct), demonstrating convergence in traits associated with reduction in the Pottiaceae:
protonema persistent, small size of gametophyte; very short seta;
spherical and cleistocarpous capsule; large spores; and papillose
calyptra (that of Encalypta is occasionally papillose).
Autapomorphic characters (e.g. the hyaline exothecial cells of
Uleobryum) were not included in the data set in that they are of
no value in determining relationship.
METHODS
The phylogenetic inference method used was that of parsimony;
this method has been successful in reconstructing a complex,
known, real phylogeny, and predicts ancestral character states
(Atchley & Fitch 1991, Hillis et al. 1992). This obtains apparently in spite of problems of present computerized techniques of
phylogenetic inference via parsimony, for example, in dealing
with treatment of missing entries representing "unknown data,
inapplicable data, and polymorphic taxa. Each of those potential
sources of ambiguity is logically (if not computationally)
different..." (Platnick et al. 1991; also see Doyle 1993, Faith
1991 and Robinson 1986a). Although, ideally, is it best to analyze the limits of genera from a cladogram in which species are
the terminal taxa (Funk 1985), this was unachievable with the
large number of species involved. The relationships of the genera reflected in the cladograms produced in this study deal with
traditional generic concepts derived from overall similarity of
the species (see above) and apprehension of morphological
"gaps" between genera. Inasmuch as only 75 morphological
characters were used, a data set with more than a thousand species would produce a very poorly resolved cladogram. It may be
hoped that the present study has resolved the family into smaller
groups related by shared apomorphic characters; these groups
may be studied separately in the future.
Maximally parsimonious trees were generated using the
program Hennig86 (Farris 1988, 1989) with an extended
branch-swapping algorithm, that is, the preliminary multiple
tree-generating command "mhennig*" followed by an extended
branch-breaking command, "bb*". The full data set consists of
75 characters and 86 taxa including ten genera not in the Pottiaceae; the "Pottiaceae" data set consists of a subset of 76 taxa.
All multistate characters were treated as additive (ordered) in
that all such characters could be viewed as having transitional
character states; character states that were not viewed as transitional (e.g. shape of propagula) and which were therefore probably governed by different genetic systems were treated as states
of different characters. A character that is variable in state
(polymorphic in the sense of Mishler 1990) is scored with a
dash ("-")as "unknown, undefined, or missing" (Farris 1988).
The particular algorithm used in Hennig86, becau'se of the large
data set, apparently produces only a "heuristic approximation,"
not a guaranteed minimal tree or trees; exact algorithms are,
apparently, impossibly time-consuming for data sets larger than
15 to 25 taxa (Sanderson 1990). On the other hand, phylogenetic analysis of this large data set offers a window on evolution in a large and complex taxonomic group.
The order in which data is presented to the computer program affects, according to Maddison (1991), the length of the
tree when heuristic tree-searching algorithms are used and
retention index is less than .67 and number of terminal taxa is
greater than 20. So, to fmd a short tree for each different
outgroup used, the data set was subjected to a minimum of 30
computer runs based on random orderings of the rows of taxon
data. A relentless search for the "shortest tree" was deemed useless, however, because of abundant homoplasy, and because the
general concurrence of critical groupings between the short
trees obtained with different outgroups was considered contrib-
EVOLUTIONARY RELATIONSHIPS
uting at least sufficient resolution of patterns for purposes of
suprageneric classification.
The number of equally parsimonious trees represented in most
strict consensus cladograms is only a fraction of the number possible because of limitations in computer memory; however, the
loss of resolution in the consensus trees based on the first 1000 or
more trees is very small compared to consensus trees based on
only the first 100 trees, and Cladograms 11 and 12 support the
fme structure of the rest of the trees, being consensus trees based
on smaller data subsets and a defmite, much smaller number of
equally parsimonious trees.
An Homoplasy-Excess-Ratio analysis (Archie 1989 a,b),
which demonstrates the difference between the length of trees
generated by the data set and that of those created by randomized
data, was not performed. This is because the data set is certainly
non-random since it reflects considerable sorting in past studies.
A data randomization study is appropriate for studies that require
identification of possibly random initial data sets (as in gene studies, cf Waters et al. 1992), or to compare two or more data sets to
judge which is more non-random.
The particular polarization of character states of the outgroups
used here supports in large part Miller's (1979) list of commonly
accepted polarizations or "principles for moss systematics"
(Miller warned of occasional reversals in these generalizations in
particular families). These polarizations were presumably derived
from non-rigorous evaluations but akin to the outgroup criterion.
Some of the 26 polarizations listed by Miller are relevant to the
present study of the Pottiaceae. Presumed primitive traits are summarized as follows: large size, well-developed stem central
strand, distinct stem scleroderrnis (vs. an undifferentiated cortex),
strong costa, epapillose laminal cells; while advanced traits
include excurrent costa, very thin- or very thick-walled laminal
cells, presence of propagula ("specialized diaspores"), axillary
hairs with brownish basal cells (vs. all hyaline hairs), monoicy,
sporophyte with short seta and immersed capsule, stomata absent,
cleistocarpy, and peristome reduced or absent. These intuitive
ideas, when compared (1) with the polarizations indicated in the
list of characters below and (2) with the character states of the
various outgroups contributing to the classification used in this
study (Polytrichum, Ptychomitrium, Timmia and Timmiella in the
Data Set) are largely acceptable at least as they apply to the
present study.
Advanced traits may be convergent across family lines. For
instance, Frahm ( 1991 b) found the following traits to be
apomorphic in the Campylopodioideae of the Dicranaceae, many
of which are likewise apomorphic in the Pottiaceae: "presence of
alar cells, leaf-borne rhizoids, incrassate laminal cells, hyaline
leaf tips and strongly differentiated perichaetialleaves, hyalocysts
in transverse section of costa .. .in the gametophyte and presence
of an annulus, fringed calyptra, filiform peristome teeth, long lid,
stomata and large spore size ... " in the sporophyte. Vitt (1984)
described an hypothetical ancestor to the Bryales as intolerant of
desiccation; perennial; acrocarpous; essentially branchless; stems
with hydroids and leptoids; leaves entire, strongly costate, spiralled, little spreading, unistratose; laminal cells thin-walled, rectangular, mostly homogeneous, alar cells not different; laminal
papillae absent; dioicous; perigonial paraphyses numerous, thinwalled, apical cell enlarged; seta elongate, with hydroids and
leptoids; capsule cutinized, with superficial stomates,
photosynthetic, annulus and operculum present, exostome
19
inwardly thickened, endostome segments oppositely placed and
partly fused to a membrane, cilia absent; spores numerous and
homogeneous in size; calyptra mitrate, hairless and epapillose.
Eddy (1991) postulated as an "archetype" for the Pottiaceae "an
erect, tufted or gregarious moss, 0.5-2 em tall, with lanceolate,
rather opaque leaves that lack either a border, teeth or specialized basal cells"; the present study has managed to clothe this
simple pro-pottiaceous structure with many additional
plesiomorphic character states.
CHARACTER STATES OF THE DATA SET
The character states scored for the data set are listed below; (R)
= reduction-related character, of which there are 22 and
weighted lower than other characters in most cladograms, (OX)
=character not used at all in Cladogram 15 (i.e., not phylogenetically informative), (lx) = character occurred once in Cladogram 15 (i.e. phylogenetically very informative), and * = the
hypothesized plesiotypic character state for the Pottiaceae.
Character numbers correspond to the those in the data matrix
used with Hennig86, and begin with zero.
0. (R, Ox) plant conflux
0. gregarious or scattered
1. *caespitose, usually in a mat or turf
1. (R) length of stem
0. short, less than 1.0 em, usually less than 0.6 em
1. *relatively long, usually 1.0 em or greater
2. (lx) stem transverse section shape
0. *rounded-pentagonal
1. triangular
3. stem central strand presence
0. absent
1. *present
4. stem scleroderrnis presence
0. *not or little differentiated from central cylinder cells
1. clearly differentiated
5. stem hyalodermis
0. *absent
1. present (sometimes small)
6. (IX) axillary hair basal cell walls
0. hyaline and all cells of hair similar
1. *of 1 or more cells with thicker or darker colored walls
?.leaves when dry
0. tubulose
1. *occasionally channeled but not distinctly tubulose
8. leaves when wet
0. appressed to weakly spreading
1. *widely spreading to squarrose
9. (Ox) leaves, size gradation
0. *gradually becoming larger distally on stem
1. rosulate (abruptly larger distally)
10. leaf shape
0. long-triangular to linear-lanceolate
1. *lanceolate
2. broadly ligulate-elliptical to spathulate
11. (R) leaf length
0. less than 1.5 mm
1. 1.5 mm to 3.0 mm
2. *3.0 mm or more
12. leaf ventral surface above rnidleaf
20
GENERA OF THE POTTIACEAE
0. *nearly plane to broadly channeled
1. keeled
13. rather deep, narrow groove along costa
0. *absent
1. present
14. margins
0. sharply incurved
1. *incurved or involute
2. plane
3. recurved to revolute
15. marginal toothing
0. entire or minutely and evenly crenulate
1. denticulate or serrulate
2. *strongly toothed
16. margins specially denticulate
0. *not denticulate only below
1. denticulate only near leaf base or at top of leaf sheath
17. (lx) upper marginal cells differentiation
0. *same number of layers as medial cells
1. always differentiated as a bistratose (or more) border
18. upper marginal cells, elongation
0. *not longer than medial cells (sometimes larger)
1. rectangular and clearly longer than medial cells
19. (lx) leaf apex
0. rounded
1. *acute to acuminate
20. leaf apex ventral surface
0. *flattened, channeled or keeled
1. cucullate
21. (R) leaf base
0. little differentiated in shape
1. ovate to elliptical
2. *sheathing the stem and commonly oblong and with
"shoulders"
22. (lx) costa
0. ending before the leaf apex
1. *percurrent or apiculate or short-excurrent as a mucro
2. excurrent as an awn
23. costal ventral cells viewed from above
0. *quadrate to very short-rectangular, occasionally in many
layers
1. longitudinally elongate 3:1 or more
24. number of rows of cells across ventral surface of costa
0. usually 2 but up to 4
1.4-6
2. 6-10
3. *usually 10 or more
25. shape of transverse section of costa
0. *round to semicircular
I. distinctly flattened, usually reniform
26. ventral stereid band
0. absent
1. *present
27. (1 x) size of ventral stereid band
0. *smaller than the dorsal or of nearly equal size
1. larger than the dorsal
28. transverse section of dorsal stereid band
0. round or semicircular
I. *clearly flattened or ventrally indented, reniform or
crescent-shaped
29. ventral costal epidermis
0. absent
1. *present
30. dorsal costal epidermis
0. absent
1. *present
31. guide cells
0. usually 0
1.2-6
2. *commonly more than 6
32. (Ox) number of guide cell layers
0. *1-2
1. 3 or more
33. hydroid strand
0. absent
1. *present
34. (R, Ox) ventral costal outgrowths
0. *absent or not a bulging pad of cells
1. a bulging pad of cells
35. (R, 1x) ventral costal filamentous outgrowths (note: exclusive of lamellae)
0. *absent or not of filaments
I. of separate filaments three or more cells in length
36. width of medial upper !aminal cells
0. *small to medium sized, 7-14j.lm
1. rather large, 15-17 j.lm
2. very large, 18-25 j.lm
3. extremely large, 25 j.lm or more
37. (Ox) length to width ratio of medial upper !aminal cells
0. *1-2:1
1. 3-4: I or more
38. (Ox) upper laminal cell walls
0 . *thin to evenly thickened, lumens quadrate to rounded
1. trigonous or porose, lumens irregularly angular or stellate
39. superficial walls of upper larninal cells
0. flat or very weakly convex on both sides
1. strongly convex to bulging on both sides
2. *ventrally bulging-mamillose, weakly convex dorsally
40. layering of upper !aminal cells
0. *medially unistratose
1. medially bistratose, at least in patches
41. (R, Ox) dorsal superficiallaminal cell walls
0. *about same thickness as the ventral or weakly thicker
near costa
1. throughout distinctly thicker than the ventral
42. (1x) upper laminal papillae
0. absent
1. *present
43. kind of papillae when present
0. simple, hemispherical
1. *bifid to multifid to columniform to capitulate
44. number of papillae per lumen when present
0. one or occasionally two
1. *2-6, usually bifid or multifid
2. many, usually 6 or more, simple or bifid
45. basal cell group
0. not or little differentiated from upper medial cells
1. *clearly differentiated approximately straight across leaf
or rising higher medially
2. differentiated as a vee, with at least laterally
EVOLUTIONARY RELATIONSHIPS
differentiated cells rising higher marginally as a border
tapering distally
46. propagula presence
0. *always absent or extremely rare and associated with
unusually moist conditions
1. often present and characteristic
47. (lx) position of propagula when present
0. not borne on rhizoids as brood bodies, or absent
1. borne on rhizoids as brood bodies
48. position of propagula when present
0. not born in leaf axils, or absent
1. borne in axils
49. (lx) position of propagula when present
0. not borne on leaf costa or lamina, or absent
1. born on leaf costa or lamina
50. shape of axillary propagula when present
0. not clavate or filamentous
1. clavate or filamentous
51. shape of axillary propagula when present
0. not branching or stellate
1. branching or stellate
52. (R) sexual condition
0. *dioicous
1. monoicous
53. (Ox) perichaetium
0. *terminal on main axis
1. lateral on main axis at ends of very short branches
54. perichaetialleaves
0. *similar to cauline leaves or occasionally smaller or somewhat enlarged
1. distinctly different in size or morphology, sometimes
strongly sheathing
55. (R) length of seta
0. nearly absent to short, less than 1 em
1. *elongate, 1 em or longer
56. (R) seta twist
0. *usually twisted clockwise
I. never twisted
2. usually twisted counterclockwise
57. (R) theca length
0. short, 1.5 mm in length or less
1. *usually between 1.5 and 3.5 mm in length
2. 3.5 mm in length or more
58. (R,lx) shape of theca
0. spherical and non-apiculate
1. spherical and apiculate
2. *ovoid to cylindrical
59. (R, lx) capsule dehiscence
0. cleistocarpous
1. *stegocarpous
60. (R, lx) capsule ornamentation
0. *surface nearly smooth
1. surface evenly mamillose or with distinct protuberances of
strongly bulging cells basally or throughout
61. (R, lx) exothecial cells
0. with walls superficially thickened centrally and lens-like
1. *with evenly thickened superficial walls
62. (R, Ox) capsule
0. cleistocarpous and rupturing mainly along weak transverse
walls at butt ends of long-rectangular exothecial cells
21
1. *rupturing irregularly if cleistocarpous, or stegocarpous
63. (lx) stomates
0. absent
1. *present
64. annulus
0. of weakly differentiated cells
1. *of vesiculose cells, often in two or more rows
65. (R) peristome teeth
0. absent
1. *present
66. (R) peristome teeth when present
0. rudimentary, a low plate or short-elliptical in shape
1. 16, often cleft to near base into two or more rami, or of
16 pairs
2. *32 similar rami
67. (Ox) ornamentation of peristome teeth
0. smooth
1. striate or ridged
2. *spiculose or papillose
68. (R) peristome teeth twist
0. untwisted or very weakly so
1. *distinctly twisted
69. (Ox) peristome teeth if twisted
0. clockwise
1. *counterclockwise
70. (Ox) calyptra ornamentation
0. *smooth or nearly so
1. papillose, distinctly roughened or strongly mamillose
71 . (R) length of calyptra
0. short, less than 1 mm
1. 1-3 mm
2. *more than 3 mm
72. (R) spore diameter
0. *8-l5Jlm
1. more than 15 Jlm
73. KOH color reaction of upper laminal cell walls
0. essentially yellow
1. essentially orange
2. *essentially red, usually a definite brick red
74. (R, Ix) calyptra shape
0. *cucullate or at least long-conic and cleft, usually more
than 1 mm in length
1. mitrate or short-conic and uncleft, commonly lobed, usually less than 1 mm in length
2. campanulate-inflated
Discussion of Character States and Polarity
Outgroups appropriate for analysis need to possess most of the
characters of the Pottiaceae, but be sufficiently distant
phylogenetically as to share a common ancestor with all genera
of the Pottiaceae. Inasmuch as closely related families are much
reduced, choosing the appropriate outgroup required considerable analysis, described below. Basically, the rationale was that
Timmia and Polytrichum were similar in morphology gametophytically to certain genera of the Pottiaceae but had peristomes
of apparently considerable evolutionary distance. The particular
haplolepideous genus used here as outgroup that placed these
two distant but related genera low in the tree when both pottiaceous and non-Pottiaceae genera were included in the dadogram would be the appropriate outgroup. Ten non-pottiaceous
22
GENERA OF THE POTTIACEAE
genera were used as outgroups to construct a series of dadograms, which were then evaluated individually and compared.
Three genera were similar in gametophyte characters to those of
particular apparently non-reduced Pottiaceae, but which have
quite different peristomes: Polytrichum (nematodontous), Timmia
(diplolepideous) and Ptychomitrium (haplolepideous, like the
Pottiaceae). These genera grouped near the base of the tree whenever one or the other of the three were used as outgroup. The
characters of these genera are taken as plesiomorphic.
It is more probable that the many character states (75 characters are scored in this study) for each character of large-statured
genera of Pottiaceae being in large part identical to that of Polytrichum, Timmia and Ptychomitrium is due to sharing those of a
common ancestor than to multiple development of these many
identical traits. There is no evidence that such traits are not
homologous. On the other hand, the probability that Polytrichum
is derived from a diplolepideous ancestor is a position not supported by Edwards' (1979, 1984) extensive evaluations of peristome structure.
In a study of this large number of terminal taxa it is doubtful
that many of the characters are globally homologous because of
their apparent lack of genetic complexity. These 75 characters
were selected for analysis because homology of the character
states is supported by similar morphological positions of the
states, the fact that the characters remain similar through transformation series intragenerically, and because other characters correlate with them intragenerically. Additional work, however, along
the lines reviewed by Wiley (1981, p. 130-158), might reduce
homoplasy in the resultant analysis.
The many quantitative characters used in this analysis have
assigned states that are not arbitrarily circumscribed, but correspond to perceived discontinuities in ranges of values, meaning
that the genera as a whole can usually be easily assigned a particular range of values (e.g. lengths, diameters) for each state of each
character. No statistical studies, however, were done beyond an
informal evaluation of the actual data for the genera as shown in
the data file compiled for the genera during the study, in which
discontinuities in measured states for the 76 genera were usually
obvious.
The list below evaluates the polarity of each character of the
Pottiaceae used in this analysis. There was no one outgroup taxon
found that had all states plesiotypic as per this analysis. The
plesiotypic state is simply stated as such when candidate
outgroups Polytrichum, Timmia and Ptychomitrium have a similar, presumed homologous feature, otherwise exceptions are discussed. Characters for Polytrichum are taken in part from G.
Smith (1971); Encalypta in part from Horton (1982 & 1983);
Timmia in part from Brassard (1979, 1980 & 1984). Discussions
of taxa other than these three are provided to better describe particular shared characters or variations that are probably apotypic
and commonly associated with reduction. A hypothetical sister
group based on this evaluation was, however, not used as
outgroup in the actual analysis.
Polarity of Character States
0. Plants growing caespitose is a plesiomorphic condition; the
gregarious or scattered state is associated with reduction.
1. A relatively long (more than I em) stem is plesiomorphic,
although Ptychomitrium has a rather short stem.
2. Rounded-pentagonal stem sections are plesiomorphic, being
found in Ceratodon, Encalypta, Polytrichum and Timmia;
triangular stem sections are rare in the Pottiaceae.
3. Presence of a stem central strand is plesiomorphic, found in
Ceratodon, Polytrichum, Timmia and Ptychomitrium but
usually absent in Encalypta; absence of a central strand is,
however, not clearly associated with reduction in the Pottiaceae.
4. A stem sclerodermis is clearly differentiated in Ceratodon
and Polytrichum but not is so in Encalypta or Timmia (outer
cortical cells are smaller and only weakly thickened); one
might assume the condition of Polytrichum is plesiomorphic
in association with the large size and complexity of the
plant. The situation is variable in Ptychomitrium.
5. Lack of a stem hyalodermis is plesiomorphic, being absent in
Ceratodon, Enca/ypta (occasionally present in patches),
Polytrichum and Timmia, but present in Ptychomitrium.
6. Basal cells of the axillary hair being colored rather than
hyaline (as is the remainder of the hair) is advanced; this is
true for Ceratodon and Timmia, but not Polytrichum. In
Enca/ypta (several species examined), the basal cells are
brownish, but entirely hyaline hairs have been reported,
Horton 1982). Ptychomitrium is variable in respect to this
character.
7. Leaves variously broadly channeled or tubulose when dry is
advanced, this being variable in Encalypta and in Timmia
and not tubulose in Ceratodon, Polytrichum or Ptychomitrium. Again, the condition of the genus with largest plant
size is taken as plesiomorphic.
8. The leaves of Polytrichum and Timmia are strongly spreading, and nearly squarrose when wet; the condition is plesiomorphic. Those of Ptychomitrium are less strongly
differentiated in this respect.
9. Leaves becoming gradually larger on the stem is the
plesiomorphic condition.
10. Polytrichum, Timmia, Ptychomitrium and Ceratodon have
lanceolate leaves; the condition is plesiomorphic. The
spathulate to ovate-lanceolate leaves of Enca/ypta are taken
as apomorphic, and, as in the Tortula group of Pottiaceae,
associated convergently with loss of the ventral stereid band
and enlargement of upper laminal cells.
11. Leaves usually relatively long, 3.0 mm or more, is the condition with Polytrichum and Timmia, and larger species of
Ptychomitrium, and is plesiomorphic. Encalypta leaves are
likewise rather large.
12. Although Ceratodon has keeled leaves, Polytrichum, Timmia, Ptychomitrium and Encalypta have broadly channeled
leaves, and the latter is the plesiomorphic condition.
13. Although Ceratodon (and many robust genera of Pottiaceae)
have leaves narrowly grooved along the costa, Polytrichum,
Timmia, Ptychomitrium and Encalypta have a ventrally
rather flat upper leaf surface, which is the plesiomorphic
state.
14. Type of marginal flexion is a difficult character to polarize
satisfactorily but is taxonomically critical. The upper leaf
margins of Polytrichum, Timmia, Ptychomitrium, Diphyscium and the Dicranaceae in general are plane to incurved
above (like those of Trichostomum in the Pottiaceae), those
of Ceratodon are recurved, tlwse of Grimmiaceae plane to
recurved. Encalypta has both plane and recurved margins.
Both incurved and recurved leaf margins are found in the
EVOLUTIONARY RELATIONSHIPS
Pottiaceae. If such flexion were equivalent evolutionarily (e.g.
both providing marginal stiffening), then both directions of
marginal recurvature would be apomorphic. Infolded margins,
however, provide additional protection in arid environments,
and this indicates an advanced condition. The state for Polytrichum, however, is taken as plesiomorphic with respect to
the Pottiaceae.
15. The upper margins of Polytrichum, Timmia, Ptychomitrium
and some species of Diphyscium are commonly strongly and
distantly toothed, and those of Ceratodon are denticulate near
the apex; Encalypta has entire margins. Robust species in the
Pottiaceae commonly have dentate margins, and, given that
robustness is here taken as plesiomorphic, the clearly dentate
condition is considered likewise plesiomorphic.
16. The marginal ornamentation of Polytrichum, Timmia, Ptychomitrium, Encalypta, Bryobartramia and Ceratodon is never of
denticulation restricted to the upper portion of the leaf base;
leaf margins not denticulate only below is plesiomorphic.
Molendoa is occasionally denticulate at the top of the leaf
sheath margin while Eucladium is nearly always so.
17. Polytrichum , Timmia, Encalypta and Ceratodon lack bistratose leaf borders, but these are common in Ptychomitrium
and Diphyscium; the unistratose margin is considered here
plesiomorphic.
18. Cells of leaf margins similar to those of the medial leaf is the
plesiomorphic condition.
19. Acute leaf apices are plesiomorphic; the apices are obtusely
acute to rounded in Encalypta, but this is correlated with an
apomorphic broadening of leaves and loss of ventral stereid
band.
20. The cucullate leaf apex is an apomorphic character.
21. While Ptychomitrium and Ceratodon have a merely ovate leaf
base, the leaf base of Polytrichum and Timmia sheathes the
stem and has "shoulders," the latter being the plesiomorphic
state. The leaf base of Encalypta is occasionally differentiated
in shape, but this is usually masked by the broadened distal
portion of the leaf.
22. A percurrent costa is plesiomorphic.
23. Polytrichum, Timmia, Ptychomitrium and Encalypta have
quadrate to very short-rectangular ventral costal cells, while
Ceratodon has elongate ventral costal cells; the former is the
plesiomorphic state.
24. Polytrichum and Timmia have a broad costa commonly more
than 10 ventral cells across, which is the plesiomorphic state.
The costa of Encalypta is (r8 ventral cells in breadth, while
Diphyscium commonly is 15 or more costal ventral cells
across. Ptychomitrium, however, has a narrower costa.
25. The costal section of Timmia is elliptical, with the ventral surface usually distinctly convex, similar to that of Encalypta;
Ceratodon and Polytrichum have a reniform costal section,
the latter shape is probably plesiomorphic. The costal section
of Ptychomitrium is semicircular and is probably derived.
26. Two stereid bands are present in Polytrichum, Timmia,
Ptychomitrium and Ceratodon, which is the plesiomorphic
character. Diphyscium commonly has two stereid bands while
Encalypta has only one.
27. The ventral stereid band being smaller than the dorsal is the
plesiomorphic state.
28. Polytrichum, Timmia, Ptychomitrium and Ceratodon have
reniform, strongly flattened ventral stereid bands. The same is
23
the case with Encalypta and Diphyscium.
29. Ventral costal epidermis being present is the plesiomorphic
state. The lack of this is also rare in the Pottiaceae.
30. Dorsal costal epidermis being present is the plesiomorphic
state; it is commonly absent in Encalypta, and is rare in
Pottiaceae. Perhaps the similar Syntrichia, for which the
lack of a dorsal costal epidermis is diagnostic, is an example
of convergent evolution with Encalypta.
31. Polytrichum and Timmia usually have more than six guide
cells (Brassard 1979 illustrates only six for Timmia);
Ptychomitrium and Ceratodon have 2-6 guide cells, and
Encalypta 6-8; a large number of guide cells is the
plesiomorphic state.
32. A single layer of guide cells is the plesiomorphic state;
Encalypta commonly has more than one; this is rare in
Pottiaceae.
33. A hydroid strand (occasionally multiple) is present in Polytrichum, Timmia and Ceratodon and is the plesiomorphic
state; a hydroid strand is variably present in Encalypta and
absent in Ptychomitrium.
34. Although the ventral surface of the costa of both Polytrichum and Timmia is distinctly convex, it does not constitute a
pad of cells (the longitudinal lamellae of Polytrichum are
not considered a pad). Lack of a bulging pad of cells is the
plesiomorphic state.
35. Ventral outgrowths absent or at least not of filaments is
plesiomorphic; the presence of lamellae in Polytrichum is
apparently advanced and is only matched in the Pottiaceae
by Pterygoneurum.
36. The upper laminal cells of Polytrichum, Timmia and Ptychomitrium are rather small, ca. 9-14 l!m in width, while those
of Ceratodon are somewhat wider, and those of Encalypta
7-181-lm in width; the first state is plesiomorphic.
37. A short length to width ratio is plesiomorphic.
38. Upper laminal cells with thin to evenly thickened walls is
the plesiomorphic state.
39. The upper laminal cells of Ceratodon, Polytrichum and
Ptychomitrium are weakly convex and those of Encalypta
strongly convex on both superficial surfaces (occasionally
somewhat more bulging ventrally than dorsally in
Encalypta), and those of Timmia are ventrally strongly
mamillose and dorsally nearly flat (except T. sibirica, in
which this difference is both poorly developed and masked
by dense papillae). An areolation similar to that of Timmia
is present in a far larger number of genera of the Pottiaceae
than previously recognized (though often masked by papil, lae). The Calymperaceae (Pottiales) commonly has ventrally
bulging upper laminal cells, which is probably the
plesiomorphic condition for the Pottiaceae.
40. The unistratose leaf condition is plesiomorphic, though variable in Polytrichum and Ptychomitrium.
41. The superficial cell walls of the upper leaf being of equal
thickness on both sides of the leaf is plesiomorphic.
Although the dorsal cell walls of the leaf are commonly
thicker than the ventral in the lower portion of the leaves of
Timmia, the ventral and dorsal walls are of similar thickness
in the majority of the limb.
42. The leaves of Ceratodon, Ptychomitrium and Polytrichum
lack papillae, those of Encalypta are papillose, and those of
Timmia are variously papillose or smooth. Inasmuch as
24
GENERA OF THE POTI1ACEAE
most taxa in the Pottiaceae, including all robust, morphologically complex taxa of Pottiaceae (excepting Streptopogon),
have papillose upper leaf cells, the papillose condition is presumed plesiomorphic and that of Polytrichum is advanced.
43. The papillae of Timmia are coarse and bifid, and those of
Encalypta are generally well developed and bi- or trifid. The
bifid to multifid state is plesiomorphic. None of the taxa was
always capitulate or columniform.
44. Both Timmia and Encalypta have 2-4 papillae per lumen, the
plesiomorphic condition. Polytrichum is scored, however, as
characteristic not present.
45. The basal cells of Polytrichum, Ptychomitrium and Timmia
are differentiated straight across the leaf base, thus being the
plesiomorphic condition. They are poorly differentiated in
Ceratodon and differentiated straight across or rising higher
medially in Encalypta.
46. Propagula are unknown in Polytrichum, Ptychomitrium and
Timmia, and are axillary and filamentous in both Ceratodon
and Encalypta. The former state is the plesiomorphic condition.
47-51. Not polarized.
52. Ceratodon and Polytrichum are dioicous, Encalypta is largely
autoicous but occasionally dioicous, and Timmia is largely
dioicous but also monoicous. Ptychomitrium is monoicous.
Monoicy, being associated with reduction in the Pottiaceae, is
here regarded as apotypic (but cf discussion of Stark and
Castetter 1987, p. 195, who find no support for the hypothesis
that monoicy is generally favored in arid lands).
53. The terminal perichaetium is the plesiotypic condition.
54. Although the perichaetial leaves are somewhat differentiated
in Ceratodon, in Polytrichum, Ptychomitrium, Timmia and
Encalypta they are little differentiated, which is considered
here the plesiomorphic state.
55. An elongate seta is the plesiomorphic state.
56. The seta is twisted clockwise in Polytrichum, Ptychomitrium,
Timmia and Ceratodon. In Encalypta it is twisted counterclockwise. The former is taken as plesiomorphic.
57. The thecae of both Polytrichum, Timmia and Ceratodon range
from 2.5 to 3.0 mm in length, which is taken to be the
plesiomorphic condition. The measurements for Encalypta are
1-3 mm, probably associated with reduction.
58. An ovoid to cylindrical capsule is the plesiomorphic state.
59. The stegocarpous condition is plesiomorphic.
60. Smooth capsule walls is the plesiomorphic state.
61. Evenly thickened superficial exothecial cell walls is the
plesiomorphic condition.
62. Capsules stegocarpous is the plesiomorphic condition.
63. Stomates present is the plesiomorphic state.
64. The annulus is of vesiculose cells in Ptychomitrium, Timmia,
Encalypta and Ceratodon, but is weakly developed in
Polytrichaceae; the latter is probably the plesiomorphic state
for the Pottiaceae.
65. Peristome teeth present is the plesiomorphic state for the
Pottiaceae.
66. There are 64 inner peristome rami in Timmia, 16 of the
homologous equivalent in Encalypta and Ptychomitrium, and
16 pairs of teeth fused at the joints in Ceratodon. Inasmuch as
there are clear reduction series in certain genera of the Pottiaceae beginning with species with 32 similar rami, this latter
condition is taken as plesiomorphic. Peristome teeth charac-
ters are scored as character not present in Polytrichum.
67. The ornamentation of the teeth of Timmia and Encalypta
(endostomes) and of Ptychomitrium and Ceratodon is
papillose to weakly spiculose in the first two and strongly
spiculose in the last two. Ridged, striate or smooth are
derived states.
68. Although the peristome teeth of Ptychomitrium, Timmia,
Encalypta and Ceratodon are straight, having twisted peristome teeth is a major, unique character state of the Pottiaceae, and occurs in morphologically disparate intrafamilial
groups; it is assumed to have been a feature of the immediate ancestry of the Pottiaceae and is here taken as
plesiomorphic.
69. Peristome teeth, when twisted, wind mostly counterclockwise in the Pottiaceae, rarely clockwise. Ptychomitrium,
Timmia, Encalypta and Ceratodon have straight teeth, while
Ditrichum tortipes (Ditrichaceae) has its peristome and cells
of the operculum clearly twisted counterclockwise. Since
the last taxon may not represent a sister group but may
instead be derived from Barbula sect. Hydrogonium, there
is no good outgroup indication of the state of the hypothetical ancestor of the Pottiaceae. A counterclockwise twist is
considered plesiomorphic based on ingroup comparisons:
twisted peristomes in various major groups of the Pottiaceae
are nearly all wound counterclockwise, and parallel derivation from a clockwise ancestor is far less probable than the
occasional reversal.
70. Calyptrae are smooth in Ptychomitrium, Timmia and Ceratodon, which is the plesiomorphic state, but variously
smooth, papillose or prorulose in Encalypta. This character
is scored as "not smooth" for Polytrichum.
71. The calyptrae of Polytrichum and Timmia are generally 5-Q
mm in length while those of Ptychomitrium and Ceratodon
are reduced in size; the former is the plesiomorphic condition. Those of Encalypta are 2-10 mm in length.
72. The spores of Timmia are commonly 12-20 Jlm in diameter,
those of Ceratodon are mostly 11-15 Jlm in diameter, and
those of Encalypta 7-80 Jlm in diameter. Because larger
spore size is generally associated with gametophytic reduction in the Pottiaceae, the smallest spore size state, also typical of Polytrichum, is taken as plesiomorphic.
73. Ceratodon is yellow in KOH; Encalypta is KOH yellow in
the upper leaf but the basal cells react red, and mature
leaves become KOH red or orange above; Buxbaumia,
Diphyscium and Polytrichum leaves are red in KOH.
Ptychomitrium is variable in KOH reactions. The upper
lamina or upper (distal on the stem) leaves of Timmia in
KOH are either yellow, orange, colorless, or faintly pink
above (in T. austriaca Hedw.-walls were examined at high
magnification under microscope to distinguish the bright
yellow-green of the cell contents), occasionally with medial
patches of brick-red blush, occasionally intensely brick-red
at damaged portions of the leaves; the leaf bases and costae
are generally a deep, clear yellow with large brick-red
blotches. The botchy condition is similar to that of certain
Tortula and Crossidium species, which are here considered
essentially yellow in KOH because most species have
strongly KOH yellow upper laminae of both young and
mature leaves. Tortula (as emended here) and Crossidium
species do not have leaves entirely with walls pink or red.
EVOLUTIONARY RELATIONSHIPS
The upper laminal cells of the more mature leaves (lower on
the stem) of Timmia species, generally, have distinctly pink
walls throughout; the mature leaf has much the color response
as have those of Bryoerythrophyllum, here considered essentially red in KOH. Some species of Bryoerythrophyllum (cf.
treatment of that genus) have KOH yellow young leaves but a
typically red reaction of more mature leaves. Timmia has an
essentially red color response of the upper laminae to KOH,
which is here considered the plesiomorphic character state for
the Pottiaceae.
74. The mitrate calyptra is characteristic only of reduced taxa in
the Pottiaceae (excepting Streptopogon, which because of its
evident lack of characters may be derived from highly
reduced ancestors, cf. Cladogram 14, with little subsequent
elaboration except in stature), and is here considered
apomorphic. Ptychomitrium has a mitrate calyptra. The
inflated campanulate calyptra typical of the Encalyptaceae is
probably an elaboration of the mitrate type.
THE CLADOGRAMS
Sixteen cladograms are presented here to summarize the cladistic
analyses. Cladograms 17 through 19 interpret diagrammatically
trees of other workers at the suprageneric level. In the cladograms
given here, dichotomous branching at a node is shown as
while multiple branching at a node is symbolized by a longer vertical line with several horizontal lines appended to the right. Thus
all terminal taxa and subclades attached to the right of a single
vertical line are branches, e.g. "G" is a multiple-branched node in
the consensus trees of Cladograms 3 and 4, and so on. The
outgroup is also shown as coming from a multiple-branched node
in all cladograms because the algorithm, apparently, cannot evaluate the relative phylogenetic position of the outgroup and the first
branch; for the purposes of this study, this can be ignored.
Cladograms 1 through 6 are based on the full data set, which
includes information on ten genera not in the Pottiaceae: Polytrichum (Polytrichaceae); Bryobartramia and Encalypta (Encalyptaceae); Ceratodon and Ditrichum (Ditrichaceae); Diphyscium
(Diphysciaceae); Grimmia (Grimmiaceae); Ptychomitrium
(Ptychomitriaceae); Syrrhopodon (Calymperaceae); and Timmia
(Timmiaceae). Cladograms 2-4 do the following: (1) compare the
cladograms of genera relatively distant from the Pottiaceae, Polytrichum and Timmia, to those of putative sister groups of the
Pottiaceae, and (2) identify the closest appropriate haplolepideous
sister group to the Pottiaceae as those two (Ditrichum and Ptychomitrium) placing the relatively distant genera Polytrichum and
Timmia closest to the base of the tree. Because Ditrichum is
-potentially a derivative of the Barbula lineage or at least may
belong to the Merceyoideae (Cladogram 4), Ptychomitrium is
probably the ideal sister for use as outgroup in analysis of the
Pottiaceae.
All the listed non-pottiaceous genera were examined individually as outgroups; Cladograms 5 and 6, with Encalypta and Grimmia respectively as outgroups demonstrate that, like cladograms
with Bryobartramia, Ceratodon, Ditrichum, Diphyscium, and
Syrrhopodon (these not shown), sister groups that are apparently
reduced and simplified in morphology create cladograms that
place Polytrichum and Timmia high in the tree. Because the data
set used is the same in Cladograms 1-6, the branching structure of
Cladograms 2-6 would be identical if they had been able to be
calculated exactly. To the extent that the actual heuristically
"--1 "
25
derived trees of Cladograms 2-6 agree, this ideal branching pattern is approximated. Branching patterns of Cladograms 2-6 (as
well as the other cladograms) that are well resolved support the
classification indicated by lettered lineages in Cladogram 16.
Cladograms 7 through 10 include only pottiaceous genera in
addition to a non-pottiaceous outgroup. Subclades that are similar throughout the 16 cladograms and which are also formally
named in Cladogram 16 are marked with letters at ancestral
nodes. Cladograms 11 and 12 analyze character state changes in
two highly evolved subclades that comprise the same species
and the same structure in both Cladograms 9 (Timmia as
outgroup) and 10 (Ptychomitrium as outgroup); these are
approximately the same as the Merceyoideae and Pottioideae of
Cladogram 16 with the major exception that the Hyophileae is
not a distinct subclade. Cladograms 13 through 16 involve only
genera in the Pottiaceae, use Timmie/la as functional outgroup,
and summarize the best hypothesis for phylogenetic relationships and of a projected suprageneric classification.
Cladograms 1, 2, 7, and 8 have Polytrichum as outgroup,
Timmia is outgroup in numbers 3 and 9, Ptychomitrium is
outgroup in numbers 4 and 10, Encalypta is outgroup in number
5, and Grimmia is outgroup in number 6.
Cladograms 1-10 and 13-14 are strict consensus trees, summarizing the information from many equally parsimonious
trees. Other cladograms are of single trees selected from sets of
equally parsimonious trees (one set for each outgroup), usually
through the use of the functional ingroup and functional
outgroup (FIG/FOG) method of Watrous and Wheeler (1981)
on various subclades that have multiple-branched nodes in the
consensus tree.
Cladograms 1, 7 and 13 weight all characters alike (at
weight 1). All other cladograms in this analysis have 22
reduction-related characters weighted at 1: 15 (i.e. reductionrelated characters are weighted at 1, non-reduction-related at
15). The reason for this particular level of weighting is that
cladograms generated at successively higher weightings but
below 1:15 are different from each other, but at 1:15 and higher
weighting ratios, they are the same. Because reduction series are
observed internal to each of many genera in the Pottiaceae,
reduction series may be expected to be several and parallel
within the family. These reduction series should be distinguished from each other on the basis of characters other than
those generally associated with reduction intragenerically. The
1: 15 weighting provides for this. Reduction-related characters
cannot be eliminated because, although they may not be
expected to be homologous globally, they may be important
clues to relationships in particular lineages.
The first cladogram (Cladogram 1) clearly shows what are
comparatively reduced taxa, such as Crossidium and Aschisma,
clustered in the same lineage. To see if such taxa were associated by characters other than those of reduction, twenty-two
characters were selected for the weighting above as either
directly associated with reduction in size and complexity, or
which are elaborations (e.g. costal ornamentation, capsule wall
ornamentation) associated with reduction in the Pottiaceae. The
characters each commonly contribute at least two additive steps
in the clade. The apotypic states of these reduction-related characters, numbered according to the main character list above, are:
0 gregarious (as opposed to caespitose); 1 stem short; 11 leaf
short; 21 leaf base not differentiated; 34 bulging pad of cells
26
GENERA OF THE POTTIACEAE
present on costa; 35 filamentous costal outgrowths present; 41
dorsal superficial !aminal cells walls clearly thicker than the
ventral; 52 monoicous; 55 seta short; 56 seta not twisted; 57 theca
short; 58 theca spherical; 59 cleistocarpous; 60 exothecial cells
bulging; 61 exothecial cells with lens-like medial thickenings; 62
capsule cleistocarpous and rupturing along weak transverse walls;
65 peristome absent; 66 teeth short when present; 68 teeth
untwisted when present; 71 calyptra short; 72 spores large; and 74
calyptra mitrate.
RESULTS (see also Summary above):
Phylogenetic Position of Non-Pottiaceous Genera
In Cladograms 1-4, Ceratodon, Grimmia and Syrrhopodon, representing three different families, are associated with what are
here termed (in Cladogram 16) the Merceyoideae (equivalent to a
combination of the traditional Barbuloideae and Leptodontioideae). Also, Bryobartramia, Diphyscium and Encalypta, representing two different families, are associated with the Pottioideae
group. This may be evidence either of derivation of these families
from Pottiaceae stock (thus a primitive member of the
Haplolepideae) or of polyphyly, or simple "long branch
attraction" of very distant relatives (cf. Albert & Mishler 1992);
distinguishing the possibilities requires extensive study at a
broader level than is appropriate in this study. The presence of the
unusual twisted peristome in three different lineages of the Pottiaceae indicates monophyly. But note support for Robinson's
( 1971 b) classification which groups all the above under the
Dicranales, eliminating the Pottiales as an order.
In the sense of Farris (1974; see Wiley 1981, p. 92), the Pottiaceae would be paraphyletic if at least one other family not sharing advanced distinctive pottiaceous characters with other genera
is included in the tree; this is a problem since the twisted peristome is commonly reduced to conditions (straight, rudimentary
or absent) indistinguishable from similar non-homologous conditions. The Pottiaceae would be also be polyphyletic if the Ditrichaceae were recognized as a separate family possessing a twisted
peristome in some members, requiring the Pottiaceae to be distinguished solely as a polythetic group. Work needs to be done to
determine the phylogenetic importance of peristome characters
through actual analysis across families.
Major Pottiaceous Groups
The various cladograms with outgroups of one of ten different
non-pottiaceous genera with three very different peristome types
(diplolepideous, nematodontous, haplolepideous) produced dadograms with much in common. It is quite possible that the three
outgroup taxa with the most similar cladograms (Polytrichum,
Timmia and Ptychomitrium) are phylogenetically closely related.
Weighting the non-reduction related characters at 15 gave greater
resolution to the cladograms, and was designed to limit artificial
pairings on the cladogram due to convergence of non-homologous
traits.
Cladograms 5 and 6 with Encalypta and Grimmia as outgroups demonstrate a similarity in many subclades with subclades
found in cladograms created with other outgroups. These dadograms indicate that the classification arrived at here reflects to a
great degree three major groups or branches that are discernible in
relatively well resolved cladograms, such as Cladograms 3, 4, 5
and possibly 2 (among those including non-pottiaceous taxa). In
Cladogram 5, the traditional Trichostomoideae (dealt with in ,
Cladogram 16 as a basal stem requiring names for otherwise
minor branches) can be seen as a major branch.
Thus the three major branches of the network shared by the
cladograms (i.e. similar Steiner minimal trees) are approximately the traditional Trichostomoideae, Barbuloideae and
Pottioideae. If Polytrichum and Timmia are not closely related
to the Pottiaceae, and these two genera are simply evolutionarily convergent to the traditional Trichostomoideae and to each
other, then the correct outgroup for the Pottiaceae might better
be a taxon similar to the pottiaceous genera near the joining of
the three major branches, which are certainly more similar to
other families in the Pottiales. No non-pottiaceous genus analyzed is close to the joining except Ditrichum in Cladogram 4.
Possible ingroups basal to such a pottiaceous tree include Leptobarbula, Hymenostyliel/a, Tetracoscinodon and Tetrapterum.
This line of attack might be pursued but, in this study, evolutionary convergence of Polytrichum, Timmia and the traditional Trichostomoideae (as exemplified by Timmiella) is
clearly less probable than derivation of the Pottiaceae from a
shared ancestor of all three. If the immediate ancestor of the
Pottiaceae were similar (in being relatively reduced) to other
families in the Pottiales, then even a fourth group, the
Ptychomitriaceae (haplolepideous), would join the taxa required
to independently evolve the large complex of morphological
and anatomical characters characteristic of robust taxa of the
traditional Trichostomoideae. Cladograms 5 and 6 are here considered improbable as hypotheses because the two outgroups for
these do not restrict Polytrichum and Timmia to the base of the
tree. If convergence of several lines of taxa with Timmiella-like
character states of the gametophyte were likely, as might be
caused by simple, single-gene desuppression of character states,
then there would probably be Polytrichum- or Timmia-like
gametophytes having sporophytes characteristic of the Dicranaceae or Grimmiaceae or Orthotrichaceae; but there are not.
Thus, convergence of complex gametophytes is unlikely while
reduction as advanced traits, common in many families, is
apparently the best justification for placing the traditional
Trichostomoideae low in the tree.
Taxa that are apparently morphologically reduced occurred
in Cladogram 1 (all characters at weight 1) either alone (e.g.
Anoectangium, Leptobarbula, Tetracoscinodon) or were clustered, i.e. Aloinella, Crossidium and Globulinella with Tortula,
and Acaulon, Aschisma, Byroceuthospora, Microbryum, Phaseapsis, Pterygoneurum, Stegonia, Stonea and Trachycarpidium
with Tetrapterum in the Pottioideae subclade. These taxa are
also either alone or clustered, respectively, in Cladogram 13, all
characters also at weight 1, but which is less resolved. The
reduced taxa remain clustered when non-reduced characters are
weighted at 15: l in generally l-3 lineages in Cladograms 2-4
and 8-10 and 14, which implies that these taxa are closely
related. The grouped reduced taxa continue to be grouped even
when the importance of reduction-related characters is
minimalized.
Two major traditional suprageneric taxa, the Pottioideae
(with spathulate leaves with a single stereid band in the costa
and margins commonly recurved) and the Barbuloideae cum
Leptodontoideae (here treated as the Merceyoideae-with
lanceolate leaves and recurved margins, the basal cells largely
reaching higher in the leaf medially), are well distinguished in
most cladograms as two large subclades terminating the tree.
EVOLUTIONARY RELATIONSHIPS
The genera traditionally comprising the Leptodontioideae clearly
belong with the Merceyoideae, as do most genera of the traditional Pleuroweisieae; the former group is well distinguished as a
subclade of the Merceyoideae, the latter genera are rather scattered within the Merceyoideae, which reflects well on my previous comment (Zander 1977c, p. 240) that the Pleuroweisieae
probably has little phylogenetic coherence. Inasmuch as the two
groups (approximately the Merceyoideae and Pottioideae of
Cladogram 16) comprise identical taxa and identical structures in
Cladograms 9 and 10 (subclades "F" and "G" of both trees), an
analysis of character state changes was made, which consisted of
analysis of the subclades alone and selection of single trees from
each of the resulting two consensus trees (Cladograms 11 and 12).
Cladogram 11 includes only those terminal taxa distal to node
Fin Cladograms 9 and 10, approximately the Merceyoideae recognized in Cladogram 16. Tetracoscinodon is the outgroup in this
analysis. Four subclades are distinguishable, Tetracoscinodon
(sole member), the Barbula group, the Leptodontium group, and
the Bryoerythrophyllum group. This is support for these groups to
be treated as tribes (see Cladogram 16) in that they are fairly stable among the various cladograms with different outgroups shown
here. Polytomies in the consensus tree occurred distal to the
immediate ancestor of Barbula. A functional ingroup and functional outgroup (FIG/FOG) analysis (Watrous and Wheeler 1981)
was done with Anoectangium as FIG and including all taxa distal
to Anoectangium as FOG. Tree 24 of the 127 trees contributing to
the consensus tree (Cladogram 11) was selected because its
Barbuleae branch matched that of the FIG/FOG result. The characters incompletely map to tree 24 (Cladogram II) in that no
character state changes were found for node "X" of Cladogram
II, the immediate ancestor of Barbula; node "X" may be taken as
an artifact signifying a multiple-branched node in its own immediate ancestor, leading directly to Barhu/a, to the Scope/ophila,
Gymnostomie/la and Didymodon lineage, and to the Gymnostomum and Molendoa lineage. Cladogram II, now tempered with
information from other cladograms, shows the general evolutionary structure of the Merceyoideae. Note that Cladogram 11 has
the Leptodontium subclade derived from a more primitive ancestor than is given in Cladogram 10. The consensus tree of Cladogram 14 shows a similar number of polytomies in the Barbuleae;
the structure of the Barbuleae of the single tree reflected in Cladograms 15 and 16 was selected from equally parsimonious trees
based on the above FIG/FOG analysis.
Cladogram 12 summarizes an analysis of the Pottioideae as
shown in Cladograms 9 and 10 with Leptobarhula as outgroup.
As FIG/FOG analysis was to no avail in further resolving this part
of Cladogram 10, but tree 5 of 8 equally parsimonious trees was
chosen, being the first tree to place Weissia and Weissiodicranum
together on a single branch, as is suggested by sufficiently
resolved Cladograms 1, 3, 7, 13 and 14 (but not Cladogram 8). In
spite of the polytomies low in the strict consensus tree, this configuration results in a map of character state changes that retains
much the same groupings for this subclade as seen in other dadograms. The structure of the Pottioideae in Cladogram 12 is much
different than that shown in Cladogram 14 although the taxa are
similar (Aschisma, Byroceuthospora and Uleobryum, among others are included in the Pottioideae in Cladogram 14). In both
cladograms, the Pottioideae subclade is not easily describable as a
morphologically integral unit; "broad leaves" is about the only
readily discernible character state applicable to most taxa. Clado-
27
gram 12 reflects apparent evolution along a continuous lineage,
while Cladogram 14 shows two distinct lineages (also cf Cladogram 8).
I agree with Mishler (1985) that the genus Tortula as traditionally viewed is paraphyletic, in that the genera into which
many of the traditional Tortula species are segregated here,
Hennedie/la, Syntrichia and Tortula s. str., are in three separate
subclades in the Pottioideae in most of the cladograms. Cladograms of Mishler's (1985b, l986a, 1990) previous work with a
number of species of Tortula s. lat. support to a considerable
extent the above three groups recognized here as genera. The
Pottioideae, as here emended with the addition of several taxa
related to Weissia and Hyophila, is a monophyletic group, the
basal synapomorphies given in Cladogram 15 being: stem
sclerodermis little differentiated, leaf base little differentiated,
and seta less than 1 em in length.
The traditional Trichostomoideae (taxa with largely lanceolate, plane leaves, often with a narrow marginal band of hyaline
cells running up from the base) is here dissected into different
lineages in many cladograms; these appear in most cases at the
base of the tree. The traditional Trichostomoideae is here split
into five subfamilies in Cladogram 16 (four are evident in
Cladogram 14 with all characters weighted equally), each of
which is also rather distinctive morphologically as a taxonomic
grade. Norris and Koponen (1989) in a recent description of the
subfamilies of Pottiaceae, found the traditional Trichostomoideae to share the following characters: most abundant at tropical
latitudes, leaf margins plane, areolation of leaf and epidermis
continuous (no leaf butress of shortened cells formed), hyalodermis commonly differentiated, basal !aminal cells often running up the margins in a vee, and a persistent felt of protonema
commonly developed. Most of these traits apply to the subclades at the base of the Pottiaceae tree, but, apparently significant dichotomies at the base of the tree require splitting (and
recognition at the subfamily level), which, by the way, is of a
higher order than the dichotomy between the Merceyoideae and
Pottioideae since such splitting is nearer the base of the tree.
As Ptychomitrium is apparently the closest of the candidate
sister groups to the Pottiaceae that keep the distant taxa Polytrichum and Timmia near the base of the tree (trees with nonpottiaceous taxa in the data set other than Encalypta and Grimmia not shown), Timmie/la is found to be the most primitive
extant genus of Pottiaceae (Cladogram 10); this is also supported in Cladogram 3, while Timmie/la is near the base of
Cladograms 1-2, 4, and 7-9. Although Aloina and Gertrudiel/a
are also found near or at the base of the last mentioned dadograms (see especially Cladogram 4), an analysis with Timmie/la
as functional outgroup was performed and taken to be the best
phylogenetic hypothesis for the family (Cladograms 13-16).
Cladogram 13 is the strict consensus tree with Timmie/la
(Pottiaceae) as functional outgroup and all characters weighted
the same. Three lineages are identical with the named lineages
in Cladogram 16, while many small branches are identical or
similar in structure and terminal taxa. Presumed reduced taxa
(e.g. four small branches including Aschisma, Phascopsis,
Luisiere/la, and Crossidium, respectively) are grouped to some
extent. As in Cladogram 14, the Pottioideae is split into (at
least) two lineages
Cladogram 14 is the strict consensus tree with Timmie/la as
functional outgroup and with non-reduction characters weighted
28
GENERA OF THE POTTIACEAE
at 15:1; it can be viewed as the "raw results" of this phylogenetic
study. Cladogram 14 is more resolved than Cladogram 13. A
striking difference between this cladogram and Cladograms 9 and
10, which agree with each other in many respects, is that the
Pottioideae (1) is split into two lineages, (2) includes the Aschisma, Byroceuthospora and Trachycarpidium group, otherwise
placed with the traditional Trichostomoideae, and (3) derives the
Aloina, Aloinella, Crossidium, Globulinella, Pterygoneurum and
Stegonia group from the Weissia branch of the Hyophila lineage
(as in Cladogram 13).
The Weissia-Hyophila group is characterized by largely
ligulate to spathulate leaves with incurved margins, upper !aminal
cells commonly strongly bulging ventrally and nearly flat dorsally. This group is distinguished in some cladograms (see especially Cladogram 8 and Cladograms 14-16), and appears at the
base of the Pottioideae lineage in others. It is here recognized at
the tribal level (as Hyophileae, see Cladogram 16) because (1) it
is a morphologically distinctive group, and (2) it is surprising that
ancestors of the end members of such a highly structurally modified group with such an unusual (tropics, especially the West
Indies) geographic distribution (see Cladogram 14) should be the
source group for the traditional Pottieae (as in Cladograms 9 and
10).
The presumed reduced taxa that are grouped in Cladogram 13
continue to be associated with each other in Cladogram 14, indicating that their relationships are not due to convergence; on the
other hand, other reduced taxa, such as Leptobarbula, Tetrapterum, Stegonia, Pterygoneurum, Microbryum, and Trachycarpidium, that are poorly resolved in Cladogram 13 are now well
resolved with higher weighting of non-reduced characters.
The Phylogenetic Hypothesis
A single tree (shown in both Cladograms 15-16) was selected
from the more than 1250 equally parsimonious trees contributing
to the consensus tree of Cladogram 14. Multiple-branching nodes
in the Merceyoideae were resolved by selecting from the 1250
trees those several trees with the structure of the Merceyoideae of
the single tree of Cladogram 11. Of these fewer trees, those that
(1) placed Aschisma, Byroceuthospora, Trachycarpidium and
Uleobryum in the same subclade and (2) which also derived
Saitoella from an earlier ancestor than the immediate ancestor of
Microbrym were selected because elsewhere in the cladogram
generally relict and southern taxa have more primitive traits than
larger and more widespread genera (this particular structure is an
example of the cladisfs "reciprocal illumination" and must not be
used as an example supporting any generalization about relict,
southern taxa). Of these yet fewer trees, one tree was selected that
( 1) put all remaining traditional Trichostomoideae that were at
multiple branches in the consensus tree into one subclade because
it allowed a classification that was both simple and did not imply
a degree of resolution greater than was warranted, (2) put Calymperastrum near the base of this subclade because Gondwana taxa
are commonly at the base of other subclades (reciprocal
illumination--do not use this particular structure to support the
generalization), (3) placed Eucladium next in evolutionary order
since this monotypic genus is isolated morphologically and may
be a relict (reciprocal illumination again), and (4) placed Trichostomum and Streptocalypta as most highly evolved because of the
comparatively larger size of the genera in numbers of species,
implying recent evolution. The single tree (tree number 1126) that
met all these above criteria is the basis for Cladograms 15 and
16 and for the present suprageneric classification.
The traits of hypothesized ancestors of named subclades in
Cladogram 15 are detailed in the following chart. These states
characterize a particular shared ancestor in a most parsimonious
tree, but may change later in each of the named lineages (subfamily names capitalized entirely).
STATES AT MAJOR ANCESTRAL NODES
TIMMIELLOIDEAE (functional outgroup)
uide cells fewer, 2-6: ERYTHROPHYLLOPSOIDEAE
ERTRUDIELLOIDEAE
Leaf base not sheathing; lamina unistratose
Leaves long-linear; ventral stereid'band larger than
dorsal: CHIONOLOMOIDEAE
Leaf hydroid strand absent
Axillary hairs completely hyaline; dorsal costal
epidermis absent: TRICHOSTOMOIDEAE
Leaves shorter, 1.5-3.0 mm; rows of cells on
ventral surface of costa fewer, 4-6; costal guide
cells fewer, 2-6; theca shorter, less than 1.5 mm
Tetracoscinodontieae
Leaf keeled above; costa grooved ventrally:
MERCEYOIDEAE
osta flattened in section; leaf hydroid
strand present; theca longer, 1.5-3.5
mm; peristome teeth of 32 similar
rami: Bryoerythrophylleae
Leaf margins recurved to revolute; costal
ventral cells elongate; rows of cells
across ventral surface of costa 2(-4)
Stem central strand absent; ventral
costal epidermis absent; upper
!aminal cells superficially flat or
weakly convex: Leptodontieae
Lamina! papillae simple; seta nearly
absent to short, less than 1 em in
length
Dorsal stereid band section round or
semicircular; peristome teeth absent:
Barbuleae
Stem hyalodermis absent; perichaetia11eaves
sheathing
Leaves tubulose when dry; leaf margins
incurved or involute; upper !aminal cell
walls ventrally bulging and dorsally nearly
flat: Hyophileae
Stem sclerodermis not or little differentiated;
leaf base little differentiated in shape; seta
nearly absent to short, less than 1 em in
length: POTTIOIDEAE
Stem short, less than 1 em in length;
peristome of 32 similar rami; peristome
distinctly twisted: Pottieae
The assumptions are several but they are reasonable and it is felt
that Cladogram 16 is the best hypothesis on which to base a
classification at this time. Again, reciprocal illumination here
EVOLUTIONARY RELATIONSHIPS
helps select that single cladogram (of many of equal length) that
is (1) most internally consistent in all subclades in geographic distribution of the terminal taxa, and (2) most consistent with other
cladograms that do resolve polytomies. Because Cladogram 14
summarizes the results of the study without modification based on
geographical ranges, the geographic data is summarized with it
(q.v.) rather than with Cladogram 16.
Cladogram 15 summarizes the character state changes of this
study's projected hypothesis of the phylogenetic relationships of
the Pottiaceae. Cladogram 16 gives a better view of the structure
of Cladogram 15, and letters the basal ancestral nodes of the
named subclades. The lettered subclades are also noted in other
cladograms when similar (or identical in the case of Cladograms
13 and 14), indicating considerable support for these relationships. Cladogram 16 is a version of Cladogram 15 that clearly
shows the basis for the classification in the taxonomic section of
this work. Note that Cladogram 16: (1) is supported in large part
by cladograms based on the non-pottiaceous outgroups Polytrichum, Timmia and Ptychomitrium, (2) makes few major changes in
traditional classification, and (3) introduces hypotheses that are of
considerable interest and that have a logical, rigorous basis.
There are 72 ancestral nodes in Cladogram 15 (on which
Cladogram 16 is based) that have synapomorphies, plus one node
to which no data on character state changes map. Of the 75 total
scored characters, 14 were unused, these being characters numbered 0, 9, 16, 32, 34, 37, 38, 41, 51, 53, 62, 67, 69, and 70. These
characters evidently have little phylogenetic importance in the
Pottiaceae, at least taking into account the assumptions (including
FIG/FOG and reciprocal illumination) on which this single tree is
based.
The 62 scored characters used in Cladogram 15 supported 97
different character state changes. Of these, 59 state changes
(61%) were true synapomorphies, i.e. only occurred once in the
cladogram. Of the 97 state changes in Cladogram 15, 38 (39%)
were homoplasous synapomorphies. Of these 38, there were 17
character state changes occurring twice in the cladogram, 14
thrice, 5 four times, I five times (character 57:0-lR), and 1 six
times (character 46:0-1). (Note that this refers to state changes in
the cladogram not the number of times the character itself occurs
in the cladogram.) The characters contributing to the many
homoplasous synapomorphies are not unimportant, and indicate
possible modes of convergence of lineages in addition to the characters selected prior to analysis as indicative of reduction.
Eighteen characters only occurred once in Cladogram 15.
These 18 are considered here the characters most important to the
taxonomy of the Pottiaceae, and are (as numbered in the list
above): 2 shape of stem section; 6 color of axillary hair basal
cells; 17 number of layers of marginal !aminal cells; 19 leaf apex
rounded or acute; 20 leaf apex cucullate or otherwise; 22 costa
ending before or at the leaf apex or excurrent; 27 ventral stereid
band larger or smaller than the dor~al; 35 ventral costal outgrowths present as filaments or otherwise; 40 upper !aminal cells
medially unistratose or bistratose; 42 upper !aminal papillose or
smooth; 47 propagula when present borne on rhizoids as brood
bodies or otherwise; 49 propagula when present borne on leaf
costa or lamina or otherwise; 58 shape of theca; 59 capsule
cleistocarpous or stegocarpous; 60 capsule smooth or mamillose;
61 exothecial cell superficial walls with central lens-like thickenings or evenly thickened; 63 stomates present or absent; 74
calyptra cucullate, mitrate or campanulate. That some of these
29
characters are among the reduction-related characters weighted
low in the analysis indicates that they are nevertheless important
in at least some lineages, and must not be eliminated from analysis.
By way of summary, traditional groupings are supported in
large part and grossly dissimilar genera are not associated, thus
the subclades make "sense" to the specialist. Such support for
classical groupings indicates that traditional methods of
"omnispection" and the like are, to a certain point, effective.
Inasmuch as the branches named here at the subfamily and
tribal level are reflected in most of the cladograms generated in
the course of this study, they may be expected to appear in at
least approximate form in future studies.
The major subclades are formally named at the subfamily
and tribal level in the taxonomic section. For these, the "sequencing convention" of Nelson (1972, 1974) is used because of the
considerable asymmetric depth of the tree; that is, the taxonomic levels recognized here-subfamilies and tribes-are
mostly not coordinate with each other according to depth in the
tree. The nine subfamilies are the Timmielloideae, Gertrudielloideae, Chionolomoideae, Erythrophylloideae, Trichostomoideae, Merceyoideae (including tribes Tetracoscinodontieae, Barbuleae, Leptodontieae and Bryoerythrophylleae), and Pottioideae (with two tribes, the Hyophileae and Pottieae).
Minor Groups
There are lesser trends in evolution in the Pottiaceae that are
shown in hypotheses presented by the cladograms. Subclades
that are similar (the same or nearly the same taxa and structure)
to those in Cladogram 16 are marked with letters on the basal
nodes in Cladograms 1-10 and 13-14. These generally support
the structure of Cladogram 16.
Certain genera always appear at the base of the tree: Gertrudiella, Timmie/la, Erythrophyllastrum and Erythrophyllopsis.
The several lineages into which the traditional Trichostomoideae are broken in the various cladograms include these "strong"
(because they appear in several of the different cladograms)
generic associations: Calyptopogon, Pleurochaete and Tortella;
Chionoloma, Pachyneuropsis and Pseudosymblepharis; and
Aschisma, Byroceuthospora, Trachycarpidium and Uleobryum
(these latter are found in the Hyophileae of the Pottioideae in
Cladograms 14-16). Taxa related to Leptodontium are presented
as a well distinguished subclade: Leptodontiella, Leptodontium,
Streptotrichum and Trachyodontium, while Hymenostylium,
Reimersia and Triquetrella are closely related. Anoectangium is
associated with the Barbula group, while Merceyoideae that are
KOH red are placed in a separate branch: Bryoerythrophyllum,
Mironia, Pseudocrossidium and Rhexophyllum, with Dialytrichia (KOH yellow) at the base of the branch.
The fact that most reduced taxa, these with no confounding
mismatches in geographical ranges (see Cladogram 14), are
grouped together (in at least two groups) indicates what are
probably monophyletic lineages, without masking convergence.
The most clearly grouped of the taxa of small stature are: (1)
Aschisma, Byroceuthospora, Trachycarpidium and Uleobryum;
and (2) Stegonia, Pterygoneurum, Aloinella, Crossidium and
Globulinella, probably including Aloina. The second group,
although of small plants, is not of highly reduced morphology in
comparison with terminal taxa with immediate ancestors inserted deeper in the subclade. These, consisting of the Weissia-Hyo-
30
GENERA OF THE POITIACEAE
phi/a group at the base of the Pottioideae lineage in Cladograms 9
and 10 and recognized as the Hyophileae in Cladogram 16, consist of plants of small stature, less simplified than the first group
of taxa above, but commonly with reduced or absent peristomes
and thus more reduced than the second group; these, too, have a
certain geographic integrity (largely tropical). Of course, if there
were more than the 22 identified reduction-related characters acting to produce false associations, these groups might be split further. This should be pursued.
Within the Pottioideae, the KOH red genera are generally segregated as a group (with Willia or sometimes Microbryum at a
basal branch), except that highly reduced taxa (e.g. Crossidium
and Globulinella) of the subfamily are generally clustered
together without regard for KOH reaction.
The position of Aloina is problematic. It is placed near Polytrichum in Cladograms 1, 4, 7 and 8, and, more traditionally, near
Aloinella in Cladograms 2, 3, 9, 10 and 14-16. One might make a
case that the absence of Palytrichum in a cladogram introduces a
false relationship of Aloina with Alaine/la since the new outgroup
(Timmia or Ptychamitrium) is not sufficiently distant from the
Pottiaceae, but this is not true for Cladogram 2 (outgroup Timmia), in which Alaina is associated with Alaine/la not Palytrichum. Pending further study, Alaina is placed with Aloinella, following Cladograms 14-16, and the association with Polytrichum
is considered anomalous, perhaps an example of convergence
together with lack of sufficient data to address the problem.
Position of Reduced Taxa
If one assumes, for the purposes of argument, that all genera
develop from a single species to a series of morphologically
diverse species in a genus and finally senesce towards extinction
by species being eliminated from the genus in response to environmental pressures, certain hypotheses can be developed from
the cladograms, notably Cladogram 14. Although one might
assume that highly morphologically reduced and simplified genera may have been derived from the immediate ancestor of extant
similar genera of greater stature and complexity, the present data
do not support this. Somewhat reduced genera appear near the
base of most subclades (Calymperastrum in the Tarte/la lineage
of Cladograms 2 and 8, Tetracascinadan in the Merceyoideae
(Cladogram 14 and elsewhere), Anaectangium at the base of the
Barbuleae (Cladogram 14 and elsewhere), Weissia in the WeissiaHyophila group of various cladograms and Cladogram 14, and
Leptobarbula at the base of the Pottioideae (Cladogram 14 and
elsewhere.
This may indicate that relatively reduced taxa were ancestors
of these lineages. If such is the case, the twisted peristome of the
Pottiaceae may have appeared several times, possibly as a result
of genetic desuppression (cf. Basile & Basile 1984). On the other
hand, simplified taxa at the base of subclades may be remnants of
ancient lineages now largely extinct. The latter seems a more
likely scenario because the possibility that more highly evolved
(morphologically reduced) taxa are more likely to survive recent
environmental change than coeval taxa retaining ancestral (nonreduced) features is more probable than multiple evolution or
even desuppression of the characteristic twisted Pottiaceae peristome. The presence of the character state change of loss of peristome in the shared ancestor of Anaectangium and the rest of the
Barbuleae (in both Cladograms 11 and 15) must be dealt with,
however, as it does not seem to be an artifact of the poor resolu-
tion of the remainder of the Barbuleae (cf. also Cladograms 2, 3,
4, 8, 9, 10).
Also, genera with complex morphology and no or few
reduced species (e.g. Tarte/la and Syntrichia) commonly occur
at the ends of lineages. These may be viewed as very recent
taxa, not having yet accumulated genetic diversity; apparently
large-statured genera of few or one species, such as Dolotortula,
Calyptapagan, Sagenotartula and Teniolaphara, are extremely
recently evolved and are just beginning evolutionary development. Some extant taxa at the middle or ends of evolutionary
lines commonly include many species and consist of plants of
both large stature and complex morphology and of smaller stature with simplified, reduced morphology, e.g. Bryaerythraphyllum, Hennediella, Pseudocrossidium, Tarte/la, Tartula and
Trichostamum (all as emended here). These genera are apparently evolutionarily more mature, having spread into habitats
that provide evolutionary pressures toward reduction and having
been in existence long enough to respond to such pressures dur~
ing speciation. Genera consisting only or largely of reduced
species, e.g. Anaectangium, Leptobarbula, Tetracoscinodon,
Tetrapterum, Trachycarpidium, Triquetrella and Weissia, are
found largely near the base of subclades or near the base of the
tree, and may be taken to be relicts of genera whose largestatured members of complex morphology have become extinct
with changing environments. Thus, among primitive genera,
presumed non-reduced species have been extinguished by evolutionary pressures before the reduced species of such genera,
leaving reduced relicts.
There are some clear exceptions to this that require explanation. For instance, large-statured taxa near the base of the tree,
e.g. Erythraphyllastrum, Erythrophyllapsis, and Gertrudiella,
may be taken to be large-statured relicts surviving in relictual
environments (i.e. Andean microhabitats). The position of, for
example, the Acaulon, Sarcaneurum and Stanea subclade seems
anomalous, but, perhaps, there the ancestral lineage consisting
of the austral genera Tetrapterum, Willia, Phascopsis and
Stonea is a line of relictual taxa that is actually separate from
the remainder of the Pottieae.
The data in general, however, is suggestive and future work
should examine whether or not the number of species and the
morphology of extant genera correlates with how deep in the
tree their immediate ancestors are, and on how the relative
harshness of relictual habitats affects phylogenetic position, the
less severe habitats preserving ancestral characters.
Geographic Trends
There appears to be considerable support for the evolutionary
hypothesis of Cladogram 16 from the geographic distributions
of the taxa given for the consensus tree of Cladogram 14, of
which there are no major mismatches. Although many of the
genera must be simply described as wide-ranging, certain distributional trends are correlated with phylogenetic groupings. The
basal-most branches are either of widespread or Andean distribution. The Merceyoideae and Pottioideae include a large number of cosmopolitan taxa, with the remainder largely restricted
to the Andes.
The members of the Chionolomoideae (see Cladogram 16)
are characteristic of lowland tropical areas, especially those of
eastern Asia. The Hyophileae (see Cladograms 8 and 14-16)
encompasses a majority of members with ranges in the lowland
EVOLUTIONARY RELATIONSHIPS
tropics, especially the West Indies. Certain highly evolved genera
(Aloine/la, Crossidium, Globuline/la, Pterygoneurum and Stegonia), considerably reduced in stature and generally grouped
together in the Hyophileae, are apparently monophyletic and
adapted for warm, highly arid habitats. Frey and Kiirschner (1983,
1988a) assigned a number of circum-Mediterranean, central Asian
and western USA Pottiaceae species to a "Circum-Tethyan" floristic zone, and indicated that these taxa may be Triassic relicts of
ancestral populations of areas around a mid-continental warm sea
formed during the breakup of Pangaea. The Hyophileae includes
several genera with species characteristic of the Circum-Tethyan
flora, and this tribe may have originated there.
The nine mainly Gondwanaland (austral) genera (Calymperastrum, Calyptopogon, Hypodontium, Phascopsis, Sarconeurum,
Stonea, Tetracoscinodon, Tetrapterum, Willia) are not grouped in
any of the subclades, excepting Phascopsis, Sarconeurum, Stonea
and Willia, which are clustered in the Pottioideae. This indicates
that the Gondwanaland ancestral group contributed to all traditional taxonomic groups and many of the presently recognized
subclades. Because Gondwana taxa occur at the base of the
Merceyoideae (see Cladogram 14 and 16), the Pottieae, and some
of the subclades making up the traditional Trichostomoideae, it is
probable that differentiation of these lineages occurred in austral
regions.
The Pottioideae, on the other hand, may have been derived
from a pantropical Hyophila-like ancestor, with austral genera
derived secondarily (cf especially Cladograms 2, 9 and 10) but do
include some austral taxa. On the other hand, the ancestral stock
contributing to the five austral genera (see Cladogram 16) of the
Pottieae may simply be extinct (certainly the nine austral genera
are reduced genera each of few species and mostly occur in marginal habitats).
The difficulty in resolving the Barbuleae may be due to continued survival of populations representative of ancestral stock,
which would add morphological variation that does not contribute
to analysis. It may be possible to better resolve the subclade by
eliminating from analysis all taxa of intermediate morphology
apparently surviving (on a geological time scale) in refugial areas.
This would have to be investigated at the species level, with due
attention to distinguishing if possible recent divergent evolution
in isolated habitats. Identification of the extant species with the
most primitive character states in each genus would be very helpful.
The most ancient stock of the Pottiaceae could be considered
Andean, as judged from the distribution of the taxa of the basal
branches, but the Andes are geologically rather recent. Possibly
the Andes have served as refugia for basal taxa (these usually well
distinguished morphologically) evolved elsewhere, probably
Gondwanaland.
Classification
Traditional classification systems presuppose that taxa can be
described as coherent morphological units that have some phylogenetic significance, most taxa sharing either at least one distinctive character state (monothetic) or possess most of a distinctive
set of character states (polythetic ). The taxonomic requirement is
that the taxa share at least to some degree these significant character states. If major subclades are considered suprageneric taxa,
however (as in Cladogram 16), such subclades are best defmed by
the character state changes at the basal ancestral node. Any shar-
31
ing of traits in the subsequently derived taxa of the subclade is
dependent on the degree of morphological divergence among
those taxa. In the present treatment, subclades named as suprageneric taxa will be identified by the character state changes at
the immediate ancestral node and will also be described morphologically to the best extent possible as though they were traditional grades. Taxonomists with some reservations about
using cladistics as a basis for classification can be reassured that
the subclades here formally recognized as subfamilies and tribes
are in most cases rather well distinguished morphologically, and
some largely match traditional groupings.
An evaluation of the cladograms indicates that, terminating
the tree, there are two highly evolved lineages of many taxa,
corresponding to two traditional subfamilies-the Merceyoideae (previously known as Barbuloideae) and the Pottioideae.
There are also several single or few-taxa lineages at the base of
the cladograms that are each given subfamily status. The
Trichostomoideae, as recognized in Cladogram 16, is an arbitrary construct from the many equally parsimonious trees contributing to the strict consensus tree of Cladogram 14, and its
structure is dependent on "reciprocal illumination."
Cladogram 16 is the basis for the classification since (1) it
provides a single tree as a hypothesis for evolutionary relationships, (2) it summarizes best the structure of Cladogram 14
while reflecting the structures of the majority of the rest of the
cladograms, and (3) because its functional outgroup, Timmie/la,
is selected largely based on Cladogram 10 with the outgroup
Ptychomitrium, a taxon that may be taken as the nearest littlemodified sister group of the Pottiaceae.
Additional Discussion
At the species level, lines of evolution involving reduction of
sporophyte and gametophyte characteristics in some cases can
be visualized as an exclamation mark(!), in which a line of species exhibiting features intermediate in a probable reduction
series terminates in a distinctive, highly reduced species or
group of species. The line may represent extant flotsam of evolution ranging from the peristomate to the eperistomate: why the
terminal group, often cleistocarpous, is commonly distinguishable at the supraspecific but seldom suprageneric level is not
presently understood but may be simply a function of general
strong modification of otherwise easily recognizable features
(e.g. of the leaf adapting to clasp the capsule). Further study is
necessary.
Examples of this reduction pattern include Weissia and its
well-characterized subgenus Phasconica; and, at the species
level, Syntrichia bartramii, a dwarf variant of S. ruralis from
the southwest of the United States, apparently trailing populations identifiable as S. intermedia in its evolutionary wake; and
Trichostomum caespitosum, distinguished from Trichostomum
crispulum by its small size and differentiated perichaetial
leaves.
Trichostomum sections may be examples of unconnected
evolutionary patterns. A cladistic evaluation of the relationships
would probably be more instructive in this case than sorting into
taxonomic categories on the basis of perceived gaps, as is done
here in the taxonomic section.
At the generic level, one might expect small-statured genera
to be at the end points of lineages of larger-statured genera, but
upon cladistic analysis (see above) this does not seem to be the
32
GENERA OF THE POTTIACEAE
case. Lineages often have apparently reduced genera at the base
of branches. Also, generalizing from the appearance of the taxa,
one might expect the following reduction pairs (the second genus
sharing ancestors with reduced species of the first genus): Tortella
and Calymperastrum; Quaesticula and Plaubelia; Barbula and
Scope/ophila; Tortula and Stegonia; Syntrichia and Stonea; Willia
and Phascopsis; Microbryum and Saitoella; but in fact, of these,
only Willia and Phascopsis (see Cladogram 14) show what might
be interpreted as reduction in series at the generic level. Clearly
what have been considered previously as minor characters are of
considerable importance as evolutionary markers. One can see,
however, a few pairs of reduced and non-reduced genera at the
end of lineages that might qualify as reduction series resulting in a
distinct genus: Bellibarbula and Gyroweisia (Cladogram 11),
Didymodon and Gymnostomiella, and Aloina and Aloinella, but
most pairs at the end of lineages are of nearly equal plant stature.
Directions of evolution may be inferred from the advanced
characters of end members of the subclades in Cladograms 11, 12
and 15. It is doubtful that genera evolve as a unit (unless one postulates sufficiently common reticulate evolution, c.f. Wyatt et a!.
1992). On the other hand, present morphology restricts and thus
may "guide" future evolutionary possibilities, i.e. evolution may
take place in species of similar morphology in similar habitats
undergoing similar environmental changes, resulting in a group of
species within the genus that has evolved similarly. Assuming
that a newly evolved genus has as its ancestor one species (or,
better, one mutated individual), not a whole genus, one might
expect that more than one extant species showing similar evolutionary directions within a genus might telegraph the next phase
in evolution at the generic level. Such is not the case.
Actually, it is not surprising that highly evolved genera are
difficult to imagine as simple extensions of the evolutionary
directions of other extant genera on the basis of easily observed
characteristics. This is because many of the most significant character state changes are anatomical as opposed to gross morphological traits.
Character weighting was employed to allow for the fact that
characters associated with possible reduction (or elaboration)
involving clearly related intrageneric taxa in transformation series
are almost certainly not globally homologous in a data set of this
size. The series are usually ascribed to reduction here, because
parallel loss or reduction is explainable with fewer assumptions
than parallel gain or elaboration.
Concentrated study of both poorly and densely speciated relatives was important to clearly distinguish states that represent
evolutionary steps. Use of unordered, multistate characters and
poorly studied characters may introduce false weighting if (1)
truly intermediate evolutionary steps are unobserved and thus
unscored or (2) certain state changes are not recognized as representing multiple synapomorphies. Character states in this study
are represented by those seen in densely speciated taxa and are
used globally on the assumption that homoplastic mutations recur
as similar states (this was observed in related densely speciated
taxa). This allows automatic weighting through use of multiple
ordered, additive states for groups in which much extinction has
taken place and intermediate states are absent. Ordered, additive
states are recognized as such by correlation with other states in
transformation series. FeatUres with similar functions that could
be treated as non-ordered states (e.g. propagula types) are here
considered non-homologous and are used as distinct characters
even when the presence of one feature appears to exclude the
others.
COMPARISON WITH THE
CLASSIFICATIONS OF PREVIOUS AUTHORS
Three authors have provided phylogenetic trees of the Pottiaceae or portions of the Pottiaceae. These may be compared
with the various cladograms developed in this study, especially
Cladograms 2, 8 and 10 as well as Cladogram 16. The similar
subclade structure of many of the cladograms developed in the
present study is given in Cladogram 16, which is a single tree
selected from many equally parsimonious trees of Cladogram
14 by FIG/FOG analysis and "reciprocal illumination."
Saito's (1975a) phylogeny (Cladogram 17) is the closest to
that of the cladograms generated in this study (assuming the
classical Trichostomoideae is a paraphyletic group). Chen
(1941; Cladogram 18) recognized the Weissia-Hyophila group
as a distinct suprageneric group but included only Hyophila and
Uleobryum in it. The Tortella group is separated by Chen from
Trichostomum at the tribal level; it is part of a lineage separate
from Trichostomum in several cladograms in the present study.
Hilpert (1933) (Cladogram 19) clearly places the Leptodontium
group near the Barbula lineage. Although there are some significant comparisons, previous authors' family trees are relatively
simple at the suprageneric level, and mostly different in structure from that recognized here.
Some authors have presented classifications without actually describing a tree. Brotherus (1924-25) divided the Pottiaceae into the Pleuroweisioideae, Cinclidotoideae, Trichostomoideae, Merceyoideae, and Pottioideae. In his treatment of the
geography of moss distributions, Herzog (1926, p. 95) recognized the Trichostomaceae and Pottiaceae as separate families.
He split the Trichostomaceae into the Pleuroweisioideae and the
Trichostomeae, the latter included the Barbuleae, Hyophileae
(Hyophi/a, Weisiopsis, and Dialytrichia), and Leptodontieae.
Walther's (1983) classification recognized the Trichostomeae
(Tortelleae, Barbuleae, Hyophileae, Eucladieae, Pleuroweisieae), Pottioideae (Pottieae, Scopelophileae), Cinclidotoideae
(Clinclidoteae), and Leptodontioideae (Leptodontieae), a classification similar to that of Saito (1975a) but the suprageneric
taxa were constituted of genera in considerably different combinations.
EVALUATION
The present phylogenetic hypothesis is in many respects at odds
with past evaluations of phylogenetic relationship and systematic position at the suprageneric level. Because of a lack of resolution and possible inaccuracies caused by abundant homoplasy,
inherent problems with the branch-swapping algorithm, and the
immaturity of cladistic methodology, the systematic classification used in the present treatment of the Pottiaceae must be seen
as reflecting only the congruent results of the several analyses
of the data set in light of these conditions.
It is here considered that cladistic analysis is the best
method at this time for organizing the information from this
large data set into the basis for a supraspecific classification.
Many lineages are identifiable with considerable certainty in the
present work (discussion of Cladogram 14), especially involving genera with one or few species.
33
Cladogram 1
Data set includes 9 non-p_ottiaceous genera in addition to the outgroup, Polytrichum. This is a strict consensus tree. All characters are weighted
the same. Tree length 557, consistency index .17, retention index .56, number of equally parsimonious trees is more than 1115. Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram 16.
POLYTRICHUM
Aloina
Timmiella
TIMMIA
r~throphyllopsis
B-Ecythrophyllastrum
Gertrudiella
_rChionoloma
~Pseudosymblepharis
Pachyneurops1s
Trichostomum
alynmerastrurn
DITRICHUM
PfYCHOMlTRIUM
HVPOdontium
:.pJeurochaete
Tortella
alyp~opogon
Tuerckhelffiia
Hyme~o~tyliella
WeiSSia
Weissiodicranum
HyQPhila
Teniolophora
Plaubeha
Tetracoscinodon
Triquetrella
ERATODON
Leptodontium
Leptodontiella
Trachyodontium
Streptotrichum
Dia}ytnchia
Rh ophyllum
rroma
Bryoerythrophyllum
Pseudocrossldium
Anoectangium
Bellibarbula
yroweisia
Hymenostylium
Reimersia
Str~topogon
Sco loP.hila
RIMMIA
SYRRHOPODON
Eucladium
Tetrapterum
M1crobryum
Trachycarpidium
Pterygoneurum
Stegonia
Phascopsis
Acaulon
Stonea
Aschisma
Bryoceuthospora
Uleobryum
Streptocalypta
uaesticula
Weisiopsis
angufeea
Luis1erella
Tortula
lobulinella
Aloinella
rossidium
BRYOBARTRAMIA
Saitoella
DIPHYSCIUM
ENCALYPTA
Leptobarbula
Didymodon
Barbula
ymnostomum
uymnostomiella
Molendoa
Sarconeurum
Willia
henia
Syntrichia
Sagenotortula
'Hilpertia
Dolotortula
rumia
Hennediella
34
Cladograml
Data set includes 9 non-pottiaceous genera in addition to the outgroup, Polytrichum. This is a strict consensus tree; 22 reduction-related characters are weighted at 1: 15. Tree lengtli 6024, consistency index .17, retention index .57, number of equally parsimonious trees is more than 1115.
Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram 16.
POLYTRICHUM
TIMMIA
Gertrudiella
A Timmiella
DITRICHUM
PTYCHOMITRIUM
Eucladium
Trichostomum
alym_perastrum
Pleurochaete
al}'l1topogon
Tortella
Hypodontium
Tuerckheimia
Trachyc!UJlidium
Aschisma
Uleobryum
Bryoceuthospora
Stre t~illypta
10noloma
Pseudosymblepharis
Pachyneurops1s
Tetral'terum
Pseudocrossidium
Bryoerythrophyllum
~throphyllopsis
~throphyllastrum
Tetracoscmodon
Dial "chia
RATODON
Mironia
.RhexoEhyllum
Tna~:~r~a
Hymenostylium
Leptodont!ella
Leptodontium
Trachyodontium
Streptotrichum
Anoectangium
Bellibarbula
yroweisia
Barbula
uymnostomum
Molendoa
Did modon
ymnostomiella
Str~topogon
s-co loP.hila
RIMMIA
SYRRHOPODON
35
3· Th' ·
· 1udes 9 non-pottlaceous
·
· add'1t1on
· to the outgroup,
CladogTr~m
· consensus tree; 22 re d uctwn-re
·
1ated ch aracters
D ata set me
~enera m
1mm1a.
1s IS a stnct
are weighted at 1:15. Tree length 60~4. consistency index .17, retention index .57, number of equally parsimonious trees is more than 1115. Lettered ancestral nodes denote lmeages that are similar to the named lineages of Cladogram 16.
TIMMIA
POLYTRICHUM
A Timmiella
Gertrudiella
,-Erythrophyllopsis
B-Erythrophyllastrum
Pachyneuropsis
Pseudosymolepharis
Trichostomum
Eucladium
al_ylllj)erastrum
PTYCHOMITRIUM
DITRICHUM
hionoloma
Streptocalypta
Pleurochaete
Tortella
alyptopogon
Leptobarbula
Molendoa
Bryoerythrophyllum
Pseudocross1d1Um
Hy()phila
Plaubelia
Teniolophora
ymnostomum
Barbula
Did)'modon
Tetracoscinooon
Anoectangium
Bellibarbula
yroweisia
Dial trichia
RATOOON
R~exophyllum
Mrroma
Triquetrella
Leptodontiella
Leptodontium
Trachyodontium
Streptotrichum
Hyme.nosty~ium
Re1mers1a
Strepto o~IA
SYRRHOPODON
Scopelophill!
ymnostom1ella
36
~
Cladogram 4
Data set includes 9 non-pottiaceous genera in addition to the outgroup, Ptychomitrium. This is a strict consensus tree; 22 reduction-related characters are weighted at 1:15. Tree length 6037, consistency index .17, retention index .57, number of equally parsimonious trees is more than
1115. Lettere<f ancestral nodes denotelineages that are similar to the named lineages of Cladogram 16.
Hy~e!lostyliella
-WeJSSia
Weissiodicranum
Hyophila
Plaubelia
Teniolophora
uaesticula
Weisiopsis
angufeea
Luis1erella
rossidium
Stegonia
Jobulinella
Aloinella
Pte!Y_goneurum
BRYOBARTRAMIA
Tortula
37
Cladogram 5
Data set includes 9 non-pottiaceous genera in addition to the outgroup, Enca/ypta. This is a strict consensus tree; 22 reduction-related characters
are weighted at 1:15. Tree length 6033, consistency index .17, retention index .57, number of equally parsimonious trees is more than 1115. Lettered ancestral nodes denote lmeages that are similar to the named lineages of Cladogram 16.
ENCALYPTA
DIPHYSCIUM
Saitoella
rumia
Hennediella
Dolotortula
Microbryum
Willia
PhascQPsis
Stonea
Sarconeurum
Acaulon
Hilpertia
Sagenotortula
S ntrichia
enia
Tortula
BRYOBARTRAMIA
Pterygoneurum
Aloinella
lobulinella
rossidium
Stegonia
Luisierella
Weisiopsis
angufeea
uaesticula
Hymenostyliella
Weissiodicranum
Weissia
HyQPhila
Teniolophora
Plaubeha
Leptobarbula
Tuerckheimia
alymperastrum
Trichostomum
Eucladium
Hyoodontium
Pleurochaete
alYP.topogon
Tortella
Trachyc!Ul'idium
Aschisma
B!)'oceuthospora
Uleobryum
Strc:ptocalypta
Pachyneuropsis
hionoloma
Pseudos~blepharis
Erythropllyllopsis
~tfirophyllastrum
Tetrapterum
TIMMIA
ertrudiella
Timmiella
PTYCHOMITRIUM
POLYTRICHUM
Aloina
H
38
Cladogram6
Data set includes 9 non-pottiaceous genera in addition to the outgroup, Vrimmia. This is a strict consensus tree; 22 reduction-related characters
are wei~hted at 1:15. Tree length 6034 1 consistency index .17, retention index .57, number of equally parsimonious trees is more than 1115. The
lettered~ancestral node is of a fmeage s1milar to a named lineage of Cladogram 16.
39
Cladogram7
Data set restricted to rottiaceous genera except for the outgroup, Polytrichum. This is a strict consensus tree; all characters are weighted the
same. Tree length 49 , consistency index .19, retention index .57, number of equally parsimonious trees is more than 1241. Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram r6.
POLYTRICHUM
A Timmiella
Aloina
Erythrophyllopsis
Gertrudiella
Erythrophyllastrum
Tuerckheimia
Hypodontium
Tetracoscinodon
yroweisia
Pseudosymblepharis
hionoloma
Trachycarpidium
Bellibarbula
Eucladium
Pachyneuropsis
uaesticula
P eurochaete
Stegonia
Streptocalypta
Leptobarbula
Pterygoneurum
Anoectangium
Trichostomum
Microbryum
Tetrapterum
alymperastrum
Tortella
alyptopogon
anguleea
Weisiopsis
Luisierella
Phascopsis
Stonea
Acaulon
Aschisma
Uleobryum
Bryoceuthospora
Saitoella
lobulinella
Tortula
rossidium
Aloinella
Didb::i~don
mnostomum
copelophila
ymnostomiella
Molendoa
Hymenostyliella
Weissiodicranum
Weissia
Hyophila
Plaubelia
Teniolophora
Tri~~i~~~~a
Hymenostylium
Trachyodontium
Streptotrichum
Leptodontiella
Leptodontium
Dialytrichia
ophyllum
roma
Bryoerythrophyllum
Pseudocross1dium
Sarconeurum
Willi a
Syntrichia
henia
Sagenotortula
-Streptopogon
Hilpert! a
Dolotortula
rumia
Hennediella
40
Cladogram8
Data set restricted to pottiaceous genera except for the outgroup, Polytrichum. This is a strict consensus tree; 22 reduction-related characters are weighted at 1:15. Tree length 5375, consistency index .19, retention index .57, number of equally parsimonious trees is more than
1241. Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram 16.
POL YTRICHUM
Aloina
A Timmiella
Erythrophyllopsis
Erythrophy llastrum
Gertrudiella
_r-Chionoloma
t___c:Pachyneuropsis
Pseudosymolepharis
Trichostomum
-Eucladium
alymperastrum
Pleurochaete
Tortella
alyptopogon
Tetracoscinodon
Triquetrella
Hymenostylium
Reimersia
Leptodontiella
Leptodontium
Trachyodontium
Streptotrichum
Anoectangium
Bellibarbula
yroweisia
uymnostomum
Molendoa
Scopelophila
ymnostomiella
Streptopogon
Baroula
Didymodon
Dialytrichia
ophyllum
rom a
Bryoerythrophyllum
Pseudocross1dium
Weissia
Hymenostyliella
HypodontiUm
Plaubelia
Teniolophora
Hyophila
Weissiodicranum
uaesticula
Weisiopsis
angufeea
Luis1erella
c-Leptobarbula
Tetrapterum
Tuerckheimia
Streptocalypta
Trachycarp1dium
Aschisma
Uleobryum
Bryoceuthospora
Iobulinella
Aloinella
Pte goneurum
rossidium
Stegonia
Tortula
Saitoella
41
Cladogram9
Data set restricted to pottiaceous genera except for the outgroup, Timmia. This is a strict consensus tree; 22 reduction-related characters are
weighted at 1:15. Tree length 5290, consistency index .19, retention index .58, number of equally parsimonious trees is more than 1241.
Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram 1o.
TIMMIA
Timmiella
ertrudiella
Erythrophy Hops is
Tetrapterum
Erytlfrophyllastrum
Pseudosymblepharis
Pachyneurops1s
hionoloma
Streptocalypta
Trichostomum
alymperastrum
Eucladium
Tuerckheimia
Trachycal'idium
Aschisma
Bryoceuthospora
Uleobryum
Hypodontium
Pleurochaete
alyptopogon
Tortella
Tetracoscinodon
Anoectangium
yroweisia
Bellibarbula
Streptopogon
Molendoa
Barbula
ymnostomum
ymnostomiella
Scopelophila
Didymodon
Triquetrella
Hymenostylium
Reimersia
Trachyodontium
Streptotrichum
Leptodontiella
Leptodontium
Dialytrichia
ophyllum
rom a
Pseudocrossidium
Bryoerythrophyllum
Leptobarbula
-Hymenostyliella
Weissiodicranum
Weissia
Hyophila
Plaubelia
Teniolophora
uaesticula
anguleea
Weisiopsis
Luisierella
Ste onia
rossidium
lobulinella
Aloina
Aloinella
Pterygoneurum
Tortula
Saitoella
42
Cladogram 10
Data set restricted to pottiaceous genera except for the outgroup, Ptychomitrium. This is a strict consensus tree; 22 reduction-related characters are weighted at 1:15. Tree length 5341, consistency mdex 9, retention index .57, number of equally parsimonious trees is more than
1241. Lettered ancestral nodes denote lineages that are similar to the named lineages of Cladogram 16.
PTYCHOMITRIUM
A Timmiella
Tetrapterum
ertrudiella
Eucladium
alymperastrum
Trichostomum
rErythrophyllopsis
B-Erythrophyllastrum
Hypodontium
Pleurochaete
aly(>topogon
Tortella
Tuerckheimia
Trachycarpidium
Aschisma
Bryoceuthospora
Uleobryum
Stre tocalypta
hionoloma
Pseudosymblepharis
Pachyneurops1s
H Tetracoscinodon
Anoectangium
yroweisia
Bellibarbula
Barbula
ymnostomiella
Streptopogon
Didymodon
Molendoa
Scopelophila
ymnostomum
Tri~~?Z~~~a
Hymenostylium
Trachyodontium
Streptotrichum
Leptodontiella
Leptodontium
Dialytrichia
Rhexophyllum
Miroma
Bryoerythrophyllum
Pseudocross1dium
Lepto~ar~ula
WeiSSia
Weissiodicranum
Hymenostyliella
Hyophila
Plaubelia
Teniolophora
uaesticula
anguleea
Weisiopsis
Luisierella
Ste onia
rossidium
lobulinella
Aloina
Aloinella
Pterygoneurum
Tortula
Saitoella
43
Cladogram 11
Data set restricted to genera in Merceyoideae "F" subclade of Cladograms 9 and 10; outgroup is Tetracoscinodon. Tree 24 of 127 equally
parsimonious trees, 22 reduction-related characters are weighted at 1:15. Tree length 1494, consistency index .48, retention index .62.
Character state changes shown for ancestral nodes. "X" denotes a node for which character state changes do not map, implying an ancestral
multiple-branched node.
TETRACOSCINODON
Triquetrella
2:0-f
Hymenostylium
54:1--0 64:1--0 65:1--0
Reimersia
29:1--0 39:1--043:1--0
Leptodontiella
46:0-1 48 :0-1 50:0-1
Leptodontium
5:0-1 15:0-1 25:0-1 30:1--0 63:1--0
Trachyodontium
11:1- 215:1-244:1-2
Streptotrium
Dialytnchia
33:0-1 66:1-2 68:0-1
Rhexophyllum
73:0-2
Mironia
24:0-1 46:0-1
Bryoerythrophyllum
23:1--0 48:0-I 50:0-1
Pseudocrossidium
3:0-1
Anoectangium
28:1--0 65 :1--0
Bellibarbula
8:1--0
44
Cladogram 12
Data set restricted to genera in Pottioideae "G" subclade of Cladograms 9 and 10, outgroup is Leptobarbula. Tree 5 of 8 equally parsimonious trees, 22 reduction-related characters are weighted at 1:15. Tree length 2247, consistency index .37, retention index .59. Character state
changes shown for ancestral nodes.
rumia
1:0--1 8:0-1R 17:0--123:0--1
36:0-1R 52:1--0R 68:0--1
Hennediella
36:1-2
Dolotortula
45
Cladogram 13
Data set restricted to pottiaceous genera. The functional outgroup is Timmiella. This is a strict consensus tree; all characters are weighted
the same. Tree length 485, consistency index .20, retention index .57, number of equally parsimonious trees is more than 1259. The lettered
ancestral nodes are of lineages identical to the named lineages of the single tree of Cladogram 16, on which the present classification is
based.
,
TIMMIELLA
ertrudiella
Trichostomum
Pleurochaete
Pachyneuropsis
Hypodontium
alymperastrum
hionoloma
Pseudosymblepharis
rErythrophyllastrum
B-Erythrophyllopsis
alyptopogon
Tortella
Tuerckheimia
Hymenostyliella
Molendoa
Weissiodicranum
Weissia
Hyophila
Plaubelia
Teniolophora
Tetracoscinodon
Anoectangium
Bellibarbula
yroweisia
Tri~~iZ~~~a
Hymenostylium
LeptodontJUm
Leptodontiella
Trachyodontium
Streptotrichum
Dialytrich1a
ophyllum
rroma
Bryoerythrophyllum
Pseudocross1dium
Stegonia
Streptocalypta
Microbryum
Leptobarbula
Eucladium
Tetrapterum
Tracliycarpidium
Pterygoneurum
Phascopsis
Acaulon
Stonea
Aschisma
Bryoceuthospora
Uleobryum
Barbula
Didymodon
ymnostomum
ymnostomiella
s.copelophila
uaestlcula
Weisiopsis
angufeea
Luis1erella
lobulinella
Tortula
Saitoella
rossidium
Aloinella
Aloina
Sarconeurum
Willi a
Syntrichia
henia
Sagenotortula
Streptopogon
Hilpert! a
Dolotortula
Hennediella
46
Cladogram 14
Data set restricted to pottiaceous genera. The functional outgroup is Timmie/la. This is a strict consensus tree; 22 reduction-related characters are weighted at 1:15. Tree length is 5270, consistency index .19, retention index .57, number of equally parsimonious trees is more
than 1250. Lettered ancestral nodes denote lineages that are identical to the named lineages of the single tree of Cladogram 16, on which
the present classification is based. Geographic ranges ("*" denotes austral taxa) are given.
A TIMMIELLA ............................................................................................................ Broad distr.
~Erythrophyllopsis
.............................................................................................................. Andes
-Eryi,~~~~ll~~~:::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: ~~~~
_r-Chionoloma ............................................................................................ S.E. Asia, Borneo
~Pseudosymblepharis .......................................................................................... Tropics
Pachyneurops1s ............................................................................................Philippines
Eucladium ....................................................................... N. Temp. Zone, India, S. Africa
alymperastrum ................................................................................................. *Australia
Trichostomum .................................................................................................. Broad distr.
Tuerckheimia ............................................................................ Neotropics, E. Asia
Streptocalypta................................................................... Mexico, Andes, S. Africa
Pleurochaete ........................................................................................... Broad distr.
alyiJtopogon ............................................... *Southern S. America, Australasia
Tortella .............................................................................................. Broad distr.
Tetracoscinodon ........................................................................... *New Zealand
Dialytrichia ................................................... Europe, N. Africa, S. & E. Asia
ophyllum ............................................... Southw. N. America, Andes
rroma .................................................... Mexico south through Andes
Bryoerythrophyllum ....................................................... Broad distr.
Pseudocross1dium .......................................................... Broad distr.
Anoectan~ium .......................................................................... Broad distr.
yroweis1a ............................................................................... Broad distr.
Bellibarbula.................................................. USA, Mexico, E. Asia, India
Barbula ................................................................................ Broad distr.
ymnostomiella .................................................................. Pantropical
Streptopogon ............................................................................. Tropics
Didymodon ......................................................................... Broad distr.
Scopelophila ........................................................................ Broad distr.
ymnostomum .................................................................... Broad distr.
Triquetrella.........................................*Mediter. climates, austral areas
Reimersia ....................................................................E. Asia, India
Hymenostylium .............................................................. Broad distr.
Trachyodontium ..................................................................... Andes
Streptotrichum ........................................................................ Andes
Leptodontiella ........................................................................ Andes
Leptodontium ................................................................. Broad distr.
Hypodontium .................................................................................. *S. Africa
Hymenostyliella ......................................................... Brazil, Philip., India
Molendoa ............................................................................ Broad distr.
Hyophila ......................................................................... Broad distr.
Plaubelia ........................................ Neotropics, S. Africa, Burma
Teniolophora ................................................................ W. Indies
Weissia .....................................................................................Broad distr.
Weissiodicranum ..................................................... W. Indies, Galapagos
uaesticula ................................................................................. W. Indies
anguleea......................................................................Brazil, India
Weisiopsis ............................................................................ Tropics
Luisierella.......................................................... Neotropics, E. Asia
Ste onia ........................................................ N. Temperate Zone
rossidium ............................................................Broad distr.
Pte goneurum ................................................. Broad distr.
lobulinella........... Southw. USA south through Andes
Aloina ..................................................... Broad distr.
Aloinella ........................................... Mexico, Andes
Leptobarbula ...................................................................... Mediterranean region
·Tetrapterum .............................................................................. *Austral areas
Trachycarpidium ........................................... S. America, Africa, Oceania
Aschisma ............................................................... N. America, Europe
Bryoceuthospora .....................................................Mexico, Angola
Uleobryum ..................................... Cent. and S. America, Australia
Tortula ................................................................................. Broad distr.
Willia .....................................:................................. *Austral islands
Saitoella .......................................................... Mexico, Andes
Microbryum .......................................................... Broad distr.
rumia ...............................................Western N. America
Hennediella .................................................Broad distr.
Dolotortula ...................... Mexico south through Andes
Phascopsis............................................................*Australasia
Stonea............................................................ *Australia
Acaulon .................................................. Broad distr.
Sarconeurum ...... *Southern S. America, Antarctica
henia .................................................... Broad distr.
Syntrichia ............................................... Broad distr.
Hilpertia ..................... Western N. America, Europe
Sagenotortula ................................... Mexico, Andes
47
Cladogram 15
Data set restricted to pottiaceous genera. The functional outgroup is Timmie/la. This is tree 1126 of more than 1250 equally parsimonious
trees, 22 reduction-related characters are weighted at 1:15. Tree fength is 5270, consistency index .19, retention index .57. Th1s trees shows
ancestral character state changes on which Cladogram 16 is based. "X" denotes a node for which character state changes do not map,
implying an ancestral multiple-branched node.
TIMMIELLA
Erythrophyllopsis
31:2-1
Erythrophyllastrum
ertrudiella
21:2-1 40:1-0
hionoloma
10:1-0 27:0-1
Pseudosymblepharis
21:1-2R 45:1-2
Pachyneuropsis
33:1-0
alymperastrum
6:1-0 30:1-0
Eucladium
55:1-0
Trichostomum
Tuerckheimia
4:1-0
Streptocalypta
45:1-2 68:0-1
Pleurochaete
11:1-2R 18:0-1 54:0-1 57:0-1R 66:1-2
alyptopogon
46:0-I
TorteII a
11:2-1 24:3-1 31:2-1 57:1-0
Tetracoscinodon
12:0-113:0-1
Dialytrichia
25:0-1 33:0-1R 57:0-1R 66:1-2 68:0-1
Rhexophyllum
21:1-2R 73:0-2
Mironia
24:0-1R 46:0-1
Bryoerythrophyllum
23:1-0R 48:0::..150:0-1
Pseudocrossidium
14:2-3 23:0-1 24:1-0
Triquetrella
2:0-f
Reimersia
54:1-0R65:1-0
Hymenostylium
3:1-0 29:1-0 39:1-0
Trachyodontium
11:1-2R 15:1-2 21:1-2R 44:1-2 57:0-1R
Streptotrichum
5:0-1R 15:0-1 24:0-1R 25:0-1
30:1-0 63:1-0 72:0-1
Leptodontiella
46:0-1 48:0-1 50:0-1
Leptodontium
43:1-055:1-0
Anoectangium
28:1-0 65:1-0
5:1-0 54:0-1
48
4:1--0 21:1--0 55:1--0
Leptobarbula
1:1--0 66:1-2 68:0--1
Tetrapterum
33:0--lR 56:0--1 72:0--1
Trachycarpidium
23:0--1 52:0--1 58:2-1 59:1--0
Aschisma
11:1--0 29:1--0
Bryoceuthospora
60:0--161:1--074:0--1
Uleobryum
30:1--0 54: l-OR 71:1--0
Tortula
10: 1-2 26:1--0 28:1--0
Willi a
73:0--2
Saitoella
14:2-3
Microbryum
30:0--lR 36:0--1
rumia
1:0--lR 8:0--lR 17:0--118:0--1
23:0--1 44:1-2 57:0--lR
Hennediella
36:1-2
Dolotortula
49
Cladogram 16
Phylogenetic arrangement of the Pottiaceae in the present study. Same as Cladogram 15 but without character state changes indicated.
Subclades recognized as suprageneric taxa are indicated by letters at ancestral nodes: "A" Timmielloideae, "B" ErythroiJhyllopsoideae, "C"
Gertrudielloideae, " D" Chiono1omoideae, "E" Trichostomoideae, "F" Merceyoideae, "G" (and all its ancestors in the family) Pottioideae,
"H" Tetracoscinodontieae, "I" Bryoerythrophylleae, "J" Leptodontieae, "K" Barbuleae, "L" Hyophileae, "M" Pottieae.
A TIMMIELLA
rErythrophyllopsis
B-Erythrophyllastrum
Gertrudiella
,e-~ionoloma
t.__c:Pseudosymblepharis
Pachyneuropsts
alJmper~strum
EucladJUm
Trichostomum
Tuerckheimia
Streptocalypta
Pleurochaete
alyptopogon
Tortella
Tetracoscinodon
Dialytrichia
ophyllum
roma
Bryoerythrophyllum
Pseudocrosstdium
Triqu~trell~
Retmersta
Hymenostylium
Trachyodontium
Streptotrichum
Leptodontiella
Leptodontium
Anoectangium
yroweisia
Bellibarbula
Streptopogon
Barbula
50
Cladogram 17
Phylogenetic arrangement of the Pottiaceae by Saito (1975).
[ ~<hooomoid<~
Barbuleae
Eucladieae
Leptodontieae
Pottieae
]Pottioid=
Cladogram 18
Phylogenetic arrangement of the Pottiaceae by Chen (1941).
Pleuroweisieae
Eucladieae
Trichostomeae
Tortelleae
inclidotoideae
Barbuleae
Hyophileae
Leptod?ntioideae
:Pott1eae
Merceyeae
l
Eucladioideae
Trichostomoideae
) Barbuloideae
) Pottioideae
Cladogram 19
Phylogenetic arrangement of the Pottiaceae by Hilpert (1933).
Leptodontioideae
Barbuloideae
ttiaceae
nclidotaceae
) Trichostomaceae
DATASET
This data set contains morphological information on the 76 genera of the Pottiaceae, and on ten (fully capitalized) non-pottiaceous _genera
used as out_groups in analysis. A dash ("-") indicates a character that is variable in state or otherwise "unknown, undefined, or missing"
(Farris 1988). There are 86 taxa and 75 characters; the first character is numbered as character "0".
0
1
2
3
4
5
6
7
012345678901234567890123456789012345678901234567890123456789012345678901234
POLYTRICHUM
1101100010120012000102103110111201000000-00--10-----0001012101110-----12020
BRYOBARTRAMIA 100-000100-0002000000110100-01-10-0000000011110-----101000100111-0----01102
CERATODON
11010011-011113100010111111011110100-00-000--00-----001-0121011-11110-0--00
DlPHYSCIUM
10000001002-002-0-0001-0201011120000000--0---10------0101121011-11-20-01021
DITRICHUM
1-01100-000-0-2-0-010--1-110111100000100000--00------0-1012101111112-101000
ENCALYPTA
0-0100--002-00-0000000-0200-11-20-00-001001111-01010-00-2-210111-1120-----2
GRIMMIA
110-00010011-1-00-0101-1000-00010-000000-00----01110-0--1-210111---20-0100PTYCHOMITIUM
0101-1-000120022010111101010111100000000100--10-----10000121011111120-01--1
SYRRHOPODON
1100000-001---2-0--10-1-1010---10000000-00--11100110100-1121011101020--1000
TIMMIA
1101001-101200-200010210301011120100000200-1110------0010121011111-20-02-20
Acaulon
000000-1001-002-000100-1000-01110-100-010--0-0000000-00010000111------00121
Aloina
-00-000-00-2001-00-012-0310-11120101200-110--1000000-00-0121011111121101020
Aloinella
1101001000-0001-00-01000210-11-1010100010--111000000001-0-21011111-20-01100
Anoectangium
11-110-1101-11-000010-110-0-0 - 110000000--0--1100000001110021011110----01000
Aschisma
-00100100010002-00-100100010-0010-00000-001112000000100010100100------10100
Barbula
1-011--1101--1--000-0-110010-1110-00000100--1111101100--00210111-1221101000
Bellibarbula
11011011001-0130000101--00100111000000--001-1000000000100021011110----01120
Bryoceuthospora
000-0001101-002200010001101000010100000100100-000000-01010101011-------0101
Bryoerythrophyllum 1-0111-1101-----00010--0101011-10-000000-01111101010-0--0-2101111--2-101020
Calymperastrum
10-11101102-0020010102101010110101ooooo1001121oooooo---------------------oCalyptopogon
110011-11012--2-00110-11ooo-110101000011001-12100101-01001210111112211110-o
Chenia
100110111-210-3-000-0---000-011101000001000--1110000000-----01111-221101-2Chionoloma
110101--100200-100110110301111020000000-001112000000---------------------oCrossidium
-00100-100-000--000--1-0100-01-1010110--0--111000000-0--0-21011111121101000
Crumia
110-001110220030011-00-1100-011100001001001-21000000000-0121011111121101-10
Dialytrichia
11011011101-1130010-0-11011011-101000001001111000000000-0121011101221101000
Didymodon
11-110111o---o3-o-o-o---o----111oooooo-1-01-1-11111ooo--0 - 210111-1---101o-o
Dolotortula
11010001112-0030011000-1000-011101002001000--0000000000101210111112211--02Erythrophyllastrum
1101-1111011-0200-0102103010011101000001-0111100000000110121011111020-01020
Erythrophyllopsis
11-101111012-020000102102110111101000001101111000000000-0121011111-20---02Eucladium
1-00-10110-1002110010-1-101011-1000000-1001-1100000000001021011101120-01000
Ganguleea
1o-o-ooo1121001ooooooo-oooo-0111ooooooo2ooo--1oooooo11100021011110------ooGertrudiella
110111011012003-ooo1o1-0300-111211ooooo2o0-011ooooooo--------------------1-
51
Globulinella
Gymnostomiella
Gymnostomum
Gyroweisia
Hennediella
Hilpertia
Hymenostyliella
Hymenostylium
Hyophila
Hypodontium
Leptobarbula
Leptodontiella
Leptodontium
Luisierella
Microbryum
Mironia
Molendoa
Pachyneuropsis
Phascopsis
Plaubelia
Pleurochaete
Pseudocrossidium
Pseudosymblepharis
Pterygoneurum
Quaesticula
Reimersia
Rhexophyllum
Sagenotortula
Saitoella
Sarconeurum
Scopelophila
Stegonia
Stonea
Streptocalypta
Streptopogon
Streptotrichum
Syntrichia
Teniolophora
Tetracoscinodon
Tetrapterum
Timmiella
Tortella
Tortula
Trachycarpidium
Trachyodontium
Trichostomum
Triquetrella
Tuerckheimia
Uleobryum
Weisiopsis
Weissia
Weissiodicranum
Willi a
-0010--100-000-00000100010--0111011000--000--110000000000-21011111120-01000
1001-0111020002-000-00-1000-00110000100100101110111000100021011100----010-0
11-10-111o1ooo-oooo1o-o-oo10-1-1ooooooo-oo1-111ooooooooooo210111-o----o-ooo
100-101100-00--0000-0--110--01-10000000--01-11110000-0-00-2101111-020-0-0-0
1-01---110--002-01110-1-100-01-101002000001-21000000-00-2-2101111--2110--20
1-01-0-1001-003-01010021000-010101002--101-01011000010100-21011111221101-20
110110100012001-0001001-1010111100000012000--100000001100-21011100----01000
11-0101110--11-oooo10-11001ooo-1ooooo--ooo1011oooooooooooo21011100----ooooo
1--11-001-2100--000-00-0101011110-00000-00-011101011-0000-21011110----01-00
11010110101-00---01-0210201-11020000000200110200000000102121011101100-11100
1001-01110100020000100-0011011110000000000111100000000100-21011111221101000
1100111110111131000101-11110100100000000001-1-101010001000210110111100011011001--1101-113-00010--111101001000000--001-11101110-01-0121011-111-0-0-100
000000101---00-0000-01001-0-100100000002000--20000001000012101111--20-010-0
000--0-1002-003-000100--100-011101001101001-1-000000100020--01110--20--0120
11011011101-113-0101021-111011110100000000111110000000110-21011111221101020
11-11-0-10--00----010---1-1011110000000-00111110101001100021011100----01000
110100111002002001010210301-1112ooooooo1001112oooooo------------~--------o-
10010-11oo-ooo2oooo-oo1-100-0101o-ooooo1001-111ooooooooo1o-oo111------00122
10011010112-001-000-00-010-001110100000200-00-10100000-00121011111120-0-000
110111011012002100110-102-1011-10000000-001112000000011101210111112211020-0
1-01--01101---30000-0--011-011110101000100111110111000--0121011111-211020-0
1101-1-110--00-000010210301111-20000000-00111200000000--012101111--20-01000
-00100-1001-001-000---20100-01110100000-00-01-00000010002-2101110-020-0-1010010000101100100000-000101001110000000-001001000000000-0021011111221101000
011010-1101-1120000102110010101100000-10000--100000000000021011100----01000
110111-11011113200010211011010110-00000110111100000000110121011110----01020
11010001002-0020000100--000-110101003001000--111000000010-21011111221102020
1001000100-000300110100-100-01-0-0100001001111000000---------------------21 - 0-0-11101-002000010-1000-0010101001000001-11100000---------------------211oooo-11021-1--ooo1o---ooo---110oooooo-ooo--110101oooooo-210111-o----ooooo
00010011001100--000-00-1100-01-101101101010--1000000100-0 - --01111--20-0-100
000000110020002-000110-0000-01-1001000010010111000000--------------------2 10010--110--002-00-----010-01102-000000-001112000000-000002101111-221101000
1-0010-1001--03-0------1000-0--1000021-1000--0100100-0-0012101110--211-1--1
1100111110121132000102-1111010010000000000102100000000101121011011120-01100
110-0--1102-----o-o----oooo--101--oooo-1-01-111101oo-oo12-210111112211o-o2o
110110101021oo- - 011-0010201011110oooooo2oo-oo1101010ooo------------------o110010-10001112000010-1-101011110oooooo1001111oooooooo1100210111111-0-o1100
1001001110110020000101102-1011110100000-001111000000101011200101------011-0
1-0111-011120022000102103-10111201000002100--1000000-0010-210111-1- --002100
110-0101101-00200001011-101011-10000000-001112100000-01-012-0111--22110--00
1-01001110--o-3-o---o---1oo-o1-101100001001-11oooooo-oo-2---o1111--211o--oo
-001001110110-20000100-1101011-10-00100-001112000000100010201111------00101
11001111101211320111021111101001ooooooo-oo1021oooooooo110221011011120----o11---1--101-002-0-o1-1101-10-1-1ooooooo1001111oooooo-oo-o-2-01111---o-o1--o
111-10111011113-000101110010-011000000--001-00000000001101210111111-0-00000
11-1oo--10-10021ooo1o-1o1o1o11-1oooooo-1oo1-2-oooooooooooo210111--o-----oo-oo1ooo11o-oo12ooo--oo11101oo-o1o1ooooo1oo1-11oooooo1ooo1o1o1o11------1o1o1
1001-01010-10-20000-01--0-0-0-11000000-2000--1000000100000210111--1-0--1000
100101101011000000010-10101011-10-00000-00111-000000-0--0---01111---0-0-100
1101oo-ooo11ooooooo10-1010101111ooooooo20011-1oooooo-oo------------------o11010011102-012-ooo1o---1oo--1o1o1ooooo1oo11110ooooooo100-2101111-22110-02-
TAXONOMIC SECTION
Family POTTIACEAE
Pottiaceae Schimp., Coroll. Bryol. Eur. 24, 1855 [1856], nom.fam. cons.
The family name is based on Pottia, which was named by Friedrich Ehrhart in his Beitrlige zur Naturkunde (1787) for "Cel. Joh.
Frid. Pott," a physician and professor of botany at Brunswick (mistakenly Braunschweig according to Muller 1853), author of an
unpublished "Flora Brunswickensis."
Plants usually turf-forming or loosely caespitose, green above and
brown below, irregularly branching. Stems short or to several em
in length, mostly pentagonal in transverse section, central strand
usually present, hyalodermis usually absent; axillary hairs several
cells in length, sometimes the basal 1-3 cells brownish. Leaves
usually appressed and often contorted when dry, spreading when
wet, ovoid to lanceolate or lingulate, ca. 1.5 to 3.5 mm in length,
margins usually recurved below, occasionally plane, entire or
sometimes dentate above, occasionally bordered by thick-walled
or elongate cells or cells in layers; apex rounded-obtuse to more
commonly narrowly acute; base usually ovate to oblong, occasionally sheathing the stem; costa ending a few cells below the
apex to short-excurrent or awned, in medial transverse section
usually with a differentiated epidermis ventrally or on both sides,
one or two stereid bands, guide cells in one or seldom more than
one layer, hydroid strand occasionally present (sometimes multiple); upper /aminal cells usually subquadrate, occasionally hexagonal or rarely short-rectangular, mostly ca. 9-16 jlm wide,
1(- 2):1, usually in one layer, walls mostly evenly thickened,
superficially flat to bulging, sometimes bulging only ventrally;
papillae usually present over the upper /aminal cells, solid or
occasionally hollow, usually bifid but occasionally simple, sometimes flattened or compound; basal cells usually differentiated,
usually clear, smooth or lightly papillose above, rectangular, generally filling the base medially, sometimes rising marginally in a
vee-shape, occasionally bulging. Asexual reproduction not
uncommon, by multicellular (rarely unicellular) propagula borne
on stalks in the leaf axils or more seldom on leaves, or obovoid
brood bodies borne on rhizoids in the soil. Perichaetia and perigonia terminal or occasionally lateral on short branchlets.
Dioicous or monoicous, occasionally apparently rhizautoicous.
Perichaetialleaves often sheathing in the lower part and then with
elongate-rhomboidal cells in lower portion, usually larger than the
stem leaves, long-oval to long-lanceolate. Perigoniate plants occasionally smaller than the perichaetiate, seldom nearly stemless
and bud-like. Sporophyte often in transformation series of apparent peristome reduction and seta shortening. Seta usually elongate, often twisted; capsules ovoid to cylindric, mostly
stegocarpous, occasionally spherical and then cleistocarpous and
rhexolytic; neck usually small or nearly absent; annulus mostly of
1-2 rows of vesiculose cells, occasionally revoluble or deciduous
in pieces; operculum short-conic to short-rostrate, cells in straight
or oblique rows; usually elongate, peristome teeth occasionally
absent, more usually erect or twisted usually clockwise, yellow,
orange or red, rudimentary or consisting of 16 mostly twice cleft,
spiculose, striate or papillose, lanceolate teeth, or 32 linear, usually densely spiculose filiform divisions, the basal membrane usually low or absent, occasionally very high and trabeculate. Calyp-
tra cucullate, smooth, occasionally mitrate. Spores ca. 10-15
jlm in diameter. Laminal color reaction yellow to orange-red or
red in two percent KOH solution. Reported chromosome number generally x = 13.
The conservation of the name Pottiaceae (Magill 1977b;
Greuter 1988) apparently was unnecessary. The original discussion of Hyophilaceae Hampe, Linnaea 4: 68, 1847, is actually
given as a footnote in which was said: "Ich habe diesen Namen
den Friiheren: Calympereae oder Syrrhopodonteae vorgezogen,
weil sich diese Namen auf die Form der Haube, oder auf das
Peristome beziehen; die Beschaffenheit des Peristoms kann dem
Familiennamen nicht zum Grunde gelegt werden, wo das
Peristom oft felt." Argument may be made that there is no
description here ("wo das Peristom oft felt" being merely rhetorical and applying to Hyophilaceae, Calympereae and
Syrrhopodonteae equally), hence Hyophilaceae was not validly
published prior to Pottiaceae; also, according to G. Zjilstra
(pers. comm.), this is not a diagnosis since the phrase including
"oft" does not cover the whole group. The name Tortulaceae,
used for the Pottiaceae in some older literature, is a synonym of
Ephemeraceae (Crosby & Magill1981). Recently, Saito (1975a)
agreed with Hilpert (1933) and Podp~ra (1954) in excluding
Cinclidotus from the Pottiaceae and placing it into a separate
family, the Cinclidotaceae Schimp. (= Ripariaceae Schimp. see
Crosby & Magill 1981), a disposition followed here. For additional information on excluded genera and species, see the discussion of Excluded Taxa.
The authorities for the names mentioned in the text are
those of the names recognized as correct in the updated list of
taxa (see Table of Contents). The specific and infraspecific
nomenclature is mainly that of the Index Muscorum (van der
Wijk et al. 1959-69), with additions and emendations based on
research since the publication of that work as summarized in the
updated list of taxa. Some of the names of subgenera and sections acknowledged as correct here may not, however, represent
taxonomically valid or properly placed groups, but are merely
starting points for revisionary study. Thorough revisionary work
is required for most genera; for this reason, no emphasis is
placed on citation of putative type specimens in that
lectotypification, inappropriate here, is required for a majority
of them; the specimens examined in the course of this study
were annotated and their herbarium designations (Holmgren et
al. 1990) are given here.
All new combinations, new names and new synonymy are
based on examination of material obtained on loan from the
herbaria cited, or in relatively few cases, are based on my judgement that the novelty is clearly warranted. The vast majority of
specimens seen during this study are "authentic" or syntype
THE POTTIACEAE
material as far as determinable with the literature immediately
available, but their exact identity as types, although probable,
remains to be more accurately ascertained through revisionary
treatments and associated lectotypification not appropriate here.
This study could not be done adequately, however, without a
great deal of name shuffling.
In the nomenclature list at the beginning of each generic treatment, names following the genus name are not indented if they
are correct (e.g. accepted infrageneric names) while synonyms (of
the genus or of infrageneric names) are indented. Synonyms that
cannot be referred to a particular infrageneric taxon are placed
immediately below the generic name. Infrageneric taxa are
grouped by category, being arranged alphabetically (after the typi-
53
cal name), and include both those recognized by the present
author and those unstudied supraspecific names given as correct
names by van der Wijk et al. (1959-69). Synonyms are grouped
by taxonomic category (subgenus, section, subsection), then
arranged alphabetically.
The citation of a species name under the heading "species
examined" does not mean that the species has been extensively
studied by me. It merely indicates that well-identified or
syntype or type or otherwise authentic specimens of the species
were examined. These fit well within the genus concept
presented in the description, or at least cannot be easily referred
to a different genus.
KEY TO SUPRAGENERIC TAXA
The subfamilies and tribes recognized here are subclades defined by character state changes at ancestral nodes, but these groups are
fairly coherent morphologically and may be described in the fashion of a standard key. The couplets of this key reflect the coordination of the subclades in Cladogram 16. See the section on morphology above or the glossary for explanation of specialized terms.
l. Upper lamina bistratose medially and the cells not vertically aligned (i.e. not directly over each other) near the costa but grading to
vertically evenly stacked towards the leaf margin, leaves broadly to linearly lanceolate ................................ Subfamily Timmielloideae
l. Upper lamina unistratose or if bistratose then cells situated directly over one another throughout.. ...... ................................................... 2
2. Upper !aminal cells ventrally mamillose medially but several rows of cells bulging on both sides marginally, costal guide cells
forming a thick-walled, multilayered cylinder........................................................................................ Subfamily Gertrudielloideae
2. Upper !aminal cells similarly bulging or not throughout leaf, guide cells either not multilayered or if so then thin-walled .............. 3
3. Leaves lanceolate, margins plane to weakly incurved, apex acute, base sheathing, upper lamina KOH red, stereid bands two,
guide cells 4-6, rows of cells across ventral surface of costa 10(-16) .................................... Subfamily Erythrophyllopsoideae
3. Not this combination of characters ................................................................................................................................................. 4
4. Leaves long-linear, margins plane, ventral stereid band larger than the dorsal.. ....................... Subfamily Chionolomoideae
4. Leaves lanceolate to spathulate, ventral stereid band absent or generally smaller than the dorsal ......................................... 5
5. Sclerodermis commonly poorly differentiated, hyalodermis commonly present, leaves lanceolate, margins plane to
weakly incurved, upper !aminal cells KOH yellow, costa lacking a differentiated dorsal epidermis, clavate axillary
propagula rare ..................................................................................................................... Subfamily Trichostomoideae
5. Not this combination of characters ............................................................................. ....................................................... 6
6. Stem sclerodermis commonly well differentiated from cells of central cylinder, which have abruptly larger
lumens, leaves usually broadly lanceolate to narrowly elliptical, usually with two costal stereid bands, leaf base
commonly differentiated in shape and ovate or rectangular, upper !aminal cells equally convex on both free surfaces, clavate axillary propagula commonly present in some genera ............................... Subfamily Merceyoideae 7
7. Stem black, leaves long-triangular, capsule with a circumstomal ring ..................... Tribe Tetracoscinodontieae
7. Not this combination of characters ........................................................................................................................ 8
8. Upper lamina usually KOH red, dorsal stereid band usually reniform ............... Tribe Bryoerythrophylleae
8. Not this combination of characters ................................................................................................................. 9
9. Stem central strand absent ........................................................................................ Tribe Leptodontieae
9. Stem central strand usually present, or if absent then costa with one stereid band ........ Tribe Barbuleae
6. Stem sclerodermis commonly not or poorly differentiated from cells of central cylinder, which generally grade in
size into the cortical cells, leaves usually broadly ligulate to spathulate, usually with one stereid band in the costa,
leaf base usually little differentiated in shape, sometimes upper laminal cell free surfaces ventrally bulging and
dorsally weakly convex, clavate axillary propagula rare ..................................................... Subfamily Pottioideae 10
10. Upper !aminal cells usually bulging ventrally and weakly convex dorsally ............................ Tribe Hyophileae
10. Upper !aminal cells equally convex on both free surfaces ............................................................ Tribe Pottieae
54
GENERA OF THE POTTIACEAE
KEY TO THE GENERA
This key was developed in part using the DELTA (DEscription Language for TAxonomy) system (Dallwitz 1974, 1980; Dallwitz &
Paine 1986; Partridge et al. 1988). Data matrices were created using the descriptions in the taxonomic section. In order to make the
key work, total variation was not scored for certain of the larger genera in that rare and unusual character states were sometimes not
included. This was done to account for problems in key creation that are due to the presence in a genus of (1) species probably
incorrectly assigned to a particular genus and requiring further study for correct disposition, (2) species possessing several of the
reliable characteristics of that genus but with one or more additional characters considered reliable for a different genus, and (3)
much reduced species with occasional secondary lack of features (e.g. ventral stereid band) whose absence was a key characteristic
of other genera (e.g. genera of Pottieae). Gametophytic characters are emphasized in the key, reflecting taxonomic importance in
the actual treatments, but closeness of taxa in the key does not imply a close phylogenetic relationship. Because artificial distinctions of sporophyte characteristics previously used to distinguish genera are not recognized, many genera necessarily key out in
more than one place. To keep the number of couplets to a manageable minimum, reliable technical characters involving sectioning
and color reactions to two percent potassium hydroxide solution are necessary at the beginning of the key. Emphasis on more immediately observable characters are practicable only in regional treatments. The key characters below are those of leaf morphology and
anatomy unless otherwise noted.
The number of times each genus appears in this key is dependent on the variation within the genus of the characters considered
reliable for identification. This is a measure of the internal complexity of the genus, whether due to taxonomic heterogeneity or simple breadth of variation of a monophyletic taxon. An index of such complexity for each suprageneric group is the number of times
of their genera occur in the key divided by the number of genera in the group, as follows: Timmielloideae, 1.0; Erythrophyllopsoideae, 1.0; Gertrudielloideae, 1.0; Chionolomoideae, 1.7; Trichostomoideae, 3.9 (Trichostomum occurs 11 times!); Merceyoideae,
2.8; tribe Tetracoscinodontieae, 1.0; tribe Bryoerythrophylleae, 2.2 (Pseudocrossidium occurs 4 times); tribe Leptodontieae, 1.5
(Triquetrel/a occurs 4 times); tribe Barbuleae, 4.2 (Didymodon and Gyroweisia each occur 9 times, Gymnostomum 7 times); Pottioideae, 2.3; tribe Hyophileae, 2.5 (Hyophila and Weissia each occur 9 times); tribe Pottieae, 2.2 (Aschisma and Trachycarpidium
each occur 4 times). It is probable that, after revision, the Trichostomoideae and Barbuleae will prove less of a source of variation in
taxonomic characters; taxa with low ratios are apparently presently well understood or at least easily characterized.
1(0).
Ventral costal stereid band absent (costa with a single stereid band) .......................................................................................... 2
Ventral costal stereid band present (costa with two stereid bands) ............................................................................................ 51
2(1).
Ventral costal outgrowths absent ................................................................................................................................................. 3
Ventral costal outgrowths present as a pad of cells, lamellae or filaments ................................................................................41
3(2).
Stem central strand absent ........................................................................................................................................................... .4
Stem central strand present ......................................................................................................................................................... l6
4(3).
KOH color reaction of upper laminal cell walls essentially yellow or orange ............................................................................. 5
KOH color reaction of upper laminal cell walls essentially red, usually a definite brick red .................................................... ll
5(4).
Superficial walls of upper laminal cells similarly shaped on both sides of lamina; leaves when dry occasionally channeled
but not distinctly tubulose .......................................................................................................................................................... 6
Superficial walls of upper laminal cells ventrally bulging-mamillose, weakly convex dorsally; leaves tubulose when dry .... 10
6(5).
Me<dial upper laminal cells small to medium sized, 7-l4Jlm in width .......................................................................................7
Medial upper laminal cells large, commonly more than l4Jlm in width ....................................................................................8
7(6).
Stem sclerodermis not or little differentiated; upper laminal papillae absent; basal cell group differentiated and rising higher
medially; length of stem usually 1.0 em or more; basal cells usually with straight walls, little wider than upper medial cells;
wide distribution ..................................................................................................................................................... Scope/ophila
Stem sclerodermis clearly differentiated; upper laminal papillae present; basal cell group differentiated approximately
straight across leaf; length of stem less than 1.0 em, usually less than 0.6 em; basal cells inflated and bulging, considerably
wider than upper medial cells; wide distribution ...................................................................................................... Gyroweisia
8(6).
Margins pllll'.e to incurved or involute; transverse section of dorsal costal stereid band clearly flattened or ventrally indented,
reniform or crescent-shaped; hydroid strand present; stem hyalodermis present (sometimes indistinct); southern South
America, Australia ................................................................................................................................................ Calyptopogon
4-39 Costa with one stereid band, costal outgrowths absent, central strand absent
THE POTTIACEAE
55
Margins recurved to revolute; transverse section of dorsal costal stereid band round to semicircular; hydroid strand absent;
stem hyalodermis absent ............................................................................................................................................................ 9
9(8).
Transverse section of costa round or ovate or elliptical; basal cell group not or little differentiated from upper medial cells;
leaves widest below midleaf; number of rows of cells across ventral surface of costa usually 2 but up to 4; stem scleroderrnis clearly differentiated; tropics ..................................................................................................................... Streptopogon
Transverse section of costa semicircular; basal cell group clearly differentiated, usually larger, less papillose, walls thinner;
leaves widest at or above midleaf; number of rows of cells across ventral surface of costa commonly 4 or more; stem
scleroderrnis not or little differentiated; western North America ................................................................................... Crumia
10(5).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped;
ventral costal epidermis absent; dorsal costal epidermis absent; leaf base clearly differentiated in shape; American tropics,
eastern Asia .......... ........................ .............................................................................................................................. Luisierella
Basal cell group differentiated straight across leaf or rising higher medially; transverse section of dorsal costal stereid band
round to semicircular; ventral costal epidermis present; dorsal costal epidermis present; leaf base little differentiated in
shape; Brazil, India .................................................................................................................................................... Ganguleea
II (4 ).
Medial upper lamina! cells small to medium sized, 7-14 Jlm in width .................................................................................... 12
Medial upper !aminal cells large, commonly more than 14 Jlm in width .................................................................................. 13
12(11).
Dorsal costal epidermis absent; basal cell group clearly differentiated, usually larger, less papillose, walls thinner; theca
ovoid to cylindrical; capsule stegocarpous; leaves usually widest at or above midleaf; wide distribution ................ Syntrichia
Dorsal costal epidermis present; basal cell group not or little differentiated from upper medial cells; theca spherical; capsule
cleistocarpous; leaves widest below midleaf; wide distribution .......................................................... .......................... Acaulon
13(11 ).
Margins plane to in curved or involute; length to width ratio of medial upper !aminal cells 1-2: 1 ...... ..................................... 14
Margins recurved to revolute; length to width ratio of medial upper lamina! cells 2-4: I or more ............... ...... ...................... 15
14(13).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped;
costal ventral cells longitudinally elongate 3:1 or more; stem sclerodermis clearly differentiated; superficial walls of upper
!aminal cells strongly convex to bulging on both sides of lamina; southern South America, Australasia ........... Calyptopogon
Basal cell group differentiated straight across leaf or rising higher medially; transverse section of dorsal costal stereid band
round to semicircular; costal ventral cells quadrate to very short-rectangular; stem sclerodermis not or little differentiated;
superficial walls of upper !aminal cells flat or very weakly convex on both sides of lamina; southern South America, Antarctica ..................................................................................................................................... ................................ Sarconeurum
15(13).
Hydroid strand absent; upper !aminal papillae absent; theca longer, usually more than 1.5 mm in length; calyptra longer, 1.0
mm or more in length; plants caespitose, usually in a mat or turf; tropics ............................................................ Streptopogon
Hydroid strand present; upper !aminal papillae present; theca short, less than 1.5 mm in length; calyptra short, less than 1.0
mm; plants gregarious or scattered; wide distribution ........................................................... ................................. .M icrobryum
16(3).
16-40 Costa with one stereid band, costal outgrowths absent, central strand present
Medial upper !aminal cells small to medium sized, 7-l4Jlm in width ..................................................................................... 17
Medial upper !aminal cells large, commonly more than l4Jlm in width ..................................................................................31
17(16).
Dorsal costal epidermis absent ....... ............................................................................................................................................ 18
Dorsal costal epidermis present .................................................................................................................................................. 23
18( 17).
KOH color reaction of upper !aminal cell walls essentially yellow or orange ........................................................................... 19
KOH color reaction of upper lamina! cell walls essentially red, usually a definite brick red ....................................................20
19(18).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; transverse section of dorsal costal stereid band clearly flattened or ventrally· indented, reniform or crescent-shaped;
stem sclerodermis not or little differentiated; costal ventral cells quadrate to very short-rectangular; guide cells commonly
more than 6; Mexico, Andes, South Africa ......................................................................................................... Streptocalypta
Basal cell group differentiated straight across leaf or rising higher medially; transverse section of dorsal costal stereid band
round to semicircular; stem sclerodermis clearly differentiated; costal ventral cells longitudinally elongate 3:1 or more;
guide cells 2-6; wide distribution ........ ..................................................................................................................... Gyroweisia
56
GENERA OF THE POTTIACEAE
20(18)0
Capsule cleistocarpous; calyptra short, less than 1.0 mm; AustralasiaooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooOOooPhascopsis
Capsule stegocarpous; calyptra longer, 1.0 mrn or more in length oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo2l
21(20)0
Dorsal superficiallaminal cell walls about same thickness as the ventral or weakly thicker near costa; basal cell group clearly
differentiated, usually larger, less papillose, walls thinner; leaves widest at or above midleaf 000000000000000000000000000000000000000000000022
Dorsal superficiallaminal cell walls throughout distinctly thicker than the ventral; basal cell group not or little differentiated
from upper medial cells; leaves widest below midleaf; western North America, central EuropeooooooooooooooooooooooooooooooooHilpertia
22(21)0
Seta nearly absent to short, less than l em; perichaetialleaves distinctly different in size or morphology, strongly sheathing
the seta; austral oceanic islands 0000000000000000 000000oooooooooooooooooooooooooooooooooooooooooo000000000000000000 oooooooooooooooooooooooooooooooooooo0000000000000000000 Willia
Seta elongate, 1 em or longer; perichaetial leaves similar to cauline leaves or occasionally smaller or somewhat enlarged;
wide distribution 00000000000000000000000000000000000000 oooooooooooooooooooooooooooooo oooooooooooooooooooooooooooooooooooooooooooooooo 000000 0000000000000000000000000000000Syntrichia
23(17)0
Superficial walls of upper laminal cells similarly shaped on both sides of laminaoooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo24
Superficial walls of upper laminal cells ventrally bulging-mamillose, weakly convex dorsally oooooooooooooooo:oooooooo oooooooooooooooooooooo29
24(23)0
Ventral costal epidermis absentoooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooOOOOooO ooooOOoooo25
Ventral costal epidermis present oooooooooooooooooooooooooooooooooooooooooooooooooooo ooooooooooooooooooooooooooooooooooooooOOoooooooooOOOOOoooooooooooo oooooooooooooooooooooooooo26
25(24)0
Leaf ventral surface above midleaf nearly plane to broadly channeled; leaves widest at or above midleaf; rather deep, narrow
groove along costa absent; perichaetium terminal on main axis; seta nearly absent to short, less than 1 em;
pantropicat. 000000000000000000 000000000000000000000000 oooooooooooooooooooooooooOOOOOOOOooOOOOoOOoOOOOOOOooOOOOOOOOOOOOOO ooooooooooooooooooooooooooooooooooooooooooooGymnostomiella
Leaf ventral surface above midleaf keeled; leaves widest below midleaf; rather deep, narrow groove along costa present;
perichaetium lateral on main axis at ends of very short ·branches; seta elongate, l em or longer; wide distribution 00000000000000000000000000000000000oooooooooooooooooo ooooooooooooooooooooooooooooooooooooooooo0ooooooooooooooOOOooooooooooooooooooooooooooooooooooooooooooooooooooooooooAnoectangium
26(24)0
Hydroid strand absentooooooooooooooooooooooooooooooooooooooooooo oooooo oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo27
Hydroid strand present; wide distribution ooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooPseudocrossidium
27(26)0
Perichaetium terminal on main axis, leaves usually broadly channeled across the ventral surfaceooooooooooooooooooooooooooo oooooooooooooooo28
Perichaetium lateral on main axis at ends of very short branches, leaves usually strongly keeled; wide distribution ooooooooooooooooooooooooooooooooooooooooooooooOOooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo Anoectangium
28(27)o
Length of stem less than 1.0 em, usually less than 006 em; basal cells inflated and bulging, considerably wider than upper
medial cells; wide distribution 0000000000000000000000000000000000000000000 000000000000oooooooooooooooooooooooo 000000oooooo ooooooooooooooooooooooooooooooooooooooo Gyroweisia
Length of stem usually 1.0 em or more; basal cells usually with straight walls, little wider than upper medial cells; wide distribution 0000 00000 00000000oo 0000000 00 000000 0Oo 000000000000 0000000000000 000000 00000000 0000 00 0000 00 000000000 00 00 000000 00000000 000000000000000000000000000000 0000000000 000 0000000Didymodon
29(23)0
KOH color reaction of upper laminal cell walls essentially yellow; margins plane to incurved or involute; transverse section
of dorsal costal stereid band round to semicircular; guide cells 2-6; stem hyalodermis absent 00000000000 000000000000000000000000000000000030
KOH color reaction of upper laminal cell walls essentially orange; margins revolute to near apex; transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; guide cells commonly more than
6; stem hyalodermis present (sometimes indistinct); AndesoooOOoooo oOOOOOO oOoOOOOOOooooOOoooooooo oooooooOoooooooooOOoooooooooooooOOOOoooooooooGertrudie/la
30(29)0
Margins plane or very weakly incurved; transverse section of costa round or ovate or elliptical; hydroid strand absent; leaf
base clearly differentiated in shape; number of rows of cells across ventral surface of costa usually 2 but up to 4; tropics oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo ooooooooooooooooooooWeisiopsis
Margins clearly incurved to involute; transverse section of costa semicircular; hydroid strand present; leaf base little
differentiated in shape; number of rows of cells across ventral surface of costa commonly 4 or more; American tropics,
South Africa, Burma ooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooOoOOOooooooooooooooooooooooOOoooooooooooooPlaubelia
31(16)0
Margins plane to incurved or involute oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo.32
Margins recurved to revolute ooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo35
32(31)0
Transverse section of dorsal costal stereid band round to semicircular; upper laminal papillae presento oooooooooooooooooooooooooooooooooo33
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; upper
laminal papillae absent; Mexico, Andes 00000000000000000000000000000000000000000000 000000000000000000000000000000000000000000 000000 000000000000000000 0Sagenotortula
33(32)0
Ventral costal epidermis absent; hydroid strand absent; superficial walls of upper larninal cells strongly convex to bulging on
both sides of lamina; pantropicalooooooooooooooooooooooooooooooooooooooooooo 000000 oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooGymnostomiel/a
THE POTTIACEAE
57
Ventral costal epidermis present; hydroid strand present; superficial walls of upper Jaminal cells flat or very weakly convex ,
on both sides ....................................................................................... ..................................................................................... 34
34(33).
Number of rows of cells across ventral surface of costa usually 2 but up to 4; upper marginal cells in same number of layers
as medial cells; upper marginal cells not longer than medial cells (sometimes larger); papillae per lumen 2--6, usually bifid
or multifid; basal cell group differentiated and rising higher medially; southern South America, Antarctica...... Sarconeurum
Number of rows of cells across ventral surface of costa commonly 4 or more; upper marginal cells differentiated as a
bistratose (or more) border; upper marginal cells rectangular and clearly longer than medial cells; papillae per lumen many,
usually 6 or more, simple or bifid; basal cell group differentiated approximately straight across leaf; wide distribution ..................................................................................................................................................... ...... ................ Hennedie/la
35(31).
KOH color reaction of upper Jaminal cell walls essentially yellow or orange ...........................................................................36
KOH color reaction of upper laminal cell walls essentially red, usually a defmite brick red .. ..................................................37
36(35).
KOH color reaction of upper laminal cell walls essentially yellow; hydroid strand present; medial upper Jaminal cells small,
7-10 jlm in width; papillae per lumen 2--6, usually bifid or multifid; wide distribution .................................. ..............Tortula
KOH color reaction of upper laminal cell walls essentially orange; hydroid strand absent; medial upper Jaminal cells medium
sized, ll-14jlm in width; papillae per lumen many, usually 6 or more, simple or bifid; western North America ....... Crumia
37(35).
Dorsal costal epidermis absent; dorsal superficiallaminal cell walls throughout distinctly thicker than the ventral; margins
revolute; perichaetialleaves distinctly larger in size; western North America, central Europe .............................. ..... Hilpertia
Dorsal costal epidermis present; dorsal superficiallaminal cell walls about same thickness as the ventral or weakly thicker
near costa; margins recurved; perichaetialleaves similar to cauline leaves or occasionally smaller or somewhat enlarged .38
38(37).
Basal cell group not or little differentiated from upper medial cells ..........................................................................................39
Basal cell group clearly differentiated, usually larger, Jess papillose, walls thinner ................................................................ .40
39(38).
Medial upper Jaminal cells 15-17 jlm in width; leaf apex obtuse or acute to acuminate; upper Jaminal papillae present; sexual
condition monoicous; upper marginal cells in same number of layers as medial cells; wide distribution ..............Microbryum
Medial upper laminal cells commonly 18 jlm in width or more; leaf apex broadly rounded; upper Jaminal papillae absent;
sexual condition dioicous; upper marginal cells differentiated as a bistratose (or more) border; Mexico, Central America,
Andes ........................................................................................................................................................................Dolotortula
40(38).
Upper Jaminal papillae absent (except sometimes along extreme margins of leaf); sexual condition dioicous; plants
caespitose, usually in a mat or turf; length to width ratio of medial upper laminal cells 1-2:1; calyptra longer, 1.0 mm or
more in length; wide distribution ..................................................................................................................................... Chenia
Upper !aminal papillae present; sexual condition monoicous; plants gregarious or scattered; length to width ratio of medial
upper lamina! cells 2-4:1 or more; calyptra short, less than 1.0 mm; wide distribution ......................................... Microbryum
41(2).
41-50 Costa with one stereid band, costal outgrowths present
KOH color reaction of upper laminal cell walls essentially yellow or orange ................................ ............ ...............................42
KOH color reaction of upper Jaminal cell walls essentially red, usually a definite brick red ................................................... .48
42(41).
Ventral costal outgrowths differentiated as filaments or lamellae .......................................................................... .................. .43
Ventral costal outgrowths differentiated as a bulging pad of cells ......................................................................... .................. .46
43(42).
Transverse section of dorsal costal stereid band round to semicircular; transverse section of costa round to semicircular ......44
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; transverse section of costa distinctly flattened, usually reniform ................................................................................................... .45
44(43).
Ventral costal outgrowths differentiated as longitudinally elongated lamellae; spore diameter more than 15 jlm; annulus of
weakly differentiated cells; wide distribution ..................................................................................................... Pterygoneurum
Ventral costal outgrowths differentiated as separate filaments three or more cells in length; spore diameter 8-15 jlm; annulus
of vesiculose cells, often in two or more rows; wide distribution ............................................................................ Crossidium
45(43).
Margins plane to incurved or involute; axillary hairs with 1 or more basal cells with thicker or darker-colored walls; leaf
apex cucullate; leaves tubulose when dry; spore diameter more than 15 jlm; Mexico, Andes ...... .............................. Aloine/la
Margins recurved to revolute; axillary hair basal cell walls hyaline and all cells of hair similar; leaf apex flattened, channeled
or keeled; leaves when dry occasionally channeled but not distinctly tubulose; spore diameter 8-15 jlm; wide distribution ................................................................................................................................................................. Pseudocrossidium
58
GENERA OF THE POTTIACEAE
46(42).
Dorsal superficial lamina! cell walls about same thickness as the ventral or weakly thicker near costa; length to width ratio of
medial upper !aminal cells 1-2:1 ............................................................................................................................................ .47
Dorsal superficial !aminal cell walls throughout distinctly thicker than the ventral; length to width ratio of medial upper
!aminal cells 2-4: 1 or more; North Temperate Zone ............................................................ ......................................... Stegonia
47(46).
Margins plane to incurved or involute; medial upper lamina! cells 7-14 J.lm in width; leaf apex cucullate; upper !aminal
papillae absent; southwestern USA south through Andes ...................................................................................... Globulinella
Margins recurved to revolute; medial upper !aminal cells commonly larger than 14 J.lm in width; leaf apex flattened, channeled or keeled; upper !aminal papillae present; wide distribution ................................................................................. Tortula
48(41).
Ventral costal outgrowths differentiated as filaments or lamellae; medial upper !aminal cells commonly larger than 14 J.1rn in
width; transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped;
leaf base clearly differentiated in shape; transverse section of costa distinctly flattened, usually reniform; wide distribution .................................................................................................................................................................................... Aloina
Ventral costal outgrowths differentiated as a bulging pad of cells; medial upper lamina! cells small to .medium sized, 7-14
Jlm in width; transverse section of dorsal costal stereid band round to semicircular; leaf base little differentiated in shape;
transverse section of costa round to semicircular ...................................................................... ............................................. .49
49(48).
Stem central strand absent; margins plane to incurved or involute; number of rows of cells across ventral surface of costa
usually 2 but up to 4; medial upper !aminal cells medium sized, 11-14 J.lm in width; leaf apex obtuse or acute to
acuminate ................................................................................................................................................................................. 50
Stem central strand present; margins recurved; number of rows of cells across ventral surface of costa commonly 4 or more;
medial upper !aminal cells small, 7-10 J.lm in width; leaf apex broadly rounded; Mexico, Andes ............................. Saitoel/a
50(49).
Basal cell group not or little differentiated from upper medial cells; leaf apex flattened, channeled or keeled; costal ventral
cells longitudinally elongate 3:1 or more; leaves widest below midleaf; sporophyte commonly present, spherical, seta short;
wide distribution .................... ........................................................................................................................................ Acaulon
Basal cell group clearly differentiated, usually larger, Jess papillose, walls thinner; leaf apex cucullate; costal ventral cells
quadrate to very short-rectangular; leaves widest at or above midleaf; sporophyte unknown; Australia ........................Stonea
51(1 ).
Sl-67 Costa with two stereid bands, upper laminal cells KOH yellow or orange, central strand absent
KOH color reaction of upper !aminal cell walls essentially yellow or orange ........................................................................... 52
KOH color reaction of upper !aminal cell walls essentially red, usually a definite brick red .................................................. l71
52(51).
Stem central strand absent .......................................................................................................................................................... 53
Stem central strand present. ........................................................................................................................................................ 68
53(52).
Margins entire or minutely and evenly crenulate ....................................................................................................................... 54
Margins denticulate or serrulate to toothed ................................................................................................................................ 60
54(53).
Ventral costal epidermis absent .................................................................................................. ................................................55
Ventral costal epidermis present .............................................................................................. ..................................................51
55(54).
Basal cell group not or little differentiated from upper medial cells; peristome teeth present; seta elongate, 1 em or longer;
theca longer, usually more than 1.5 mm in length; annulus of vesiculose cells, often in two or more rows; austral areas and
Mediterranean climates in the North Temperate Zone ............................................................................................ Triquetrel/a
Basal cell group clearly differentiated, usually larger, Jess papillose, walls thinner; peristome teeth absent; seta nearly absent
to short, less than 1 em; theca short, less than 1.5 mm in length; annulus of weakly differentiated cells ............................... 56
56(55).
Transverse section of dorsal costal stereid band round to semicircular; medial upper laminal cells small, 7-10 J.lm in width;
upper laminal papillae present; calyptra short, less than 1.0 mm; wide distribution ......... ................................. Hymenostylium
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; medial
upper !aminal cells medium sized, 11-14 J.lm in width; upper !aminal papillae absent; calyptra longer, 1.0 mm or more in
length; eastern Asia, India ................................................. ................................................ .......................................... Reimersia
57(54).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; wide distribution ...................................................................................................................................................... Tort ella
Basal cell group differentiated straight across leaf or rising higher medially ............................................................................ 58
THE POTTIACEAE
59
58(57).
Leaf ventral surface above midleaf nearly plane to broadly channeled ..................................................................................... 59
Leaf ventral surface above midleaf keeled; New Zealand .................................................................................. Tetracoscinodon
59(58).
Costal ventral cells quadrate to very short-rectangular; stem hyalodermis present (sometimes indistinct); length of stem usually 1.0 em or more; basal cells usually with straight walls, little wider than upper medial cells; wide distribution ..... .......................... .... ...................................................... ...... .................................................................. ...... .Trichostomum
Costal ventral cells longitudinally elongate 3: I or more; stem hyalodermis absent; length of stem less than 1.0 em, usually
less than 0.6 em; basal cells inflated and bulging, considerably wider than upper medial cells; wide distribution . Gyroweisia
60(53).
Margins denticulate only near leaf base or at top of leaf sheath; North Temperate Zone, India, South Africa ........... Eucladium
Margins denticulate or serrulate to toothed near apex or throughout ......................................................................................... 61
61(60).
Margins plane to incurved or involute; leaf ventral surface above midleaf nearly plane to broadly channeled; rather deep,
narrow groove along costa absent ............... ........................................................................ ..................................................... 62
Margins recurved; leaf ventral surface above midleaf keeled; rather deep, narrow groove along costa present .......................63
62(61).
Ventral costal epidermis absent; hydroid strand present; theca spherical; superficial exothecial cell walls with a central and
lens-like superficial thickening; stem hyalodermis absent; Mexico, Angola ................... ............................... Bryoceuthospora
Ventral costal epidermis present; hydroid strand absent; theca ovoid to cylindrical; superficial exothecial cell walls evenly
thickened; stem hyalodermis present (sometimes indistinct); wide distribution ............................................. .....Trichostomum
63(61).
Basal cell group not or little differentiated from upper medial cells .............................. ............................................................ 64
Basal cell group clearly differentiated, usually larger, less papillose, walls thinner ............. .................................................... 65
64(63).
Dorsal costal epidermis absent; stem hyalodermis present (sometimes indistinct); number of rows of cells across ventral surface of costa commonly 4 or more; transverse section of costa distinctly flattened, usually reniform; stomates absent;
Andes ............................ ............... ........................................................................................................................ Leptodontiella
Dorsal costal epidermis present; stem hyalodermis absent; number of rows of cells across ventral surface of costa usually 2
but up to 4; transverse section of costa round to semicircular; stomates present; austral areas and Mediterranean climates of
the North Temperate Zone ...... ............................................ .............................. .................. ..................................... Triquetrella
65(63).
Papillae per lumen 2-6, usually bifid or multifid ... ........ ................ ....... ..................................... ............................ .................... 66
Papillae per lumen many, usually 6 or more, simple or bifid ........... ......................................................................................... 67
66(65).
Theca short, less than 1.5 mm in length; Andes ........................................................................ ........................ ..... Leptodontiel/a
Theca longer, usually more than 1.5 mm in length; wide distribution .................................................................... Leptodontium
67(65).
Upper marginal cells in same number of layers as medial cells, not longer than medial cells (sometimes larger); seta nearly
absent to short, less than 1 em; theca usually 1.5-3.5 mrn in length; marginal cell walls not thicker than those of medial
cells; Andes ................................................... .......................................... .................. ............. ............. .................Streptotrichum
Upper marginal cells differentiated as a bistratose border, rectangular and clearly longer than medial cells; seta elongate, 1
em or longer; theca more than 3.5 mm in length; marginal cell walls thicker than those of medial cells;
Andes ................................................................................................................ ....................................... ......... Trachyodontium
68(52).
68-170 Costa with two stereid bands, upper !aminal cells KOH yellow or orange, central strand present
Margins entire or minutely and evenly crenulate ............ ...........................................................................................................69
Margins denticulate or serrulate to toothed .............................................................................................................................. 148
69(68).
Hydroid strand absent ............. ...... .............................. ...... ........ ............. ............................................................... ............. .........70
Hydroid strand present ................................ ......................... .... .................. .............................................................................. 130
70(69).
Inflated, banana-shaped alar cells absent or alar cells merely swollen and rounded ........... ...... ................................................ 71
Inflated, banana-shaped alar cells present and decurrent as a pad on the stem; Mexico, West Indies .... .......... Weissiodicranum
71 (70).
Superficial walls of upper lamina! cells similarly shaped on both sides of lamina .. .. ............................... ....................... ..........72
Superficial walls of upper !aminal cells ventrally bulging-mamillose, weakly convex dorsally ..... ........................................ 119
72(71).
Ventral costal stereid band smaller than the dorsal or of nearly equal size ..... .................. ...... ............ ....................... ............... 73
Ventral costal stereid band distinctly larger than the dorsal .................................................................................................... 116
60
GENERA OF THE POTTIACEAE
73(72).
Margins plane to incurved or involute ....................................................................................................................................... 74
Margins recurved to revolute .................................................................................... .............................. .................. ............... 107
74(73).
Margins plane or very weakly incurved ............. ...... .................................... .............................................................................. 75
Margins clearly incurved to involute .... .............................................................................. ...................................................... ! 05
75(74).
Dorsal costal epidermis absent ............................ .............................................................................. ............ .............................76
Dorsal costal epidermis present .................................................................................................................................................. 89
76(75).
Capsule cleistocarpous, i.e., rupturing (usually) irregularly ......................................................................................................77
Capsule stegocarpous, i.e., dehiscing by an operculum ......................................................................... .................................... 80
77(76).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border ... .............................................................................................................................................................................. ........... 78
Basal cell group differentiated straight across leaf or rising higher medially (occasionally rising weakly along extreme basal
margin as a line of elongate cells); wide distribution ............ ........................................................................ .......Trichostomum
78(77).
Ventral costal epidermis absent; theca spherical; stomates absent; calyptra papillose, distinctly roughened or strongly mamillose; central North America, Europe ........................................................................................................................... Aschisma
Ventral costal epidermis present; theca ovoid to cylindrical; stomates present; calyptra smooth or nearly so ......................... 79
79(78).
Leaf base little differentiated in shape; axillary hairs with I or more basal cells with thicker or darker colored walls; capsule
surface evenly mamillose or with distinct protuberances of strongly bulging cells basally or throughout; stem hyalodermis
absent; theca short, less than 1.5 mm in length; South America, Africa, Oceania ........ .................................. Trachycarpidium
Leaf base clearly differentiated in shape; axillary hair cell walls hyaline and all cells of hair similar; capsule surface nearly
smooth; stem hyalodermis present (sometimes indistinct); theca longer, usually more than 1.5 mm in length; wide distribution ....................................................................................................................... .................................... .................. ..... Tortella
80(76).
Transverse section of dorsal costal stereid band round to semicircular ..................................................................................... 81
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped ............ 84
81(80).
Costal ventral cells quadrate to very short-rectangular ........................... ................................................................................... 82
Costal ventral cells longitudinally elongate 3:1 or more ....................................................................... ..................................... 83
82(81 ).
Costa ending before the leaf apex; wide distribution ......................................................... ....................................Gymnostomum
Costa percurrent to excurrent; wide distribution .......................... ............ ............................................................... Trichostomum
83(81).
Stem sclerodermis not or little differentiated; length of stem usually 1.0 em or more; basal cells usually with straight walls,
little wider than upper medial cells; wide distribution ......................................................................................... Gymnostomum
Stem sclerodermis clearly differentiated; length of stem less than 1.0 em, usually less than 0.6 em; basal cells inflated and
bulging, considerably wider than upper medial cells; wide distribution ........................................ .................. ........ Gyroweisia
84(80).
Guide cells 2-6; length of stem usually 1.0 em or more .................................................................................... ........................ 85
Guide cells commonly more than 6; length of stem less than 1.0 em, usually less than 0.6 em; Mexico, Andes, South
Africa ................................................................................................................................................................... Streptocalypta
85(84).
Stem hyalodermis absent ......................................................................................................... ................................................... 86
Stem hyalodermis present (sometimes indistinct) ...................................................................................................................... 87
86(85).
Medial upper !aminal cells small, 7-10 Jlm in width; leaf apex obtuse to broadly acute; costa ending before the leaf apex;
wide distribution .............................................................................................................. .................................... Gymnostomum
Medial upper !aminal cells medium sized, 11-14 Jlm in width; leaf apex narrowly acute to acuminate; costa percurrent to
excurrent; American tropics, eastern Asia .............................................................................................................Tuerckheimia
87(85).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; perichaetialleaves distinctly different in size or morphology, sometimes strongly sheathing; wide distribution .. Tortella
Basal cell group differentiated straight across leaf or rising higher medially; perichaetial leaves similar to cauline leaves or
occasionally smaller or somewhat enlarged ............................................................................................................................88
THE POTTIACEAE
61
88(87).
Costa ending before the leaf apex; wide distribution .............................. ...............................................................Gymnostomum
Costa percurrent to excurrent; wide distribution ................................................. ....................................................Trichostomum
89(75).
Transverse section of costa round to semicircular .....................................................................................................................90
Transverse section of costa distinctly flattened, usually reniform ........................................................................................... 103
90(89).
Guide cells 2-6 ................................................................. ..........................................................................................................91
Guide cells commonly more than 6; Philippines ................. ................................................................................. Pachyneuropsis
91(90).
Capsule cleistocarpous, i.e., rupturing (usually) irregularly ...................................................................................................... 92
Capsule stegocarpous, i.e., dehiscing by an operculum ............................................................................................................. 94
92(91).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border ... ........................................................................................................................ ................................................................. 93
Basal cell group differentiated straight across leaf or rising higher medially; wide distribution ...................... ~ .... Trichostomum
93(92).
Axillary hairs with 1 or more basal cells with thicker or darker colored walls; stem hyalodermis absent; capsule surface
evenly mamillose or with distinct protuberances of strongly bulging cells basally or throughout; theca short, less than 1.5
mm in length; South America, Africa, Oceania ............................................................................................... Trachycarpidium
Axillary hair basal cell walls hyaline and all cells of hair similar; stem hyalodermis present (sometimes indistinct); capsule
surface nearly smooth; theca longer, usually more than 1.5 mm in length; wide distribution ....................................... Torte /Ia
94(91).
Stem sclerodermis not or little differentiated ............................................................................................................................. 95
Stem sclerodermis clearly differentiated .................................................................................................................................... 99
95(94).
Stem hyalodermis absent ......................................................... ........................................................................ ........................... 96
Stem hyalodermis present (sometimes indistinct) ................................ ...................................................................................... 97
96(95).
Medial upper laminal cells small, 7-10 Jlm in width; leaf apex obtuse to broadly acute; costa ending before the leaf apex;
wide distribution ............................................................................................................................................ ...... Gymnostomum
Medial upper laminal cells medium sized, 11-14 Jlm in width; leaf apex narrowly acute to acuminate; costa percurrent to
excurrent; American tropics, eastern Asia ............... .............................................................................................. Tuerckheimia
97(95).
Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; perichaetialleaves distinctly different in size or morphology, sometimes strongly sheathing; wide distribution .. Tortella
Basal cell group differentiated straight across leaf or rising higher medially; perichaetial leaves similar to cauline leaves or
occasionally smaller or somewhat enlarged ............................................................................................................................ 98
98(97).
Costa ending before the leaf apex; wide distribution ............................................................................................. Gymnostomum
Costa percurrent to excurrent; wide distribution ................................ ..................................................................... Trichostomum
99(94 ).
Perichaetium terminal on main axis .......................... ...................................................... .................. ....................................... 100
Perichaetium lateral on main axis at ends of very short branches; wide distribution ................................................... Molendoa
100(99).
Costal ventral cells quadrate to very short-rectangular ............................................................................................................ 101
Costal ventral cells longitudinally elongate 3:1 or more ................................................................................... ....................... l02
101(100). Leaf base little differentiated in shape; leaves widest at or above midleaf; wide distribution ........................................ Hyophila
Leaf base clearly differentiated in shape; leaves widest below midleaf; wide distribution ..................... ............... Trichostomum
102(100). Basal cells usually with straight walls, little wider than upper medial cells; wide distribution ........................................Barbu/a
Basal cells inflated and bulging, considerably wider than upper medial cells; wide distribution ...................... ...... .. Gyroweisia
103(89).
Perichaetium terminal on main axis .................................................. .......................................... .............................. ...... ......... 104
Perichaetium lateral on main axis at ends of very short branches; wide distribution ................................................... Molendoa
104(103). Peristome twisted; leaf base (of sterile leaves) little differentiated in shape; superficial walls of upper laminal cells flat or
very weakly convex on both sides of lamina; stem hyalodermis absent; basal cell group differentiated and rising higher
medially; length of stem less than 1.0 em, usually less than 0.6 em; Mediterranean region ................................. Leptobarbula
62
GENERA OF THE POTTIACEAE
Peristome straight or absent; leaf base commonly clearly differentiated in shape; superficial walls of upper lamina! cells
strongly convex to bulging on both sides of lamina; stem hyalodermis present (sometimes indistinct); basal cell group
differentiated approximately straight across leaf; length of stem usually 1.0 em or more; wide distribution ......Trichostomum
105(74).
Transverse section of dorsal costal stereid band round to semicircular ..................................................................... Quaesticula
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 106
106(105). Upper laminal margins not sharply and narrowly infolded; leaf base little differentiated in shape; leaves widest at or above
midleaf; stem sclerodermis clearly differentiated; axillary hair basal cell walls hyaline and all cells of hair similar; wide
distribution ........................................................................................................................................ ........................... .Hyophila
Upper laminal margins sharply and narrowly infolded; leaf base clearly differentiated in shape; leaves widest below midleaf;
stem sclerodermis not or little differentiated; axillary hairs with 1 or more basal cells with thicker or darker colored walls;
wide distribution ............................................................................................................................................................. Weissia
107(73).
Stem sclerodermis not or little differentiated; wide distribution ............................................................................. Gymnostomum
Stem sclerodermis clearly differentiated ...................................................................................... ............................................ ! 08
108(1 07). Ventral costal epidermis absent; austral regions and Mediterranean climates of the North Temperate Zone ........... Triquetrella
Ventral costal epidermis present .............................................................................................................................................. 109
109(108). Perichaetium terminal on main axis ................................................................................ .................. ....................................... 110
Perichaetium lateral on main axis at ends of very short branches; wide distribution ................................................... Molendoa
110(109). Transverse section of dorsal costal stereid band round to semicircular ................................................................................... 111
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 114
111(110). Leaf plane to broadly channeled ventrally, costa lacking a ventral groove .............................................. ...... ...... ................... 112
Leaf channeled or keeled, costa with a rather deep, narrow groove ventrally ......................................................................... 113
112(111). Length of stem less than 1.0 em, usually less than 0.6 em; basal cells inflated and bulging, considerably wider than upper
medial cells; wide distribution .................................................................................................................................. Gyroweisia
Length of stem usually 1.0 em or more; basal cells usually with straight walls, little wider than upper medial cells; wide distribution ................................................................................................................................... .................................. Didymodon
113(111 ). Basal cells usually with straight walls, little wider than upper medial cells; wide distribution ............... .................. ...... .Barbula
Basal cells inflated and bulging, considerably wider than upper medial cells; wide distribution .............................. Gyroweisia
114(110). Leaf plane to broadly channeled ventrally, costa lacking a ventral groove ............................................................................. 115
Leaf channeled or keeled, costa with a rather deep, narrow groove ventrally ................................................................. .Barbula
115( 114). Leaves widest at or above midleaf; axillary hair basal cell walls hyaline and all cells of hair similar; peristome teeth absent;
leaves tubulose when dry; wide distribution ............................................................................... ..................................Hyophila
Leaves widest below midleaf; axillary hairs with 1 or more basal cells with thicker or darker colored walls; peristome teeth
present; leaves occasionally channeled but not distinctly tubulose when dry; wide distribution ............................. Didymodon
116(72).
Stem hyalodermis absent ................................................................................................... ....................................................... 117
Stem hya1odermis present (sometimes indistinct); tropics ............................................................................Pseudosymblepharis
117(116). Stem sclerodermis not or little differentiated; Philippines ................................................................................... Pachyneuropsis
Stem sclerodermis clearly differentiated .................................................................................................................................. 118
118( 117). Length of stem less than 1.0 em, usually less than 0.6 em; basal cells inflated and bulging, considerably wider than upper
medial cells; wide distribution ........................................................................................... .................................... ... Gyroweisia
Length of stem usually 1.0 em or more; basal cells usually with straight walls, little wider than upper medial cells; wide distribution ..................................................................................................................................................................... D idymodon
119(71). . Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border ....................................... .................................................................................................................. ................................. 120
Basal cell group differentiated straight across leaf or rising higher medially .......................................................................... 125
THE POTTIACEAE
63
120(119). Ventral costal stereid band smaller than the dorsal or of nearly equal size ............................................................................. 121
Ventral costal stereid band distinctly larger than the dorsal .................................................................................................... 124
121(120). Ventral costal epidermis absent; stomates absent; capsule cleistocarpous and rupturing mainly along weak transverse walls at
butt ends of long-rectangular exothecial cells; central North America, Europe .......................................................... Aschisma
Ventral costal epidermis present; stomates present; capsule rupturing irregularly if cleistocarpous, or stegocarpous ........... 122
122(121). Upper !aminal margins not sharply and narrowly infolded; guide cells commonly more than 6 ............................................ 123
Upper !aminal margins sharply and narrowly infolded; guide cells 2-6; wide distribution ............................................. Weissia
123(122). Medial upper lamina! cells small, 7-10 11m in width; papillae per lumen one or occasionally two; calyptra papillose, distinctly roughened or strongly mamillose; number of rows of cells across ventral surface of costa usually 6 or more; South
Africa ..................................................................................................................................................................... Hypodontium
Medial upper lamina! cells medium sized, 11-14 11m in width; papillae per lumen usually two or more; calyptra smooth or
nearly so; number of rows of cells across ventral surface of costa 4-6; Mexico, Andes, South Africa.............. Streptocalypta
124(120). Calyptra smooth or nearly so; papillae per lumen usually two or more; spore diameter 8-15 11m; upper marginal cells not
longer than medial cells (sometimes larger); tropics ..................................................................................Pseudosymblepharis
Calyptra papillose, distinctly roughened or strongly mamillose; papillae per lumen one or occasionally two; spore diameter
more than 15 11m; upper marginal cells rectangular and clearly longer than medial cells; South Africa .............. Hypodontium
125(119). Transverse section of dorsal costal stereid band round to semicircular; Mexico, West Indies .................................. Quaesticula
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 126
126(125). Transverse section of costa round or ovate or elliptical ........................................................................................................... 127
Transverse section of costa semicircular .................................................................................................................................. 128
127(126). Upper lamina! margins not sharply and narrowly infolded, with an intramarginal border of elongate cells; leaf base little
differentiated in shape; stem sclerodermis clearly differentiated; leaves widest at or above midleaf; stem hyalodermis
absent; West Indies ................................................................................................................................................ Teniolophora
Upper !aminal margins sharply and narrowly infolded, leaf not bordered intramarginally by differentiated cells; leaf base
clearly differentiated in shape; stem sclerodermis not or little differentiated; leaves widest below midleaf; stem hyalodermis
present (sometimes indistinct); wide distribution ........................................................................................................... Weissia
128(126). Perichaetium terminal on main axis; upper !aminal cell walls thin to evenly thickened, lumens quadrate to rounded ........... l29
Perichaetium lateral on main axis at ends of very short branches; upper !aminal cell walls clearly trigonous or porose, lumens
irregularly angular or stellate; Brazil, India, Philippines ...................................................................................Hymenostyliella
129(128). Upper lamina! margins not sharply and narrowly infolded; leaf base little differentiated in shape; stem sclerodermis clearly
differentiated; axillary hair basal cell walls hyaline and all cells of hair similar; leaves widest at or above midleaf; wide distribution ......................................................................................................................................................................... Hyophila
Upper !aminal margins sharply and narrowly infolded; leaf base clearly differentiated in shape; stem sclerodermis not or little differentiated; axillary hairs with 1 or more basal cells with thicker or darker colored walls; leaves widest below
midleaf; wide distribution ............................................................................................................................................... Weissia
130(69).
Ventral costal outgrowths absent ............................................................................................................................................. 131
Ventral costal outgrowths present as a pad of cells, lamellae or filaments .............................................................................. 147
131(130). Dorsal costal epidermis absent ................................................................................................................................................. 132
Dorsal costal epidermis present ................................................................................................................................................ 138
132(131). Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border .......................................................................................................................................................................................... 133
Basal cell group differentiated straight across leaf or rising higher medially .......................................................................... 135
133(132). Ventral costal epidermis absent; stomates absent; calyptra papillose, distinctly roughened or strongly mamillose; leaf apex
obtuse to broadly acute; capsule cleistocarpous and rupturing mainly along weak transverse walls at butt ends of longrectangular exothecial cells; central North America, Europe ...................................................................................... Aschisma
Ventral costal epidermis present; stomates present; calyptra smooth or nearly so; leaf apex narrowly acute to acuminate; capsule rupturing irregularly if cleistocarpous, or stegocarpous ................................................................................................. 134
64
GENERA OF THE POTTIACEAE
134(133). Margins plane or very weakly incurved; upper !aminal margins not sharply and narrowly infolded; leaf base little
differentiated in shape; costal ventral cells longitudinally elongate 3: 1 or more; stem hyalodermis absent; South America,
Africa, Oceania ............................................................................................................................... ................. Trachycarpidium
Margins clearly incurved to involute; upper !aminal margins sharply and narrowly infolded; leaf base clearly differentiated
in shape; costal ventral cells quadrate to very short-rectangular; stem hyalodermis present (sometimes indistinct); wide distribution ................................................................................... ........................................................................................ Weissia
135(132). Transverse section of dorsal costal stereid band round to semicircular; Central and South America, Australia ......... Uleobryum
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 136
136(135). Leaf ventral surface above midleaf nearly plane to broadly channeled; margins plane to incurved or involute; narrow groove
along ventral surface of costa absent; costal ventral cells quadrate to very short-rectangular; transverse section of costa
round to semicircular ................................................................................................................................ ............................. 137
Leaf ventral surface above midleaf keeled; margins recurved; rather deep, narrow groove along ventral surface of costa
present; costal ventral cells longitudinally elongate 3:1 or more; transverse section of costa distinctly flattened, usually
reniform; Europe, North Africa, southern and eastern Asia ...................................................................... ."........ ..... Dialytrichia
137(136). Margins plane or very weakly incurved; upper !aminal margins not sharply and narrowly infolded; leaves widest at or above
midleaf; axillary hair basal cell walls hyaline and all cells of hair similar; stem sclerodermis clearly differentiated; Australia ........................................................................................................ ............................................................... Calymperastrum
Margins clearly incurved to involute; upper !aminal margins sharply and narrowly infolded; leaves widest below midleaf;
axillary hairs with 1 or more basal cells with thicker or darker colored walls; stem sclerodermis not or little differentiated;
wide distribution .............................................................................................................. .............................. ................. Weissia
138(131). Capsule cleistocarpous ............................................................................................... ....................... ......................... .............. 139
Capsule stegocarpous ..................................................... ............................................................ .......................................... .... 142
139(138). Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border ... .............................. .......................................................................................... .................. .................. ........................... 140
Basal cell group differentiated straight across leaf or rising higher medially ................ .................. ...... .................................. 141
140(139). Margins plane or very weakly incurved; upper !aminal margins not sharply and narrowly infolded; leaf base little
differentiated in shape; costal ventral cells longitudinally elongate 3:1 or more; stem hyalodermis absent; South America,
Africa, Oceania ................................................................................................................................................ Trachycarpidium
Margins clearly incurved to involute; upper !aminal margins sharply and narrowly infolded; leaf base clearly differentiated
in shape; costal ventral cells quadrate to very short-rectangular; stem hyalodermis present (sometimes indistinct); wide distribution ..................................................................................................................................................... ............ .......... Weissia
141(139). Margins plane or very weakly incurved, not sharply and narrowly infolded above; stem hyalodermis absent; capsule cleistocarpous and rupturing mainly along weak transverse walls at butt ends of long-rectangular exothecial cells; austral
areas ...................... ........................................................................................................................ ........................... Tetrapterum
Margins clearly incurved to involute, sharply and narrowly infolded above; stem hyalodermis present (sometimes indistinct);
capsule rupturing irregularly if cleistocarpous, or stegocarpous; wide distribution ....................................................... Weissia
142(138). Costal ventral cells quadrate to very short-rectangular ............................................................................................................ 143
Costal ventral cells longitudinally elongate 3:1 or more .......................................................................................................... 146
143(142). Transverse section of dorsal costal stereid band round to semicircular; American tropics, South Africa, Burma ........ Plaubelia
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 144
144(143). Upper lamina! margins not sharply and narrowly infolded; axillary hair basal cell walls hyaline and all cells of hair similar ........................................................................................................................................................................................... 145
Upper lamina! margins sharply and narrowly infolded; axillary hairs with 1 or more basal cells thicker or darker colored;
wide distribution .......................................................................................................................... ................................... Weissia
145(144). Transverse section of costa round to semicircular; peristome teeth absent; leaves widest at or above midleaf; leaves tubulose
when dry; calyptra 1.0-3 .0 mm; wide distribution ............... .................................................................. ...................... Hyophila
Transverse section of costa distinctly flattened, usually reniform; peristome teeth present; leaves widest below midleaf;
leaves when dry occasionally channeled but not distinctly tubulose; calyptra more than 3.0 mm; wide distribution ..... ........................................................................................................................ .................. .................. Pseudocrossidium
THE POTTIACEAE
65
146(142). Transverse section of costa round to semicircular; theca short, less than 1.5 mm in length; upper marginal cells in same number of layers as medial cells; wide distribution ....... ................ ..... ......................................... ...... .. ............................. .....Barbula
Transverse section of costa distinctly flattened, usually reniform; theca longer, usually more than 1.5 mm in length; upper
marginal cells differentiated as a bistratose (or more) border; Europe, North Africa southern and eastern Asia ... Dialytrichia
147(130). Ventral costal outgrowths differentiated as filaments or lamellae; margins recurved to revolute; transverse section of dorsal
costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; leaf apex flattened, channeled or
keeled; transverse section of costa distinctly flattened, usually reniform; wide distribution ........................ Pseudocrossidium
Ventral costal outgrowths differentiated as a bulging pad of cells; margins plane to incurved or involute; transverse section
of dorsal costal stereid band round to semicircular; leaf apex cucullate; transverse section of costa round to semicircular;
southwestern USA south through Andes ........................................................... .................. ................ ................... Globulinella
148(68).
Margins denticulate only near leaf base or at top of leaf sheath .............................................................................................. 149
Margins denticulate or serrulate to toothed near apex or throughout ................... ................................. ....... ............................ l50
149(148). Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; dorsal costal epidermis absent; superficial walls of upper !aminal cells ventrally bulging-mamillose, weakly convex
dorsally; stem sclerodermis not or little differentiated; peristome teeth present; South Africa ............................ Hypodontium
Basal cell group differentiated straight across leaf or rising higher medially; dorsal costal epidermis present; superficial walls
of upper !aminal cells similarly shaped on both sides of lamina; stem sclerodermis clearly differentiated; peristome teeth
absent; wide distribution .................................................................................................................. ........................... Molendoa
150(148). Dorsal costal epidermis absent ................................................................................................................... ,....... ...................... 151
Dorsal costal epidermis present ................................................................................................................................................ l57
151(150). Ventral costal stereid band smaller than the dorsal or of nearly equal size; upper lamina! margins not sharply and narrowly
infolded ............................ ..... .. ................. ............................... ......................... .................................... .................................. 152
Ventral costal stereid band distinctly larger than the dorsal; upper !aminal margins sharply and narrowly infolded near apex;
Southeast Asia, Borneo ................. ............... ................... .. ....................................................................................... Chionoloma
152(151). Ventral costal epidermis absent; theca spherical. .......................................... ......... ................................. ................................. l53
Ventral costal epidermis present; theca ovoid to cylindrical .... .......................................................... ...................... ............... 154
153(152). Superficial exothecial cell walls with a central and lens-like thickening; costal ventral cells longitudinally elongate 3:1 or
more; number of rows of cells across ventral surface of costa commonly 4 or more; axillary hair basal cell walls hyaline
and all cells of hair similar; papillae per lumen one or occasionally two ......... ................... ............... .............. Bryoceuthospora
Superficial exothecial cell walls thickened; costal ventral cells quadrate to very short-rectangular; number of rows of cells
across ventral surface of costa usually 2 but up to 4; axillary hairs with 1 or more basal cells with thicker or darker colored
walls; papillae per lumen usually two or more; central North America, Europe ........... .......................................... ... Aschisma
154(152). Perichaetium terminal on main axis; perichaetial leaves similar to cauline leaves or occasionally smaller or somewhat
enlarged ........... ................................................................................................................ ... ...... .............................................. l55
Perichaetium lateral on main axis at ends of very short branches; perichaetial leaves distinctly different in size or morphology, strongly sheathing the seta; wide distribution ................................................. ..... ............... .......... ................. Pleurochaete
155(154). Guide cells 2-6; length of stem usually 1.0 em or more .......................................... .................. ............ .................................. l56
Guide cells commonly more than 6; length of stem less than 1.0 em, usually less than 0.6 em; Mexico, Andes, South
Africa .. ...... .................. ...... .................. .................................................................................................. ....... ........ Streptocalypta
156(155). Stem hyalodermis absent; American tropics, eastern Asia ........ ....... ......... ....................... .......... .. ........................ ...Tuerckheimia
Stem hyalodermis present (sometimes indistinct); wide distribution .................. ................ ...................................Trichostomum
157(150). Superficial walls of upper !aminal cells similarly shaped on both sides of lamina ................ ......... ......................................... 158
Superficial walls of upper !aminal cells ventrally bulging-mamillose, weakly convex dorsally ...... ......... ........ ................ ...... 165
158(157). Rather deep, narrow groove along costa absent .... ............ ....................... .................................... ............................................ 159
Rather deep, narrow groove along costa present. ............. ....... .............................. ................. .................................................. 164
159(158). Margins plane to incurved or involute ..................................................... ......... ................................................ ....................... 160
Margins recurved to revolute ..................................... ,.................... ................................. ........................................................ 163
66
GENERA OF THE POTTIACEAE
160(159). Perichaetium terminal on main axis; upper marginal cells not longer than medial cells (sometimes larger); perichaetialleaves
similar to cauline leaves or occasionally smaller or somewhat enlarged .............................................................................. 161
Perichaetium lateral on main axis at ends of very short branches; upper marginal cells rectangular and clearly longer than
medial cells; perichaetial leaves distinctly different in size or morphology, strongly sheathing the seta; wide distribution ................................................................................................................................... ...................................... Pleurochaete
161(160). Leaves widest at or above midleaf; wide distribution ..................................................................................................... Hyophila
Leaves widest below midleaf ......................................................................................................................................... .......... 162
162(161). Stem hyalodermis absent; American tropics, eastern Asia ...................................................................................... Tuerckheimia
Stem hyalodermis present (sometimes indistinct); wide distribution ............................ .................................... ..... Trichostomum
163(159). Leaves widest at or above midleaf; axillary hair basal cell walls hyaline and all cells of hair similar; peristome teeth absent;
leaves tubulose when dry; wide distribution ............................................................. .................................................... Hyophila
Leaves widest below midleaf; axillary hairs with I or more basal cells with thicker or darker colored walls; peristome teeth
present; leaves when dry occasionally channeled but not distinctly tubulose; wide distribution ............ ....... :.... ..... Didymodon
164(158). Ventral costal epidermis absent; basal cell group not or little differentiated from upper medial cells; theca longer, usually
more than 1.5 mm in length; calyptra short, less than 1.0 mm; stem transverse section triangular; austral areas and Mediterranean climates of the North Temperate Zone ......................................................................................................... Triquetrella
Ventral costal epidermis present; basal cell group clearly differentiated, usually larger, less papillose, walls thinner; theca
short, less than 1.5 mm in length; calyptra longer, 1.0 mm or more in length; stem transverse section rounded-pentagonal;
wide distribution .................................................................................................................... .........................................Barbula
165(157). Hydroid strand absent. .................................................................................................................. ............................................ l66
Hydroid strand present ......................................................................................................................... .................................... 169
166(165). Basal cell group differentiated as a vee, with at least laterally differentiated cells rising higher marginally as a tapering border; leaf base clearly differentiated in shape; wide distribution .................................. ........ .................................. Pleurochaete
Basal cell group differentiated straight across leaf or rising higher medially; leaf base little differentiated in shape ............ 167
167(166). Transverse section of costa round or ovate or elliptical; upper marginal cells differentiated intramarginally as a bistratose
border; upper marginal cells rectangular and clearly longer than medial cells; marginal cell walls thicker than those of
medial cells; West Indies ................................................................................................................. ...................... Teniolophora
Transverse section of costa semicircular; upper marginal cells in same number of layers as medial cells; upper marginal cells
not longer than medial cells (sometimes larger); marginal cell walls not thicker than those of medial cells ....................... 168
168(167). Leaves widest at or above midleaf; upper !aminal cell walls thin to evenly thickened, lumens quadrate to rounded; axillary
hair basal cell walls hyaline and all cells of hair similar; perichaetium terminal on main axis; annulus of vesiculose cells,
often in two or more rows; wide distribution ........ ........................ .............................. .................................. ................ Hyophila
Leaves widest below midleaf; upper !aminal cell walls clearly trigonous or porose, lumens irregularly angular or stellate;
axillary hairs with I or more basal cells with thicker or darker colored walls; perichaetium lateral on main axis at ends of
very short branches; annulus of weakly differentiated cells; Brazil, India, Philippines .................................... Hymenostyliella
169(165). Leaf base little differentiated in shape; guide cells 2-6; seta nearly absent to short, less than 1 em; upper !aminal cells medially unistratose ....................................................................................................................................................................... 170
Leaf base clearly differentiated in shape; guide cells commonly more than 6; seta elongate, 1 em or longer; upper !aminal
cells medially bistratose, the cells staggered, not one directly above the other in section; wide distribution ............ Timmiella
170(169). Transverse section of dorsal costal stereid band round to semicircular; axillary hairs with 1 or more basal cells with thicker
or darker colored walls; peristome teeth present; American tropics, South Africa, Burma ........................................ Plaubelia
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped; axillary
hair basal cell walls hyaline and all cells of hair similar; peristome teeth absent; wide distribution ............. .............. Hyophila
171(51).
171-181 Costa with two stereid bands, upper !aminal cells KOH red
Margins entire or minutely and evenly crenulate .............................................................................. ...... ................................. 172
Margins denticulate or serrulate to toothed .............................................................................................................................. 179
172(171 ). Transverse section of dorsal costal stereid band round to semicircular ................................................................................... 173
Transverse section of dorsal costal stereid band clearly flattened or ventrally indented, reniform or crescent-shaped .......... 176
THE POTTIACEAE
67
173(172). Margins plane to incurved or involute; costal hydroid strand present ..................................................................................... 174
Margins recurved to revolute; costal hydroid strand absent..................................................................................................... 175
174(173). Dorsal costal epidermis absent; medial upper larninal cells commonly larger than 14 I.J.m; transverse section of costa round or
ovate or elliptical; number of rows of cells across ventral surface of costa usually 2 but up to 4; superficial walls of upper
laminal cells flat or very weakly convex on both sides of lamina; southern South America, Antarctica ............. Sarconeurum
Dorsal costal epidermis present; medial upper laminal cells 7-141.J.m in width; transverse section of costa semicircular; number of rows of cells across ventral surface of costa commonly 4 or more; superficial walls of upper laminal cells strongly
convex to bulging on both sides of lamina; Andes ...................................................................................... Erythrophyllastrum
175(173). Rather deep, narrow groove along costa absent; peristome teeth present; spore diameter 8-15 j.l.m; wide distribution ............................................................................................................................................................................ Didymodon
Rather deep, narrow groove along costa 'present; peristome teeth absent; spore diameter more than 15 j.l.m; USA, Mexico,
India, eastern Asia ........................................................................................................................................... ,....... Bellibarbula
176(172). Stem sclerodermis not or little differentiated; Andes ........................................................................................ Erythrophyllopsis
Stem sclerodermis clearly differentiated .................................................................................................................................. 177
177(176). Stem hyalodermis absent ......................................................................................................................................................... 178
Stem hyalodermis present (sometimes indistinct); wide distribution ........................................................... Bryoerythrophyllum
178( 177). Hydroid strand absent; rather deep, narrow groove along costa absent; number of rows of cells across ventral surface of costa
usually 2 but up to 4; superficial walls of upper laminal cells strongly convex to bulging on both sides of lamina; wide distribution ..................................................................................................................................................................... Didymodon
Hydroid strand present; rather deep, narrow groove along costa present; number of rows of cells across ventral surface of
costa commonly 4 or more; superficial walls of upper laminal cells flat or very weakly convex on both sides of lamina;
Mexico south through Andes ..........................................................................................................................................Mironia
179(171). Ventral costal epidermis absent; southwestern USA, Mexico, Andes .................................................................... Rhexophyllum
Ventral costal epidermis present .............................................................................................................................................. 180
180(179). Stem hyalodermis absent .......................................................................................................................................................... 181
Stem hyalodermis present (sometimes indistinct); wide distribution ........................................................... Bryoerythrophyllum
181 (180). Hydroid strand absent; rather deep, narrow groove along costa absent; number of rows of cells across ventral surface of costa
usually 2 but up to 4; superficial walls of upper laminal cells strongly convex to bulging on both sides of lamina; wide distribution ..................................................................................................................................................................... Didymodon
Hydroid strand present; rather deep, narrow groove along costa present; number of rows of cells across ventral surface of
costa commonly 4 or more; superficial walls of upper laminal cells flat or very weakly convex on both sides of lamina;
Mexico south through Andes ..........................................................................................................................................Mironia
Subfamily TIMMIELLOIDEAE
Timmielloideae Zand., subfam. nov. Type: Timmie/la (De Not.) Limpr.
Plantae rosulatae; filum centrale robustum hyalodermidem evolvens; folia in statu sicco tubulosa marginibus planis, serrulatis
praedita; costa percurrens, lata, stratis stereidarum duobus et filo hydroideo praedita; cellulae supernae laminales in regione folii
mediana bistratosae, juxta costam altero strato cellularum supra alterum excentrico praeditae, ventra/iter tumescentes, dorsa/iter
subconvexae; dentes peristomii stricti vel sinistrorsum extus visi subtorti.
Plants rosulate; central strand strong, hyalodermis present; leaves tubulose when dry, margins plane, serrulate; costa percurrent,
broad, with two stereid bands, hydroid strand present; upper !aminal cells bistratose in the middle portion of the leaf and not directly
stacked over each other near the costa, superficially bulging ventrally and weakly convex dorsally; peristome teeth straight or
twisted weakly clockwise.
Of all terminal taxa, Timmie/la, the sole member of this subclade, shares the most plesiomorphic characters with the outgroups
Polytrichum, Timmia and Ptychomitrium (see cladograms and discussion above). The distinguishing autapomorphy is !aminal cells
medially bistratose but not situated directly over one another near the costa (as seen either from above or in transverse section). This
unusual feature is matched in Diphyscium (Diphysciaceae), a genus with many of the characteristics of the Pottiaceae (see discussion above). The following major traits in combination aid in distinguishing the genus: costa very wide, with multiple hydroid
strands; leaf cells epapillose, ventrally bulging and dorsally nearly flat; peristome twisted clockwise (rare in the Pottiaceae) or
straight. The distribution of the subfamily is rather general, being most common in temperate areas.
Plates 1-2
1. TIMMIELLA
Timmie/la (De Not.) Limpr., Laubm. Deutchl. 1: 590, 1888. Type:
Timmie/la anomala (BSG) Limpr.
Trichstomum subg. Eutrichum Schimp., Corol. 28, 1856.
Barbula subg. Timmie/la (De Not.) Kindb., Eur. N. Amer.
Bryin. 2: 245, 1897.
Barbula sect. Anomalae BSG, Bryol. Eur. 2: 75, 107 (fasc.
13-15 Mon. 13. 45), 1846.
Trichostomum sect. Timmiella De Not., Cronac. Briol. Ital. 1:
14, 1866.
From Timmia, a genus + -ella, diminutive; resembling the genus
Timmia.
Plants loosely caespitose to cushion-forming, often rosulate,
green or often dark green above, brown below. Stems branching
irregularly or simple, to 1.5 em in length, transverse section
rounded-pentagonal, often flattened, central strand very strong,
sclerodermis of 1-3 layers (occasionally substereid), hyalodermis
present, weak; axillary hairs of ca. 7 cells, all hyaline or basal cell
frrm-walled; sparsely radiculose. Leaves incurved and tubu/ose
when dry, spreading when moist, long-elliptical to ligulate or
broadly /anceo/ate, often wasp-waisted, 3.5-5.0 mm in length,
upper lamina broadly channeled across leaf, margins plane to
weakly incurved, distantly denticulate or serrulate to near base,
usually strongly serrate near apex, lamina bistratose except along
margins; apex acute; base often broadened, elliptical, somewhat
sheathing, occasionally with distinct shoulders; costa percurrent,
tapering to apex and much broadened below midleaf, superficial
cells ventrally quadrate and bulging, dorsally elongate and
smooth, 6-19 rows of cells across costa ventrally at midleaf,
costal transverse section flattened, reniform or elliptical, two
stereid bands present, ventral epidermis present, this bistratose or
occasionally tristratose, dorsal epidermis present, unistratose,
guide cells 6-16 in !layer, hydroid strand present, often multiple,
often on both ventral and dorsal sides of guide cells; upper
/aminal cells quadrate to rounded-hexagonal, 9-12 J..lm in width,
1:1, walls mostly evenly thickened, lumens occasionally rounded,
superficially bulging ventrally and nearly flat dorsally, cells
bistratose medially but near the costa not situated directly over
each other; papillae absent; basal cells differentiated straight
across leaf, bulging-rectangular, 10-18 J..lm in width, mostly
3-4: 1, walls thin, hyaline to yellowish. Dioicous or monoicous
(autoicous, synoicous or apparently rhizautoicous). Perichaetia
terminal, inner leaves scarcely different from cauline leaves,
base often shortly and broadly clasping. Perigonia lateral on
archegoniophore stem, as somewhat flattened buds, or very
small and terminal on perigoniate plant. Seta ca. 1.0-2.0 em in
length, 1 per perichaetium, yellowish to reddish brown, twisted
clockwise; theca 3.0-5.0 mm in length, brown, long-elliptical to
long-cylindrical, occasionally somewhat ventricose, exothecial
cells 13-20 J..lm in width, ca. 4-6:1, evenly thick-walled,
stomates phaneropore, on neck of theca, annulus of weakly
differentiated cells or of up to several layers of highly
vesiculose cells, revoluble or deciduous in pieces; peristome
teeth of 16 paired teeth or 32 evenly spaced rami, linearlanceolate to filamentous, papillose to closely branchingspiculose, occasionally also striate below, rudimentary or to
300-700 J..lm, with several articulations, straight or twisted
clockwise weakly once or less, basal membrane absent or very
low, essentially smooth. Operculum long-conic to rostrate,
0.6-1.7 mm in length, cells in straight rows or twisted clockwise. Calyptra cucullate, smooth, ca. 3.5 mm in length. Spores
10-13 J..lm in diameter, yellowish brown, weakly papillose.
Lamina/ color reaction to KOH yellow. Reported chromosome
numbern= 13, l3+m, 14, l4+m, 14+m+lacc, 15.
Found on rock and soil; in arid lands and mountainous areas
of North and South America, Europe, Asia and Africa.
Timmiella is characterized by the following combination of
distinguishing features: very strong stem central strand (Pl. 1, f.
7); lamina! margins plane to weakly incurved, denticulate to
dentate; costa very wide, with multiple hydroid strands; !aminal
cells medially bistratose but not vertically aligned near the
costa, epapillose, ventrally bulging and dorsally nearly flat (Pl.
I, f. 2); peristome straight (Pl. 2, f. 6) or twisted clockwise (Pl.
l, f. 5).
Timmie/la is one of only two genera in the Pottiaceae with
TIMMIELLOIDEAE
69
Plate 1. Timmiella. 1-9. T. anomala. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Clustered lateral perigoniate buds. 9. Peristome. 10-13. T. barbuloides. 10-11. Two leaves. 12. Leaf apex. 13. Peristome.
70
GENERA OF THE POTTIACEAE
peristome teeth (in at least some species) twisted clockwise
(Leptodontiella, with one species, has teeth that are straight or
twisted weakly clockwise). Characters important in Timmie/la are
variations in sexual condition, annulus development, and peristome size and degree of torsion. Timmie/la corniculata is
dioicous, contrary to Brotherus' (1924-25) key. Propagula are
only doubtfully present in the genus; small, clavate, multicellular
propagula, ca. 50-70 J.lm in length, were found loosely associated
with gametophores in the type ofT. brevidens, but may be due to
admixture of other species.
Additional literature: Castaldo-Cobianchi et al. (1982),
Chopra and Kapur (1989), Gambardella et al. (1980), Kapur
(1989, 1991), Ligrone (1986), Ligrone et al. (l980a,b).
Number of accepted species: 13.
Species examined: T. anomala, T. barbuloides, T. brevidens
(BM), T. corniculata (NY), T. diminuta (NY), T. flexiseta (NY),
T. pelindaba (NY), T. subintegra (BM), T. umbrosa (NY).
New heterotypic synonymy: Timmie/la argentinica Broth. =
Timmie/la acaulon (C. Miill.) Zand.
New combinations: Timmie/la acaulon (C. Miill.) Zand.,
comb. nov. (Trichostomum acaulon C. Miill., Linnaea 42: 320,
1879; Tortella acaulon (C. Miill.) Broth.).
Plate 2. Timmiella. 1-4. T. brevidens. 1-2. Two leaves. 3. Leaf apex. 4. Peristome. 5. T. corniculata. 5. Capsule (theca, peristome and
operculum). 6. T.flexiseta. 6. Capsule. 7-9. T. pelindaba. 7-8. Two leaves. 9. Leaf apex.
Subfamily ERYTHROPHYLLOPSOIDEAE
Erythrophyllopsoideae Zand., subfam. nov. Type: Erythrophyllopsis Broth. in Herz.
Plantae magniusculae, altitudine 2.0-3.5 cm;folia lanceo/ata marginibus planis, apicibus acutis, basibus vaginantibus, laminis in
solutione KOH rubrescentibus, costae cellulas ducium 4-6 et trans superficiem ventra/em cel/ulas 10-16 evolventi praedita.
Plants rather large, 2.0-3.5 em in height, leaves lanceolate, with plane margins, apices acute, bases sheathing; upper lamina
KOH red, costa with 4-6 guide cells and 10-16 cells across the ventral surface.
With Timmie/la as outgroup, this subclade is distinguished from lower branches on the tree by the advanced character of costal
guide cells fewer, 2-6. The general appearance of the two included genera, Erythrophyllastrum and Erythrophyl/opsis, is intermediate between that of the traditional Trichostomoideae and the Barbuloideae, especially in the lanceolate leaves with plane margins.
Other distinguishing traits are the acute leaf apex, sheathing leaf base, KOH red upper lamina, two costal stereid bands, 4-6 guide
cells, and many (10-16) rows of cells across ventral surface of costa. The species of this small subfamily are rare and restricted to
the Andes.
2. ERYTHROPHYLLOPSIS
Plate 3.
Erythrophyllopsis Broth. in Herz., Biblioth. Bot. 87: 41, 1916.
Type: Erythrophyllopsis boliviano Broth., Bolivia, Cerros de
Malaga, Herzog 4371, syntype, M, also Tunarisee, Herzog
4765, syntype, NY.
From Erythrophyl/um, a genus + ~'ljf<n~. -£~, appearance;
resembling the genus Erythrophyllum.
Plants forming cushions, dark green to reddish brown above,
reddish brown below. Stems branching occasionally, to 2-3 em in
length, transverse section rounded-triangular to pentagonal, central strand strong, sclerodermis weakly developed, substereid,
hyalodermis of 1 layer of usually collapsed cells; axillary hairs ca.
10 cells in length, the basal 1-2 cells more flllll walled or brown,
otherwise hyaline; sparsely radiculose. Leaves appressed below
and incurled above when dry, squarrose when moist, longlanceolate, 3.5-4.5 mm in length, upper lamina broadly channeled to keeled, margins plane, entire, occasionally denticulate at
leaf apex; apex narrowly acute; base strongly sheathing, with distinct shoulders, bordered by 4-6 rows of narrower, often thickerwalled cells; costa percurrent or ending in an apiculus, rather
broad below midleaf, lamina inserted about 45', costal superficial
cells quadrate and papillose, but becoming smooth and elongate
in distal 1/4 of leaf ventrally, dorsally elongate, ca. 10 rows of
cells across costa ventrally at midleaf, costal transverse section
reniform, stereid bands ventrally distinct, the dorsal crescentshaped, epidermis present ventrally and dorsally, guide cells 4-6
in l layer, l-3 hydroid strands usually present, weakly distinguishable; upper lamina/ cells bistratose across the leaf, quadrate,
7-12 J.l.m in width, l(-2):1(-2), walls evenly thickened, superficially weakly convex on both sides of lamina; papillae bifid, 2-6
per lumen , low, scab-like, flattened, crowded, solid or hollow;
basal cells sharply differentiated nearly across the leaf base but
rising highest medially, rectangular, ca. 9-12 j.lm (to 16 j.lm
above) in width, mostly ~5:1, walls thin. Dioicous. Perichaetia
terminal, inner leaves little different from the cauline, sheathing
the seta. Perigonia terminal, weakly gemmate, inner leaves shortly
lanceolate. Seta 0.5-1.5 em in length, 1 per perichaetium, reddish
brown, twisted weakly clockwise; theca ca. 3.3 mm in length, reddish brown, cylindrical, exothecial cells rectangular, 3-5: l, ca.
13-18 j.lm in width, thin-walled, stomates at base of capsule,
phaneropore, annulus of ca. 3-4 rows of vesiculose cells, persistent; peristome teeth much reduced or 16, truncate to shortly
lanceolate, papillose, 50-130 j.lm in length, with up to 5 articulations, straight, basal membrane low or absent in height, papillose. Operculum conic-rostrate, ca. 1 mm in length, cells
straight. Calyptra not seen. Spores 13-15 J.l.m in diameter, light
brown, essentially smooth. Lamina/ KOH color reaction red.
A monotypic genus of the Andes of Bolivia and Argentina,
found on moist rocks at high elevations.
Erythrophyllopsis, known from one species, E. fuscula , is
characterized by a combination of characters: red coloration in
KOH, leaves bistratose above (Pl. 3, f. 5), with plane margins
and a strongly differentiated sheathing leaf base having
"shoulders" (Pl. 3, f. 3-4) at the top of the base, and peristome
rudimentary or short (illustrated by Herzog 1916). It is similar
to Bryoerythrophyllum by the red coloration, leaves with
differentiated leaf base and upper lamina! cell walls evenly
thickened, with papillae crowded and bifid to multiplex. My
previous treatment of this genus (Zander 1977a) included
Trichostomum andinum, a morphologically similar taxon here
placed in a separate genus, Erythrophyllastrum, (q .v. The upper
lamina of Erythrophyllopsis, which has been figured by previous authors (Herzog 1916; Brotherus 1924-25; Hilpert 1933), is
entirely bistratose. Mironia is similar but has leaves bistratose
only along the upper margins, and has a well developed peristome. Leptodontium has a similarly lanceolate leaf with a
broad, sheathing base, but is yellow in KOH and lacks a stem
central strand, among other differences. There may be a confusion with Didymodon sect. Vineales because of the lanceolate,
KOH-red leaves with dorsally quadrate costal cells; the leaf
apex has ventrally elongate costal cells in a boat-shaped groove
and is occasionally abruptly constricted, cf D . occidentalis.
This last taxon, however, has a quite different costa, it being
dorsally rounded and protuberant, and the guide cells commonly
in two layers. Hilpert (1933) placed Erythrophyllopsis in close
relationship with what are here dealt with as Didymodon sect.
Asteriscium, Gertrudiella and Bryoerythrophyllum.
Erythrophyllopsis challaensis is here transferred to Didymodon ((q.v.). It differs from E. fuscula in slightly larger upper
lamina! cells, weakly serrulate basal marginal cells, shorter and
broader basal cells which are transversely slit through cell wall
resorption of the medial portion of the longitudinal cell walls,
and a yellow color reaction to KOH.
Number of accepted species: I.
Species examined: E. fuscula (M, NY).
72
GENERA OF THE POTIIACEAE
New heterotypic synonymy: Didymodon semivaginatus
(Britt.) Broth. = Erythrophyllopsis fuscula (C. Miill.) Hilp.
3. ERYTHROPHYLLASTRUM
Plate 4.
Erythrophyllastrum Zand., gen. nov. Type: Erythrophyllastrum
andinum (Sull.) Zand. Holotype: Peru, Andes, U.S. Expl. Exp.
Wilkes, 1838-1842, FH.
Plantae magnae, usque altitudine 3.5 em; caules in sectione
interdum triangulares, filo centrali robusto praediti, hyalo-
dermidem e stratis 1-3 texturae laxae compositam in maturitate
subcol/apsam evolventes; folia in statu madido late patentia,
late lanceolata, longitudine 3.5-4.5 mm, marginibus planis,
integris, uni- vel bistratosis, apice acuto, base subvaginanti,
lamina superna in solutione KOH rubrescenti, costa in sectione
semicirculari strata stereidarum duo evolventi praedita.
From Erythrophyllum, a genus + -astrum, an ending implying deception; mimicking Erythrophyllum.
Plants forming cushions, green to reddish brown above,
reddish brown below. Stems branching irregularly, to 3.5 em in
Plate 3. Erythrophyllopsis. 1-7. E.fuscula. l. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Transverse section at midleaf. 6. Leaf
apex. 7. Basal cells.
. ERYTHROPHYLLOPSOIDEAE
73
Plate 4. Erythrophyllllstrum. 1-13. E. andinum. I. Habit. 2. Transverse section of stem. ~. Four leaves, one enlarged. 7-8. Two transverse
sections, midleaf. 9. Upper lamina! papillae. 10. Leaf apex. 11. Basal cells. 12. Rudimentary peristome. 13. Operculum.
74
GENERA OF THE POITIACEAE
length, transverse section rounded-pentagonal, central strand
strong, sc/erodermis weakly developed, substereid, hya/odermis
of 1(-3) eel/layers of lax cells, often not collapsed; axillary hairs
long, to 20 cells in length, basal 1-4 cells yellowish; radiculose
below. Leaves appressed-incurved and curled when dry, widely
spreading when moist, /anceo/ate, 2-3 mm in length, upper lamina broadly channeled to keeled, broadly grooved along costa,
margins plane, entire, upper lamina unistratose or variably
bistratose in large patches or throughout; apex acute; base
broadly sheathing below, with weak shoulders; costa percurrent,
with lamina inserted laterally, superficial cells quadrate, papillose,
often short-rectangular near apex ventrally, dorsally usually quadrate near apex, rectangular below, 10-16 rows of cells across
costa ventrally at mid/eaf, costal transverse section semicircular,
stereid bands two and strong, the dorsal semi-circular in shape,
ventral and dorsal epidermises present, the dorsal medially weak,
guide cells (4-)6 in 1 layer, hydroid strand(s) very weak, often
multiple; upper /aminal cells small, subquadrate, 7-10 Jlm in
width, 1:1, walls thin to evenly thickened, superficially weakly
convex on both sides of lamina; papillae bifid, 2-3 per lumen to
multiplex, generally hollow, crowded; basal cells differentiated
across the sheathing base, rectangular, ca. 13 Jlm in width, mostly
3-4:1, walls thin. Propagula not seen. Dioicous. Perichaetia terminal, inner leaves lanceolate, somewhat larger than the cauline, to
4.5 mm in length, sheathing the seta in lower 1/3, lower cells rectangular, walls weakly porose. Perigonia terminal, weakly
gemmate. Seta ca. 1 em in length, 1 per perichaetium, reddish
brown, twisted clockwise; theca 1.5-2.0 mm in length, reddish
brown, cylindrical, exothecia1 cells short-rectangular, ca. 2: 1,
thin-walled, stomates at base of capsule, phaneropore, annulus of
1-2 rows of vesiculose cells; peristome teeth rudimentary, consisting of a few low plates, to 30-45 Jlm in height. Operculum
long-rostrate, oblique, ca. 1.2 mm in length, cells straight. Calyp-
tra conic-cucullate, smooth, ca. 1.5 mm in length. Spores ca. 13
Jlm in diameter, light brown, weakly papillose. Lamina/ KOH
color reaction red.
A monotypic genus found in the Andes of Colombia and
Peru, on moist rock at high elevations (ca. 3700-4000 msm).
This taxon was previously placed with Erythrophyllopsis
(Zander 1977a), which is superficially similar by its red coloration; stem with large central strand; leaves with plane margins,
broad, flat costae and sheathing leaf bases; and upper lamina
bistratose. Erythrophyl/astrum differs from Erythrophyllopsis
by the former's 1(-3) layers of stem hyaloderm cells (Pl. 4, f.
2), comprising loose tissue little collapsed with maturity; stems
sometimes triangular in section; the leaves commonly shorter
and more broadly lanceolate, sometimes reflexed above the base
and widely spreading to squarrose; the leaf base less strongly
sheathing, sometimes without highly differentiated "shoulders"
(Pl. 4, f. 4); upper !aminal cells variably entirely bistratose or
bistratose in patches or entirely unistratose (Pl. 4, f. 7-8); both
costal section and dorsal stereid band semicircular or nearly so
(Pl. 4, f. 7-8, both being reniform in Erythrophyllopsis); dorsal
costal cells usually quadrate near the leaf apex; and peristome
even more reduced, to a few small plates (Pl. 4, f. 12).
Through its plane, reflexed, broadly keeled leaves with
sometimes weakly sheathing base, small, dense upper !aminal
cells, and basal cells usually differentiated completely across
the leaf, Erythrophyllastrum may have much the general aspect
of species of Barbula sect. Convolutae (e.g. B. amplexifolia).
Number of accepted species: 1.
Species examined: E. andinum (BUF).
New combination: Erythrophyllastrum andinum (Sull.)
Zand., comb. nov. (Trichostomum andinum Sull., U.S. Expl.
Exp. Wilkes Musci 5, 1859; Erythrophyllopsis andina (Sull.)
Zand.).
Subfamily GERTRUDIELLOIDEAE
Gertrudielloideae Zand., subfam. nov. Type: Gertrudiella Broth.
Plantae filum centra/em et hyalodermidem caulis evo/ventes; folia in statu madido squarrosa, magniuscula, longitudine 3.5-4.5
mm, apice acuto, base vaginanti, costa e strato stereidarum unico composita et cylindro multistratoso e cellulis ducium parietibus
crassis praeditis composito, cellulis laminae supernae so/urn in margine folii papillosis, in regione mediano ventra/iter
tumescentibus dorsa/iter planis sed in margine folii in aliquot seriebus utrinsecus tumescentibus praedita.
Plants with strong stem central strand, hyalodermis present; leaves squarrose when wet, rather large, 3.5-4.5 mm in length, apex
acute, base sheathing, costa with one stereid band and a multilayered cylinder of thick-walled guide cells; upper !aminal cells papillose only marginally, medially bulging ventrally and flat dorsally but marginally bulging on both sides in several rows.
This tribe is more highly evolved than the Timmielloideae by the advanced traits of leaf base not sheathing and upper lamina
unistratose. There are two autapomorphies: the upper !aminal cells medially ventrally bulging and dorsally flat but the marginal
cells bulging on both sides in several rows, and the costal guide cells forming a thick-walled, multilayered cylinder. This is a rare,
monotypic, Andean subfamily.
4. GERTRUDIELLA
Plate 5.
Gertrudiel/a Broth., Nat. Pfl. ed. 2, 11: 528, 1925. Type: Gertrudiel/a validinervis (Herz.) Broth.
Gertrudia Herz., Biblioth. Bot. 87: 44, 1916, hom. illeg. non K.
Schumann, 1900. Type: Gertrudia validinervis Herz.
From Gertrude, a name + i + -ella, diminutive; T. Herzog's
(1916) original name for this genus was a dedication to his wife.
Plants in cushions, green above, brown to red-brown or
blackish-brown below. Stems branching occasionally, to 2.0 em
in length, transverse section rounded-pentagonal to elliptical,
central strand very strong and well differentiated, sclerodermis
present, hyalodermis present but firm-walled; axillary hairs of
5-7 clear cells. Leaves crowded, weakly spreading, weakly to
GERTRUDIELLOIDEAE
strongly contorted or curled when dry, squarrose from a short,
sheathing base when moist, lanceolate, 3.5-4.5 mm in length,
upper lamina broadly channeled across leaf, margins strongly
revolute to near apex, entire or occasionally weakly dentate near
apex; apex narrowly acute, occasionally narrowly obtuse; base
long-elliptical; costa percurrent to excurrent as a short, denticulate awn, superficial cells quadrate ventrally, elongate dorsally,
10-15 rows of cells across costa ventrally at midleaf, costal
transverse section semicircular, stereid band single, strong and
75
reniform, epidermis of 1 layer of strongly hollow-papillose cells
ventrally, epidermal cells weakly developed dorsally, guide
cells 15-20 in 3-4 layers, hydroid strand present; upper
/aminal cells subquadrate to irregularly hexagonal, small, 8-10
jlm in width, 1:1, walls evenly thickened, superficially bulgingmamillose ventrally, flat dorsally in medial portion of lamina,
bulging on both sides in 10-20 rows marginally; papillae
present on marginal cells, low, simple, hollow; basal cells
differentiated medially, rectangular, little wider than the upper,
3
Plate 5. Gertrudiel/a. 1-6. G. validinervis. l. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Upper marginal cells. 6. Basal cells. 7.
Transverse section at midleaf. 8. G. validinervis var. se"atopungens. 8. Leaf apex.
76
GENERA OF THE POTTIACEAE
3-4: 1, walls thin, occasionally transversely slit across longitudinal
walls. Apparently dioicous. Perichaetia unknown. Perigonia
appearing as axillary buds near stem apex, inner leaves deltoid,
outer leaves elongate. Sporophyte unknown. Lamina! KOH color
reaction orange.
Rare, restricted to Bolivia where it has been collected on stone
in rather dry areas, at low to high (300-3200 m) elevations.
This monotypic genus is well characterized, mainly by the
ventrally mamillose medial cells of the lamina, several rows of
marginal cells bulging on both sides and low-papillose, epidermal
cells on the ventral surface of the costa numerous and strongly
hollow-papillose, and guide cells rather thick-walled and forming
a multilayered cylinder (Pl. 5, f. 7). Hilpert (1933) included
Trichostomum ferrugineum Herz. in Gertrudiella on account of
the several layers of guide cells. This species is actually a Didymodon (with much the same general appearance of D .
occidentalis of western North America) by the orange reaction to
KOH, lanceolate leaves deeply and widely channeled ventrally
along the stout costae, and upper !aminal margins bistratose, and
is here treated as Didymodon herzogii (nom. nov. in treatment of
Didymodon), differing from related taxa of sect. Vineales by the
microstomous, eperistomate capsule and medial basal cells transversely resorbed across longitudinal walls to form slits. The last
character has also been described for Kingiobryum Robins. (see
Zander & Cleef 1982) of the Dicranaceae. Although G.
validinervis shows some slitting of basal lamina! cells, it and D.
herzogii are not closely related. Kingiobryum, which is
eperistomate, may well belong to the Pottiaceae; further study is
needed.
Gertrudiella shares several characters with various species
of Pseudocrossidium, among them revolute margins, differentiation of medial upper !aminal cells (especially P. leucocalyx)
and presence of ventral costal epidermal cells, a single stereid
band, guide cells in more than one layer, and small upper
!aminal cells. It is, however, apparently not closely related.
Streptocalypta has a similar arrangement of thick-walled guide
cells but has quite different areolation. The guide cell cylinder
of Gertrudiella is probably mainly of structural importance;
sections reveal that there is no direct connection with the
equally well-differentiated stem central strand (Pl. 5, f. 2).
Biblography: Herzog (1916), Hilpert (1933).
Number of accepted species: 1.
Taxa examined: G. validinervis (BUF and isotypes at JE, L,
W), G. validinervis var. serratopungens (JE, L).
New combination: Gertrudiella validinervis (Herz.) Broth.
var. serratopungens (Herz.) Zand., comb. nov. (Gertrudia
validinervis Herz. var. serratopungens Herz., Biblioth. Bot. 87:
45, 1916).
Subfamily CHIONOLOMOIDEAE
Chionolomoideae Zand., subfam. nov. Type: Chionoloma Dix.
Plantae magniusculae; caules plerumque usque altitudine 4 em; folia longilinearia, fiagelliformia, apice anguste acuto, base
plerumque vaginanti, marginibus planis, costa stratum ventrale stereidarum maius quam stratum dorsale evolventi et filum
hydroideum effacienti praedita.
Plants relatively large; stems often to 4 em in length; leaves long-linear, whip-like, apex narrowly actute, base often sheathing,
margins plane; costa with ventral stereid band larger than the dorsal, and hydroid strand absent.
This subclade is distinguished from the Timmielloideae, Erythrophyllopsoideae and Gertrudielloideae by the advanced character
of hydroid strand absent. The genera share two advanced character states: leaves long-linear and ventral stereid band larger than the
dorsal. The three included genera have long, whip-like leaves with plane margins, and are characteristic of tropical areas of low or
medium elevation.
Plate 6.
5. CHIONOLOMA
Chionoloma Dix., J. Bot. 60: 102, 1922. Type: Chionoloma
induratum Dix.
From XtCOV, Xt6vo~. snow + o + A.l\la., -a.n~. fringe, hem, border.
Plants growing in a dense turf, yellowish green above, light
reddish brown below. Stems branching occasionally, ca. 4.0 em in
length, transverse section rounded-pentagonal, central strand distinct, often strong, sclerodermis not differentiated from central
cylinder, which is of incrassate, large-lumened cells, hyalodermis
present, usually collapsed in mature parts of stem; axillary hairs
of several hyaline cells, basal 1-2 also hyaline, but with somewhat thicker walls; radiculose and also closely invested with a
verrucose or papillose red tomentum. Leaves incurved, curled and
often tubulose when dry, spreading when moist, long-linearlanceolate, ca. 6-7(-8) mm in length, upper lamina broadly channeled to shallow-grooved along costa near apex, margins plane to
weakly incurved, often sharply incurved near apex, distantly
weakly denticulate along border, bordered in lower 112-314 of
leaf by 2-7 rows of unistratose, rectangular, epapillose, thickwalled, porose cells; apex very narrowly acute; base longelliptical, weakly sheathing; costa excurrent as a cylindrical
smooth mucro, superficial cells quadrate and papillose ventrally,
elongate dorsally, 10-12 rows of cells across costa ventrally at
midleaf, costal transverse section elliptical to roundedtriangular, stereid bands two, the ventral dorsal band stronger
than the reniform, epidermis present ventrally, entirely absent
dorsally, guide cells 8-10 in 1 layer, hydroid strand absent;
upper /aminal cells small, rounded-quadrate, often shortrectangular, 6-9 l!m in width, 1-2: 1, walls thick, obscured by
papillae, medially free walls bulging ventrally and weakly convex dorsally, bulging marginally on both sides of lamina; papillae massive, multiplex-capituliform, crowded, with many small
salients; basal cells differentiated (but not sharply so) across
leaf, not or weakly grading into marginal border. rectangular,
CHIONOLOMOIDEAE
77
Plate 6. Chionoloma. 1-7. C. induratum . 1. Habit. 2. Transverse section of stem. 3-4. Two leaves and tomentum. 5. Leaf apex. '6. Basal cells.
7. Transverse section at midleaf. 8-11. C. lalifolium. 8. Transverse section of stem. 9-10. Two leaves. 11. Transverse section at midleaf. 12-14.
C. longifolium . 12. Leaf. 13. Transverse section at midleaf. 14. Transverse section near leaf apex.
78
GENERA OF THE POTTIACEAE
Plate 7. Pseudosymblepharis. 1-7. P. schimperillna. 1. Habit. 2. Transverse section of stem. 3. Leaf. 4. Leaf apex. 5. Basal cells. 6. Transverse
section at midleaf. 7. Peristome teeth. 8-10. P. angustata. 8-9. Two leaves. 10. Leaf apex. 11-13. P. duriuscula. 11-12. Two leaves. 13. Leaf
apex. 14-16. P. mauiensis. 14-15. Two leaves. 16. Leaf apex. 17-20. P. subduriuscula. 17-18. Two leaves. 19. Leaf apex. 20. Transverse
section at midleaf. 21-23. P. sy"hopodontoides. 21-22. Two leaves. 23. Leaf apex.
CHIONOLOMOIDEAE
mostly 10-13 j.lm in width, 3-5:1, thin-walled and hyaline near
insertion grading to incrassate and porose in upper part of leaf
base. Perichaetia, perigonia and sporophyte unknown. Laminal
KOH color reaction deep yellow or yellowish orange above, reddish brown below.
Found on limestone in Burma, Thailand, Malaysia and Borneo.
This genus has much the appearance of Tortella, but differs in
having four areas of very distinct laminal areolation: quadrate
upper laminal cells; a marginal border of long-rectangular,
porose, incrassate cells (Pl. 6, f. 5); short-rectangular, incrassate
and porose medial upper leaf base cells (similar to those characteristic of Trichostomum subg. Oxystegus), and hyaline, thinwalled rectangular cells of the lower leaf base (Pl. 6, f. 6). The
border of Chionoloma is similar to that of Pleurochaete in that it
does not clearly meld with the basal cells to reach the costa
below, and the upper medial !aminal cells are also often bulging
strongly ventrally but weakly so dorsally; but Pleurochaete differs from Chionoloma by the costal section reniform, the ventral
stereid band smaller than the dorsal, the leaf margin plane or
broadly channeled near the apex, and the marginal leaf border of
relatively thin-walled cells. Pleurochaete is unusual in being
pleurocarpous, and the sexual position in Chionoloma is
unknown. The incurved upper margins (Pl. 6, f. 14), the rather
thick papillae (best seen in section), and the leaf border of thickwalled cells of Chionoloma are similar to those of Hypodontium.
Only the quadrate upper leaf cells, the upper ventral costal
cells, and cells of the uppermost portion of the leaf base are papillose. The stem section (Pl. 6, f. 2) is distinctive and similar to that
of many Trichostomum species, with all cells incrassate except
those of the central strand and the hyalodermis. The leaf section
shows an unusually large number of ventral epidermal cells (in
costal sections) and of guide cells (but both are each in one layer),
and the section is otherwise similar to that of Pseudosymblepharis
in the ventral stereid band being stronger than the dorsal and the
costal section often rounded-triangular. The red tomentum (Pl. 6,
f. 3) is also a distinctive feature, reminiscent of that of some species of Leptodontium, a genus which, however, differs in the lack
of a central strand and of a differentiated ventral costal epidermis.
The extreme upper !aminal margins are sometimes sharply and
narrowly incurved like those of Weissia, but that genus does not
have the marginal border or characteristic costal section; W.
jamaicensis is similar in size, the distinct rectangular leaf base,
and its ventral stereid band is commonly larger than the dorsal,
but the costal section is round and the upper !aminal cells are
mainly isodiametric, with walls not so highly thickened. The three
species of Chionoloma are not especially different from one
another. The genus was synonymized with Pseudosymblepharis
by Eddy (1990, cf Menzel 1992) as merely "exceptionally robust
plants in which the pellucid leaf border ascends high into the leaf
limb, but the latter character is extremely variable, even within a
single species." The denticulate border of Chionoloma will immediately distinguish this genus from Pseudosymblepharis.
Additional literature: Dixon ( 1922a).
Number of accepted species: 3.
Species examined: C. induratum (BM), C. latifolium (BM), C.
longifolium (BM).
79
6. PSEUDOSYMBLEPHARIS
Plate 7.
Pseudosymblepharis Broth., Nat. Pfl. ed. 2, 10: 261, 1924.
Lectotype: Pseudosymblepharis papillosula Card. & Ther.
fide Saito, J. Hattori Bot. Lab. 39: 439, 1975.
From \jf£UO'J1c;, falsehood + o + Symblepharis, a genus; resembling the genus Symblepharis.
Plants growing in clumps or turf, green above, brown or
reddish brown below. Stems branching often and irregularly,
often long, to 2-4(-6) em in length, transverse section roundedpentagonal, central strand usually present, occasionally small,
cells of central cylinder often thick-walled, sc/erodermis usually
weakly developed and composed of substereid cells, hyalodermis present, sturdy, collapsed only in very mature parts of
stem; axillary hairs to 16 cells in length, all hyaline or basall-3
cells yellow; rhizoids sparse or stem occasionally weakly redtomentose. Leaves incurved, contorted to spiralled when dry,
spreading to squarrose from a usually sheathing base when
moist, lanceolate to linear, 1-7 mm in length, upper lamina
often narrow, flat to weakly tubulose, margins plane, entire or
occasionally weakly dentate at apex, margins often eroded by
fragmentation, occasionally decurrent; apex subulate, sharp,
occasionally narrowly obtuse, often fragile and broken; base
usually strongly sheathing, ovate to rectangular, with
"shoulders"; costa excurrent as a sharp, cylindrical mucro or
occasionally percurrent, superficial cells quadrate to shortrectangular, occasionally elongate near apex ventrally, elongate
dorsally, (4-)8-15 rows of cells across costa ventrally at
midleaf, costal transverse section semicircular to ovate, stereid
bands strong, the dorsal usually larger than the ventral, epidermis ventrally present, dorsally present, weak or absent, guide
cells (4-)6-9 in 1 (occasionally partially bistratose) layer,
hydroid strand absent; upper laminal cells subquadrate to shortrectangular, 8-10 IJ.m in width, 1(-2):1, walls evenly thickened,
superficially weakly convex on both sides, occasionally
strongly bulging ventrally and weakly convex dorsally; papillae
bifid, 3-5 crowded over each lumen or fused into one capitulate
papilla, or cells occasionally nearly smooth; differentiated basal
cells filling leaf base, often reaching up margins in a vee, rectangular to bulging-rectangular, 13-25 IJ.m in width, 3-5:1,
walls thin, becoming thicker above, occasionally porose medially. Dioicous. Perichaetia terminal (these occasionally appear
lateral because of subperichaetial elongate, overtopping
branches), inner leaves little different from cauline leaves or
high-sheathing. Perigonia terminal, inner leaves little different
from the cauline leaves. Seta 0.7-2.0 em in length, 1 per
perichaetium, brown to reddish brown, twisted clockwise
below, counterclockwise above; theca (0.5-)2.0-2.5 mm in
length, reddish brown, cylindrical, exothecial cells rectangular,
20-30 j.lm in width, 2-4:1, walls thin, stomates present at base
of theca, phaneropore, annulus usually of 2-4 rows of
vesiculose cells, deciduous in pieces; peristome teeth 16, rather
short, occasionally absent, yellow to whitish, triangular to linear, cleft and perforated to near base, spiculose, 125-300 IJ.m in
length, with many articulations, straight, basal -membrane when
present 35-50 IJ.m in height, spiculose. Operculum rostrate,
(0.4-)1.2 mm in length, cells straight. Calyptra cucullate,
80
GENERA OF THE POTTIACEAE
smooth, ca. 2 mm in length. Spores 11-15 J..Lm in diameter,
yellow-brown, papillose. Lamina! KOH color reaction golden
yellow-orange, occasionally deep yellow. Reported chromosome
number n = 13, 14.
A genus found in most tropical and warm-temperate areas,
occurring on soil, rock (mostly calcareous) and bark.
This genus is like both Tortella and Trichostomum subg.
Oxystegus in the plane or broadly incurved leaf margins (Pl. 7, f.
20) and well differentiated, often inflated basal cells that usually
extend up the margins in a more or less distinct vee (Pl. 7, f. 9,
18). The broadly sheathing leaf base with distinct "shoulders" has
been used in the past as a major character, but is actually found to
some extent in other genera, and is poorly or not at all developed
in some species of Pseudosymblepharis. An additional and possibly better character is the relative size of the ventral stereid band
(Pl. 7, f. 6, 20). In Pseudosymblepharis the ventral stereid band is
almost always distinctly larger than the dorsal, and is often
strongly bulging ventrally. Tortella may be additionally distinguished by the usually better developed peristome, while Trichostomum subg. Oxystegus differs, gametophytically, by the vee of
basal cells poorly developed or absent. There may well prove to
be no acceptable distinction between Pseudosymblepharis and
Trichostomum subg. Oxystegus, excepting the color of the peristomes, yellow to white in the former and red in the latter (also
see discussion of Norris and Koponen 1989). In leaf shape, P.
indica might be recognized in Trichostomum subg. Oxystegus (cf
Hilpert 1933) but the costa is quite thick, with the ventral stereid
band larger than the dorsal (at least in the larger, mature leaves).
Problematically, T. (Oxystegus) hybernicus has the sheathing leaf
base of Pseudosymblepharis and a ventral stereid band equal to or
greater than the dorsal, but the rectangular upper !aminal cells and
thin-walled cells of the central cylinder associate it clearly with
Trichostomum tenuirostris. The genus Pseudosymblepharis has
considerable resemblance to Symblepharis Mont. (Dicranaceae),
but gametophytes of the latter genus may be distinguished by
their smooth leaf cells. The generitype of Trichostomum, T.
brachydontium, has the ventral stereid band much larger than the
dorsal in most very robust specimens, much as is the case in
Pseudosymblepharis. Weissia jamaicensis has a leaf shape and
costal structure similar to that of Pseudosymblepharis but the
sharply and narrowly incurved upper lamina! margins are characteristic only of Weissia s. lat. and Chionoloma. This entire relationship needs careful revision.
Tortella has 32 filamentous teeth, longer than 500 J..Lm, these
strongly twisted. Trichostomum (including subgenus Oxystegus)
has 16 teeth cleft variously to near that base, shorter than 400 J..Lm,
and straight to very weakly twisted. Future evaluation may find
that Tortella and Trichostomum (s. lat.) may better be divided by
characters other than those of the sporophyte and basal cell area
shape of the leaves. A similar situation has occurred in the Barbula-Didymodon group, which was in the past distinguished along
similar lines of peristome morphology, but which is now (Saito
1975a) separated into three genera (Bryoerythrophyllum being
segregated from Didymodon) by a combination of several
gametophytic characters (see treatments below).
There may be quite a difference in stature between various
collections of the same species, with certain variation correlated
with plant size. In P. schimperiana, for instance, small plants may
be only weakly sheathing at the leaf base, the hyaline basal cells
of such collections may reach up the margins to half the length of
the leaf, and the leaves are more commonly fragile. Examined
collections of the widespread species Old World species P.
angustata (holotype, NY!) are nearly identical to the New
World P. schimperiana, but differ in lacking a stem central
strand (weak in P. schimperiana). A revision of this genus,
however, may yet demonstrate synonymy and a pan-tropical
distribution of P. angustata.
Number of accepted species: 11.
Additional literature: Crum (1952a).
Species examined: P. angustifolia (MICH), P. angustata
(BUF, NY), P. cavernarum (H), P. circinnatula (H), P.
duriuscula (BUF, NY), P. indica (BM, NY), P. khasiana (NY),
P. mauiensis (DUKE, NY), P. perlongifolia (NY), P.
schimperiana (BUF, CANM, DUKE, NY, SPA, TENN), P.
subduriuscula (BUF, NY), P. syrrhopodontoides (BM), P.
verrucosa (H).
New heterotypic synonymy: Tortella grossiretis Bartr. =
Pseudosymblelpharis schimperiana (Par.) Crum. Pseudosymblepharis bartramii Ther. ex Bartr. = Pseudosymblepharis
schimperiana (Par.) Crum.
New combinations:
Pseudosymblepharis cavernarum (Broth.) Zand., comb. nov.
(Trichostomum cavernarum Broth., Denkschr. Ak. Wiss.
Wien Math. Nat. Kl. 83: 283, 1926).
Pseudosymblepharis circinnatula (Broth. in VOltzk) Zand.,
comb. nov. (Trichostomum circinnatulum Broth. in Voltzk,
Reise Ostafr. 3: 54, 1908).
Pseudosymblepharis khasia!Ja (Mitt.) Zand., comb. nov. (Tortuta khasiana Mitt., J. Linn. Soc. Bot. Suppl. 1: 29, 1859;
Barbula khasiana (Mitt.) Jaeg.; Trichostomum khasianum
(Mitt.) Broth.; Oxystegus khasianus (Mitt.) Gangulee).
Pseudosymblepharis perlongifolia (Frohl.) Zand., comb. nov.
(Trichostomum perlongifolium Frohl., Rev. Bryol. Lichenol.
31: 92, 1962).
Pseudosymblepharis syrrhopodontoides (Dix.) Zand., comb.
nov. (Tortella syrrhopodontoides Dix., J. Bot. 76: 228,
1938.).
Pseudosymblepharis verrucosa (Broth. & Par.) Zand., comb.
nov. (Trichostomum verrucosum Broth. & Par., Ofv. Finsk.
Vet. Soc. Forh. 51A(17): 12, 1909; Oxystegus verrucosus
(Broth. & Par.) Hilp.).
7. PACHYNEUROPSIS
Plate 8.
Pachyneuropsis H. Mill., Taxon 19: 822, 1970, nom. nov. for
Pachyneurum Bartr. Type: Pachyneuropsis bartlettii
(Bartr.) H. Mill.
Pachyneurum Bartr., Philippine J. Sci. 68: 100, 1939, hom.
il/eg. non Amann, 1912.
From Pachyneurum, a genus+ VEUpov, nerve, sinew, tendon+
3\lf<n<;. -EO><;. appearance; resembling the genus Pachyneurum.
Plants in cushions, light green above, tan below. Stems
branching often, ca. 1.5 em in length, transverse section
rounded-pentagonal, central strand present, sclerodermis
present but weakly developed, hyalodermis present; axillary
hairs 6-15 cells in length, becoming swollen and firm-walled
throughout; rhizoids sparse. Leaves incurved, somewhat twisted
when dry, spreading when moist, long-linear, ca. 5 mm in
81
CHIONOLOMOIDEAE
7
Plate 8. Pachyneuropsis.l-10. P. bartlettii. I. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8-9. Two
transverse sections near midleaf. 10. Upper !aminal papillae . .
82
GENERA OF THE POTTIACEAE
length, lamina usually very narrow, the blade on each side seldom
as wide as the costa and occasionally absent above midleaf, often
narrower on one side than the other, upper lamina broadly channeled but costa bulging ventrally, margins plane, entire,
bi(tri)stratose from just above leaf base to near apex; apex narrowly acute; base abruptly ovate, with distinct shoulders; costa
very stout, short-excurrent as a sharp, smooth mucro or smooth
hyaline apiculus, superficial cells quadrate and papillose on both
sides, ca. 10-13 rows of cells across costa ventrally at midleaf,
costal transverse section circular, ventral stereid band stronger
than dorsal, dorsal band reniform, epidermis present on both
sides, guide cells ca. 8 in 1 layer, hydroid strand absent; upper
!aminal cells rounded-quadrate, 8-l 0 ).lm in width, l (-2): l, walls
evenly thickened, lumens somewhat angular, superficially bulging
on both sides; papillae bifid, 3-4 per lumen, to multiplex,
crowded, solid; basal cells differentiated across leaf, reaching
somewhat higher along margins, inflated rectangular, bulging, to
25 ).lm, much wider than upper cells, 2-4: l, walls thin-walled,
hyaline. Gametoecia and sporophyte unknown. Lamina! KOH
color reaction yellow except leaf insertion red.
Endemic to the Philippines. It is known from a single collection: Luzon, Rizal Province, Mantalban, Bartlett 14392,
holotype, FH, without indication of substrate.
Pachyneuropsis is morphologically close to Pseudosymblepharis in the linear leaf shape with abruptly broadened base (Pl.
8: 3-5) and a tendency to extreme length, the thick costa with
ventral stereid band larger than the dorsal (Pl. 8, f. 8-9), among
other characters. It differs in the very narrow or absent upper
lamina with bi- to tristratose margins, and the costa even thicker
than is usual in the latter genus. Bartram (1939) suggested a
passing similarity to Barbula pachyloma Broth., which he had
not seen, but this latter species (transferred to Cinclidotus as C.
involutus Hilp., nom. nov., by Hilpert 1933) is actually a good
species of Barbula.
Number of accepted species: l.
Species examined: P. bartlettii (FH).
Subfamily TRICHOSTOMOIDEAE
Trichostomoideae (BSG) Limpr. in Broth., Nat. Pfl. 1(3): 381, 1902 "Trichostomeae" used as a subfamily name.
Trichostomaceae Schimp., Syn. 141, 1860 "Trichostomeae." Type: Trichostomum Bruch, nom. cons.
Trichostomeae BSG in Schimp., Syn. 1: 168, 1872, apparently used as a family name, see Saito, J. Hattori Bot. Lab. 39: 416,
1975.
Trichostomataceae Crosby & Magill, Diet. Mosses 42, 1977, orth . err pro Trichostomaceae Schimp., 1860.
Trichostomeae Limpr., Laubm. Deutsch. 1: 519, 1888, rank not given.
Eucladioideae Chen, Hedwigia 80: 40, 1941.
Trichostomeae Dix., Stud. Handb. Brit. Moss. 205, 1924.
Tortelleae Chen, Hedwigia 80: 142, 1941.
Eucladieae Chen, Hedwigia 80: 55, 1941.
This subclade is distinguished from lower branches on the tree by the advanced characters of leaves shorter, 1.5 to 3.0 mm in
length; rows of cells across the ventral surface of the costa at midleaf fewer, 4-6; costal guide cells fewer, 2--6; and theca shorter,
less than 1.5 mm in length. The traits at the immediate ancestral node are axillary hairs completely hyaline and dorsal costal epidermis absent. The eight included genera may be characterized generally by at least combinations of the most of the following characters: stem sclerodermis poorly differentiated from the central cylinder, hyalodermis commonly present; leaves lanceolate; margins
plane or weakly recurved below and plane or erect, occasionally incurved above; upper !aminal papillae crowded; basal cells occasionally reaching much higher along the margins to form a differentiated vee; upper lamina KOH reaction yellow (seldom yellowish
orange or reddish brown); costa lacking a differentiated dorsal epidermis; clavate propagula absent. This subfamily is widely distributed; the basal taxon, Calymperastrum, is austral.
Saito (1975a) cited the number of amphithecial cell layers at the mouth of the capsule as one of the major characters distinguishing the Trichostomoideae (including Timmie/la, Tortella and Weissia according to Saito) from the Pottioideae (including Barbuleae,
Eucladieae, Leptodontieae and Pottieae all sensu Saito), the former with five layers and the latter with only four. He illustrated these
conditions for eleven species in eight (Barbula, Bryoerythrophyllum, Desmatodon, Didymodon, Tortel/a, Tortula s. str., Trichostomum and Weissia) different peristomate genera of the Japanese moss flora. This anatomical difference certainly appears promising
but should be reevaluated in the light of cladistic study of all characters. Although Catcheside (1980) has echoed, with reservations,
the use of Saito's amphitheciallayer distinction for the traditional Trichostomoideae, a thorough study of taxa worldwide is needed
using techniques of the plant anatomist. Sectioning capsules lengthwise freehand is a considerable challenge and wasteful of capsules if not immediately successful. It has been impossible to use this character in practice. Saito and Hirohama (1974a) pointed out
that "multistalked verrucae" as spore ornamentation are unique to but not always present (as seen with the SEM) in the Trichostomoideae (sensu Saito) for the 19 species in 14 genera of Japanese Pottiaceae that they studied.
TRICHOSTOMOIDEAE
8. CALYMPERASTRUM
Plate 9.
Calymperastrum Stone, J. Bryol. 14: 315-318, 1986. Type:
Calymperastrum latifolium (Hampe) Stone.
From Calymperes, a genus+ -astrum, an ending implying deception; mimicking the genus Calymperes.
Plants in a low turf, yellowish green above, yellowish brown
below. Stems not branching, ca. 6 mm in length, transverse section rounded-pentagonal to triangular, central strand distinct, generally with an satellite strand just below the sclerodermis, sclerodermis of 1-2 layers of thick-walled cells, hyalodermis weakly
developed in patches; axillary hairs ca. 6 cells in length, all
hyaline; papillose, reddish brown tomentum on lower part of
stem. Leaves spreading-incurved and conduplicate from an
appressed base when dry, spreading when moist, long-spathulate,
2.0-3.1 mm in length, upper lamina broadly concave, margins
83
plane to weakly incurved above, entire, bistratose in 1(-3) rows
from shoulder to 314 length of leaf, cell walls of border slightly
thickened; apex rounded-acute and bluntly apiculate; base
strongly differentiated in shape in lower 1/3 of leaf, cuneate,
with distinct shoulders, basal cells narrower along margins;
costa strong and tapering above, percurrent, superficial cells
quadrate to short-rectangular and papillose ventrally, elongate
and weakly papillose dorsally, 4-6 rows of cells across costa
ventrally at midleaf, costal transverse section nearly round,
stereid bands two, usually strong ventrally (the cell walls often
thicker than those of dorsal band, especially in sections made
through leaf base) and generally larger than the dorsal, distinct
to nearly absent dorsally, epidermis well differentiated ventrally
but absent dorsally, guide cells small and 4 in 1 layer, hydroid
strand strong, in a central position just dorsal to the guide cells
and often also ventrally; upper /aminal cells 'subquadrate to 5-
Plate 9. Calymperastrum. 1-9. C.latifolium. 1. Habit. 2. Transverse section of stem showing a satellite strand. 3-5. Three leaves. 6. Leaf apex.
7. Marginallaminal cells showing bistratose border. 8. Basal cells. 9. Transverse section at midleaf.
84
GENERA OF THE POTTIACEAE
or 6--sided, often slightly longer than wide, 9-13 Jlm in width,
l(-2):1, walls thin to weakly evenly thickened, superficially
strongly bulging ventrally, less strongly bulging dorsally; papillae
small, crowded, bifid, solid, ca. 8 per lumen; basal cells strongly
differentiated from upper cells, rising higher and larger medially,
weakly inflated, rectangular, ca. 20 Jlm in width, 2-4: 1, walls
thin, hyaline, lacking internal or superficial pores. Sexual structures and sporophyte unknown. Lamina[ KOH color reaction yellow.
A monotypic genus endemic to western Australia, growing on
cycads.
This genus can be excluded from the Calymperaceae mainly
by the hydroid strand (Pl. 9, f. 9) in the leaf. It appears to have
many of the characteristics of Bryoerythrophyllum and Mironia
(especially those of the areolation), but it differs markedly in
being yellow in KOH. From Dialytrichia, which also has a
bistratose leaf margin (Pl. 9, f. 7), it differs in its broadly concave
ventral laminal surface (not deeply grooved along the costa), its
plane leaf margins (not recurved below midleaf), its round (not
semicircular) costal section and semicircular (not crescentshaped) section of dorsal stereid band. Calymperastrum is also
somewhat like Trichostomum especially in the plane to incurved
margins but a hydroid strand is rare in that genus, and the only
Trichostomum species seen with a bistratose leaf margin, T.
marginatum, only doubtfully belongs to Trichostomum .
Calymperastrum is also like certain species of Leptodontium
(e.g. L. stoloniferum) in the leaf shape and strongly differentiated
basal cells, but the presence of a central strand, a hydroid strand
and a differentiated costal epidermis are not characters of Leptodontium. Transverse sections of the stem show satellite strands
just interior to the sclerodermis, usually one per section and correlated with a broad stem ridge decurrent from each costa. The satellite strand appears to be a leaf hydroid strand (these being
unusually strong in Calymperastrum) extending into the central
cortex but not reaching the stem central strand (Pl. 9, f. 2). Aloinella, an unrelated taxon, has a similar stem morphology. A more
thorough description of the genus is given by Stone (1986), who
saw additional material.
This monotypic genus appears to be, like Hypodontium, transitional in morphology between the Pottiaceae and the Calymperaceae, approaching the latter through the following combination
of characters: arboreal habitat, leaves with rQunded apices and
plane, thickened margins, costa percurrent, ending in an apiculus,
the ventral stereid band of cells with walls more strongly thickened that those of the dorsal (cf Syrrhopodon richardsii Dix.),
upper laminal cells more strongly bulging ventrally than dorsally,
basal cells strongly differentiated from the upper cells (but not as
sharply demarcated as is usual in Calymperes and Syrrhopodon),
and laminal KOH reaction yellow. The ventral stereid cells being
more strongly thickened than those of the dorsal band is a character not found elsewhere in the Pottiaceae, but is apparently occasional in the Calymperaceae. The only unique character to my
knowledge distinguishing Calymperastrum from genera of the
Calymperaceae is the presence of a leaf hydroid strand. Other
characters cited in the original description (Stone 1986) as indicative of Pottiaceae are also found in various genera of both the
Calymperaceae and the Pottiaceae, including the stem central
strand, papillae on the upper cells of the sheathing leaf base, bifid
papillae on both surfaces of laminal cells, and absence of foliar
gemmae. The significance of the presence of seriate papillae on
the dorsal costal surface cannot be evaluated here, and information on the sporophyte is lacking.
Number of accepted species: I.
Species examined: C. latifolium (MELU).
9. EUCLADIUM
Plate 10.
Eucladium B.&S. in BSG, Bryol. Eur. 1: 93, 1846 (fasc. 33-36
Mon. 1). Type: Eucladium verticillatum (Brid.) B&S.
Mollia subg. Eucladium (B .&S.) Lindh., Musci Scand. 21,
1879.
Weissia subg. Eucladium (B.&S.) Kindb., Eur. N. Amer.
Bryin. 2: 283, 1897.
Weissia sect. Saxicolae Nees & Hornsch., Bryol. Germn.
2(2):P 26, 97, 183l,p.p.
Mallia sect. Eucladium (B.&S.) Braithw., Brit. Moss A. 1:
230, 439, 1885.
Weissia sect. Eucladium (B.&S.) Dix., Stud. Handb. Brit.
Moss. 210, 1896.
From eti, well, very, true, good+ KA.dooc;, branch, twig, stem,
repeatedly branched or forked stem + -ium, characteristic of.
Plants growing in turfs or cushions, bright to dark green
above, pale green to yellowish brown below. Stems branching
irregularly, 0.5-2.0 em in length, transverse section elliptical,
central strand absent, sclerodermis usually absent, hyalodermis
present, inflated; axillary hairs of 5-10 clear cells; occasionally
weakly radiculose. Leaves appessed to erect-spreading from
base, incurved above when dry, weakly spreading to spreadingrecurved when moist, oblong- to linear-lanceolate, 1.5-2.5 mm
in length, upper lamina broadly channeled, margins plane,
denticulate on lower margins, rarely entire; apex broadly to narrowly acute or subulate; base scarcely differentiated in shape to
ovate; costa strong, usually excurrent as a stout mucro,
decurrent at base, superficial cells quadrate to elongate ventrally, elongate dorsally, ca. 6 rows of cells across costa ventrally at midleaf, costal transverse section semicircular to elliptical, 2 stereid bands present, epidermis present ventrally, usually
present dorsally, guide cells 4--7 in 1(-2) layers, hydroid strand
absent; upper /aminal cells subquadrate (occasionally rectangular medially), smaller at leaf margins, 8-10 Jlm in width,
1: 1(-2), walls moderately thick-walled, often irregularly thickened, somewhat bulging superficially; papillae low, indistinct,
simple to occasionally multifid, scattered to centered, 2-5 per
cell; basal cells usually strongly differentiated across leaf or
medially, bulging-rectangular to rhomboidal, 12-15 Jlm in
width, 2-5:1, walls hyaline, thin. Dioicous. Perichaetia terminal,
inner leaves ovate-lanceolate, to 2.5 mm in length, sheathing,
lower cells rhomboidal in lower half. Perigonia terminal, inner
leaves long-lanceolate, similar to cauline. Seta ca. 5-6 mm in
length, 1 per perichaetium, yellow, twisted little or not at all;
theca ca. 1 mm in length, yellow-brown, ovoid to cylindrical,
exothecial cells short-rectangular, thin-walled, bulging,
stomates phaneropore, present at base of theca, annulus of ca. 2
rows of weakly vesiculose cells; peristome teeth 16, lanceolate,
entire to variously cleft, yellow, papillose, rudimentary or to
300 Jlm, with up to several articulations, straight, basal membrane low, papillose. Operculum conic-rostrate, ca. 0.5-0.8 mm
in length, cells in straight rows. Calyptra cucullate, smooth, ca.
2.5 mm in length. Spores 9-14 Jlm in diameter, pale, essentially
TRICHOSTOMOIDEAE
smooth. Laminal KOH color reaction yellow. Reported chromosome number n = 13.
Found on calcareous rock in wet places, especially in spray of
waterfalls, across Europe and Asia, northern and southern Africa,
and North America (including Mexico).
Brotherus (1924-25) accepted the combination Eucladium
irroratum (Mitt. in Hook.) Jaeg., as did Dixon (1923). The rigid,
erect leaves and the wet, calcareous habitat of this New Zealand
species certainly suggest this genus, but technical characters indicate that that species belongs to a monotypic genus, Tetracoscinodon. Eucladium may now be considered a monotypic genus.
The most distinctive characters of Eucladium are the bright
green leaves, lack of a stem central strand or sclerodermis and
presence of a somewhat inflated hyalodermis (Pl. 10, f. 2), the
broad costa, plane leaf margins, which are denticulate below (Pl.
10, f. 6) except in certain Bermuda populations, laminal cells generally larger medially than at the margins above midleaf (Pl. 10, f.
5) and bulging-hyaline at the leaf base (Pl. 10, f. 6), laminal papil-
85
lae simple (Pl. 10, f. 8), perigonial leaves little different from
the cauline, and exothecial cells thin-walled and bulging. Eucladium has several morphological features that are similar to
those of Leptodontium and Hymenostylium: lack of a stem central strand, lanceolate leaves, trigonous upper laminal cells,
inflated basal laminal cells, peristome similar to that of L.
viticulosoides. The denticulate lower leaf margins are matched
in Molendoa hornschuchiana. The narrowly lanceolate leaf
shape, broad costa and large laminal cells ornamented with
rather large, simple papillae are reminiscent of Tuerckheimia.
The generic relationships are, on analysis, however, with
Trichostomum.
Additional literature: Dalby (1966), Dixon (1912), Dunk
and Dunk (1973), Glowacki (1909), Nagano (1959), Osada
(1958), Saito (1972a).
Number of accepted species: 1.
Species examined: E. verticillatum.
Plate 10. Eucladium. 1-9. E. verticil/atum. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Papillae. 9. Peristome.
86
GENERA OF THE POTTIACEAE
Plates 11-13.
10. TRICHOSTOMUM
Trichostomum Bruch, Flora 12: 396, 1829, nom. cons. non Hedw.,
1801. Lectotype: Trichostomum brachydontium Bruch.
Subg. Trichostomum (Hedw.) Turn., Muse. Hib. Spic. 35, 1804
(as autonym).
Trichostomum subg. Trichostomum Lor., Bryol. Notizb. 20:
1865, nom. i/leg.
Didymodon subg. Trichostomum Kindb., Eur. N. Amer. Bryin.
2: 272, 1897. Type: Trichostomum Hedw., nom. rej., p.p.
Bryum sect. Trichostomum (Hedw.) Relh., Fl. Cantabr. ed 2:
422, 1802, nom. illeg.
Trichostomum sect. Lancifolia B.&S. in BSG, Bryol. Eur. 2:
119 [fasc. 18-20 (Trichost.): 5], 1843.
Trichostomum sect. Pycnophyllum C. Miill., Syn. 1: 567, 1849.
Tortula sect. Trichostomum Mitt., J. Linn. Soc. Bot. 12: 142,
146, 1869, nom. illeg.
Subg. Crispuliformes (Kindb.) Zander, comb. et stat. nov. see
below.
Didymodon sect. Crispuliformes Kindb., Eur. N. Amer. Bryin.
2: 272, 1897. Type: Didymodon crispulus (Bruch) Wils.
Subg. Laminanchium Zander, subg. nov. see below. Type: Trichostomum tortelloides (Broth. & Dix.) Zand.
Subg. Oxystegus Limpr., Laubm. Deutsch!. 1: 569, 1888. Type:
Trichostomum cylindricum (Bruch ex Brid.) C. Miill. fide
Saito, J. Hattori Bot. Lab. 39: 436, 1975, hom. illeg.
Oxystegus (Limpr.) Hilp., Beih. Bot. Centralbl. 50: 666, 1933.
Stephanodictyon Dix., J. Linn. Soc. Bot. 50: 86, 1935. Type:
Stephanodictyon borneense Dix.
Paraleptodontium Long, J. Bryol. 12: 181 , 1982. Type:
Paraleptodontium recurvifolium (Tayl.) Long.
Didymodon subg. Oxystegus (Limpr.) Roth, Eur. Laubm. 1:
304, 1904.
Sect. Campy/opus Arnott, Mem. Soc. Linn. Paris 5: 244, 1827.
Sect. Leptomitrium Wallr., Fl. Crypt. Germ. l : 170, 1831.
From 6p!x. 'tplxoc;. hair + o + a'toJ.la, -a'toc;, mouth; a peristome of filiform teeth.
Plants turf-forming, yellowish green above, medium brown to
light brown below. Stems branching irregularly, to ca. 3 em in
length, transverse section rounded-pentagonal, occasionally
rounded-triangular, central strand present or absent, cells of central cylinder thin- or thick-walled, sclerodermis variously developed but often weak, of substereid cells or of one or two layers of
stereid cells, hyalodermis present or seldom absent; axillary hairs
ca. 10 cells in length, basal 1-3 cells thicker walled or all hyaline;
sparsely radiculose, or occasionally with a thin red tomentum.
Leaves incurved, occasionally tubulose and often catenulate or
reflexed at top of appressed base when dry, spreading to squarrose
when moist, oblong, elliptical or ligulate to long-lanceolate,
1.5-2.5(-5.0) mm in length, upper lamina flat or broadly channeled, leaves seldom keeled, occasionally grooved along costa,
margins usually plane, seldom broadly incurved to tubulose,
entire to crenulose-notched or occasionally dentate in the upper
1/2-3/4, occasionally with a narrow, less papillose border above
or throughout, rarely 2-3 rows of marginal cells bistratose; apex
narrowly to broadly acute or rounded, occasionally fragile and
broken, occasionally silarply reflexed or constricted, seldom
cucullate; base elliptical to rectangular, occasionally sheathing or
not differentiated in shape; costa usually excurrent as a smooth,
sharp mucro, occasionally ending 2-3 cells below apex, seldom
short-awned, superficial cells quadrate or occasionally elongate
(especially near apex) ventrally, elongate or seldom shortrectangular dorsally, 2-6(-8) rows of cells across costa ventrally at midleaf, costal transverse section semicircular, reniform
or ovate, two stereid bands present, occasionally very weak ventrally, ventral stereid band smaller or about same size as dorsal
stereid band, epidermis present ventrally, weak or absent dorsally, guide cells 2-4(-6) in 1 layer or seldom 1-3 scattered
bistratose pairs, hydroid strand absent or very seldom present;
upper !aminal cells rounded-quadrate, occasionally rectangular
or hexagonal, seldom transversely elliptical along margin,
6-12(-18) J.lm in width, 1(-2):1, seldom bistratose along margins or in small medial patches, walls usually evenly thickened,
superficially usually strongly convex on both sides or seldom
somewhat more bulging ventrally than dorsally; papillae usually bifid, crowded, 2-6 per lumen, occasionally single, multiplex and completely covering lumen; basal cells differentiated
across leaf or occasionally only medially or rising weakly along
margins, rectangular, seldom bulging superficially, 7-15 J.lm in
width, 3-5:1, walls thin to somewhat thickened, occasionally
porose. Propagula rare, on rhizoids or ventral surface of costa,
of several cells, vermiform to irregular in shape, occasionally
branching. Dioicous or occasionally autoicous. Perichaetia terminal, inner leaves usually little different from cauline leaves,
rarely oblong-sheathing and constricted at the apex, lower basal
cells inflated rectangular to rhomboid, hyaline. Perigonia terminal, outer leaves often similar to· the cauline, inner leaves
elongate-triangular, or occasionally present in leaf axils of
archegoniate plants as flattened buds. Seta 0.4-1.5 em in length,
I or seldom 2-3 per perichaetium, yellowish brown, occasionally reddish, twisted clockwise, in cleistocarpous species with
an abscission layer just below capsule; theca ca. 1-3 mm in
length, yellowish brown, occasionally orange, cylindrical or
elliptical, occasionally slightly curved, exothecial cells thinwalled, rectangular, stomates phaneropore at base of theca,
stegocarpous or seldom cleistocarpous, annulus of 1-4 rows of
vesiculose cells, persistent or seldom revoluble; peristome teeth
16, usually rather short, occasionally rudimentary or absent,
ligulate to filamentous, entire or occasionally irregularly cleft
2-3-fid or perforate, papillose, striate or spiculose, occasionally smooth, occasionally rather distant, to 400 J.lm, with several
articulations, straight or seldom weakly twisted counterclockwise, basal membrane absent or low but distinct, papillose to
spiculose, mouth of capsule occasionally closed by a
hymenium. Operculum long-conic to rostrate, ca. 0.4-0.7 mm in
length, cells straight, seldom weakly twisted counterclockwise
or not differentiated. Calyptra cucullate, smooth, ca. 2.0-2.5
mm in length. Spores 8-20 J.lm in diameter, yellow to brown,
essentially, smooth to strongly papillose. Laminal KOH color
reaction medium orange to yellowish orange, occasionally yellow. Reported chromosome number n = 12, 13, 13+m.
A large genus found on soil, rock (often calcareous) or
organic material on all continents except Antarctica.
Trichostomum has been a large, "wastebasket" genus of species that cannot be easily assigned to other genera. It differs
from similar Barbula (Merceyoideae) species with plane or
erect upper leaf margins (B. sect. Convolutae) by the cells of the
outer layer of the stem usually with large lumens (Pl. 11, f. 14;
TRICHOSTOMOIDEAE
87
Plate 11. Trichostomum. 1-6. T. tenuirostre. 1. Habit. 2-3 . Two leaves. 4. Leaf apex. 5. Propagulum. 6. Peristome. 7-12. T. aequilorillle. 7.
Transverse section of stem. 8-9. Two leaves. 10. Leaf apex. 11. Basal cells. 12. Transverse section at midleaf. 13-16. T. contractum. 13. Habit.
14. Transverse section of stem. 15-16. Two leaves. 17. Leaf apex.18-21. T. crispulum. 18-19. Two leaves. 20. Leaf apex. 21. Peristome.
88
GENERA OF THE POITIACEAE
13, f. 11), lower leaf margins plane, upper leaf margins seldom
dentate (but commonly minutely serrulate by projecting cell
walls), costal hydroid strand seldom present (Pl. 11, f. 12), dorsal
costal epidermal cells sometimes not differentiated (Pl. 12, f. 21),
ventral stereid band generally large, almost or about the thickness
of the dorsal band, perichaetial leaves generally not differentiated
from the cauline, and propagula (Pl. 11, f. 5) very rarely present.
The genus differs from Hyophila by the mostly narrower leaves
that are broadest at the base, !aminal cells usually somewhat bulging on both exposed surfaces (rather than just one surface though
this is variable in Hyophila), propagula usually lacking, and peristome usually present (among other characters, see Hyophila);
however, the thick-walled exothecial cells of Hyophila (forming a
series of concatenated semicircles as seen in transverse section of
the theca) are matched in some eperistomate species of Trichostomum.
I have found no good distinctions between Trichostomum and
Oxystegus at the generic level. There are no characters of the
sporophyte that acceptably distinguish the two; for instance,
Trichostomum (subg. Trichostomum) brachydontium may have
the spiral peristome tooth ornamentation typical of T. (subg.
Oxystegus) tenuirostre). Gametophytically, subg. Oxystegus differs only weakly from subg. Trichostomum in (l) generally plane
(occasionally tubulose when dry) upper leaf laminae, (2) a tendency toward broadly sheathing leaf bases with the differentiated
marginal cells somewhat running up the margins as in Tortel/a,
(3) medial upper basal cells thick-walled and rectangular (forming
a group sensibly different from the upper cells and lower basal
cells), and (4) the upper !aminal cells often distinctly enlarged and
rounded rectangular, 1-2:1. The first two of these characters are,
however, also present to some degree in Trichostomum sect.
Trichostomum, and the last is neither constant nor easily gauged.
Crum and Anderson (1981) likewise proposed' that Trichostomum
include Oxystegus. Saito (1975a) recognized Oxystegus but did
not discuss it as a genus. His key to genera of the Trichostomoideae separated Oxystegus and Pseudosymblepharis from Trichostomum by the presence of a central strand in the last; Eddy (1990)
used the same character to separate the two. This distinction does
not, however, hold outside of Japan or, apparently, Malesia.
Stoneburner ( 1985) expressed the opinion that differences
between Trichostomum and Oxystegus were probably artificial.
Norris and Koponen (1989) distinguished between Oxystegus and
Trichostomum by "a suite of differing, although somewhat overlapping characters": Trichostomum having leaves with lengths
less than 6: 1, blunt papillae grouped over the lumens, and !aminal
marginal cells thick-walled near the shoulders; Oxystegus having
leaves longer than 6:1, sharp papillae grouped around the lumens,
and cells of the shoulders thin-walled. These differences are recognized here at the subgeneric level. They also indicated that
although Oxystegus and Pseudosymblepharis are morphologically
close gametophytically, the red peristome of the former contrasts
with the whitish peristome of the latter.
There are trends in subg. Oxystegus towards three
morphotypes of robust stature: a lingulate leaf as in Trichostomum
recurvifolium (= Paraleptodontium Long 1982b, a synonym), a
lanceolate leaf (e.g. Trichostomum hibernicum), and a linear leaf
(e.g. Trichostomum bomeense, originally published as the type of
Stephanodictyon). Such tendencies are duplicated in infraspecific
variation in the central species of the Oxystegus complex-T.
tenuirostre. Eddy (1990) made Stephanodictyon a synonym of
Pseudosymblepharis (along with Chionoloma, here recognized
as a good genus).
Of species common in North Temperate Zone climates,
Trichostomum (subg. Oxystegus) tenuirostre is rather different
from T. crispulum and T. brachydontium (both of subg. Trichostomum) in its plane margins, vaguely vee-shaped area of
differentiated basal cells, and less well-developed peristome; as
a genus, however, Oxystegus cannot be maintained at the world
level because of considerable variation in expression and combination of characters among the species. For instance, the difference between the peristomes of T. crispulum (often 32 filamentous rami like of those of Tortella, but sometimes reduced)
and T. brachydontium (16 variously cleft or perforated lanceolate teeth) is greater than that between the peristomes of T.
brachydontium (generitype of Trichostomuin) and T. (subg.
Oxystegus) tenuirostre.
Tortella can be separated from Trichostomum on the basis
of a single relatively constant character, the vee-shaped basal
cell region, with the usually long and twisted peristome and
generally larger upper !aminal cells being associated trends; see
Cladogram 15 for analysis of phylogenetically important character state changes. Species of Trichostomum may also have
basal cells differentiated in a (less distinctive) vee-shape, and
those with flattened autoicous buds (e.g. Trichostomum
fragilifolium, Pl. 12, f. 14) differ gametophytically from certain
Tortel/a species (like Tortel/a humilis) with similar buds only in
the basal cells being differentiated evenly across the leaf rather
than running higher up the margins. Quite probably, Trichostomum, Pseudosymblepharis and possibly also Tortel/a represent
a complex of several transformation series. This needs to be
studied. Future revision of the Trichostomum complex may tum
up a satisfactory generic classification recognizing several separate phyletic series, with perhaps Trichostomum brachydontium,
T. crispulum, Weissia jamaicensis or even species of Tortel/a
variously at the unreduced end of each.
Species of Weissia (and of Tetrapterum) with clearly plane
or erect-incurved upper !aminal margins (including subg.
Hymenostomum and subg. Astomum) examined during this
study are here transferred to Trichostomum s. lat. Groups of
Trichostomum species very similar to each other in areolation
and certain other gametophytic characters may be construed
(until analysis demonstrates otherwise) as separate reduction
series toward smaller stature of gametophyte, shorter seta or
capsule (the latter occasionally almost spherical), short or
absent peristome, and monoicy, and are recognized below as
subgenera.
Trichostomum subg. Trichostomum
Leaves ovate to long-lanceolate, apex usually rounded, plane;
upper margins plane; costa thick and excurrent in a stout, sharp
mucro; basal cells moderately differentiated across leaf base.
Occasionally the upper margins in some leaves are rather
narrowly incurved, like W eissia species. A reduction series
(large to small stature with concomitant reduction of morphological details) might include: Trichostomum brachydontium, T.
p/anifolium, T. urceolare, T. sinaloense (Pl. 13, f. 16-19), T.
termitarum, T. williamsii, T. unguiculatum and T. austrocrispum. Also belonging here is T. incertum.
TRICHOSTOMOIDEAE
89
Plate 12. Trichostomum. 1-3. T. bombayense. l-2. Two leaves. 3. Leaf apex. 4-8. T. duidense. 4-5. Two leaves. 6. Leaf apex. 7. Transverse
section at midleaf. 8. Capsule. 9-16. T. fragilifolium. 9-10. Two leaves. 11. Perichaetialleaf. 12. Leaf apex. 13. Basal cells. 14. Autoicous bud.
15. Capsule. 16. Peristome. 17-21. T. pallidens. 17. Transverse section of stem. 18-19. Two leaves. 20. Leaf apex. 21. Transverse section at
midleaf.
90
GENERA OF THE POTTIACEAE
Trichostomum subg. Crispuliformes (Kindb.) Zander, comb. and
stat. nov. Basionym: Didymodon sect. Crispuliformes Kindb.,
Eur. N. Amer. Bryin. 2: 272, 1897. Type: Trichostomum
crispulum Bruch ex F. A. MUller.
Leaves elliptical to long-lanceolate; apex rounded-acute, often
somewhat cucullate; upper margins broadly incurved; costa
evenly tapering and generally ending before the apex or
percurrent; basal cells moderately differentiated across leaf base.
This taxon includes the series: Tortel/a injlexa, Trichostomum
crispulum (Pl. 11, f. 18-21), T. castaneum, T. caespitosum, T.
brittonianum, T. suhangustifolium, T. atrocaule, T. perligulatum,
and possibly Weissia crispa. Also belonging here are T. connivens
and T. pulicare.
Trichostomum subg. Laminanchium Zander, suhg. nov. Type:
Trichostomum tortelloides (Broth. & Dix.) Zand.
Folia longiligulata , et plerumque in regione supra hasem
subvaginantem angustata et regione sub apice , apice acuto,
plano, marginibus supernis et interdum infernis planis, costa
decrescenti excurrentique, mucronem formanti, cel/ulis basalibus
valde in regione mediana distinctis et in marginali cellulis
subangustatis praedita.
Leaves long-ligulate, often pinched just above the somewhat
sheathing base and also just below the apex; apex acute, plane;
upper margins and sometimes the lower margins plane; costa
tapering and excurrent as a mucro; basal cells strongly
differentiated medially with narrower cells marginally.
The upper medial superficial cell walls of most species of this
subgenus are ventrally bulging and dorsally nearly flat, and are
also commonly papillose. Few sporophytes have been seen, but
these appear to uniformly lack peristomes and sometimes have
rather short setae. Species included here are similar to Barbula
sect. Convolutae by the mainly medially differentiated basal cells,
but the basal cells of species of sect. Convolutae are rather thickwalled and not strongly differentiated. Species of subg.
Laminachium are: Trichostomum bombayense (Pl. 12, f. 1-3, and
see discussion of Townsend 1983), T. contractum (Pl. 11 , f.
13-16), T. criotum (autoicous, with flattened axillary perigonia),
and T. tortel/oides (Pl. 13, f. 20-23). See also discussion of
Calymperastrum.
Trichostomum subg. Oxystegus Limpr. Type: Trichostomum
tenuirostre (Hook. & Tayl.) Lindb.
Leaves lanceolate to linear-lanceolate, occasionally with a sharply
dilated base; apex rounded to narrowly acute, plane; upper margins plane; costa tapering and excurrent as a short mucro; basal
cells moderately differentiated, often running up the margins
somewhat as in Tortel/a, and bordered distally by a region of
thick-walled rectangular cells.
Species of this subgenus could include the following series:
Trichostomum tenuirostre (Pl. 11, f. 1-6), T. duidense (Pl. 12, f.
4-8), T. melanostomum, T. spirale and T. abyssinicum. There are
certainly many other names that might go here.
Trichostomum caespitosum, having leaves with small upper
laminal cells and broadly incurved margins, and being autoicous,
was recognized in Pottia recently by both Smith (1978) and
Corley et al. (1981), but clearly fits in a reduction series involving
T. crispulum. Its distinctly differentiated perichaetial leaves are,
however, unusual in the genus.
The idea that Trichostomum might include taxa with much
reduced or absent peristomes has been entertained recently by
several bryologists (pers. comm.) with interest in the group.
Magill (1981) pointed out that Trichostomum brachydontium
has rudimentary peristomes or gymnostomous capsules both in
South Africa and South America. He also noted that the
gametophyte of the gymnostomous and rhexolytically
operculate species Phasconica tisserantii (= T. unguiculatum cf
Crundwell & Nyholm 1974) resembles that of T .
brachydontium. Brotherus (1924-25) avoided the problem of
the relationship of species with Trichostomum-like gametophytes and cleistocarpous capsules by referring all these to
Tetrapterum .
Tuerckheimia is quite like Trichostomum in general appearance but the leaf bases of the former genus are not differentiated
in shape, the basal cells are merely short-rectangular in a small
area near the insertion, the upper laminal cells have a massive,
multiplex papilla over the center of each lumen (but not covering the lumen as in Oxystegus), and the leaves (often widest
near the middle) are pale yellow in KOH solution.
Non-type specimens from India collected by Ramskuhl
(BM) and identified as Desmatodon longirostris (Griff.) Mitt.
(= Merceyopsis longirostris (Griff.) Broth. & Dix.) proved to be
the superficially similar Scopelophila cataractae; the correct
name for D. longirostris is now Trichostomum contractum,
nom. nov. (see below).
Additional literature: Bartram (1924c), Blumrich (1916),
Brown (1897), Crum and Anderson (1958a), Dixon (1911b),
Frisvoll (1978), Hammerschmid (1915), Herzog (1907), Lett
(1901), Redfearn (1976), Zander (1978d,h, 1982a,c,i, 1985a).
Number of accepted species: 130, plus 2 combinations
remaining in Oxystegus.
Species examined: T. abyssinicum (PC), T. acutiusculum
(H), T. aequitoriale (BM), T. arcticum (BUF, C, CANM, NY),
T. atrocaule (TNS), T. austrocrispum (NY), T. bombayense
(NY-typified by Townsend 1983), T. borneense (BM), T.
brittonianum (NY), T. castaneum (NY), , T. connivens (NY), T.
criotum (BUF), T. cylindrotheca (NY), T. distans (NY), T.
duidense (BUF, NY), T. eckelianum (NY), T. exulatum (NY),
T. fa/lax (L), T. fragilifolium (NY), T. hyalinoblastum (NY), T.
imshaugii (NY), T. incertum (NY), T. laticostatum (PC), T.
lindigii (NY), T. melanostomum (NY), T. mitteneanum (NY), T.
orthodontum (NY), T. ovatifolium (H), T. pallidens (BM), T.
perannulatum (BM), T. perligulatum (COLO), T. planifolium
(MICH. NY), T. platyphyllum (NY), T. portoricense (BUF,
NY), T. pulicare (PC), T. recurvifolium (BUF), T. sinaloense
(FH , MICH. NY, TENN), T. spirale (BUF, PC), T.
subangustifolium (PC), T. subintegrum (H), T. tenuirostre, T.
termitarum (NY), T. tortelloides (FH), T. unguiculatum (PRE),
T. urceolare (BM), T. wagneri (H), T. wi/liamsii (NY).
New heterotypic synonymy: Barbula wollei Aust. (Didymodon wollei (Aust.) Aust.) = Trichostomum tenuirostre var. holtii
(Braithw.) Dix. Hyophila usambarica Broth. = Trichostomum
brachydontium Bruch. Phasconica tisserantii P. de Ia Yarde =
Trichostomum unguiculatum (Mitt.) Zand. Stephanodictyon
obscurirete Dix. = Trichostomum brachydontium Bruch.
Trichostomum molariforme Zand. = Trichostomum portoricense
Crum & Steere. Trichostomum schlimii C. Mtill. = Trichostomum tenuirostre (Hook. & Tayl.) Lindh. Weissia tortivelata
Williams = Trichostomum tenuirostre var. gemmiparum
TRICHOSTOMOIDEAE
(Schimp.) Zand.
New names, combinations and statuses (see also discussion
section above):
Trichostomum abyssinicum (Ther.) Zand., comb. nov. (Weissia
91
abyssinica Ther., Bull. Mus. Hist. Nat. Paris 34: 116, 1928).
Trichostomum acutiusculum (Broth.) Zand., comb. nov. (Hyophila acutiuscula Broth., Bot. Jahrb. 20: 183, 1894).
Trichostomum arboreum (Mitt.) Zand., comb. nov. (Weissia
Plate 13. Trichostomum. 1-5. T. hyalinoblastum. 1-2. Two leaves. 3. Leaf apex. 4. Basal cells. 5. Transverse section at midleaf. 6-10. T.
perannulatum. 6-8. Three leaves. 9. Leaf apex. 10. Basal cells. 11-15. T. platyphyllum. 11. Transverse section of stem. 12-13. Two leaves. 14.
Leaf apex. 15. Transverse section at midleaf. 16-19. T. sinaloense. 16. Habit. 17-18. Two leaves. 19. Leaf apex. 20-23. T. tortel/oides. 2Q-21.
Two leaves. 22. Leaf apex. 23. Sporophyte.
92
GENERA OF THE POTTIACEAE
arborea Mitt., J. Linn. Soc. Bot. 12: 138, 1869; Hyophila
arborea (Mitt.) Jaeg.).
Trichostomum atrocaule (Saito) Zand., comb. nov. (Weissia
atrocaulis Saito, J. Hattori Bot. Lab. 39: 425, 1975).
Trichostomum austrocrispum (Beck.) Zand., comb . nov.
(Phascum austrocrispum Beck., Trans. New Zealand lnst. 26:
274, 1894; Astomum austrocrispum (Beck.) Broth.).
Trichostomum austrocrispum var. longifolium (R. Br. ter) Zand.,
comb. et stat. nov. (Phascum longifolium R. Br. ter, Trans.
New Zealand Inst. 26: 308, 1894).
Trichostomum borneense (Dix.) Zand., comb. nov. (Stephanodictyon borneense Dix., J. Linn. Soc. Bot. 50: 86, 1935).
Trichostomum brittonianum Zand., nom. nov. (Hymenostomum
flavescens E. Britt. in N. Britt. & Millsp., Bahama Fl. 485,
1920; Weissia flavescens (Britt. in N. Britt. & Millsp.) Reese,
Bryologist 94: 54, 1991; non Trichostomumflavescens Dix.).
Trichostomum castaneum (Crum & Steere) Zand., comb. nov.
(Hymenostomum castaneum Crum & Steere, Amer. Midland
Nat. 60: 12, 1959).
Trichostomum criotum Zand., nom. nov. (Hyophila perannulata
Ren. & Card., Bull. Soc. R. Bot. Belg. 34(2): 60, 1896; non
Trichostomum perannulatum Dix. et P. Yard.).
Trichostomum contractum Zand., nom. nov. (Gymnostomum
longirostre Griff., Calcutta J. Nat. Hist. 2: 480, 1842; Merceyopsis longirostris (Griff.) Broth. & Dix.; Merceya longirostris
(Griff.) Wijk & Marg. Type: India, Khasia Hills, "123-a 1,"
lectotype, BM; non Trichostomum longirostre (Web. & Mohr)
Hartm.).
Trichostomum deciduaefolium (Saito) Zand., comb. nov. (Weissia
deciduaefolia Saito, J. Hattori Bot. Lab. 39: 429, 1975).
Trichostomum eckelianum Zand., nom. nov. (Trichostomum
cirrhatum Hampe~ Icon. Muse. 28, 1844, hom. illeg.; Torre/la
cirrhata Broth., Nat. Pfl. 1(3): 397, 1902).
Trichostomum exulatum Zand., nom. nov. (Phascum vernicosum
C. Miill. ex Roth, Aussereur. Laubm. 212, 1911 ; Tetrapterum
vernicosum (Roth) Broth. Type: Brazil, Santa Catharina, Ule
8, isotype, NY; non Trichostomum vernicosum Ren. & Card.).
Trichostomum finukamactum Zand., nom. nov. (Stephanodictyon
angustinerve Frohl., Rev. Bryol. Lich. 31: 92, 1962; non
Trichostomum angustinerve Card.), not seen.
Trichostomum imshaugii (Yitt) Zand., comb. nov. (Barbula
imshaugii Yitt, Bryologist 74: 464, 1971 [ 1972]); the leaf
shape, the variably but often strongly excurrent costa, and a
red KOH reaction make this species curiously reminiscent of
the genus Willia, but·it is placed here by the plane leaf margins and the two stereid bands in some leaves.
Trichostomum incertum (Mitt.) Zand., comb. nov. (Weissia
incerta Mitt., Phil. Trans. Roy. Soc. London 168: 389, 1879;
Tortula incerta (Mitt.) Broth.).
Trichostomum (subg. Oxystegus) ligulaefolium (Broth. & Par.)
Zand., comb. nov. (Hyophila ligulaefolia Broth. & Par., Rev.
Bryol. 31: 44, 1904).
Trichostomum lindigii (Hampe) Zand., comb. nov. (Systegium
lindigii Hampe, Ann. Sc. Nat. Bot. ser. 5, 3: 337, 1865;
Astomum lindigii (Hampe) Jaeg.).
Trichostomum melanostomum (Mitt.) Zand., comb. nov. (Weissia
melanostoma Mitt., J. Linn. Soc. Bot. 12: 138, 1869; Hyophila melanostoma (Mitt.) Jaeg.).
Trichostomum mitteneanum Zand., nom. nov. (Weissia umbrosa
Mitt., J. Linn. Soc. Bot. 12: 133, 1869; Hymenostomum
umbrosum (Mitt.) Kindb.; non Trichostomum umbrosum C.
Miill.).
Trichostomum ovatifolium Zand., nom. nov. (Hymenostomum
anomalum Broth. in Herz., Biblioth. Bot. 87: 29, 1916; non
Trichostomum anomalum (BSG) Schimp.).
Trichostomum pallidens (Dix.) Zand., comb. nov. (Pseudosymblepharis pallidens Dix., J. Bombay Nat. Hist. Soc. 39: 776,
1937).
Trichostomum perligulatum (Flow. ex Crum) Zand., comb. nov.
(Weissia perligulata Aow. ex Crum, Bryo1ogist 76: 291,
1973); this species is monoicous, and is not a synonym ofT.
crispulum as per Stoneburner and Wyatt (1985).
Trichostomum planifolium (Dix.) Zand., comb. nov. (Weissia
planifolia Dix., Rev. Bryol. Lichenol. 1: 179, 1928).
Trichostomum pulicare (Besch.) ZaQd ., comb. nov.
(Hymenostomum pulicare Besch., Ann. Sc. Nat. Bot. ser. 6,
9: 299, 1880).
Trichostomum recurvifolium (Tayl.) Zand., comb. nov. (Bryum
recurvifolium Tayl., Ann. Mag. Nat. Hist. 11 : 208, 1843;
Oxystegus recurvifolius (Tayl.) Zand., comb. nov .;
Paraleptodontium recurvifolium (Tayl.) Long).
Trichostomum sinaloense (Bartr.) Zand., comb. nov. (Weissia
sinaloensis Bartr., Bryo1ogist 28: 64, 1925; Hyophila
sinaloensis (Bartr.) Bartr.).
Trichostomum soulae (C. Miill. in Ren. & Card.) Zand., comb.
nov. (Ptychomitrium soulae C. Mtill. in Ren. & Card., Bull.
Soc. R. Bot. Be1g. 33(2): 118, 1895; Oxystegus soulae (C.
Miill. in Ren. & Card.) Wijk & Marg.).
Trichostomum soulae var. corticicola (Ren. & Card.) Zand.,
comb. et stat. nov. (Barbula corticicola Ren. & Card., Bull.
Soc. R. Bot. Belg. 35(1): 309, 1897; Oxystegus soulae var. '
corticicola (Ren. & Card.) Wijk & Marg.).
Trichostomum subangustifolium (Ther.) Zand., comb. nov.
(Hyophila subangustifolia Ther., Smiths. Misc. Coli. 85(4):
14, 1931; Weissia subangustifolia (Ther.) Zand.).
Trichostomum tenuirostre var. gemmiparum (Schimp.) Zand.,
comb. nov. (Didymodon cylindricus var. gemmiparus
Schimp. , Syn. Muse . ed. 2 165, 1876; Oxystegus
tenuirostris var. gemmiparus (Schimp.) Zand.).
Trichostomum termitarum (C. Mtill.) Zand., comb. nov. (Weissia termitarum C. Mtill., Hedwigia 39: 267, 1900).
Trichostomum tisserantii (P. de Ia Yarde) Zand., comb. nov.
(Phasconica tisserantii P. de Ia Yarde, Rev. Bryol.
Lichenol. 7: 231, 1934).
Trichostomum tortelloides (Broth. & Dix.) Zand., comb. nov.
(Ca/ymperes tortelloides Broth. & Dix., J. Bot. 48: 306,
191 0; suggested as being Pottiaceae by Reese, Bryologist
88: 106, 1985)
Trichostomum unguiculatum (Mitt.) Zand., comb. nov.
(Systegium unguiculatum Mitt., J. Linn. Soc. Bot. 22: 304,
1886; Astomum unguiculatum (Mitt.) Broth.; Weissia
unguiculata (Mitt.) Crundw. & Nyh.).
Trichostomum urceolare (Hampe) Zand., comb. nov. (Hyophila
urceolaris Hampe, Yid. Medd. Naturh. For. Kjoebenh. ser.
3,2: 269, 1870; Hymenostomum urceolare (Hampe)
Hampe).
Trichostomum williamsii Zand., nom. nov. (Astomum chilense
Williams, Bull. Torrey Bot. Club 42: 393, 1915; non
Trichostomum chilense Mont.).
TRICHOSTOMOIDEAE
93
Plate 14. Tuerckheimia. 1-7. T. svihloe. l. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7. Papillae.
8-13. T. guatemalensis. 8. Heterotrichous protonema. 9-10. Two leaves. 11. Upper !aminal cells. 12. Theca. 13. Remnants of peristome. 14-15.
T. robusta. 14. Leaf apex. 15. Transverse section at midleaf. 1~17. T. valeriana. 16. Leaf. 17. Upper marginal cells.
94
GENERA OF THE POTTIACEAE
11. TUERCKHEIMIA
Plate 14.
Tuerckheimia Broth., Ofv. F. Vet.-Soc. Forh. 52A (7): 2, 1910.
Type: Tuerckheimia guatemalensis Broth.
Named for H. von Tiirckheim, who collected plants in Guatemala
from 1877 to 1908 and in the West Indies from 1909 to 1910.
Plants growing in a turf, light to dark green and somewhat
glaucous throughout, seldom to commonly branching. Stem to 2.5
em in length, in transverse section rounded-pentagonal or triangular, central strand present, sclerodermis absent or weakly developed, hyalodermis rarely differentiated; axillary hairs up to 16
cells in length, the basal cell occasionally brown; red tomentum
sometimes present; heterotrichous, persistent protonema occasionally present, with green, much branched, aerial chloronemata
and green to red-brown caulonemata. Leaves about equal-sized to
near base, spreading-incurved from the insertion and subtubulose, weakly twisted to crisped or catenulate when dry, widely
spreading when moist, oblong- to linear-lanceolate, to 3.2 mm in
length; margins plane to weakly incurved, entire or sometimes
deeply dentate, occasionally bistratose in patches; apex narrowly
acute; base not or weakly differentiated in shape to short-ovate;
costa percurrent and ending in an apiculus to stoutly excurrent as
a short mucro, ventral superficial cells quadrate to short-rectangular, papillose, 4-6 cells across costa at midleaf, costa in
transverse section circular or ovate, ventrally commonly bulging,
with two stereid bands, both usually strong, the dorsal crescentshaped, two or more guide cells in one layer, a ventral and sometimes a dorsal epidermis present, hydroid strand absent; upper
/aminal cells subquadrate to hexagonal or elliptical, ca. 10-14
jlm in width, 1:1, somewhat bulliform, walls evenly thickened or
occasionally trigonous or irregularly thickened, lumens angular or
rounded; papillae usually massive, simple, bifid or multifid, usually centered over the lumens, (1-)2-4(-6) salients per lumen;
basal cells differentiated in a small group across the leaf base,
smooth, rectangular, yellowish to hyaline. Dioicous. Perichaetia
terminal, leaves abruptly sheathing below, otherwise little different from cauline leaves, somewhat larger or smaller. Perigonia
terminal, gemmate. Seta ca. 4-8 mm in length, reddish to yellowish brown, twisted clockwise above; theca ellipsoidal to shortcylindrical, brown, ca. 1.0-1.5 mm in length, stomates phaneropore, at base of theca, annulus not vesiculose to strongly so
but of persistent cells; peristome absent or present but eroded (or
possibly rudimentary), consisting of a basal membrane 20-25 Jlm
in height, papillose, retaining the extreme basal portions of one or
two narrow papillose teeth. Operculum long-rostrate, ca. I mm in
length, cells in straight rows. Calyptra not seen. Spores ca. 8-13
Jlm in diameter, essentially smooth. Lamina/ KOH color reaction
yellow.
Found on mostly calcareous substrates in moist areas in
U.S.A. (southeastern states and Alaska), Mexico, Central America, and Eastern Asia.
The major characteristics of Tuerckheimia are the narrow,
stoutly mucronate leaves with broadly channeled ventral surfaces,
acute apices, plane margins, and generally massive papillae centered over the lumens of the often rather large and bulliform upper
lamina! cells (Pl. 14, f. 6, 7, 15). The basal lamina! cells are
poorly differentiated (Pl. 14, f. 5) and do not run up the margins
as a vee, as in Tortella. Fruiting plants are rare and reproduction
in some species may take place largely through fragile leaf apices.
The deeply dentate leaves of Tuerckheimia valeriana are sugges-
tive of Leptodontium but the central strand and short cells of the
ventral surface of the costa are features distinguishing Tuerckheimia; T. valeriana, for which sporophytes are unknown, may
actually be a Ptychomitrium with anomalous papillae.
There is a resemblance to Eucladium, with its longlanceolate leaves, large upper !aminal cells generally larger
medially and with simple papillae, and the generally stoutly
excurrent mucro. The upper lamina! cells of Tuerckheimia
robusta have the angular lumens of Hymenostylium. The narrow, plane leaves of Tuerckheimia are like of those of Trichostomum, but species of that genus generally have small cells and
crowded, multiplex, flattened papillae covering most of the cell.
K. Saito annotated the type specimen of T. guatemalensis (Pl.
14, f. 8-13) with a new combination (unpublished) in
Trichostomopsis (a genus here referred to Didymodon sect.
Asteriscium), apparently based on bistratose upper margins.
These last were made much of by Brotherus (191 0) and Bartram
( 1949), but are merely patches of bistratose cells not comparable to the completely bistratose margins of, e.g. Didymodon
umbrosus, which has similar long-lanceolate leaves. Didymodon
sect. Asteriscium is easily distinguished by the much
differentiated leaf base and general lack of a ventral stereid
band. Quaesticula is similar in areolation but has infolded upper
!aminal margins, and a rounded-acute, often cucullate leaf apex.
Remnants of a papillose basal membrane that were found at
a capsule mouth (Pl. 14, f. 13) in the single known specimen
(Guatemala, Livingston, Tuerckheim, 1908, holotype, H) ofT.
guatemalensis indicate the presence of a peristome in this one
species. Additional material is necessary to ascertain the exact
morphology of this feature.
Crosby et al. (1992) found Gymnostomum angustifolium
Saito and thus its combination Tuerckheimia angustifolia
(Saito) Zand. to be invalid names because a single element was
not cited as holotype. The correct name for that taxon is now T.
svihlae (Bartr.) Zand.
Additional literature: Iwatsuki and Sharp (1958), Zander
(1978f).
Species recognized: 4.
Species examined: T. guatemalensis (H, NY), T. robusta
(BM), T. svihlae (BUF, DUKE, MEXU, TENN), T. valerianum
(DUKE, FH, MICH).
New heterotypic synonymy: Tuerckheimia angustifolia
(Saito) Zand. = Tuerckheimia svihlae (Bartr.) Zand.
New combinations: Tuerckheimia robusta (Dix.) Zand.,
comb. nov. (Merceyopsis rofmsta Dix., Ann. Bryol. 3: 59,
1930). Tuerckheimia svihlae (Bartr.) Zand., comb. nov. (Trichostomum svihlae Bartr., Rev. Bryol. Lichenol. 23: 245, 1954;
Oxystegus svihlae (Bartr.) Gangulee).
12. STREPTOCALYPTA
Plate 15.
Streptocalypta C. Miill., Linnaea 42: 353, 1879. Type: Streptoca/ypta /orentziana C. Miill.
Barnesia Card., Rev. Bryol. 37: 122, 1910. Type: Barnesia
tortelloides Card.
From atpE7tt6<;, twisted+ o + KCiA.U7tt6<;, covered, a calyptra,
a veil, cover, lid.
Plants loosely caespitose, green above, light brown below.
~
TRICHOSTOMOIDEAE
95
Plate 15. Streptocalypta. 1-8. S. lorentziana. 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6-7. Transverse sections from midleaf. 8.
Upper lamina! papillae. 9-12. S. pulchriretis. 9-10. Two leaves. 11. Leaf apex. 12. Transverse section at midleaf. 13-17. S. santosii. 13-14.
Two leaves. 15. Leaf apex. 16. Transverse section at midleaf. 17. Peristome. 18-20. S. tortelloides. 18. Perigoniate branch (autoicous bud). 19.
Leaf. 20. Leaf apex.
96
GENERA OF THE POTTIACEAE
Stems to 0.6 em in length, rounded pentagonal in transverse section, central strand present or occasionally absent, sclerodermis
not or little differentiated, hyalodermis occasionally
differentiated, axillary hairs of up to 10 hyaline cells or the basal
cells brownish; weakly radiculose. Leaves incurved and contorted
when dry, spreading when wet, ligulate-lanceolate, long-elliptical
or spathulate, to ca. 3.0 mm in length, upper lamina flat to
broadly channeled across leaf; margins plane or weakly incurved,
entire or distantly weakly denticulate above, spmetimes bordered
by elongate or less papillose cells; apex acute to rounded, usually
apiculate, occasionally somewhat cucullate; base little
differentiated in shape to ovate; costa ending up to 4 cells below
the apex or percurrent to short-excurrent, superficial cells ventrally quadrate and dorsally elongate, ca. 5-7 cells across costa ventrally at midleaf, costal transverse section with ventral stereid
band absent or present but weak, 1-4 layers of many (totalling ca.
7-20) guide cells (these essentially of equal size and often with
thickened walls), dorsal stereid band strong, reniform, dorsal epidermis not or little differentiated, hydroid strand absent; upper
lamina! cells quadrate to hexagonal, ca. 10-13 jlm in width, 1:1,
superficially bulging on both sides or more strongly ventrally than
dorsally; papillae bifid to sometimes multiplex, often rather small,
generally centered and 2-4 over each lumen; basal cells sharply
differentiated, extending up the leaf margins in a vee, bulgingrectangular, 10-35 jlm in width, 3-4: I, walls thin, smooth.
Dioicous or monoicous (synoicous, paroicous, autoicous or
rhizautoicous). Perichaetia terminal, inner leaves little
differentiated or narrower. Seta elongate, to I em in length,
brown, twisted clockwise, 1 per perichaetium; theca longelliptical to cylindrical, to 1.5 mm in length, yellowish brown,
exothecial cells rectangular to rhomboidal, walls thin to evenly
thickened, stomates phaneropore, at base of theca; annulus of
vesiculose cells; peristome absent or of 32 straight or weakly
twisted counterclockwise or rarely clockwise at the base, filamentous, densely spiculose teeth up to 500 jlm in length, basal
membrane low, spiculose or weakly papillose. Operculum longconic to rostrate, cells straight or twisted weakly clockwise,
0.5-1.2 mm in length. Spores (8-)10-15 jlm in diameter, yellow
to brown, weakly papillose. Calyptra cucullate, smooth, 1.5-2.0
mm in length. Lamina[ KOH color reaction yellow or light olive.
Found on soil and rock in mountainous regions of Latin
America and Sol)th Africa. Quite rare; known only from two
states in Mexico and single localities in Bolivia, Argentina and
South Africa (Natal).
This genus is similar to Tortella in the plane leaf margins,
basallaminal cells strongly differentiated in a vee-shape (Pl. 15, f.
5), small upper lamina! cells, and spiculose, generally twisted
peristome (slightly and probably secondarily twisted clockwise at
the base-Pl. 15, f. 17-in S. santosii as its opercular cells are in
straight rows), but Streptocalypta is distinguished by its internal
costal structure of several layers of guide cells, general lack of a
ventral stereid band (Pl. 15, f. 6, 7, 12, 16), and absence of a peristome in at least one species. Gertrudiella likewise has a cylinder
of mostly equal-sized guide cells but has strongly recurved upper
leaf margins and lacks the basal vee. Species of certain taxa in
other tribes (e.g. Tortula and Didymodon sect. Asteriscium) do
have more than one layer of guide cells, but are easily distinguished by other characters. The character of multistratose guide
cells is distinctive for the genus Streptocalypta, but is less pronounced inS. pulchriretis than in other species. The peristome of
S. pulchriretis is extremely fragmentary in the single known
collection; this species is very similar to S. lorentziana.
Although a few stereid cells are commonly present ventrally
in S. lorentziana (Pl. 15, f. 7) and usually several ventral
stereids inS. pulchreretis (Pl. 15, f. 12), the usual absence of a
ventral stereid band in the genus may be viewed as a derived
character, and is duplicated in Pseudocrossidium of the Barbuleae. Species of Tortula, of the Pottieae, may occasionally
exhibit a few ventral stereid cells.
A misprint in my previous treatment of the genus (Zander
1983a) gives lengths of the setae of S. santosii and S.
tortelloides in millimeters; it should have read "em".
Additional literature: Magill (1981, discussion as Weisiopsis
pulchriretis).
Number of accepted species: 3.
Species examined: S. lorentziana (BUF, F, H), S.
pulchriretis (BM), S. santosii (FH, MICH), S. tortelloides (NY,
PC, TENN).
New combinations: Streptocalypta pulchriretis (Dix.)
Zander (Weisiopsis pulchriretis Dix., Trans. Roy. Soc. S. Afr.
18: 252, 1930).
13. PLEUROCHAETE
Plate 16.
Pleurochaete Lindh., Oefv. K. Vet. Ak. Foerh. 21: 253, 1864.
Type: Pleurochaete squarrosa (Brid.) Lindh.
Barbula subg. Pleurochaete (Lindh.) Schimp., Syn. ed. 2:
220, 1876.
Tortella subg. Pleurochaete (Lindh.) Limpr., Laubm.
Deutsch!. 1:607, 1890.
Barbula sect. Pleurochaete (Lindh.) C. Mtill., Linnaea 39:
400, 1875.
Barbula sect. Squarrosae Lesq. & James, Man. N. Am. Moss.
130, 1884, nom. illeg. incl. sect. prior. Type: Barbula
squarrosa Brid.
Barbula sect. Squarrosa Lazaro e Ibiza, Bot. Oeser. Comp. A .
Esp. 1: 586, 1896, nom. illeg. incl. Barbula sect. Pleurochaete (Lindh.) C. Miill. Type: Barbula squarrosa Brid.
Barbula subsect. Squarrosae C. Mtill., Linnaea 39: 402, 1875.
From 1tAeUpd, side, or 1tAeUp6v, rib + 0 + xa{'tTJ, long hair,
mane; referring to the laterally borne sporophytes.
Plants forming a deep or sprawling turf, green above,
brown below. Stems branching irregularly, to 4 .0 em in length,
transverse section rounded-pentagonal, central strand small,
walls of central cylinder cells thin to weakly thickened, sclerodermis of 2-4 rows thick-walled cells, hyalodermis present;
axillary hairs to 15 cells in length, cells all hyaline;
indumentum weakly radiculose. Leaves spreading and strongly
contorted when dry, squarrose-recurved above a sheathing base
when moist, oblong-lanceolate, 3-5 mm in length, upper lamina
broadly channeled, margins plane but occasionally recurved to
revolute along leaf base, denticulate in upper 1/3 of leaf to
throughout, 1-2 marginal rows often weakly papillose distally
beyond a border of hyaline, thin-walled rhomboidal cells not
merging he/ow with inner basal cells but instead extending as a
distinct hyaline strip to leaf insertion, 4-7 cells in width below,
narrowing upwards, reaching barely higher than shoulder of leaf
base to 3/4 leaf length; apex usually sharply acute, occasionally
broadly acute; base ovate to rectangular, often broadly
~
TRICHOSTOMOIDEAE
97
Plate 16. Pleurochaete. 1-10. P. squarrosa var. squarrosa 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Upper marginal cells. 6. Upper lamina!
papillae. 7. Lamina! cells at leaf shoulder. 8. Transverse section of upper part of leaf. 9. Perigoniate bud. 10. Peristome. 11-16. P. squarrosa
var. luteola. 11. Transverse section of stem. 12. Leaf. 13. Midleaf marginal cells. 14. Basal cells. 15. Perichaetiate branch. 16. Perigoniate bud.
17-20. P. malacophylla. 17-18. Two leaves. 19. Leaf apex. 20. Transverse section at midleaf.
98
GENERA OF THE POTTIACEAE
sheathing and with distinct shoulders; costa short-excurrent as a
sharp mucro, superficial cells quadrate to short-rectangular and
papillose from top of sheathing leaf base to near apex ventrally,
elongate and smooth dorsally, 6-8 rows of cells across costa ventrally at midleaf, costal transverse section semicircular or
reniform, stereid bands strong ventrally and dorsally, larger dorsally, epidermis differentiated in one layer ventrally, present or
absent dorsally, guide cells 4-6 in l layer, hydroid strand absent;
upper laminal cells subquadrate, 8-11 jlm in width, l (-2): l, walls
thin or evenly weakly thickened, superficially bulging on both
exposed sides or only ventrally and then weakly convex dorsally;
papillae bifid, l-4 per lumen; basal cells differentiated and rising
higher medially, distinct from the 4-6 rows of border cells, rectangular to rhomboidal, 10-20 jlm in width, 2-5:1, walls thin to
evenly thickened or porose. Dioicous. Perichaetia on very short
lateral branches, inner leaves long-lanceolate, narrower than
cauline leaves, to 5 mm in length, sheathing the seta, lower cells
rhomboidal to rectangular and porose, reaching to 3/4 length of
inner leaves. Perigonia lateral on the stem, small. Seta 1.3-1.7 em
in length, 1 per perichaetium, orangish to reddish brown, twisted
clockwise; theca 2.0-2.8 mm in length, light yellowish or reddish
brown, cylindrical, exothecial cells rectangular to rhomboidal,
20-25 jlm in width, 3-4: 1, walls thin, stomates phaneropore, at
base of theca, annulus of 3-5 rows of vesiculose cells, persistent;
peristome teeth 32, filamentous, branched spiculose, 300-1000
jlm in length, often broken, with several articulations, twisted
once counterclockwise, basal membrane apparently absent or to
35 jlm high in height, low-spiculose. Operculum long-conic,
0.9-1.7 mm in length, cells twisted counterclockwise. Calyptra
cucullate, smooth, ca. 4 mm in length. Spores 10-13 jlm in diameter, yellowish brown, papillose. Laminal KOH color reaction
deep yellow to orange. Reported chromosome number n = 13.
Found in dry areas on soil and rock (generally calcareous),
occasionally on tree roots; North, Central and South America,
Europe, North and Central Africa, the Middle East and Asia.
Crum and Anderson (1981) pointed out that Pleurochaete is
quite like Tortella in several respects but differs in that the
differentiated thin-walled marginal cells, which extend up from
the insertion often to above midleaf, do not form a coherent basal
vee extending medially to the costa (Pl. 16, f. 7, 14). The inner
basal cells, instead, form a distinct region, much as in the case
with some but not all Pseudosymblepharis species. Pleurochaete
is much like the Asian Chionoloma in leaf shape, marginal strip
of elongate cells (Pl. 16, f. 13), and upper medial cells often bulging much greater ventrally than dorsally, but the latter has thickwalled border cells and the upper laminal margins are sharply
incurved as in Weissia .
Variation in the degree of extension of the basal marginal
cells up the leaf margins and in the length of the inner basal cells
is not quite continuous, but is correlated with plant stature.
Pleurochaete squarrosa has a largely Temperate Zone and
paleotropical facies (P. squarrosa s. str., found in Europe, Ethiopia, the Congo, southern U.S.A., Mexico) and a neotropical facies
(P. luteola, found in southeastern U.S.A., e.g. Tennessee, Zander
4333, BUF, and Arkansas, Crum & Anderson's exsiccat Mosses
of North America 951 as P. squarrosa; and Latin America).
These morphotypes are distinguishable in most specimens examined. Crum and Anderson (1981 ), however, felt that the
neotropical variant is only a "robust expression," not worthy of a
separate name; Crum (1951) indicated that, where growing
sympatrically, these may sometimes be difficult to name as one
or the other. In any case, a varietal name is provided here for
workers like myself who wish to distinguish among the two.
The var. luteola (Pl. 16, f. 11-16) differs in being more robust,
with a more highly sheathing leaf base, and the border of thinwalled marginal cells being strongly denticulate and extending
farther up the leaf margins. This parallels the distinction
between Molendoa hornschuchiana and M. sendtneriana, the
former differing in stature (largest in the genus) and denticulate
lower leaf margins; however, the larger species of Molendoa is
distributed primarily across the Alps and Himalayas (one station
is in Alaska), while the latter ranges from the Andes and North
American Cordillera across the Arctic and throughout the Eurasian continent in arctic-montane situations. Thus, if the each of
these two genera developed intraspecifically through reduction,
then Pleurochaete probably originated in Gondwanaland and
Molendoa is Laurasian, but vice versa if large stature and marginal denticulation are elaborations.
African material identified as Pleurochaete beccarii at BM
and NY is P. squarrosa var. squarrosa; type material of P.
beccarii at TR could not be obtained on loan but is probably
also synonymous. Pleurochaete malacophylla is doubtfully distinct from P. squarrosa var. squarrosa.
Additional literature: Nebel (1990), Quarterman (1956),
Wyatt and Stoneburner (1982).
Number of accepted species: 4.
Species examined: P. malacophylla (BM), P. squarrosa.
New combinations and statuses: Pleurochaete squarrosa
(Brid.) Lindh. var. luteola (Besch.) Zand., comb. et stat. nov.
(Trichostomum luteolum Besch., Mem. Soc. Sc. Nat. Cherbourg
16: 178, 1872; Pleurochaete luteola (Besch.) Ther.).
14. CALYPTOPOGON
Plate 17.
Calyptopogon (Mitt.) Broth., Nat. Pfl. 1(3): 419, 1902.
Streptopogon sect. Calyptopogon Mitt., Phil. Trans. R. Soc.
London 168: 33, 1879. Type: Calyptopogon mnioides
(Schwaegr.) Broth.
From K~U7tt6<;, covered + o + 1t<6yrov, -rovoc;, beard; referring to the ragged margin of the calyptra.
Plants growing in tufts or mats, yellowish green above,
brown below. Stems branching occasionally, 2-4 em in length,
transverse section rounded-pentagonal, central strand absent,
sclerodermis present, hyalodermis present, occasionally only in
patches; axillary hairs to 7 cells in length, all hyaline or 1-2
basal cells thick-walled; sparsely to densely radiculose. Leaves
incurved or catenulate, margins extremely undulate when dry,
widely spreading and reflexed when moist, long-ovate to ovatelanceolate, 3.5-4.5 mm in length, upper lamina flat to somewhat keeled, margins plane, nearly entire, minutely serrulate
with projecting transverse cell walls and papillae, bordered
intramarginally throughout length of leaf by 3-4 rows of rectangular epapillose porose cells as an extension of the region of
differentiated basal cells; apex acute, often narrowly channeled,
extreme cells bordering the mucro rhomboid; base little
differentiated in shape but tubulose-clasping; costa evenly
tapering, excurrent as a papillose mucro, superficial cells shortrectangular and distantly papillose ventrally, elongate and distantly papillose with forward-pointing cell ends dorsally, ca. 4
~
TRICHOSTOMOIDEAE
99
Plate 17. Calyptopogon. 1-17. C. mnioides. 1. Perichaetiate plant. 2. Perigoniate plant. 3. Transverse section of stem. 4-5. Two leaves. 6. Leaf
apex showing papillae. 7. Basal cells. 8. Transverse section at midleaf. 9. Transverse section near leaf base. 10. Upper leaf margin. 11.
Propagula. 12-13. Two perichaetialleaves. 14. Cells of perichaetialleaves. 15. Peristome teeth. 16. Operculum. 17. Calyptra.
100
GENERA OF THE POTTIACEAE
rows of cells across costa ventrally at midleaf, costal transverse
section ovate, stereid band absent ventrally, strong dorsally and
crescent-shaped, lumens of stereid cells smaller towards dorsal
surface, substereid near guide cells, ventral epidermis of bulging
or conic cells, dorsal epidermis absent, guide cells 2-4(--6) in
1(-2) layers, hydroid strand very large, ventral costal cells often
protuberant as scattered, ovate, thick-walled cells; upper /aminal
cells irregularly hexagonal, occasionally longitudinally or transversely elliptic, ca. 12-16 J.Lm in width, ca. 1:1, walls trigonous,
porose, superficially weakly convex on both sides; papillae small,
simple to bifid, conic-spiculose, solid, 3-4 per lumen, evenly scattered over the lamina; basal cells differentiated across leaf, rising
higher marginally and fusing with the inframarginal border, rectangular, little wider than upper cells, to 5-6:1, walls evenly
thickened to porose. Propagula multicellular, caltrop-shaped,
20-30 J.Lm in diameter, borne on the ventral surface of the costa.
Dioicous. Perichaetia terminal, inner leaves strongly
differentiated, linear-lanceolate, to 12-13 mm in length, strongly
sheathing, convolute, cells entirely rectangular to longrhomboidal, thick-walled and porose. Perigonia born terminally
on an equal-sized perigoniate plant as gemmate buds, often several per plant, each ending a branch. Seta short, ca. 0.5 em in
length, 1 per perichaetium, brown, twisted clockwise; theca
1.8-2.2 mm in length, brown, short-rectangular, occasionally
curved, exothecial cells rectangular, evenly thickened, ca. 20 J.Lm
in width, 2-4: 1, stomates phaneropore, at base of capsule, annulus
of 3-4 rows of somewhat vesiculose, persistent cells; peristome
often coming off with the operculum, teeth 32, filamentous,
densely low-spiculose, to 500 J.Lm, with many articulations,
twisted once counterclockwise, basal membrane high, to 300 J.Lm
in height, with round, thin-walled windows, densely lowspiculose. Operculum conic, ca. 1.5 mm in length, cells twisted
counterclockwise. Calyptra cucullate, somewhat rough above,
ragged and longitudinally undulate below, smooth, ca. 3 mm in
length. Spores ca. 13-15 J.Lm in diameter, brown, essentially
smooth. Lamina! KOH color reaction red, often blotchy, or yellow with red blotches, or yellow-orange.
This monotypic rupestral genus is restricted to southern South
America, New Zealand, Tasmania and Australia.
Sainsbury (1955) described the calyptra as mitriform, but it is
actually long-cucullate, weakly pleated and strongly frayed at the
base (Pl. 17, f. 17). The genus is distinctive in the combination of
the trigonous upper !aminal cells with small, simple to bifid, solid
papillae (Pl. 17, f. 6); intramarginal border of porose, rectangular,
epapillose cells (Pl. 17, f. l 0); single costal stereid band (Pl. 17, f.
8-9); nerve-borne, short-branching propagula (Pl. 17, f. ll );
convolute-sheathing perichaetial leaves of thick-walled porose
cells (Pl. 17, f. 12-13); and 32 filamentous peristome teeth (Pl.
17, f. 15). Sections of the costa near the base show more than one
hydroid strand and a clear differentiation of tissues into a crescent
of 2 or 3 superficial rows of stereids dorsally, and substereid cells
just ventral to that crescent, the significance of which is presently
unclear. Older axillary hairs apparently become thick-walled basally; hairs near the apex are entirely hyaline.
The ventral stereid band is absent, as is characteristic of
highly evolved taxa of the Pottioideae. This genus is strongly
reminiscent of Syntrichia papillosa (Pottieae), a species of broad
distribution fruiting, however, only in Australia (cf. Catcheside
1980 and Dixon 1923). Syntrichia papillosa shares the following
features with Calyptopogon: large, collenchymatous upper
!aminal cells; papillae simple (occasionally bifid); stereid band
crescent-shaped; no dorsal epidermis; hydroid strand strong;
propagula clavate with bulging cells, borne on ventral surface of
costa; perichaetial leaves strongly sheathing the costa, !aminal
cells rhomboidal and hyaline throughout; calyptra large; KOH
red. Syntrichia papillosa was thought by Kramer (1980) to possibly form its own section of Tortula. It differs from Calyptopogon by its stem having a (weak) central strand; awned, concave
leaves lacking a leaf border and not strongly undulate marginally when dry; upper lamina! cells smooth or only singly papillose; apically dorsally spinose costa; and perichaetial leaf cell
walls not thickened. Syntrichia pagorum has, like C. mnioides, a
semicircular costal section, which lacks dorsal epidermal cells
and bulges laterally over the dorsal surface of the leaf, and the
species likewise fruits only in austral regions (cf. Catcheside
1980; Stone 1971); S. pagorum differs, however, in the only
weakly differentiated perichaetial leaves and the relatively
small, multiplex-papillose upper lamina! cells characteristic of
Syntrichia. The cladistic study, however, places Calyptopogon
in the Trichostomoideae.
Number of accepted species: 1.
Species examined: C. mnioides (NY, US).
15. TORTELLA
Plates 18-19.
Tarte/la (Lindb.) Limpr., Laubm. Deutsch!. 1: 599, 1888, nom.
cons. Lectotype: Tortella caespitosa (Schwaegr.) Limpr.
Mallia subg. Tarte/la Lindb., Musci Scand. 21, 1879.
Tortella subg. Tortella Limpr., Laubm. Deutsch!. 1: 600, 1888, nom. illeg.
Barbula subg. Tortella (Lindb.) Kindb., Eur. N. Amer. Bryin.
2: 245, 1897.
Tortella subg. Eutortella Roth, Eur. Laubm. l : 344, 1903,
nom. illeg.
Trichostomum subg. Tortelloidea Roth, Eur. Laubm. 1: 315,
1903.
Tortula sect. Tortuosae De Not., Mem. R. Ace. Sc. Torino 40:
288, 1838. Type: Tortula tortuosa Hedw.
Tortula sect. Caespitosae De Not., Mem. R. Ace. Sc. Torino
40: 287, 1838. Type: Tortula caespitosa Schwaegr.
Barbula sect. Tortuosae BSG, Bryol. Eur. 2: 86, 1842 (fasc.
13-15 Mon. 24). Type: Barbula tortuosa (Hedw.) Web. &
Mohr.
Barbula sect. Tarte/la C. Miill., Syn. 1: 599, 1849, nom. illeg.
incl. sect. prior.
Barbula sect. Fragiles Schimp., Syn. 181, 1860. Type: Barbulafragilis (Hook. & Wits.) BSG.
Tortula sect. Fragiles (Schimp.) Lindb., Oefv. K. Svensk.
Vet. Ak. Foerh. 21: 214, 1864. Type: Tortella fragilis
(Hook. & Wits.) C . Hartm.
Mallia sect. Tortella (Lindb.) Braithw., Brit. Moss Fl. 1: 230,
247, 1885.
From tortus (a participle), twisted+ -ella, diminutive.
Plants loosely caespitose or forming turfs or cushions, green
to dark green above, brown to tan below. Stems branching occasionally or often, usually short but up to 4 em in length, in transverse section rounded-pentagonal, central strand absent, weak
or strong, cells of central cylinder often thick-walled, scleradermis present but usually weak, hyalodermis present, occa-
TRICHOSTOMOIDEAE
sionally weak or only in patches, composed of cells that are not or
little collapsed when mature; axillary hairs long, ca. 10-20 cells
in length, all hyaline; weakly radiculose, occasionally matted with
reddish brown rhizoids. Leaves often crowded, when dry
appressed, erect or incurved, often twisted or contorted, occasionally spiralled about stem, spreading when moist, ligu/ate to longlanceolate, rarely spathulate, 1.5-7.0 mm in length, upper lamina
flat, broadly channeled across leaf or weakly channeled along
costa, margins plane, occasionally erect or narrowly incurved or
tubulose, entire, weakly crenulate with projecting papillae or seldom weakly dentate near apex or below midleaf, rarely bordered
with elongate cells near apex; apex subulate or narrowly to
broadly acute, occasionally rounded or cucullate; base oblong,
elliptical or short-ovate, occasionally long-sheathing or not
differentiated in shape; costa ending 1-4 cells below apex,
percurrent or more usually short-excurrent as a mucro, superficial
cells quadrate or elongate, occasionally only quadrate at midleaf
ventrally, elongate dorsally, 2-6(-10) rows of cells across costa
ventrally at midleaf, costal transverse section semicircular or
ovate, stereid bands two, weak to strong ventrally, strong dorsally, epidermis present ventrally and present, weak or absent dorsally, guide cells 4-6(-8) in I layer, hydroid strand absent; upper
lamina! cells quadrate to hexagonal, occasionally bistratose, 7-15
J.lm in width, 1(-2):1, walls thin to evenly thickened, superficially
convex on both sides; papillae spiculose, bifid or occasionally
simple, 3-6 per lumen, seldom capituliform and centered; basal
cells differentiated across leaf in a vee-shape, usually sharply
differentiated, occasionally forming only a low vee or weakly
differentiated from the upper cells, rectangular, occasionally bulging, ca. 8-30 J.lm in width, 4-6:1, walls thin or evenly thickened
to porose, usually smooth. Asexual reproduction occasional,
apparently by fragile leaf apices or portions of lamina. Dioicous
or occasionally autoicous. Perichaetia terminal, inner leaves usually long-lanceolate, to 6 mm in length, usually sheathing the seta,
lower cells long-rhomboidal in lower 1/2. Perigonia terminal,
weakly gemmate, inner leaves smaller, or occurring as flattened,
stalked, buds in leaf axils of perichaetiate plants. Seta ca. 0.7- 3.0
em in length, I per perichaetium, yellowish, reddish or light
brown, twisted clockwise below, occasionally counterclockwise
above, seldom straight; theca ca. (1.0-)1.5-3.0 mm in length, yellowish to reddish brown, cylindrical or occasionally elliptical,
exothecial cells rectangular, 20-45 J.lm in width, 3-5: l, seldom
hexagonal, walls thin, stomates at base of theca, phaneropore,
annulus of 1-4 rows of weakly or strongly vesiculose cells, persistent; peristome teeth 32, linear, spiculose, occasionally
branching-spiculose, ca. 550-1400 J.lm, with many articulations,
twisted counterclockwise, usually two or three times, basal membrane absent or low, papillose to spiculose, or rarely absent, or
capsule rarely cleistocarpous and then elliptical and longapiculate. Operculum long-conic to rostrate, ca. 1.0-2.5 mm in
length, cells twisted counterclockwise, seldom undifferentiated.
Calyptra cucullate, smooth, 2.5-3.5 mrn in length. Spores 8-20
J.lm in diameter, yellowish brown, essentially smooth or papillose.
Lamina! KOH color reaction usually yellow, sometimes yellowish
orange or reddish brown, leaves often yellow when immature but
orange lower on the Stem. Reported chromosome number n = 7,
13, 13+m, 14, 15, 26, 30, 52.
Found on soil, rock or organic substrates on all continents.
Tortella and Trichostomum (inCluding subg. Oxystegus) are
distinguished at present by characters that are somewhat variable.
101
A thorough reevaluation is needed through revision of both genera together. Tortel/a simplex, for instance, has much the
appearance of Trichostomum brachydontium, especially in the
characteristic weakly reflexed apex ending in a stout mucro, and
its basal cells are only weakly differentiated as a vee; this species may be more closely related to Trichostomum than to Tortel/a in spite of its somewhat twisted peristome.
The vee-shaped area of basal cells is generally easily distinguished in most species of Tortel/a, yet, in some, the
differentiated basal cells do not rise very high along the margins
(Pl. 18, f. 6, forming a rather low vee) or they are not sharply
different in size and shape or thickness of cell walls from the
upper !aminal cells. Some species with a vee-shaped basal cell
region (e.g. Tortel/a flavovirens) have untwisted, often short
peristomes, similar to those of Trichostomum s. lat.
Two autoicous species, Tortella lilliputanum (Pl. 19, f. 1-3)
and T. fruchartii, have cleistocarpous capsules and a distinctly
vee-shaped basal !aminal cell area, plus a stem section with a
characteristic well developed tortellaceous hyalodermis. The
gametophytes are clearly Tortel/a; the two taxa are much alike
and may be conspecific. Species of Tortella with reduced
sporophytes (T. eckendorffii Jacks a peristome) are presently
few, but more should be added to Tortel/a when Hyophila and
other eperistomate genera are revised, e.g. Tortel/a walkeri is
eperistomate and stegocarpous.
Pseudosymblepharis is usually distinguishable by the narrow upper laminae and sharply broadened and sheathing leaf
bases, but specimens of Tortel/a tortuosa with cirrhate leaves
have much the appearance of Pseudosymblepharis. Some species of Pseudosymb/epharis, also, are much like Tortel/a in the
broad upper laminae. Trichostomum hibernicus has these characters of Pseudosymblepharis but is obviously closely related to
Trichostomum (subg. Oxystegus) tenuirostre by the upper
!aminal areolation of evenly thickened, rectangular cells,
although the well developed sclerodermis is a character more
common in subg. Trichostomum. Collections of Pseudosymblepharis schimperianum of small stature have been in the past
assigned to Tortella (e.g. the synonyms Tortel/a mollissima,
Tortel/a richardsii and Tortella subfragilis) because of a tendency for such small plants to have less well-marked sheathing
leaf bases. The names Tortel/a, Trichostomum, Pseudosymblepharis and Oxystegus may actually represent only one genus,
characterized by a usually vee-shaped basal cell arrangement, a
vee-shaped transverse section of the dorsal stereid band (in most
species), an often thick-walled central cylinder, and sturdy, noncollapsing hyaloderm cells, but further research is necessary.
Barbu/a, Didymodon, Bryoerythrophyllum, and even Pseudocrossidium have occasionally been treated even in recent times
(e.g. Nyholm 1989) as a single genus under the name Barbula s.
lat., and it has been a temptation to similarly unify Tortella and
Trichostomum s. lat. here pending further study. As a possible
parallel to be pursued in future studies of Tortella and Trichostomum, the above four groups of Merceyoideae have been successfully separated in recent studies (Saito 1975a; Zander
1978e, 1978g, 1979f, 198la, 198lc) as genera each with considerable variation in the development of the peristonie, but taxonomically distinguishable by gametophytic features . Because it
is thought, however, that good gametophytic characteristics will
eventually be used to place at least the type species of Tortella
and Trichostomum in separate genera, these two generic names
102
GENERA OF THE POTTIACEAE
Plate 18. Tortel/Q. 1-10. T. humilis. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells 7. Transverse section
at midleaf. 8. Perichaetialleaf. 9. Perigonial bud. 10. Peristome. 11-13. T. bryotropica. 11-12. Two leaves. 13. Leaf apex. 14-16. T. fragilis.
14-15. Two leaves. 16. Transverse section of stem.
TRICHOSTOMOIDEAE
103
Plate 19. Tortella. 1-3 T.lilliputana. 1. Habit. 2-3. Two leaves. 4-6. T.japonica. 4-5. Two leaves. 6. Leaf apex. 7-10. T. rubripes. 7-9. Three
leaves. 10. Leaf apex. 11-13. T. tortuosa. 11-12. Three leaves. 13. Leaf apex. 14-17. T. xanthocarpa. 14-15. Two leaves. 16. Leaf apex. 17.
Transverse section at midleaf.
104
GENERA OF THE POTTIACEAE
are accepted here as representing different taxa, albeit with the
counsel that the present descriptions are based on species many of
which will probably be redistributed at some future date among
two or more generic names. It is emphasized here that Tortella
cannot be adequately revised taxonomically unless Trichostomum
s. lat. and Hyophila are reviewed at the same time.
Species of Torte/la and some related genera may usually be
distinguished from many other plane-margined taxa, mainly those
of the Barbuleae, by the stem section. Although presence of a central strand is variable, as a rule the hyalodermis is present, composed of rectangular (in transverse section) cells that are seldom
collapsed in mature parts of the stem, and the sclerodermis is generally only weakly developed, often of substereid cells only
slightly smaller than those of the central cylinder or of a few scattered stereid cells. Often, the cells of the central cylinder are
thick-walled. Taxa of Merceyoideae with a hyalodermis have it
composed of rounded cells, these usually collapsed except in the
region of the extreme stem apex, and have a distinct sclerodermis
of one or more layers of stereid cells.
Pleurochaete has a similar vee of basal cells, but in that genus
the thin-walled basal marginal cells form a group easily distinguished from the thicker-walled inner basal cells.
Selected bibliography: Crundwell and Nyholm (1962, 1963),
Dixon (1900), Gorton and Eakin (1958), E. J. Hill (1913),
Nicholson (1910), Zander and Hoe (1979).
Number of accepted species: 53.
Species examined: T. acaulon (NY), T. alpicola (BUF,
CANM, NY), T. bryotropica (BUF), T. cirrifolia (NY), T.
cryptocarpa (NY), T. cyrtobasis (BM), T. densa (DUKE, NY), T.
eckendorffii (PC), T. flavovirens (BUF), T. fragilis, T. fruchartii
(H, NY), T. germainii (NY), T. goniospora (NY), T. hildebrandtii
(NY), T. humilis, T. inclinata (BUF), T. injlexa (BUF), T.
japonica (BUF), T. knightii (MO, NY), T. lilliputana (NY, S), T.
linearis (NY), T. nitida (BUF), T. novae-valesiae (H), T.
pseudocaespitosa (NY), T. rigens (DUKE, NY), T. rubripes
(NY), T. simplex (US), T. somaliae (NY), T. tortuosa, T.
xanthocarpa (H, NY).
New heterotypic synonymy: Astomum latifolium Broth. in
Roth= Torte /Ia fruchartii (C. Miill.) Zand. Tort ella tortel/oides
(Greene) Robins. in Llano. = Tortella alpicola Dix. Trichostomum sitkanum Card. & Ther. = Tortel/a tortuosa var. arctica
(Arnell) Broth. in Fedch.
New combinations: Torte/la cryptocarpa (Broth.) Zand.,
comb. nov. (Astomum cryptocarpum Broth., Bih. K. Svensk.
Vet. Ak. Hand!. 26 Afd. 3(7): 19, 1900).
Tortella eckendorffii (P. Yarde) Zand., comb. nov .
(Hymenostomum eckendorffii P. Yarde, Rev. Bryol. Lichenol.
11: 170, 1939).
Torte/la fruchartii (C. Miill.) Zand., comb. nov. (Phascum
fruchartii C. Miill., Flora 71: 4, 1888; Astomum fruchartii (C.
Miill.) Broth.).
Tortel/a goniospora (C. Miill.) Zand., comb. nov. (Barbula
goniospora C. Miill., Hedwigia 37: 131, 1898).
Tortel/a lilliputana (C. Miill. ex Roth) Zand., comb. nov.
(Phascum lilliputanum C. Miill. ex Roth., Aussereur. Laubm.
212, 1911 "liliputanum"; Tetrapterum lil/iputanum (C. Miill. ex
Roth) Broth.).
Tortel/a linearis (Web. & Mohr.) Zand., comb. nov. (Barbula linearis Web. & Mohr, Ind. Mus. Pl. Crypt. 2, 1803;
Tuerckheimia linearis (Web. & Mohr) Britt.; Oxystegus linearis
(Web. & Mohr.) Hilp.).
Tortel/a somaliae (C. Miill.) Zand., comb. nov. (Hyophila~
somaliae C. Miill., Linnaea 40: 293, 1876).
Tortella walkeri (Broth.) Zand., comb. nov. (Hyophila
walkeri Broth., Rec. Bot. Surv. India 1: 317, 1899).
Subfamily MERCEYOIDEAE
Merceyoideae Broth., Nat. Pfl. ed. 2, 10: 246, 1924.
The Merceyoideae subclade (as constituted in Cladograms 14-16) is readily discernible in general fonn and composition of tenninal
taxa in Cladograms 2-5 and 8-10; while Tetracoscinodon is the basal taxon of the Merceyoideae in Cladograms 2, 5 and 8-10. The
traits of the ancestral node distinguishing the Merceyoideae (and the Pottioideae) from branches lower in the tree (i.e. Timmielloideae, Erythrophyllopsoideae, Gertrudielloideae, Chionolomoideae and Timmielloideae) are stem hyalodennis absent and perichaetial
leaves sheathing. The character states at the immediate ancestral node of the Merceyoideae lineage are leaves keeled above midleaf
and costa grooved ventrally. The Merceyoideae may be generally described as taxa with stem sclerodennis usually differentiated
from the inner cylinder of cells, leaves lanceolate, basal portion of leaf commonly differentiated in shape and ovate to elliptical,
laminal margins usually recurved below, upper laminal cells comparatively small, costa usually with two stereid bands and often
grooved along the ventral surface which in most taxa is only 2(-4) rows of cells across, and clavate propagula commonly present in
certain genera (rarely present in other subfamilies).
Tribe TETRACOSCINODONTIEAE
Tetracoscinodontieae Zand., tribus nov. Type: Tetracoscinodon R. Br.
Plantae elongatae, longitudine 1-3 em, tenues; caules nigri in sectione sclerodermide praediti, cellulis amplificatis cylindri
centra/is et filo centrali carentes; folia longitriangularia vel linearia, apicem angustatum, setaceum evolventia, costa stratis
,
stereidarum duobus praedita; theca annulum crassum, circumstomalem evolvens.
Plants elongate, 1-3 em in length, thin; stem black, in section with sclerodennis, enlarged cells of the central cylinder, and central strand absent; leaves long-triangular to linear, with a narrow, setaceous apex; costa with two stereid bands; theca with a thick
circumstomal ring.
The tribe Tetracoscinodontieae is monotypic; it is apparently the most primitive member of the Merceyoideae. It has the characters of the single species, Tetracoscinodon irroratus, notably the black stem with hyalodennis absent, comparatively large central
cylinder cells below a distinct sclerodennis, and central strand absent; a long-triangular or linear leaf shape and setaceous apex;
superficially bulging upper laminal cells with crowded, bifid papillae; mouth of capsule with a thickened circumstomal ring, and
laminae with a yellow color in KOH. Known only for New Zealand from very wet habitats.
Plate 20.
16. TETRACOSCINODON
Tetracoscinodon R. Br. ter, Trans. & Proc. New Zealand Inst. 29:
532, 1897. Type: Tetracoscinodon hectorii R. Br. ter.
From 'tE'tpci-, four+ Ko<:nctvov, sieve + d~ouc; (d~rov),
oMvtoc;. tooth.
Plants growing in clumps or turf, green above, tan below.
Stems blackish brown, branching occasionally, 1-3 em in length,
transverse section rounded-pentagonal, central strand absent,
cells of the central cylinder large, 30--45 Jlm in diam., sclerodennis present, of 1-2 layers of stereids, hyalodennis absent;
axillary hairs ca. 10 cells in length, all hyaline or basal cell thicker
walled; rhizoids sparse. Leaves when dry appressed-incurved,
when moist rigid, weakly spreading from the insertion, longtriangular to linear, 2-3 mm in length, upper lamina broadly and
deeply channeled across leaf, margins plane or seldom weakly
recurved, entire (minutely crenulate by projecting cell walls and
papillae); apex setaceous and very narrowly obtuse; base triangular, broadest at insertion; costa percurrent, superficial cells on
both sides of costa variously papillose and quadrate or elongate
and smooth, 4-6 rows of cells across costa ventrally at midleaf,
costal transverse section semicircular, two stereid bands present,
epidermis present on both sides of costa, guide cells 4-6 in 1
layer, hydroid strand absent; upper !aminal cells quadrate to
short-rectangular, 8-12 Jlm in width, 1-2:1, walls evenly thickened, superficially bulging on both sides; papillae bifid,
crowded, 3-6 per lumen; differentiated basal cells filling the triangular leaf base, rectangular, little wider than upper cells,
3-5:1, walls evenly thickened, smooth, bulging more strongly
dorsally than ventrally, somewhat longer on the margins.
Dioicous. Perichaetia tenninal, inner leaves lanceolate, little
longer than the cauline, convolute-sheathing in lower half,
lower cells long rhomboidal in sheathing portion. Perigonia terminal, not gemmate, inner leaves sheathing below, paraphyses
linear. Seta ca. 1 em in length, 1 per perichaetium, brown,
twisted clockwise; theca 0.6-1.5 mm in length, dark brown,
short-cylindrical, exothecial cells rectangular, thin-walled,
stomates phaneropore, at base of theca, mouth of capsule rather
thick-walled with a somewhat swollen circumstomal ring several layers of cells in thickness bearing above it 4-6 rows of
dark brown, vesiculose annular cells; peristome teeth ca . 16,
flat, short-triangular, strongly incurved, much perforated and
often cleft into 2-4 rami, smooth, covered externally by a
hyaline membrane, 150-200 Jlm in length, with ca. 6-10 articulations, straight, basal membrane absent or present but low,
smooth. Operculum long-conic, usually inclined, 0.5-1.0 mm in
length, cells straight. Calyptra cucullate, smooth, ca. 2 mm in
length. Spores large, 25-35 Jlm in diameter, brown, papillose.
~
106
GENERA OF THE POTTIACEAE
Plate 20. Tetracoscinodon. 1-16. T. irroratus. 1. Plant with sporophyte. 2. Habit of sterile plant. 3. Transverse section of stem. 4-7. Four
leaves. 8. Leaf apex. 9. Upper marginal cells. 10. Basal cells. 11. Transverse section through upper leaf. 12. Transverse section through leaf
base. 13. Upper laminal papillae. 14. Perichaetialleaf. 15. Thecal mouth (vertical view) showing portion of annulus, inner cells of circumstomal
ring and peristome teeth. 16. Peristome teeth (lateral view).
MERCEYOIDEAE
Lamina/ KOH color reaction yellow.
This is an infrequent taxon restricted to New Zealand where it
is found usually encrusted with limestone deposits on dripping
rocks.
Dixon (1923) was correct in synonymizing Brown's Tetracoscinodon hectorii with Eucladium irroratum as evidenced by
authentic material of both names at NY, but this species differs
widely from Eucladium ((q.v.) in important features that require
recognition of a separate genus. Tetracoscinodon shares many
apparently significant characters with Leptobarbula and Barbula,
such as the stem section showing rather large central cylinder
cells and a strong sclerodermis (Pl. 20, f. 3), a long-triangular leaf
shape (Pl. 20, f. 4-7), superficially bulging upper !aminal cells
with crowded bifid papillae (Pl. 20, f. 13), and yellow color in
KOH. It shares with Tridontium (Grimmiaceae, Scoulerioideae)
and Dialytrichia a hygric habitat and relatively pachydermous
capsule. Like Tridontium (see Excluded Taxa), the peristome (Pl.
20, f. 15-16) of Tetracoscinodon has an adherent hyaline membrane, but in the former genus the teeth are closely spiculose. The
peristome of Tetracoscinodon, in being smooth, extremely thin
107
periclinally, and in having transverse joints that often do not go
all the way across the tooth but instead angle down to intersect
the next upper or lower joint, has the appearance of derivation
from the outer rather then the inner peristomial layer. This
needs further study. The most salient characters of Tetracoscinodon are its black stems with large cells in the central cylinder and lacking a central strand; the rigid, long-triangular to linear leaves with narrowly obtuse apices and papillose, quadrate
to short-rectangular superficial cells dorsally; the rather large
and crowded bifid upper !aminal papillae; the rather thickwalled basal cells (Pl. 20, f. 10); the circumstomal ring of the
capsule; and the calcareous incrustation generally found on the
lower parts of the plants.
Additional literature: Brotherus (1924-25), Sainsbury
(1955).
Number of accepted species: 1.
Species examined: T. irroratus (NY).
New combination: Tetracoscinodon irroratus (Mitt. in
Hook. f.) Zand., comb. nov. (Weissia irrorata Mitt. in Hook. f.,
Handb. New Zealand Fl. 404, 1867).
Tribe BRYOERYTHROPHYLLEAE
Bryoerythrophylleae Zand., tribus nov. Type: Bryoerythrophyllum Chen.
Plantae plerumque magnae; folia plerumque lanceolata, lamina superna plerumque in solutione KOH rubra, marginibus recurvis
vel revolutis, saepe denticulatis vel dentatis, costa ventraliter e 2(-4) seriebus cel/ularum composita, in sectione complanata, strata
stereidarum duo evolventi, strata dorsali saepe in sectione remformi, fila hydroideo praedita; peristomium e 32 dentibus
similaribus compositum.
Plants often large, leaves usually lanceolate, upper lamina usually red in KOH, margins recurved or revolute and often denticulate or toothed, costa with 2(-4) rows of cells ventrally, flattened in section, with two stereid bands, dorsal stereid band often
reniform in section, hydroid strand present, and peristome teeth of 32 similar rami.
This subclade of the Merceyoideae is distinguished from the Tetracoscinodontieae by the leaf margins recurved to revolute
below midleaf, costal ventral cells elongate, and 2(-4) rows of cells across ventral surface of costa. The traits of the immediate
ancestral node are: costa flattened in section and possessing a hydroid strand, theca 1.5-3.5 mm in length, and peristome teeth of 32
similar rami. The Bryoerythrophylleae consists largely of taxa that have laminae red in KOH, upper margins commonly denticulate
or toothed, the dorsal costal epidermis often absent, and a dorsal stereid band that is reniform in section. Ancestors of this tribe were
probably adapted to hygric environments. The tribe's distribution is nearly worldwide, generally in mountainous areas.
17. DIAL YTRICHIA
Plate 21.
Dialytrichia (Schimp.) Limpr., Laubm. Deutsch!. 1: 691, 1888.
Type: Dialytrichia brebissonii (Brid.) Limpr.
Barbula subg. Dialytrichia Schimp., Flora 47: 211, 1865.
Barbula sect. Dialytrichia (Schimp.) Schimp., Syn. ed. 2: 222,
1876.
From BtaA.t5-, separated, disbanded (BtaA.t5etv, to part asunder)
+ 9pfx. tplxoc;, hair; referring to the fine peristome teeth
Plants hydrophilic, forming a turf, yellowish green above,
light brown below. Stems branching occasionally, to 2-3 em in
length, transverse section pentagonal, central strand strong,
sclerodermis weak, hyalodermis absent; axillary hairs of 6-15
cells, basal 1-4 cells yellow and thicker walled; radiculose below.
Leaves appressed to contorted, conduplicate when dry, weakly
spreading when moist, oblong to oblong-lanceolate, 2.5-3.5 mm
in length, upper lamina narrowly grooved along costa, margins
recurved below midleaf, entire, thickened as a 2-4-stratose border 2-3 cells in width, border cells quadrate and walls weakly
thickened; apex broadly acute to rounded; base oblong, scarcely
differentiated in shape, weakly sheathing; costa tapering, excurrent as a short, stout mucro, superficial cells elongate, weakly
papillose Ventrally, with small, simple scattered papillae dorsally, 2-4 rows of cells across costa ventrally at midleaf, costal
transverse section reniform, two stereid bands present (of
substereid cells), dorsal band lunate, ventral epidermis present
but weakly differentiated, dorsal absent or very weakly
differentiated, guide cells 4 in 1 layer, hydroid strand present,
often more than one; upper /aminal cells subquadrate, small,
8-10 J.lm in width, 1: 1, walls thin to weakly evenly thickened,
superficially strongly bulging on both sides of lamina; papillae
small, bifid, solid, 3-4 per lumen; basal cells differentiated
across leaf, rising higher medially, rectangular, little wider than
upper cells, 2-4:1, walls thin to evenly weakly thickened.
Dioicous. Perichaetia terminal, inner leaves oblong-lanceolate,
to 3 mm in length, sheathing in lower l/2-2/3, cells rhomboidal
and thin-walled in sheathing portion. Perigonia terminal as a
single, small, inconspicuous gemmate bud. Seta ca. 0.8-1 .0 em
~
108
GENERA OF THE POTTIACEAE
Plate 21. Dialytrichia. 1-12. D. mucronata. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8.
Transverse section at midleaf. 9. Inner perichaetial leaf. 10. Exothecial cells. 11. Transverse section of thecal wall. 12. Peristome teeth.
MERCEYOIDEAE
109
Plate 22. Rhexophyllum. 1-9. R. subnigrum. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7.
Transverse section at midleaf. 8. Upper !aminal papillae. 9. Perichaetialleaf.
110
GENERA OF THE POTTIACEAE
in length, 1 per perichaetium, reddish brown, twisted clockwise;
theca 2.5-3.0 mm in length, yellowish to orangish brown, cylindrical, occasionally curved, thecal walls fleshy, 3-4 cells in thickness, exothecial cell walls evenly thickened, 20-28 J.lm in width,
2: I, stomates phaneropore, on the fleshy neck, annulus of 2-4
rows of small, weakly vesiculose cells; peristome teeth 32, somewhat paired, filamentous, finely spiculose, ca. 700 J.lm, with many
articulations, weakly twisted counterclockwise, basal membrane
75-90 J.lm in height, fmely spiculose. Operculum conic, ca. 1.3
mm in length, cells twisted counterclockwise. Calyptra cucullate,
smooth, ca. 3 mm in length. Spores 13-15 Jlm in diameter, light
brown, finely papillose. Lamina/ KOH color reaction yellow to
yellowish orange.
A monotypic genus found on wet calcareous rock (occasionally submerged); Europe, northern Africa, and eastern and southern Asia.
Corley et al. (1981) regard the single species of this genus as
belonging to Cinclidotus, which Saito (1975a) regards as comprising a separate family in its own right, the Cinclidotaceae; the peristome (Pl. 21, f. 12) and laminal areolation (Pl. 21, f. 6) of Dialytrichia, however, are those of the Pottiaceae. The yellow KOH
reaction; oblong leaves (conduplicate when dry) narrowly
grooved along the costa ventrally; fme, solid, crowded, bifid
papillae; mucronate apex; presence of two stereid bands in the
costa (Pl. 21, f. 8); and a striking superficial resemblance to Barbuta unguiculata are characters indicating that Dialytrichia is correctly placed with the Merceyoideae; cladistic study puts it with
the Bryoerythrophylleae, however, rather than the Barbuleae.
It is probable that Dialytrichia represents a largely palearctic
representative of a now largely extinct, possibly Andean assemblage (Mironia, for instance, has three species, two Andean, one
widely distributed along the Latin American Cordillera), paralleling the situation with Pseudocrossidium, a large, essentially
Andean genus that likewise has a few northern and in its case
much reduced outlier species in the Northern Hemisphere, including Europe. Dialytrichia has the KOH yellow color reaction that
is plesiotypic in its lineage (see Cladogram 15), and the one
extant species may have survived through adaptation to a specialized habitat.
Additional literature: Bizot and Roux (1968), Gyorffy (1911),
Sergio and Sim-Sim (1984).
Number of accepted species: I .
Species examined: D. mucronata (BUF, DUKE, NY).
18. RHEXOPHYLLUM
Plate 22.
Rhexophyllum Herz., Biblioth. Bot. 87: 38, 1916. Type: Rhexophyllum laciniatum Herz.
Neocardotia Ther. & Bartr. in Ther., Smiths. Misc. Coli. 85(4):
12, 1931. Type: Neocardotia subnigra (Mitt.) Ther. & Bartr.
in Ther.
From p..,~t~, -Em<;, a breaking+ o + qn5A.A.ov, leaf; the upper leaf
margins of this genus split easily.
Plants forming a turf, green to blackish green above, reddish
brown below. Stems often branching, ca. 2-4 em in length, transverse section rounded-pentagonal, central strand present, scleradermis strong, hyalodermis present; axillary hairs of ca. 10 cells,
all hyaline or basal 1-3 cells brownish; rhizoids often present,
reddish brown. Leaves conduplicate, twisted, appressed-incurved
when dry, widely spreading to squarrose when moist, lanceolate, mostly 2.0-3.0 mm in length, keeled above, margins
recurved in lower 114-112, sharply dentate to erose-dentate in
upper 1/4-1/2; apex acute, fragile, often absent; base rectangular, somewhat sheathing, basal margins short-decurrent; costa
excurrent as a sharp, smooth mucro, superficial cells elongate
ventrally, 3-4 across costa at midleaf, short-rectangular to quadrate above, serrulate dorsally by projecting papillae at distal
ends of cells, costal transverse section reniform, two stereid
bands present, the dorsal crescent-shaped, epidermis absent or
weakly differentiated ventrally, present dorsally as cells with
enlarged lumens, guide cells 2-4 in 1 layer, hydroid strand
sometimes present; upper /aminal cells subquadrate to roundedhexagonal, mostly 7-10 Jlm in width, 1:1, walls thin, superficially bulging, bistratose in patches across leaf; papillae
crowded, multifid or 1-4 irregular plates per lumen, solid or
hollow; basal cells differentiated across leaf base, rectangular,
8-15 Jlm in width 3-5:1, walls thin. Dioicous. Perichaetia terminal, inner leaves highly differentiated, long-lanceolate, 4-6
mm in length, convolute-sheathing, lower cells long-rhomboidal
in lower 1/2-7/8. Perigonia terminal, leaves little differentiated
from the cauline. Seta ca. 1 em in length, 1 per perichaetium,
brown, twisted clockwise above; theca 2.5-3.5 mm in length,
brown, ellipsoidal to cylindrical, exothecial cells shortrectangular, 1-3:1, stomates present at base of theca; annulus of
ca. 2 rows of strongly vesiculose cells, deciduous in pieces;
peristome absent, operculum short-conic to conic-rostrate,
0.5-1.0 mm in length, cells straight. Calyptra cucullate, smooth,
ca. 3.0 mm in length. Spores 13-15 Jlm in diameter, weakly
papillose, brown. Lamina/ KOH color reaction red.
Known from southwestern U.S.A., Mexico, and the Andes
of Bolivia and Peru, where it is found on soil, rock and trees in
montane situations.
The genus is readily recognized by the plant coloration:
blackish green above and reddish brown below; leaves lanceolate with an oblong base (Pl. 22, f. 3-4), squarrose when moist,
upper margins sharply and deeply dentate (Pl. 22, f. 5); upper
laminal cells bistratose in patches across the leaf (best seen in
section, Pl. 22, f. 7); dorsally serrulate costa; and eperistomate
capsule. It is similar to Leptodontium in the leaf shape and
stance, elongate cells on the ventral costal surface (Pl. 22, f. 5),
reniform costal section, and much differentiated perichaetial
leaves, but differs mainly in the red color reaction with KOH,
bistratose laminae and a hydroid strand (at least occasionally) in
the costa.
Theriot and Bartram (Theriot 1931) felt that the genus was
related to Leptodontium because of the habit of the plant, the
dentate cauline leaves, and the strongly sheathing perichaetial
leaves. Hilpert (1933) placed Rhexophyllum near Leptodontium
and Triquetrella, considering the eperistomate condition to be a
reduction from a leptodontioid peristome structure. He suggested that the small leaf cell size, the dense papillae, and the
patchy bistratose upper lamina are xeromorphic adaptations.
Cladistic analysis, however, places it with the Bryoerythrophylleae.
Additional literature: Zander (1976).
Number of accepted species: 1.
Species examined: R. subnigrum (BUF, FH, JE, MEXU,
NY, TENN, US).
MERCEYOIDEAE
111
Plate 23. Mironia. 1-11. M. stenotheca. l. Transverse section of stem. 2. Habit of perichaetiate plant. 3. Habit of perigoniate plant. 4-6. Three
leaves. 7. Leaf apex. 8. Basal cells. 9. Transverse section at midleaf. 10. Upper laminal papillae. 11. Peristome at capsule mouth. 12-14. M.
crassicuspis. 12-13. Two leaves. 14. Propaguloid leaf apex. lS-18. M. ehrenbergiana. 15-17. Three leaves. 18. Leaf apex, lateral view.
112
GENERA OF THE POTTIACEAE
Plate 23.
19.MIRONIA
Mironia Zander, nom. nov.
Morinia Card., Rev. Bryol. 37: 124, 1910. Type: Morinia
trichostomoides (Besch.) Card., Mexico, D.F., Desierta Vieja,
Bourgeau 1335, holotype, PC; isotype, NY. (Non Morinia
Berlese & Bresadola 1889 = Rinomia Nieuwl. , nee Morinia
Linn. 1753.)
Named for L'Abbe F. Morin, author of "Anatomie comparee et
experimentale de la feuille des Muscinees. Anatomie de la
nervure appliquee a la classification," 1893, These, Rennes,
France. Like the fungus name Rinomia, the new name is a simple
anagram.
Plants in cushions, yellowish or reddish green above, reddish
brown below. Stems occasionally branching, to 4.0 em in length,
rounded-pentagonal in transverse section, central strand strong,
sclerodermis present, hyalodermis absent; axillary hairs of ca.
6-10 cells, the basal 1-2 yellowish or brownish. Leaves twisted,
appressed to erect when dry, erect-spreading to squarrose from
top of sheathing base when wet, ovate- to oblong-lanceolate,
1.5-3.5 mm in length; margins narrowly recurved in lower 3/4,
entire or dentate, narrowly bistratose in upper 113; apex broadly
to narrowly acute, occasionally thickened as a fragile or deciduous multistratose propagulum; base oblong and sheathing; costa
percurrent or excurrent as a short, sharp mucro, superficial cells
ventrally quadrate or elongate, dorsally elongate, ca. 4--Q cells
across ventral surface of costa, costal transverse section reniform,
with two stereid bands (the ventral often weak), epidermis present
on both sides (dorsally weak), guide cells 2-5 in 1 layer, hydroid
strand present; upper !aminal cells subquadrate, occasionally
short-rectangular, ca. 10 ~-tm in width, 1(-2):1, walls thin to
evenly thickened, superficially flat; papillae solid, low, flattened,
granular to multifid, 2-6 per lumen; basal cells differentiated
across leaf, filling the sheathing leaf base and reaching somewhat
higher medially, rectangular, ca. 10 ~-tm in width, 3-5 :1, walls
moderately thickened to porose. Dioicous. Perichaetia terminal,
inner leaves oblong-lanceolate to long-oval, sheathing, often to 6
mm in length, basal cells long rhomboidal in lower half to most of
leaf. Perigonia terminal, gemmate, inner leaves ovate-triangular.
Seta ca. 1-2 em in length, yellow- to red-brown, twisted counterclockwise above, clockwise below; theca (1-)3-6 mm in length,
red-brown, long-cylindrical; exothecial cells rectangular, walls
thin; stomates present at base of theca, phaneropore; annulus of
1-3 rows of strongly vesiculose cells, deciduous in strips; peristome of 32 filamentous, red, densely spiculose teeth, up to 1.2
mm in length, of many articulations, twisted counterclockwise
about 2.5 times, basal membrane low but distinct, spiculose.
Operculum long-conic, ca. 1.3 mm in length, cells in counterclockwise twisted rows. Calyptra cucullate, smooth, ca. 4 mm in
length. Spores ca. 8-11 ~-tm in diameter, light brown to yellowish,
weakly papillose. Lamina/ KOH color reaction red, seldom olive
or yellow.
The genus is restricted to Mexico, Central America and the
northern Andes (Venezuela, Ecuador, Colombia), generally found
at high elevations on a variety of substrates, including soil, rock
and bark.
Mironia is similar to Bryoerythrophyllum in the red coloration
of the plants (especially in reaction to KOH) , the well
differentiated leaf base; the upper leaf cells with characteristically
thin to evenly but rather weakly thickened walls, these often
somewhat sinuose; the low, generally solid (in mature leaves)
and bifid !aminal papillae, 4--Q scattered over each lumen (Pl.
23, f. 10); and the often reniform costal transverse section generally with one or more hydroid strands (Pl. 23, f. 9). It differs
in the leaves commonly keeled, with narrowly bistratose margins, and leaf apices sometimes thickened, fragile or deciduous
(Pl. 23, f. 14). The peristome is very long and twisted; likewise,
some species recently transferred to Bryoerythrophyllum from
Barbula have well-developed peristomes (Zander 1980a).
The comparatively large, lanceolate, occasionally dentate,
keeled, often squarrose leaves with highly differentiated leaf
bases and the much modified perichaetial leaves are also characters of Leptodontium, but Mironia differs in the presence of
an epidermis and hydroid strand(s) in the costa and of a central
strand in the stem (Pl. 23, f. 1). The general leaf shape is similar
to that of Barbula, but this last genus has unistratose leaf margins and, like Leptodontium, a yellow response to KOH solution.
Additional literature: Hilpert (1933), Zander (1978g,
1983d), Zander et al. (1980).
Number of accepted species: 3.
Species examined: M. crassicuspis (BUF, FH, MICH.
TENN), M. ehrenbergiana (BM, BUF, DUKE, FH, NY, PC,
TENN, US), M. stenotheca (BUF, MEXU, TENN).
New heterotypic synonymy: Didymodon killipii Williams =
Mironia ehrenbergiana (C. Mtill.) Zand.
New combinations: Mironia crassicuspis (Robins.) Zand.,
comb. nov. (Barbula crassicuspis Robins., Bryologist 67: 446,
1964; Morinia crassicuspis (Robins .) Zand.). Mironia
ehrenbergiana (C . Mtill.) Zand., comb. nov. (Barbula
ehrenbergiana C. Mtill., Synop. Muse. 1: 636, 1849; Morinia
ehrenbergiana (C. Mtill.) Ther.). Mironia ehrenbergiana var.
elongata (Wils. in Mitt.) Zand., comb. nov. (Barbula elongata
Wils. in Mitt., Kew J. Bot. 3: 51, 1851; Morinia ehrenbergiana
var. elongata (Wils. in Mitt.) Zand.). Mironia stenotheca
(TMr.) Zand., comb. nov. (Barbula stenotheca Ther., Smiths.
Misc. Coll. 85(4): 21, 1931; Morinia stenotheca (Ther.) Zand.).
20. BRYOERYTHROPHYLLUM
Plates 24-25.
Bryoerythrophyllum Chen, Hedwigia 80: 4, 1941. Type: Bryoerythrophyllum recurvirostrum (Hedw.) Chen.
Globulina C. Mtill., Nuov. Giorn. Bot. !tal., N.S. 4: 39, 1897,
hom. illeg. non Link in Nees 1820, nee Turpin 1827, nee
Velen. 1834, nee Spegaz. in Sacc. 1891. Type: Globulina
boliviano C. Mtill.
Erythrophyllum (Lindb. in Braithw.) Loeske, Hedwigia 47:
175, 1908, hom. illeg. non J. Ag., 1872.
Erythrobarbula Steere, Bryologist 54: 199, 1951 , nom illeg.
incl. gen . prior.
Didymodon subg. Erythrophyllum Lirnpr., Laubm. Deutschl.
1:405, 1887. Type: Didymodon rubellus BSG.
Barbula subg. Erythrophyllum (Lindb. in Braithw.) C. Jens.,
Skand. Bladmfl. 243, 1939.
Barbula subg. Erythrobarbula Szafr., A. Polska Mchy 1: 213,
1957 [1958], nom. inval. descr. polon.
Barbula sect. Erythrophyllum Lindb. in Braithw., Brit. Moss
A. 1: 260, 1887. Type: Barbula rubella Mitt. in Lindb.
Didymodon sect. Orthocarpae Broth., Nat. Pfl. 1(3): 405,
1902.
MERCEYOIDEAE
Didymodon sect. Amblystegioideae Broth., Nat. Pfl. 1(3): 405,
1902.
From ~puov, a moss + o + £pu6p6~. red + o + <puA.Aov, leaf;
referring to the red cast of older leaves.
Plants turf-forming or loosely caespitose, usually green above
113
and red-brown below. Stems occasionally branching, short or to
several em in length, rounded-pentagonal in transverse section,
usually with central strand, sclerodermis usually present, hyalodermis usually absent; axillary hairs several cells in length,
sometimes with 1-3 brownish basal cells. Leaves appressed
when dry, spreading when wet, ovate, elliptical or lanceolate,
Plate 14. Bryoerythrophyllum. 1-10. B. inaequa/ifolium. 1. Habit, perichaetiate plant. 2. Habit, perigoniate plant. 3-4. Two leaves. 5. Leaf
apex. 6. Basal cells. 7. Transverse section at midleaf. 8. Propagula. 9. Perichaetialleaf. 10. Peristome. 11-15. B. bolivianum. 11. Habit. 12-13.
Two leaves. 14. Leaf apex. 15. Perichaetialleaf. 16-18. B.ferruginascens. 16-17. Leaf. 18. Leaf apex.
114
GENERA OF THE POTTIACEAE
1-5 mm in length, often grooved along the costa, marins usually
recurved below or to near apex, seldom plane, entire or more
often dentate above, often with a border of 3-4 rows of thickwalled cells; apex rounded-obtuse to acute; base usually ovate to
oblong and sheathing; costa ending a few cells below the apex to
short-excurrent, with quadrate, papillose cells ventrally, cells
elongate dorsally, in transverse section showing a differentiated
epidermis ventrally or on both sides, ca. 2-6 rows of cells
across costa ventrally at midleaf, two stereid bands present,
guide cells 2-4 in 1(-2) layers, one or more hydroid
Plate 25. Bryoerythrophyllum. 1-8. B. binsii. 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7.
Perichaetialleaf. 8. Peristome. 9-10. B. rubrum. 9. Leaf. 10. Leaf apex. 11-13. B. wallichii. 11-12. Two leaves. 13. Leaf apex.
MERCEYOIDEAE
strands occasionally present; upper laminal cells subquadrate to
short-rectangular, ca. 10-15 Jlm in width, 1(-2):1, walls evenly
thickened, weakly sinuose, superficially nearly flat, occasionally
bulging; papillae solid, or occasionally hollow, bifid, or occasionally flattened or compound, mostly 4-6 per lumen, obscuring the
lumens; basal cells usually hyaline and rectangular, filling most
of the base medially, little wider than upper cells, occasionally
bulging, 2-5: l. Asexual reproduction occasional, by unicellular
propagula in masses in the leaf axi/s or obovoid brood bodies
borne on rhizoids in the soil. Dioicous or occasionally monoicous.
Perichaetia and perigonia terminal. Perichaetial leaves usually
sheathing in the lower half, larger than the stem leaves, long-oval
to long-lanceolate, lower cells long rhomboidal in lower half.
Perigoniate plants occasionally small, bud-like. Seta elongate,
twisted clockwise below, often counterclockwise above, usually 1
per perichaetium; theca ellipsoidal to cylindric, usually redbrown, 0.7-2.5(-4.0) mm in length, sometimes curved; stomates
phaneropore, at base op theca, annulus of 1-2 rows of vesicu/ose
cells, often revoluble or deciduous in pieces; operculum shortconic to short-rostrate, cells in straight or oblique rows; peristome
none, rudimentary, or consisting of 16 or 32 linear rami, erect to
twisted to 2-4 times counterclockwise, usually densely spiculose,
basal membrane low or absent. Operculum short-conic to shortrostrate, 0.2-1.2 in length. Calyptra cucullate, smooth, ca. 2 mm
in length. Spores ca. 8-15 J.1m in diameter, light brown, lightly
papillose. Lamina/ KOH color reaction red to orange-red.
Reported chromosome number n = 13, l3+m, 14, 26+2m.
An essentially cosmopolitan genus. More extensive treatments
of this taxon for the New World have been done by Zander
(1978g, 1980a, 1981a).
Bryoerythrophyllum has much the appearance of Barbula or
Didymodon species, but is easily distinguished by a combination
of features, most of which are usually present: red coloration (at
least in older parts of stems-Bryoerythrophyllum fuscinervium,
for instance, generally having immature leaves at the stem apex
with a yellow KOH reaction), the red color usually evident even
without KOH; bifid, crowded laminal papillae obscuring the
upper laminal cell lumens; usually flat superficial laminal cell
walls (easily seen in section, Pl. 24, f. 7; 25, f. 6); and welldifferentiated leaf basal cells (Pl. 24, f. 6; 25, f. 5). Like Barbula
and Didymodon, Bryoerythrophyllum exhibits a full range of peristome development, some species eperistomate (e.g. B. boliviano,
Pl. 24, f. 11), some with rudimentary peristomes (e.g. B.
rotundifolia), others with straight, well-developed teeth (e.g. B.
binsii, Pl. 25, f. 8, and B. recurvirostrum), and finally those with
long, twisted peristomes (e.g. B. inaequa/ifolium, Pl. 24, f. 10).
Only recently has it been recognized (Saito 1975a; Zander 1978g,
1980a) that this genus is rather large and includes many species
previously placed in Barbula and Didymodon. Bryoerythrophyllum fuscinervium and the similar B. co/umbianum have a rather
distinctive broad and bulging ventral costal epidermis of a single
layer of cells and may be confused with Pseudocrossidium species (such as P. chi/ense) but the red KOH reaction and distinct
ventral stereid band are diagnostic. Bryoerythrophyllum differs
from Syntrichia, which is also KOH red, in having two stereid
bands, a usually narrower leaf shape and dorsal costal epidermis
commonly differentiated. The upper areolation of rather large
cells with evenly thickened walls is similar to that of Trichostomum subg. Oxystegus but the red color in KOH, presence of a
stem central strand and costal hydroid strand(s), and the basal
115
cells extending farthest up the leaf medially distinguish Bryoerythrophyllum (e.g. cf B . chimborazense). The genus Mironia
is very similar, but differs in the keeled leaves with bistratose
upper margins (the upper lamina of Bryoerythrophyllum, in one
species, B. sharpii, may be bistratose throughout) and, in two
species of Mironia, the presence of propaguloid leaf apices.
The genus Globulina was published validly by C. MUller
(1897a) in a combined genus and species description. Although
Miiller indicated that Seligeria globifera Hampe (= Globulinella
globifera (Hampe) Steere) was related to and was probably also
a member of the genus ("Globulina mihi"), he left the genus
monotypic with G. boliviana. Later, Miiller (1901) established
Seligeria subg. G/obulina C. Miill., with S. globifera the only
species, being thus the type of that, different, taxon. Steere
(Steere & Chapman 1946) proposed Globulinella as a new
genus, with Globulina C. Miill. non Link in Nees as a synonym,
but excluded the type of G/obulina, G. boliviano, later in the
publication. Steere apparently did not intend Globulinella as a
nom. nov. for Globulina as was suggested by Magill (1977a).
This confusion is reflected in the Index Nominum Genericorum
(Farr et al. 1979). Actually, Globulina C. Mtill., hom. illeg., is
not a synonym of Globulinella Steere, but is instead a synonym
of Bryoerythrophyllum Chen. Seligeria subg. Globulina C.
Mtill. is a synonym of the valid genus Globulinella Steere, both
taxa having the same type species, G. globifera (lectotype fide
van der Wijk et al., 1959-69).
The type of Husnotiel/a glossophylla Herz. is not at L, JE or
Wand may have been destroyed during the Second World War
(pers. comm. F. K. Meyer at JE). Specimens collected by
Hosseus in Argentina (FH, W) identified by Bartram as this or
as H. baueri are eperistomate collections of the rather variable
Didymodon tophaceus. The description and illustrations of H.
glossophylla itself indicate that it is probably the same as Bryoerythrophyllum calcareum.
Additional literature: Chen (1941), Crum (1957a), Long
(1982a, 1982b), Ratkowsky (1980).
Number of accepted species: 27.
Species examined: B. andersonianum (BUF, TENN), B.
binnsii (BUF), B. bolivianum (BUF, JE, TENN), B. byrdii
(Bartr.) Zand. (NY), B. ca/careum (BUF, FH, MICH, TENN,
US), B. campylocarpum, B . chimborazense (NY), B.
columbianum (UBC, US), B. ferruginascens, B. fuscinervium
(BUF, NY), B. hostile (JE), B. inaequalifolium (BUF, DUKE,
NY, TENN), B. jamesonii, B. ligulare (NY), B. recurvirostrum,
B. rotundatum (H), B. rubrum (BUF, NY), B. sharpii (BUF), B.
wallichii (NY, US).
New heterotypic synonymy: Bryoerythrophyllum columbianum var. atacamense Zand. & Lewis in Lewis = Bryoerythrophyllum fuscinervium (Mitt.) Zand. Didymodon integrifolius
Broth. in Mildbr. var. paucidentatus Ther. = Bryoerythrophyllum campylocarpum (C. Mtill.) Crum. Didymodon luzonensis
Bartr. = Bryoerythrophyllum ferruginascens (Stirt.) Giac. (the
type of the former at FH is of fruiting material and includes the
characteristic propagula). Didymodon merceyoides Broth. in
Herz. = Bryoerythrophyllum campylocarpum (C. Mtill.) Crum.
Didymodon pelichucensis Williams = Bryoerythrophyllum
campylocarpum (C. Mtill.) Crum. Hyophila calymperoides
Ther. & Nav. = Bryoerythrophyllum campylocarpum (C. Mtill.)
Crum.
New combinations: Bryoerythrophyllum byrdii (Bartr.)
116
GENERA OF THE POTTIACEAE
Zand., comb. nov. (Barbula byrdii Bartr., Ann. Missouri Bot.
Gard. 25: 720, 1938). Bryoerythrophyllum chimborazense (Mitt.)
Zand., comb. nov. (Tortula chimborazensis Mitt., J. Linn. Soc.
Bot. 12: 163, 1869; Didymodon chimborazensis (Mitt.) Broth. in
Par.). Bryoerythrophyllumfuscinervium (Mitt.) Zand., comb. nov.
(Tortulafuscinervia Mitt., J. Linn. Soc. Bot. 12: 154, 1869; Barbuta fuscinervia (Mitt.) Jaeg. Bryoerythrophyllum ligulare (Mitt.)
Zand., comb. nov. (Tortula ligularis Mitt., J. Linn. Soc. Bot. 12:
156, 1869; Barbula ligularis (Mitt.) Jaeg.).
21. PSEUDOCROSSIDIUM
Plate 26-27.
Pseudocrossidium Williams, Bull. Torrey Bot. Club 42: 396,
1915. Type: Pseudocrossidium chilense Williams.
Barbula sect. Revolutae BSG, Bryol. Eur. 2: 89, 1842 (Fasc.
13-15 Mon. 27). Type: Barbula revoluta Brid. in Schrad.
Tortula sect. Revolutae (BSG) Spruce, Ann. Mag. Nat. Hist. ser.
2, 3: 377, 1849.
Barbula sect. Platyneuron Kindb., Eur. N. Amer. Bryin. 2: 246,
1897. Type: Barbula platyneura C. Miill. & Kindb.
Barbula sect. Pseudocrossidium (Williams) Nyholm, Ill. Fl.
Nordic Mo. 2: 102, 1989.
Barbula subsect. Revolutae (BSG) Chen, Hedwigia 80: 209,
1941.
From 'ljfEUOT{~. false + o + Crossidium, a genus; a false Crossidium.
Plants growing in cushions or turf, yellowish green to brown
above, brown to reddish brown below. Stems branching often, ca.
0.3-2.0 em in length, transverse section rounded-pentagonal, central strand usually present, often strong, sclerodermis weakly
differentiated, hyalodermis absent or weakly differentiated;
axillary hairs of 5-8 cells, all hyaline or occasionally basal 1-3
cells thicker walled; weakly radiculose. Leaves appressed and
often spiralled when dry, weakly or widely spreading when moist,
ovate or ligulate to lanceolate, 0.5-3.0 mm in length, upper lamina channeled or grooved along costa, margins recurved to
broadly revolute or spiralled, entire or occasionally weakly
denticulate near apex, the rolled margins occasionally
differentiated as cylindrical photosynthetic organs of thin-walled,
hollow-papillose cells; apex acute to rounded; base scarcely
differentiated in shape to oblong; costa often broad and flat, often
swollen medially, excurrent as a mucro or short, smooth awn,
occasionally long-awned, superficial cells quadrate to shortrectangular, papillose ventrally, often differentiated as a pad of
papillose, thin-walled photosynthetic filaments, elongate, smooth
or papillose to rough or weakly toothed dorsally, 2-5 rows of
cells across costa ventrally at midleaf, costal transverse section
reniform to circular, stereid bands present or absent ventrally,
present and usually strong and flattened crescent-shaped (occasionally semicircular) dorsally, ventral and dorsal epidermis
present, the latter often weak, guide cells 2-4(-9) in 1-2 layers,
hydroid strand present, often multiple; upper /aminal cells
subquadrate to hexagonal, often transversely elongated,
8-16(-18) j!m in width, 1:1(-3), walls evenly thickened, occasionally weakly trigonous, superficially weakly convex to bulging
on both sides; papillae rarely absent, crowded, usually hollow,
occasionally plate-like or bifid to multiplex, usually crowded,
occasionally capitulate and solid; basal cells differentiated medially, occasionally across leaf, rectangular, 11-13(-23) j!m in
width, 2--6: 1, walls thin to evenly thickened, occasionally
porose, hyaline or occasionally orange. Propagula occasionally
present, borne on ventral surface of costa or in leaf axils, clavate
or spherical, 40-50 j!m in length. Dioicous. Perichaetia terminal, inner leaves little different from the cauline leaves or more
commonly highly differentiated, enlarged, often awned, often
convolute-sheathing, lower cells not differentiated or rectangular to rhomboidal throughout. Perigonia gemmate. Seta 1.0-1.7
in length, 1 per perichaetium, yellowish to reddish brown,
twisted clockwise; theca 1.6-3.0(-3.6) in length, yellowish to
reddish brown, elliptical to cylindrical, occasionally curved,
exothecial cells short-rectangular, 16-20 J.lm in width, 2-3:1,
thin-walled to evenly thickened, stomates phaneropore, at base
of theca, annulus of 2-4 rows of vesiculose cells, persistent;
peristome teeth 16, cleft to base or 32, linear, densely spiculose,
350-1000 j!m in length, with many articulations, twisted to
once twisted counterclockwise, occasionally straight, basal
membrane low or absent, weakly spiculose. Operculum shortto long-conic or conic-rostrate, 0.6-2.1 mm in length, cells
counterclockwise. Calyptra cucullate, smooth, 3.2-3.5 mm in
length. Spores 8-15 j!m in diameter, yellow to light brown,
essentially smooth to weakly papillose. Lamina! KOH color
reaction yellow to orange, occasionally with red blotches.
Reported chromosome number n = 13.
A large genus largely growing on soil and rock at high elevations; present in mountainous regions of the New World,
Europe, the Middle East, Africa, and Australasia.
Important characters for this genus are the usual small size
or absence of the ventral stereid band and the broadly crescentshaped dorsal stereid band with a clearly differentiated dorsal
epidermis of cells often with semicircular lumens through differential thickening of the walls (Pl. 26, f. 7, 21; 27, f. 8, 13,
17). Other important characters are not consistent in appearance
through the genus, but are often striking when found: leaves
often ending in a short or long awn (Pl. 26, f. 5); differentiation
of photosynthetic tissue either as a ventral pad of costal filaments (Pl. 26, f. 20, 21) or within rolled margins (Pl. 27, f. 17)
or both (Pl. 27, f. 13); medial cells more papillose and thicker
through (the distance between the two superficial walls) than
the marginal cells (Pl. 26, f. 7; 27, f. 8); and perichaetialleaves
abruptly enlarged and sheathing the seta (Pl. 26, f. 8).
A few stereid cells of a second stereid band may be found in
occasional specimens of what has been called P. aureum in
Mexico and southwestern U.S.A. This is synonymized (independently by Sollman 1990) with P. crinitum, which usually has
two stereid bands. The North American material is quite like the
type of the South American Barbula arenicola, specimens of
which have a single stereid band but which is also here placed
in the synonymy of P. crinitum. Pseudocrossidium crinitum (Pl.
26, f. 1-8) is an essentially Gondwanaland taxon, being found
in southern parts of Africa, South America and Australasia,
which may indicate a southern origin for the genus. The fact
that P. crinitum also occurs in Mexico and southwestern U.S.A.,
albeit as a sterile, depauperate (shorter awned and smaller stature) population, shows that there can be considerable northward
extension of the ranges of species of ultimately southern derivation. A simple explanation of the evolutionary history of the
genus, following the thread of a previous discussion (Zander
1979f) and discounting long-distance dispersal as a factor,
would be that ancestors essentially identical to modern P.
MERCEYOIDEAE
117
Plate 26. Pseudocrossidium. 1-8. P. crinilum. I. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7.
Transverse section at midleaf. 8. Perichaetialleaf. 9-12. P. apiculatum. 9-11. Three leaves. 12. Leaf apex. 13-17. P. chilense. 13-16. Four
leaves. 17. Leaf apex. 18-22. P. elatum. 18-19. Two leaves. 20. Leaf apex. 21. Transverse section at midleaf. 22. Perichaetialleaf.
118
GENERA OF THE POTTIACEAE
crinitum spread across Gondwanaland to be later isolated in austral areas through tectonic plate separation. In migrating northward along the Andes, ancestors of P . crinitum developed into the
several species now there, these characterized by loss of the
ventral stereid band and elaboration of photosynthetic tissue along
the leaf margins or ventral costal surface. In North America,
descendants of the derived species P. replicatum lost many of the
characters of Pseudocrossidium while developing into the essentially high-northern latitude taxa P . revolutum and P .
hornschuchianum (Pl. 27, f. 1-5), which may be referred to the
genus by the general lack of a ventral stereid band, the highly
revolute leaf margins with walls somewhat thinner and more
papillose than the medial portion of the leaf, and the presence of
propagula on the ventral surface of the costa (possibly a vestige of
the ventral costal elaboration). Thus, P. cnmtum and P .
revolutum, though once both regarded as Barbula species, are
actually at opposite ends of a complex north-south evolutionary
series through mid-Andean taxa of Pseudocrossidium. The
northernmost taxa subsequently became established in southern
Africa and Australia probably through human agency. Evidence
for anthropogenic disjunction is the discovery of P.
hornschuchianum in North America in Massachusetts in the
U.S.A. (Mishler & Miller 1983) and in British Columbia, Canada (Tan et al. 1981), associated with parks or gardens; Arts
(1988) has reported rhizoidal tubers in the closely related P.
revolutum. In the southern hemisphere, P. crinitum can be separated into two morphotypes, which may rate recognition as taxa
(if so, types of synonyms, cf Catcheside 1980, Magill 1981,
Plate 27. Pseudocrossidium. l-5. P. hornschuchianum. 1-3. Three leaves. 4. Leaf apex. 5. Perichaetialleaf. 6-8. P. leucocalyx. 6. Leaf. 7.
Leaf apex, lateral view. 8. Transverse section at midleaf. 9-13. P. pachygastrellum. 9-11. Three leaves. 12. Leaf apex. 13. Transverse section at
midleaf. 14-19. P. replicatum. 14-15. Two leaves. 16. Leaf apex, lateral view. 17. Transverse section at midleaf. 18. Capsule. 19. Operculum.
MERCEYOIDEAE
Weber 1972, should be consulted for the earliest name, and this
would require a careful revision of the complex): one has a
recurved leaf margin, coarse upper !aminal papillae, dorsal costal
epidermis absent or weakly developed, and ventral stereid band
strong, while the second has revolute upper !aminal margins, delicate upper lamina! papillae, dorsal costal epidermis well developed, and ventral stereid band absent or weakly developed.
Awned species of Syntrichia may have much the same appearance
as P. crinitum (obtusely and broadly short-lanceolate leaves with
densely papillose upper laminal cells) but differ in their distinctive red KOH color reaction, P. crinitum reacting deeply yellow.
Pseudocrossidium leucocalyx (Pl. 27, f. 6-8) is unusual for
the genus in its costa ending in a distinctive conical, smooth cell
and rough dorsally with both sharp prorulae and solid simple
papillae; the upper lamina! cells medially strongly bulging on
both sides, with high, solid, capitulate and spiculose papillae, but
marginal cells smaller in several rows, weakly bulging and
smooth; and lamina bright yellow in KOH. It may belong elsewhere, possibly in a monotypic genus of its own. It is superficially similar to Hypodontium species, which differ significantly,
however, in their strongly incurved upper laminal margins and
two strong stereid bands in the costa. Pseudocrossidium
porphyreoneurum is a South African species having a longmucronate costa and strongly revolute leaf margins, but is unusual
in its semicircular (not crescent-shaped) stereid band and basal
lamina! cells not differentiated from the upper cells; it is placed
here only tentatively (cf Magilll981, p. 213). The genus requires
a thorough revision for adequate evaluation.
Although P. e/atum (Pl. 26, f. 18-22) is similar to Crossidium
in its smooth upper larninal cells and rather rounded section of the
costa, it is recognized here in Pseudocrossidium (following
Delgadillo 1975a) because of its elongate stem, lanceolate
leaves,a few stereid cells occasionally differentiated immediately
below the ventral costal filaments, and the well-differentiated dorsal costal epidermis. The species remains, however, uncomfortably intermediate in morphology between Pseudocrossidium and
Crossidium (especially C. spiralifolium of South Africa), while
differing from both by the quite elongate patch of ventral costal
filaments and poorly differentiated basal laminal cells. Eventual
recognition of this species in a monotypic genus may be the best
reflection of relational distances.
Additional literature: Churchill (1990), Delgadillo and Zander
119
(1984), Frey and Kiirschner (1988c).
Number of accepted species: 16.
Species examined: P. apiculatum (BUF, NY, US), P.
austrorevolutum (NY), P. carinatum (NY), P. chilense (BUF,
US), P. crinitum (NY, SPA), P. e/atum (F, NY), P. excavatum
(NY), P. hornschuchianum, P. leucocalyx (CU, FH, NY, US),
P. mendozense (NY), P. pachygastrellum (L), P. perrevolutum
(NY), P. porphyreoneurum (NY), P. replicatum, P. revolutum,
P. steerei (BUF).
New heterotypic synonymy: Barbula arenicola Dus. =
Pseudocrossidium crinitum (Schultz) Zand.
New homotypic synonymy: Pseudocrossidium obtusulum
(Lindh.) Crum & Anderson = Pseudocrossidium revolutum var.
obtusulum (Lindh.) Tan, Zand. & T. Tayl.
New combinations:
Pseudocrossidium austrorevolutum (Besch.) Zand., comb. nov.
(Barbula austrorevoluta Besch. in Britt., Bull. Torr. Bot. Cl.
23: 480, 1896).
Pseudocrossidium carinatum (Gill. ex Grev.) Zand., comb. nov.
(Tortuta carinata Gill. ex Grev., Edinburgh J. Nat. Geogr.
Sc. 2:2, 1830), near P. crinitum.
Pseudocrossidium crinitum (Schultz) Zand., comb. nov. (Barbuta crinita Schultz, Nov. Act. Ac. Leop. Car. 11(1): 226,
1823).
Pseudocrossidium mendozense (Mitt.) Zand., comb. nov. (Tortuta mendozensis Mitt., J. Linn. Soc. Bot. 12: 154, 1869;
Barbuto mendozensis (Mitt.) Jaeg.).
Pseudocrossidium perrevotutum (C. Miill.) Zand., comb. nov.
(Barbula perrevotuta C. Miill., Linnaea 43: 486, 1882).
Pseudocrossidium perrevotutum var. acutifotium (C. Miill.)
Zand., comb. nov. (Barbuto subrevotuta var. acutifotia C.
Miill., Linnaea 42: 335, 1879; Barbuto perrevoluta var.
acutifotia (C. Miill.) Par.), not seen.
Pseudocrossidium perrevotutum var. linearifolium (C. Miill.)
Zand., comb. nov. (Barbuto subrevotuta var. linearifolia C.
Miill., Linnaea 42: 335, 1879; Barbuto perrevoluta var.
linearifolia (C. Miill.) Par.), the isotype at NY seems to be
different at the species level and needs evaluation.
Pseudocrossidium porphyreoneurum (C. Miill. ex Vent.) Zand.,
comb. nov. (Barbuto porphyreoneura C. Miill. ex Vent. in
Nuovo Giorn. Bot. !tal. 4: 13, 1872; Tortula
porphyreoneura (C. Miill.) Townsend).
Tribe LEPTODONTIEAE
Leptodontieae Herz., Geogr. d. Moose 101, 1926.
Leptodontioideae (Herz.) Hilp., Beih. Bot. Centralbl. 50: 679, 1933.
This subclade of the Merceyoideae, including Leptodontium and other genera once recognized as Leptodontioideae, is distinguished
from the Tetracoscinodontieae by the following advanced character states of the node common to the Leptodontieae and Barbuleae:
!aminal papillae simple; seta nearly absent to short, less than I em in length. The traits of the immediate ancestral node of the
Leptodontieae are: stem central strand absent, ventral costal epidermis absent, and upper !aminal cells superficially flat or weakly
convex. The tribe may be further characterized generally by upper !aminal margins commonly denticulate or toothed, costa flattened
in section, and dorsal costal epidermis often absent. Analysis shows Hymenostylium, previously treated in the Pleuroweisieae
(Zander 1977c), to be a member of this group. Although the geographic distribution of the tribe is nearly worldwide, certain genera
are restricted to the Andes.
Plates 28-29.
22. TRIQUETRELLA
Triquetrella C. Mtill., Ost. Bot. Zeit. 47: 421, 1897. Lectotype:
Triquetrella tristicha (C. Mull.) C. Mull. fide Grout, Moss Fl.
N. Amer. 1: 170, 1938.
From triquetrus, three-edged, three-angled + -ella, diminutive;
referring to the leaves commonly arranged in three rows.
Plants growing in mats, yellowish or blackish green above,
brown or blackish brown below. Stems branching occasionally, to
7 em in length, transverse section rounded-triangular, central
strand absent or present and small, sclerodermis present, of 1-3
layers of stereid cells, hyalodermis absent; axillary hairs to 9 cells
in length with cells weakly bulging, basal 1-2 cells thick-walled;
weakly radiculose below. Leaves in 3 distinct rows, the rows
straight or weakly spiralling in either direction, appressed- to
spreading-incurved when dry, widely spreading to squarrose and
strongly reflexed when moist, triangular to ovate-triangular, ca. 2
mm in length, upper lamina keeled, narrowly channeled along
costa, margins recurved in middle 3/4-4/5 of leaf, entire to
weakly serrulate near apex; apex narrowly acute to shortacuminate; base ovate, basal leaf margins broadly longdecurrent; costa percurrent, superficial cells elongate and weakly
papillose ventrally, rhombic-quadrate near apex and papillose
dorsally, 2(-4) rows of cells across costa ventrally at midleaf,
costal transverse section ovate to semicircular, 2 stereid bands
present, ventral epidermis absent, the dorsal weakly
differentiated, guide cells 2-4 in I layer, hydroid strand absent;
upper /aminal cells rounded rhombic to quadrate, ca. 9-11 J.lm in
width, 1: 1, occasionally elongate transversely, walls thickened,
weakly trigonous, superficially weakly convex to distinctly bulging on both sides; papillae spiculose, mostly simple to bifid, 1 per
lumen; basal cells differentiated weakly in a very small group
medially, occasionally across insertion, rectangular, little wider
than upper cells, 2-4:1, walls thick-walled, somewhat porose.
Dioicous. Perichaetia both terminal and lateral in the same species and occasionally on the same plant (acrocarpous and
pleurocarpous), often in clusters, inner leaves ovate, rounded or
short-acuminate to apiculate, to 2.5 mm in length, strongly
sheathing seta, lower cells rhombic, thick-walled throughout.
Perigonia terminal and lateral in clusters. Seta ca. 1.5-2.3 em in
length, 1 per perichaetium, yellow, twisted clockwise; theca
1.6-2.1 mm in length, yellowish brown, elliptical, exothecial cells
short-rectangular, 20-25 J.lm in width, thin-walled, stomates
phaneropore, at base of theca, annulus of ca. 3 rows of smaller,
vesiculose cells; peristome teeth 16, variously cleft (usually to
base) or occasionally perforate, subulate, transparent, smooth
or very weakly spirally striate, 110-180 J.lm in length, with ca. 6
articulations, straight, basal membrane absent. Operculum
conic, ca. 0.7-0.8 mm in length, cells straight. Calyptra
cucullate, smooth, ca. 2.6-2.8 mm in length. Spores ca. 10-13
J.lm in diameter, light brown, papillose. Lamina! KOH color
reaction orange to yellowish orange.
A small genus found on soil or rock generally in or near dry,
Mediterranean climates; Europe (Spain), South Africa, western
North America (California), southern South America, Australia,
Tasmania and New Zealand.
Triquetrella papillata (Pl. 28, f. l-11, NY!) has perichaetia
that in some plants are terminal and in others are distinctly lateral on the stem, often clustered below a terminal perichaetium
(cf. Ganguleea). Perigonia, too, are clustered both terminally
and laterally near the stem apex on separate, entirely perigoniate
plants. The "acropleurocarpous" condition may be an anomaly,
but whether this also occurs elsewhere in the genus cannot be
immediately determined since perichaetia-bearing herbarium
specimens are uncommon. Catcheside ( 1980) indicated that
sporophytes are rare in South Australia, and Brotherus
(1924-25) found the genus mostly sterile.
Triquetrella is similar to Leptodontium in the 16 smooth or
nearly smooth peristome teeth, mostly cleft to the base (Pl. 28,
f. 11; 29, f. 7); strongly differentiated perichaetialleaves (Pl. 28,
f. 10); stem usually without a central strand (Pl. 28, f. 3); squarrose, strongly reflexed cauline leaves; and absence of a
differentiated epidermal layer on the ventral surface of the costa
(Pl. 28, f. 9). Triquetrella differs from Leptodontium by the
occasional presence of a stem central strand (in T. californica,
Pl. 28, f. 12-14, which also lacks the otherwise characteristic
triangular section of the stem); the short, triangular cauline
leaves forming three distinct rows (Pl. 28, f. l-2); and sharp,
spiculose upper !aminal papillae (blunt and columnar in sect.
Leptodontium). An intermediate taxon is Leptodontium
paradoxicum, which shares with Triquetrella two characters
rare in Leptodontium: entire leaves and dorsal costal epidermis
differentiated. On the other hand, Eddy (1990) said about Triquetrella that "the sum of its gametophytic and sporophytic
characters strongly suggests that the genus is misplaced in the
Pottiaceae," without further explanation.
There are differences between the species of Triquetrella in
length of the theca, leaf shape, acumination of the leaf apex,
length of the decurrencies, and in presence or absence of the
stem central strand, that might be used to distinguish taxa, but,
MERCEYOIDEAE
121
Plate 28. Triquetrella. 1-11. T. papillata. I. Habit. 2. Stem with clustered terminal and lateral perichaetia. 3. Transverse section of stem. 4-6.
Three leaves. 7. Leaf apex. 8. Leaf base and decurrency. 9. Transverse section at midleaf. 10. Perichaetial leaf. 11. Peristome. 12-14. T.
californica. 12. Transverse section of stem. 13. Leaf. 14. Leaf apex.
122
GENERA OF THE POTTIACEAE
Plate 29. Triquetrella. 1-1. T. filicaulis. 1. Transverse section of stem (with attached leaf decurrencies). 2. Leaf. 3. Leaf apex. 4. Leaf base and
decurrency. 5. Transverse section at midleaf. 6. Perichaetialleaf. 7. Peristome. 8-12. T. tristicha. 8-11. Four leaves. 12. Leaf apex.
MERCEYOIDEAE
in general, the species are much alike. Casas et al. (1993) summarized the differences between most of the species in their study of
T. arapilensis. Triquetrella preissiana of Australia has been
synonymized with T. papillata by Watts and Whitelegge (1902)
according to Stone and Scott (1976); the type of the former name
(BM!) is indeed taxonomically T. papillata.
Additional literature: Cour (1955), Moore et al. (1982),
Miiller ( 1897b), Stark ( 1980), Zander ( 1980a).
Number of accepted species: 9.
Species examined: T. arapilensis (FH), T. californica (FH,
NY), T. ftlicaulis (NY), T. papillata (BM, DUKE, FH, NY), T.
patagonica (FH), T. tristicha (NY).
23. REIMERSIA
Plate 30.
Reimersia Chen, Hedwigia 80: 62, 1941. Type: Reimersia
inconspicua (Griff.) Chen.
Named for Hermann J. 0. Reimers, 1893-1961, German bryo1ogist and professorial advisor of P.-c. Chen.
Plants in cushions, light green above, light brown below.
Stems branching irregularly, to 6 em in length, transverse section
distinctly triangular, central strand absent, cells of central cylinder thick-walled, sclerodermis present, hyalodermis absent;
axillary hairs of ca. 10 cells, all clear or basal 1-3 cells yellow;
weakly radiculose. Leaves trifarious, widely spreading when dry,
sharply squarrose-recurved when moist, lanceolate, ca. 3.0 mm in
length, upper lamina strongly keeled above, margins plane,
entire; apex narrowly acute to acuminate; base broadly ovate,
sheathing the stem, strongly convex in 2 small circular areas on
both sides of costa at top of leaf base, strongly decurrent at margins; costa slender, excurrent as a sharp mucro, superficial cells
elongate ventrally and dorsally, 2-4 rows of cells across costa
ventrally at midleaf, costal transverse section semicircular, 2
stereid bands present (the ventral weak), epidermis... absent ventrally, present dorsally, guide cells 2-4 in 1 layer, hydroid strand
absent; upper /aminal cells quadrate to short-rectangular or
rhomboidal, 10-13 jlm in width, 1-3:1, thick-walled, trigonous or
porose, superficially flat, marginal cells narrower than medial;
papillae absent; basal cells differentiated across the leaf, rectangular, scarcely wider than upper cells, to 4-5:1, walls thickwalled, porose. Dioicous. Perichaetia terminal, inner leaves similar to the cauline and sheathing the seta, !aminal cells long
rhomboidal to near apex. Perigonia terminal, gemmate. Seta
0.5-0.6 em in length, 1 per perichaetium, yellowish brown,
twisted clockwise; theca ca. 1.0 mm in length, yellowish brown,
ovoid, macrostomous, neck differentiated, exothecial cell walls
thick, stomates present on neck, annulus weakly differentiated, of
1-2 rows of weakly vesiculose cells; peristome absent. Operculum rostrate, erect or oblique, ca. 1.0-1.2 mm in length, cells in
straight rows. Calyptra cucullate, smooth, ca. 2.0 mm in length.
Spores ca. 10 jlm in diameter, brown, essentially smooth. Lamina/
KOH color reaction lemon yellow. Reported chromosome number
n = 13+"f'.
Found on soil and calcareous rock in wet areas at moderately
high elevations (2000 to 3000 m) in India, Nepal, China and the
Philippines.
Reimersia is clearly like Hymenostylium in its hygric habitat;
lack of a stem central strand (Pl. 30, f. 2) or of a ventral costal
epidermis (Pl. 30, f. 7); trigonous or porose and generally rectan-
123
gular upper !aminal cells, which are larger medially (Pl. 30, f.
5); and eperistomate capsule with rostrate operculum. It resembles most closely Hymenostylium recurvirostrum var. insigne
(Zander & Eckel 1982). The genus is easily recognized by the
combination of the trifarious, epapillose leaves, which are
sharply squarrose from a sheathing base; the triangular stem
section; the elongate cells on the ventral surface of the costa;
and the strongly porose areolation. The two concavities at the
upper part of the sheathing base are distinctive, but not immediately evident on superficial examination. The capsule is not
systylious, but this is not a character found in all taxa of
Hymenostylium and is also not constant in the widespread H.
recurvirostrum.
Chen (1941) provided a through review of the history (combinations in Gymnostomum, Hymenostylium, Zygodon, Triquetrella, Pottia and Didymodon) and morphological characters of
the single species of this rare genus.
Number of accepted species: 1.
Species examined: R. inconspicua (BM, F, FH, NY, US).
24. HYMENOSTYLIUM
Plates 31-32.
Hymenostylium Brid., Bryol. Univ. 2: 81, 1827. Type: Hymenostylium xanthocarpum (Hook.) Brid.
Gymnoweissia Mont. in Orbigny, Diet. Univ. Hist. Nat. 7:
402, 1849.
Hymenostelium Engl., Syllab. 48, 1892, nom. inval. err. pro
Hymenostylium Brid.
Barbula subg. Hymenostylium (Brid.) Lindb., Musci Scand.
22, 1879.
Weissia subg. Hymenostylium (Brid.) Kindb., Eur. N. Amer.
Bryin. 2: 283, 1897.
Gymnostomum sect. Hymenostylium (Brid.) Griff., Calcutta J.
Nat. Hist. 2: 480, 1842.
Gymnostomum sect. Vera Griff., Calcutta J. Nat. Hist. 2: 478,
1842, p.p., nom. il/eg.
Pottia sect. Hymenostylium (Brid.) C. Miill., Syn. 1: 562,
1849.
Weissia sect. Hymenostylium (Brid.) Mitt., J. Linn. Soc. Bot.
12: 134, 1869.
Barbula sect. Hymenostylium (Brid.) Braithw., Brit. Moss Fl.
1: 258, 1887.
From {)llllV, -~vo<;, membrane+ o + ot~o<;, pillar, column+
-ium, characteristic of.
Plants growing in turfs or cushions, often flagellate, green,
often glossy, occasionally glaucous above, light brown below.
Stems often branching, to 3(-8) em in length, occasionally
papillose, transverse section rounded-pentagonal to triangular,
central strand usually absent, sclerodermis present, hyalodermis usually absent; axillary hairs ca. 8 cells, basal 1-2 cells
brownish or occasionally all hyaline; often with a red tomentum.
Leaves often distant on stem, appressed-incurved, sometimes
twisted or lax when dry, spreading, occasionally squarrose
when moist, ligulate to lanceolate or linear-lanceolate, ca.
2.0(-3.5) mm in length, upper lamina keeled, margins plane to
broadly recurved (occasionally revolute) along 1 or both sides,
entire or rarely serrulate near apex by projecting cell walls,
rarely bistratose in patches along margins or medially; apex
acute, occasionally obtuse or rounded; base scarcely
124
GENERA OF THE POTTIACEAE
differentiated in shape to oval or rectangular, occasionally narrowly decurrent; costa often stout, sometimes ending l-2 cells
below apex or percurrent or more usually excurrent as a broad
mucro, often "scalloped" along margins by projecting cell walls,
superficial cells usually elongate ventrally, short- to longrectangular dorsally, 2-4 or occasionally several rows of cells
across costa ventrally at midleaf, costal transverse section semi-
circular to round, two stereid bands usually present, the dorsal
crescent-shaped, epidermis usually absent ventrally, often
absent dorsally, guide cells 2-4 in I layer, hydroid strand
absent; upper /aminal cells usually heterogeneous in size and
shape, quadrate to rectangular or rhomboidal, ca. 8-10 Jlm in
width, 1-3:1, walls thin-walled to trigonous, often porose,
superficially flat to somewhat convex, seldom bistratose in
Plate 30. Reimersia. 1-8. R. inconspicua. l. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Capsule.
MERCEYOIDEAE
125
Plate 31. Hymenostylium. 1-8. H. dicranelloides. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Young leaf. 6. Leaf apex. 7.
Transverse section at midleaf. 8. Upper lamina! papillae. 9-11. H. congoanum. 9-10. Two leaves. 11. Leaf apex. 12-16. H. crassinerve. 12.
Transverse section of stem. 13-14. Two leaves. 15. Leaf apex. 16. Transverse section at midleaf. 17-19. H.filiforme. 17-18. Two leaves. 19.
Leaf apex. 20-22. H. papillinerve. 20--21. Two leaves. 22. Leaf apex.
·
126
GENERA OF THE POTTIACEAE
patches; papillae low, simple to granular, not obscuring lumens,
centered to scattered, rarely absent; basal cells differentiated
across leaf, rectangular, little wider than upper cells, 2-4:1, walls
thin to porose. Dioicous. Perichaetia terminal, inner leaves weakly
differentiated, lanceolate, somewhat longer than the cauline,
sometimes sheathing and lower cells inflated-rectangular in lower
third. Perigonia terminal, gemmate. Seta to 1 em in length, 1 per
perichaetium, reddish or yellowish brown, twisted clockwise; capsule occasionally systylious, theca ca. 1 mm in length, yellowish
or reddish brown, ovoid to short-rectangular, exothecial cells
thin- to thick-walled, 1-4:1, stomates phaneropore, at base of capsule, annulus weakly vesiculose; peristome absent. Operculum
narrowly rostrate, occasionally long-conic from a flaring base,
oblique, ca. 0.5-1.0 mm in length, cells straight. Calyptra
cucullate, smooth, ca. 1.0-1.5 mm in length. Spores ca. 13 Jlm in
diameter, brownish, weakly papillose. Laminal KOH color reaction yellow. Reported chromosome number n = 12+m, 13.
Widespread in the world (North, Central and South America,
Europe, Asia, Africa and Australasia), found generally on
calcareous rock, rarely trees, in seepage, along streams and rivers,
near waterfalls, at various elevations.
Hymenostylium is characterized by the usual lack of a stem
central strand (Pl. 31, f. 2, 12; 32, f. 1, 10); mostly keeled,
ligulate-lanceolate leaves with a tendency to trigonous cell
walls and longitudinally elongate median leaf cells; costa with
two stereid bands and the ventral epidermis usually lacking (Pl.
31, f. 7, 16; 32, f. 5, 13); upper laminal papillae mostly low,
simple, not obscuring the lumens (Pl. 31, f. 8); capsules ovoid to
short-rectangular, sometimes systylious; and peristome lacking.
As with many hygrophiles, there is much morphological variation. In fact, in addition to presumed ecotypic differentiation,
collections of this genus may be commonly found with leaves
of rather different shape and areolation on the same stem. The
amount of variation recognized for one species, Hymenostylium
recurvirostrum (Pl. 32, all figures), in the New World (Zander
1977c; Zander & Eckel 1982) is inclusive of t!J.at seen in almost
all species of the genus examined in the course of this study,
indicating that the genus, on revision, should be found to
include far fewer species than are presently accepted. The
systylious capsule is not a constant feature in H. recurvirostrum.
Plate 32. Hymenostylium. 1-S. H. recurvirostrum var. recurvirostrum. 1. Transverse section of stem. 2-3. Two leaves. 4. Leaf apex. 5.
Transverse section at midleaf. 6-9. H. recurvirostrum var. cylindricum. 6. Transverse section of stem. 7-8. Two leaves. 9. Leaf apex. 10-13.
H. recurvirostrum var. insigne. 10. Transverse section of stem. ll. Leaf. 12. Leaf apex. 13. Transverse section at midleaf.
MERCEYOIDEAE
Hymenostylium has many of the gametophyte characters of
Leptodontium, including the usual absence of a stem central
strand (always absent in Leptodontium); lanceolate, carinate
leaves that are strongly recurved when moist; lower leaf margins
often recurved; and ventral costal cells elongate (because the
ventral epidermis is absent and the ventral stereid band is thus
exposed). Hymenostylium is particularly similar to L.
viticulosoides in the usual absence of a stem hyalodermis, often
trigonous or porose lamina! cell walls, upper medial lamina! cells
often longitudinally elongate, and !aminal papillae simple. Some
collections of H. recurvirostrum var. cylindricum from the West
Indies with denticulate upper leaf margins bear a striking resemblance to L. viticulosoides. Leptodontium differs from Hymenostylium, however, in the dry to mesic habitat; broad, reniform
costal section; convolute-sheathing perichaetial leaves; and longcylindrical, peristomate capsule with well-developed annulus.
Certain Trichostomum species (T. tortelloides and T. contractum)
are also similar to Hymenostylium in the short, eperistomate capsules with long-conic opercula (these often falling off with the
columella) and plane-margined leaves, but differ in the presence
of a stem central strand and a ventral costal epidermis, broad
acumination, and upper !aminal cell walls thin and evenly bifidpapillose. There is, also, a similarity with Didymodon sect.
Fa/laces, which, in addition to having a similarly broad geographic range, also has simple !aminal papillae and an exposed
ventral stereid band; D. waymouthii and D. brotheri, which also
grow in hygric habitats, have elongated, somewhat porose medial
!aminal cells, and lack a central strand, and are apparently the
closest morphologically to Hymenostylium in Didymodon. Cladistic study shows a more distant relationship between Hymenostylium and Didymodon than between the former and Leptodontium.
Additional study is needed of West Indian populations of H.
recurvirostrum of great morphological variability (Zander 1977c)
with respect to similar variation often recognized at the specific
level in Asia.
Additional literature: Andrews (1926, 1943), Azziz and Vohra
(1988), Dixon (1927), Gyorffy (1905a), Khanna (1976).
Number of accepted species: 18.
Species examined: H. congoanum (BM), H. contextum (L), H.
crassinervium (NY), H. dicranelloides (BM, NY), H. filiforme
(BM), H. hildebrandtii (NY), H. papillinerve (BM), H. recurvirostrum, H. rigescens (H).
New heterotypic synonymy: Barbula svihlae Bartr. =
Hymenostylium recurvirostrum (Hedw.) Dix. var. recurvirostrum.
Gyroweisia tophicola (C. Miill.) Kindb. = Hymenostylium
recurvirostrum (Hedw.) Dix. Hymenostylium firmum (C. Miill.)
Broth. in Bartr. = Hymenostylium recurvirostrum (Hedw.) Dix.
var. recurvirostrum. Hymenostylium luzonense Broth. (H.
recurvirostrum var. luzonense (Broth.) Bartr.) = Hymenostylium
recurvirostrum var. cylindricum (Bartr.) Zand. in Zand. & Eckel
(based on Hymenostylium glaucum var. cylindricum Bartr., 1936).
Weissia venezuelensis C. Miill. (Gymnostomum venezuelense (C.
Miill.) Kindb.) = Hymenostylium recurvirostrum (Hedw.) Dix.
Molendoa burmensis Bartr. (FH) = Hymenostylium
recurvirostrum var. cylindricum (Bartr.) Zand. in Zand. & Eckel.
Molendoa sordida (Mitt.) Steere (NY) = Hymenostylium
recurvirostrum (Hedw.) Dix.
New combination: Hymenostylium hildebrandtii (C. Miill.)
127
Zand., comb. nov. (Weissia hildebrandtii C. Miill., Linnaea 40:
298, 1876; Gyroweisia hildebrandtii (C. Miill.) Kindb.).
25. TRACHYODONTIUM
Plate 33.
Trachyodontium Steere, Bryologist 89: 17, 1986. Type:
Trachyodontium zanderi Steere.
From 'tpdxu~. rough + 0 + OOoU~ (O~OlV), o&5v'to~. tooth
+ -ium, characteristic of; referring to the ornamentation of the
peristome teeth.
Plants loosely caespitose to forming cushions, yellow-green
above, light brown below. Stems often branching, to 3 em in
length, in transverse section rounded-pentagonal, central strand
absent, sclerodermis differentiated but thin, hyalodermis
strongly differentiated, of thin-walled, bulging cells, collapsed
in older parts of stem; axillary hairs ca. 12 cells in length, basal
1-3 cells brownish; occasionally with knotty tomentum below.
Leaves twisted-flexuose and spreading when dry, spreadingrecurved to squarrose when moist, oblong-lanceolate, to 4 mm
in length, keeled; margins recurved in lower 112 to 113, dentate
in upper 112 to 113, bordered by 1-3 rows of bistratose, elongate, thick-walled cells in upper 213; apex acute; base oblong,
sheathing; costa percurrent, superficial cells elongate and finely
papillose on both surfaces, 4-6 rows of cells across ventral surface at midleaf, costa in transverse section reniform, with two
stereid bands, lacking differentiated epidermal cells, hydroid
strand absent, guide cells 4 in one layer; upper lamina! cells
quadrate to hexagonal or short-rectangular, ca. 12-14 jlm in
width, 1(-2): 1, walls evenly thin, superficially convex but not
strongly bulging; papillae simple, small, scattered, 6-8 per
lumen; basal cells strongly differentiated across leaf base,
reaching higher medially, rectangular, 10-14 )lm in width,
4-7:1, thin-walled, mostly weakly papillose. Propagula absent.
Dioicous. Perichaetia terminal, inner leaves long-lanceolate, to
10-ll mm in length, sheathing the seta, porose-rectangular,
thick-walled cells reaching up 1/2 to 2/3 leaf length. Perigonia
terminal, gemmate, inner leaves deltoid. Seta 1.2-1.4 em in
length, brown, twisted clockwise above; theca 5.5-6.0 mm in
length, brown, smooth, long-cylindrical; exothecial cells rectangular, thick-walled; annulus of deciduous, vesiculose cells,
stomates absent; peristome of 64 linear teeth, arranged in 16
groups of 4, to 600 jlm in length, up to 13 articulations, straight,
red-orange, densely spiculose grading below to low ridges,
basal membrane absent. Operculum and calyptra not seen.
Spores rather variable in size, 13-33 jlm in diameter. Lamina/
KOH color reaction yellow with occasional red patches.
Found on large grass culms and small tree trunks, ca. 2650
m elevation, endemic to Ecuador.
The salient distinguishing features of this genus are the
cartilaginous leaf border (Pl. 33, f. 8), simple !aminal papillae
(Pl. 33, f. 9), elongate cells on both surfaces of the costa (Pl. 33,
f. 6), two stereid bands, and spiculose peristome teeth arranged
in groups of four each (Pl. 33, f. 11). It has a superficial resemblance to Calyptopogon by the leaf shape and border, the simple
papillae, and the lack of a stem central strand, but Calyptopogon
is easily distinguished from Trachyodontium by its semicircular
costal transverse section showing a single stereid band and a
hydroid strand.
128
GENERA OF THE POTTIACEAE
Plate 33. Trachyodontium. 1-11. T. zanderi. 1. Habit, perichaetiate plant. 2. Habit, perigoniate plant. 3. Transverse section of stem. 4-5. Two
leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse section at midleaf. 9. Upper !aminal papillae. 10. Perichaetialleaf. 11. Peristome.
MERCEYOIDEAE
129
Plate 34. Streptotrichum. 1-11. S. ramicola. I. Habit of perichaetiate plant. 2. Perigoniate plant. 3. Transverse section of stem. 4-5. Two
leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse section at midleaf. 9. Upper laminal papillae. 10. Perichaetialleaf. II. Portion of peristome
(note perpendicular aspect of teeth formed from inner anticlinal walls).
130
GENERA OF THE POTTIACEAE
Trachyodontium agrees with salient features of closely related
members of the Leptodontieae in its large size, the convolutesheathing perichaetial leaves (Pl. 33, f. 1), elongate-cylindrical
capsule lacking stomates, and spreading-recurved to squarrose,
dentate leaves with reniform costal transverse section showing
two stereid bands (Pl. 33, f. 8). It is particularly close to Leptodontium through the characteristic !aminal KOH color reaction
and the lack of both a stem central strand and differentiated costal
epidermal cells. Leptodontium, unlike Trachyodontium, has
spirally grooved peristome teeth, but a closely related genus,
Streptotrichum, has spiculose teeth. Another related genus, Leptodontiella, has 16 peristome teeth similarly divided into three or
four rami (these not spiculose, however).
Spindle-shaped propagula are common in the material of
Trachyodontium examined, but evidently belong to a Zygodon
species occurring in mixture.
Literature: Steere (1986).
Number of accepted species: 1.
Species examined: Trachyodontium zanderi (BUF).
Plate 34.
26. STREPTOTRICHUM
Streptotrichum Herz., Biblioth. Bot. 87: 37, 1916. Type: Streptotrichum ramicolum Herz.
From a'tp£1t't6~, twisted + o + 8pf;~.• 'tplx6c;, hair; the peristome
teeth are, however, not sensibly twisted.
Plants in loose mats, green above, tan below. Stems branching
often, to 4 em in length, transverse section rounded-pentagonal,
central strand absent, sclerodermis present, hyalodermis present;
axillary hairs of ca. 7 cells, the basal 2-3 brownish and fmnwalled; knotty tomentum present but sparse. Leaves spreadingtwisted when dry, squarrose-recurved when moist, lanceolate, ca.
3.0-3.3 mm in length, upper lamina keeled, margins recurved
below, dentate above midleaf, apex acute; base sheathing in lower
1/4 to 1/3 of leaf; costa ending 2-4 cells below the apex, superficial cells elongate on both sides of costa, 4-7 rows of exposed
stereid cells across costa ventrally at midleaf, costal transverse
section reniform, two stereid bands present, epidermis absent ventrally and dorsally, guide cells 4 in I layer, hydroid strand absent;
upper !aminal cells quadrate to short-rectangular, ca. 9-12 jlm in
width, 1-2: 1, walls evenly thickened, superficially flat to weakly
convex; papillae very small, simple, solid, ca. 6- 7 per lumen;
basal cells differentiated across leaf base, rectangular, little wider
than upper cells, 3-4:1, walls evenly thickened. Dioicous. Perichaetia terminal, or subterminal by innovations, inner leaves
long-lanceolate, to 8 mm in length, convolute-sheathing, hiding
greater portion of seta, cells long-rhomboidal, thick-walled and
porose. Perigonia gemmate, terminal or subterminal by repeated
innovations. Seta rather short, ca. 5-7 mm in length, 1(-2) per
perichaetium, yellowish brown, not twisted; theca 2-3 mm in
length, yellowish brown, cylindrical, occasionally somewhat
ventricose, exothecial cells thin-walled, short-rectangular,
stomates absent, annulus of 1-2 rows of vesiculose cells, persistent; peristome teeth 16, bright orange-red, linear, cleft into two
rami from near base, occasional interpolated rami arise from
anticlinal walls at margins of regular teeth, densely branchingspiculose, 500-1000 jlm in length, with many articulations,
straight or very weakly twisted counterclockwise, basal membrane low, ca. 70 Jlm high in height, sharply granulose to
spiculose-papillose, occasionally striate in patches. (Operculum
reportedly conic-rostrate, not seen.) Calyptra cucullate, smooth,
ca. 3 mm in length. Spores large, ca. 20 jlm in diameter, brown,
low, spiculose-papillose. Lamina/ KOH color reaction yellow,
with red mottling.
Endemic to Bolivia, where it is known from a single collection from a joint of a bamboo-like grass at 3400 m elevation.
Streptotrichum is close to Leptodontium in the lack of a
stem central strand (Pl. 34, f. 3); the knotty tomentum; the
squarrose-recurved, lanceolate leaves (Pl. 34, f. 4-5) with a
reniform costal section lacking differentiated epidermal layers
(Pl. 34, f. 8); and the highly differentiated perichaetial leaves
(Pl. 34, f. 10). It differs from Leptodontium, as does the similar
Leptodontiella, in having an unusual peristome, in this case of
16 deeply bifid, densely spiculose teeth, each of which occasionally bears an additional ramus or two formed from an interior anticlinal wall (Pl. 34, f. 11). These extra teeth are similar
to the other teeth except they are shorter and appear edge on
when viewed laterally from outside the capsule. Streptotrichum
has much the same lax habit (Pl. 34, f. 1-2) and short capsules
(though this is variable) as does Leptodontiella, but the peristome is quite different.
Literature: Herzog (1916).
Number of accepted species: I.
Species examined: S. ramicola (Bolivia, Waldgrenze tiber
Tablas, Herzog 2844, syntypes, JE, W).
27. LEPTODONTIELLA
Plate 35.
Leptodontiella Zand. & E. Hegew. Type: Leptodontiella
apiculata (Zand.) Zand. & E. Hegew.
From Leptodontium, a genus + i + -ella, diminutive; resembling
the genus Leptodontium.
Plants in loose mats, slender, often branching, greenishbrown. Stems to 3.0 em in length, rounded-pentagonal in section, central strand lacking, sclerodermis strong, hyalodermis
distinct, collapsed; axillary hairs of ca. 6 cells, basal 2 fmnwalled, yellowish. Leaves distant, appressed andjlexuose when
dry, spreading-recurved when moist, distinctly trifarious,
oblong-lanceolate to lanceolate, to 2.5 mm in length, carinate
above; apex narrowly acute, usually ending in a long, pellucid
cell or f ine apiculus; margins weakly recurved below,
decurrent, distantly dentate in the upper 113; base ovate in
shape; upper !aminal cells rounded-quadrate to shortrectangular, 9-12 Jlm in width, 1(-2): I; costa subpercurrent to
percurrent, cells elongate and smooth ventrally and dorsally, ca.
4 cells across surface ventrally, with two stereid bands, the dorsal crescent-shaped, lacking differentiated epidermal cells on
both sides of the costa, guide cells ca. 4(-6) in one row, hydroid
strand absent; upper !aminal papillae simple to bifid, spiculate
or scablike, several scattered over each lumen; walls thin or
collenchymatous, superficially flat; basal cells quadrate to
short-rectangular and chlorophyllose to abruptly enlarged and
hyaline medially, little wider than the upper cells, to 1-4: 1, bordered marginally by several rows of quadrate to shortrectangular cells. Propagula occasionally present, borne on the
stem, cylindrical, 70-120 Jlm in length, of ca. 3-5 uniseriate
cells, occasionally with longitudinal internal walls, dark brownish green excepting a single transparent basal stalk cell.
MERCEYOIDEAE
131
Plate 35. Leptodontiella. 1-11. L. apiculata. 1. Habit of perichaetiate plant. 2. Perigoniate plant. 3. Transverse section of stem. 4-5. Two
leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse section at midleaf. 9. Papillae. 10. Propagula. 11. Peristome.
132
GENERA OF THE POTTIACEAE
Dioicous. Perichaetia terminal, perichaetial leaves to 6 mm in
length, long-sheathing, concealing most or all of the seta; perigonia lateral on the stem, gemmate. Seta short, to 2.0 mm in
length, brown, twisted clockwise; theca short-elliptical to ovoid,
brown, wide-mouthed, 1.0-1.2 mm in length, without stomata;
exothecial cells 33-40 11m in width, relatively short, 2: I, walls
thickened; annulus 2-4 cells in height, vesiculose, persistent;
operculum conic-rostrate, ca. 0.5 men in length, cells in straight
rows; peristome teeth 16, inserted below the annulus, straight or
somewhat twisted clockwise, criss-crossing perpendicularly when
dry, forming an open cone or dome when moist, to 500 !liD in
length, yellow, cleft to base in (2-)3-4 linear to filiform rami,
variously weakly fused below, indistinctly spirally striate, with
7-11 articulations. Calyptra cucullate, smooth, ca. 2 mm in
length. Spores 15-20 !liD in diameter, lightly papillose. Lamina/
KOH color reaction yellow to yellow-orange.
Found on rock, trees and shrubs at 600 to 4236 m elevation;
endemic to Peru.
The genus is closely related to Leptodontium by the enlarged,
sheathing perichaetial leaves (Pl. 35, f. 1); lateral perigonia (Pf.
35, f. 2) as in some species of Leptodontium (e.g. Leptodontium
wallisii); the nearly straight, striate peristome teeth (Pl. 35, f. 11);
the unistratose, carinate cauline leaves; and the costa with a ventrally exposed stereid band (Pl. 35, f. 8). Leptodontiella differs,
however, in the very short seta (rare in Leptodontium), the short
urn, the 16 peristome teeth each cleft into three or four rami
(occasional in the polymorphic L. viticulosoides), and the propagula being cylindrical, uniseriate, and dark brownish green.
Sainsbury (1955) indicated that Brotherus' (1924--25) illustration of very regularly trifid peristome teeth for Tridontium
tasmanicum Hook. f. (here placed in the Grimmiaceae, see
Excluded Taxa) were somewhat misleading in that they are actually irregularly cleft hi-trifid, and are often not cleft to the base,
although perforate below. The multi-fid peristome teeth of Leptodontiella are clearly cleft to the base, although there are marginal
fusions.
Number of accepted species: 1.
Species examined: Leptodontiella apiculata (BUF, FH, H,
US, herb. E. & P. Hegewald).
Additional literature: Zander (1972), Zander and Hegewa1d
(1976).
28. LEPTODONTIUM
Plates 36-37.
Leptodontium (C. Miill.) Hampe ex Lindh., Ofv. K. Vet. Ak.
Forh. 21: 227, 1864. Type: Leptodontium squarrosum (Hook.)
Hampe in Lindh.
Sect. Leptodontium
Didymodon subg. Leptodontium (C. Miill.) Lor., Bryol. Notizb.
18, 1865.
Trichostomum sect. Leptodontium C. Miill., Syn. 1: 577, 1849.
Sect. Verecunda Zand., Bryologist 75: 230, 1972. Type: Leptodontiumflexifolium (Dicks. ex With.) Hampe in Lindh.
Didymodon subg. Leptodon BSG, Bryol. Eur. 2: 2, 1851 (fasc.
46-47 Consp. 2: IV), p.p.
Sect. Crassicostata Zand., Bryologist 75: 256, 1972. Type: Leptodontium pungens (Mitt.) Kindb.
Sect. Coronopapillata Zand., Bryologist 75: 264, 1972. Type:
Leptodontium longicaule Mitt.
Williamsia Broth., Nat. Pfl. 1(3): 1190, 1909, hom. illeg. non
Merrill, 1908. Type: Williamsia tricolor Williams.
Williamsiella Britt., Bryologist 12: 62, 1909, nom. nov. for
Williamsia Broth.
From A.ett·t<f ~. peeled, fine, small, thin, delicate +
+ -ium, characteristic of.
Generally robust plants, in thick mats or short turf, greenish
yellow- to orange-brown above, yellow- to red-brown below.
Stems seldom to often branching, 1-5(-20) em in length, transverse section rounded-pentagonal, central strand absent, outer
cortex usually of thick-walled cells, hyalodermis often present,
usually collapsed in mature parts of stem; axillary hairs of 6-16
cells, cells often bulging, with hyaline walls or basal 1-2 cells
brownish; tomentum often present, red to brown, knotty or
kinky. Leaves erect to spreading, twisted to contorted when dry,
spreading to squarrose-recurved when wet, ovate- to longlanceolate, occasionally lingulate or oblong, (1-)2-5(-8) men in
length, keeled above or channeled along costa, margins
recurved in lower 1/3-3/4, rarely to near insertion, usually
dentate in upper 1/3-1/2, rarely to near insertion, occasionally
bordered above by 1-5 rows of less papillose, thick-walled
cells; apex acute, occasionally rounded to narrowly obtuse; base
commonly rectangular, often sheathing, basal margins sometimes long-decurrent; costa shortly excurrent, percurrent or ending 1-6(-15) cells below apex, superficial cells elongate both
ventrally and dorsally, costal transverse section reniform, occasionally elliptical or semicircular, two stereid bands present,
epidermis absent both ventrally and dorsally, guide cells 2-4 in
1 layer, hydroid strand absent; upper laminal cells subquadrate,
mostly 11-15 11m in width, 1: 1, walls thin or evenly to
collenchymatously thickened, often trigonous, occasionally
strongly bulging superficially; papillae variously simple, bifid,
multifid, often hollow (oh- or cee-shaped in optical section),
occasionally simple- or branching-columnar; basal cells generally strongly differentiated medially or across base, often
sharply demarcated and hyaline, rectangular (occasionally
somewhat inflated), slightly wider than upper cells, 2-5:1, walls
thin to evenly or laterally thick-walled, occasionally porose.
Propagula multicellular, clavate to obovate, borne on short
stalks in leaf axils, occasionally on leaf apices or leafless
branchlets. Flagellate branchlets occasionally present in axils.
Usually dioicous, rarely autoicous or possibly rhizautoicous.
Perichaetia terminal, inner leaves usually long-lanceolate, to
7-8 mm in length, usually convolute-sheathing, lower cells
long-rhomboidal, porose or thin-walled. Perigonia terminal or
lateral (as autoicous buds), or both lateral and terminal on
antheridiate plants. Seta 0.3-3.0 em in length, 1(-2) per
perichaetium, twisted usually clockwise above; theca cylindrical, 2.0--3.5 mm in length, exothecial cells short-rectangular,
moderately thick-walled; stomates absent or present at base of
theca; annulus of 2-7 rows of yellowish or reddish vesiculose
cells, persistent or irregularly deciduous; peristome teeth 16,
linear, occasionally rudimentary (a preperistome rarely
present), yellowish brown to reddish orange, smooth to deeply
striate, usually 300--500 !liD in length, of several articulations,
straight, basal membrane absent or very low. Operculum conic
to conic-rostrate, 0.5-1.0 mm in length, cells in straight rows.
Calyptra cucullate, smooth, 2.5-4.0 mm in length. Spores
homogeneous or occasionally heterogeneous, i.e., in one capsule half larger and chlorophyllose and half smaller and
ooou~ (o&ov), o&5v'to~. tooth
133
MERCEYOIDEAE
~uu
Plate 36. Leptodontium. 1-10. L. pungens. l. Habit. 2-3. Two transverse sections of stem; cells of hyalodermis bulge in immature stem and are
collapsed in mature stem. 4-5. Two leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse section at midleaf. 9. Perichaetialleaf. 10. Peristome.
134
GENERA OF THE POTTIACEAE
Plate 37. Leptodontium. 1-4. L. flexifolium. 1-2. Two leaves. 3. Leaf apex. 4. Propagula. 5-6. L. luteum. 5. Leaf. 6. Leaf apex. 7-9. L.
stoloniferum. 1. Stem apex with propaguliferous branchlets. 8. Leaf. 9. Leaf apex. 10-15. L. viticulosoides var. viticulosoides. 10. Transverse
section of stem. 11-12. Two leaves. 13. Leaf apex. 14. Transverse section at midleaf. 15. Perichaetialleaf.
MERCEYOIDEAE
brownish, weakly papillose, mostly ca. 17-20 11m in diameter.
Lamina/ KOH color reaction usually strong, yellow or less often
orange to yellowish orange, occasionally yellow with red
blotches. Reported chromosome number n = 13.
This is a large genus found mainly in tropical, mountainous
areas of the world, especially characteristic of high elevation fog
forests, growing on soil, acidic rock, trees and shrubs.
The genus Leptodontium is superficially well distinguished by
the robust size of the plants, squarrose-recurved and carinate
leaves, sheathing leaf base, elongate and sheathing perichaetial
leaves (Pl. 36, f. 9), and the straight peristome teeth (Pl. 36, f. 10),
but there is significant variation in many of these characters
among the species. More constant features include the absence of
a stem central strand (Pl. 36, f. 2-3), absence of a differentiated
epidermis over the costa (Pl. 36, f. 8), and smooth or striate peristome teeth (Pl. 36, f. 10). Although Leptodontium is, all in all, a
surprisingly well-delimited genus (for the family), Leptodontiella
and Streptotrichum are rather similar in morphology when sterile,
and their descriptions and illustrations should be carefully studied
to avoid confusion. These last two genera lack stomates in their
capsules, as do many species of Leptodontium. Hollow papillae
reported (Newton & Boyce 1987) as "mamillae" in British L.
flexifolium are actually rather common in tropical collections of
that species.
A revision for the New World (Zander 1972) reviewed the
nomenclatural history, morphology and geographic distribution of
the genus in detail, including much discussion of extra-American
species. Sloover (1987) provided a key to the tropical African
species of the genus.
Didymodon sect. Fa/laces is rather similar in aspect to Leptodontium. Didymodon erosodenticulatus has many of the characteristics of L. viticulosoides (Pl. 37, f. 10--15), including reflexed,
keeled leaves with serrate upper leaf margins and simple papillae,
but differs significantly in the presence of a stem central strand,
brownish basal cells of the axillary hairs, costal ventral epidermal
layer present in at least some leaf sections, perichaetialleaves not
strongly differentiated, and red-orange !aminal KOH color reaction. Hymenostylium ((q.v.) has many of the characters of Leptodontium . The presence of upper !aminal cell wall trigones in
Hymenostylium and the related genus Reimersia, plus the keeled
leaves, general absence of a dorsal costal epidermis and of a stem
central strand are important characters indicating an ancestral link
with Leptodontium, and this link is supported by the cladistic
study. Trigones are variously present in L. wallisii (Peru,
Hegewald 6948, BUF; Colombia, Van Cleef 249, BUF) and L.
viticulosoides.
Leptodontium stellatifolium of Brazil has somewhat the
appearance of Barbula sect. Convolutae and has an unusual bright
orange coloration of its basal cells, but may be retained in Leptodontium by its striate peristome teeth.
Crum and Anderson (1981) recognized L. excelsum (Sull.)
135
Brit. (given as a synonym of L. viticulosoides var. panamense
[= var. sulphureum] by Zander 1972) as a good species endemic
to the Appalachian Mountains of the southeastern United States,
based largely on a uniformity of small size and flagellate
appearance. Robust collections from Jackson Co., North Carolina (BUF), which they may not have seen, are quite like L.
viticulosoides from Mexico, and their report of an autoicous
sexual condition would require transference of this name to the
synonymy of L. viticulosoides var. viticulosoides; the material
available to me lacks perigonia. Actually, the var. sulphureum is
commonly dimorphic (cf Mexico, Oaxaca, Smith et al. 3028,
TENN), the perigoniate plants with hair-like stems and distant
leaves and the perichaetiate not flagellate.
Leptodontium stoloniferum (Pl. 37, f. 7-9) has the appearance of species of Calymperaceae, but it differs in the shape of
the !aminal papillae, in the stalk bearing the propagula being
branch-like (not a modified leaf), and the propagula with internal longitudinal cross walls (Calymperaceae apparently have
only uniseriate cells in their propagula). A specimen from Ecuador (Steere 27591, BUF, NY) has centered, capituliform papillae, which places the species in sect. Coronopapillata.
The anisosporous condition (heterogeneous spores, with two
size classes) characteristic of the spores of L. viticulosoides var.
viticulosoides (Anderson & Zander 1986) was also found, at
least rarely, in L. wallisii; in one Bolivian collection
(Cochabamba, Lewis 79-2217, F), about half the spores in a
capsule were brown and collapsed, the other half green and turgid. Recognized in my (Zander 1972) revision, L. viticulosoides
var. exasperatum is characterized by high, columnar upper
!aminal papillae and is variably isosporous or anisosporous in
different collections. Two additional collections (Mexico, Bowers et al. 5252, 5264-f, both BUF and TENN) have capsules
showing both conditions in the same collection; the latter specimen is clearly dioicous. Perhaps the lethal factors proposed by
Mogensen (1978, 1981, cf Andrews 1929, Wettstein 1928) in
his discussion of false anisospory are the source of variation in
spore size in Leptodontium; this should be investigated.
Additional literature: Frahm (1973, 1986), Herzog (1932),
Janssens and Zander (1980), Long (1982b), Newton and Boyce
(1987), Schumacker and de Zuttere (1981), Sloover (1987),
Zander (1982b, 1983d), Zander and Hegewald (1976), Zander
& Vitt (1979).
Number of accepted species: 39.
Species examined: see revision (Zander 1972) of species in
North, Central and South America, also L. aggregatum (NY), L.
gemmascens (NY), L. hyalinum (FH), L. insolitum (FH), L.
interruptum (FH), L. joannis-meyeri (NY), L. latifolium (H), L.
paradoxicum (BUF), L. pumilum (US), L. styriacum (NY), L.
taiwanense (US).
New heterotypic synonymy: Leptodontium ramosum Crum
& Richards= Leptodontium stoloniferum Zand.
Tribe BARBULEAE
Barbuleae Herz., Geogr. Moose 98, 1926.
Anoectangiaceae Schimp., Coroll. Bryol. Eur. 7, 1855 [1856].
Merceyaceae Casares-Gil, Fl. Iberica, Musgos 247, 1932.
Gymnoweisieae Limpr., Laubm. Deutschl. 1: 222, 1888, rank not given. Lectotyp. nov.: Gymnostomum Hedw.
Barbuloideae (Herz.) Hilp., Beih. Bot. Centralbl. 20: 612, 1933.
Merceyeae (Broth.) Chen, Hedwigia 80: 265, 1941.
The Barbuleae is distinguished at the immediate ancestral node by two advanced features, the dorsal stereid band section round or
semicircular in section and the peristome teeth absent. As per the discussion of Cladograms 11 and 15 in the section on phylogenetic analysis, the exact interrelationships of the genera of the Barbuleae are at best little more than guesswork. Both cladograms
also indicate that loss of a peristome is an advanced character state in the immediate ancestor of the Barbuleae; it is difficult to suppose re-evolution of the peristome in this lineage unless it involved some simple suppression and desuppression mechanism.
Four members of the Barbuleae heretofore placed elsewhere-Anoectangium (Zander 1977c) with the traditional Pleuroweisieae
and Gymnostomiel/a, Scopelophila, and Streptopogon with the Pottioideae (but cf Walther 1983 who places Gymnostomiel/a with
the Hyophileae)-are placed with the Merceyoideae in Cladograms 2-5, 8-10 and 14 in spite of the lack of a ventral stereid band in
these genera. This last feature was previously considered a crucial character distinguishing the traditional Pottioideae (one stereid
band) and Barbuloideae (= Merceyoideae here, two stereid bands). Apparently, like Calyptopogon in the Trichostomoideae, these
four genera are examples of characteristically lanceolate-leaved lineages that have simply lost the ventral stereid band.
The distribution of the Barbuleae is nearly worldwide.
Plate 38.
29. ANOECTANGlUM
Anoectangium Schwaegr., Spec. Muse. Suppl. 1(1): 33, 1811,
nom. cons. non Anoectangium Rohl, 1809, nom. rejic. Type:
Anoectangium compactum Schwaegr.
Anictangium Hedw., Spec. Muse. 40, 1801, nom. rejic.
Anyctangium Hedw. in Lam. & Cand., Fl. Franc. 2: 444, 1805,
orthogr. var.
Pleurozygodon Lindh., Utkast Nat. Grupp. Eur. Lavmoss. 35,
1878. Type: Pleurozygodon aestivus (Hedw.) Lindh.
Anoectangium subg. Euanoectangium Roth, Eur. Laubm. 1:
171, 1904, nom. illeg. Type: Anoectangium compactum
Schwaegr.
Anoectangium subg. Pleurozygodon (Lindb.) Kindb., Eur. N.
Amer. Bryin. 2:317, 1897.
Gymnostomum sect. Anictangium (Hedw.) Leman, Diet. Sc.
Nat. 20: 151, 1821.
Zygodon sect. Anoectangium C. Mi.ill., Syn. 1: 683, 1849.
From dvoex-ro~. opened + dyye'lov, diminutive of &yyo~. vessel or container, capsule; with a wide- or open-mouthed capsule.
Plants growing in turf or mats, green to yellow-brown above,
light to dark brown below. Stems seldom branching, to 1(-3) em
in length, transverse section oval to rounded-triangular or pentagonal, central strand present, strong, outer cortex with smaller
lumens, usually thick-walled, hyalodermis absent; axillary hairs
of 3-10 cells, all hyaline or basal 1-2 brownish and with thicker
walls; weakly radiculose or with red-brown tomentum. Leaves
often distant or crowded, occasionally secund, appressedincurved, often twisted when dry, spreading when moist, ligulate
to lanceolate, occasionally triangular or acuminate, 1.0-1.5(-2.0)
mm in length, upper lamina strongly keeled, deeply grooved
along costa, margins plane or weakly recurved in lower 112,
entire, occasionally finely crenulate or weakly denticulate above;
apex broadly obtuse to sharply acute, usually apiculate, occasionally acuminate or somewhat cucullate; base scarcely differentiated
in shape or ovate; costa ending at the apiculus or sometimes a
few cells below apex or becoming short-excurrent as a mucro, sel-
dom stoutly excurrent, superficial cells elongate or occasionally
short-rectangular to quadrate near apex ventrally, elongate dorsally and generally papillose, 2-3 cells across costa ventrally at
midleaf, costal transverse section oval to reniform, stereid band
single, strong, semicircular to oval, epidermis absent ventrally
(the guide cells superficially exposed) to weakly developed, epidermis usually distinct dorsally, guide cells 2-4 in 1 layer,
hydroid strand absent; upper !aminal cells subquadrate, occasionally hexagonal or elongate transversely or longitudinally in
patches, (5-)7-9(-15) jlm in width, 1(-2):1(-2), walls thin to
greatly thickened, superficially flat to bulging, occasionally in
extraplanar rows (very seldom bistratose); papillae multifid,
often massively so, centered over the lumens or simple to bifid,
scattered, rarely absent; basal cells differentiated in a small
group at base of costa, short-rectangular, little wider than upper
cells, 2-4:1, walls usually thick-walled. Dioicous. Perichaetia
terminal on short lateral branches, inner leaves ovateacuminate, 1.0-1.5 mm in length, convolute-sheathing, lower
cells short-rhomboidal to near apex. Perigonia lateral. Seta
0.3-0.8 em in length, 1 per perichaetium, yellow-brown, twisted
clockwise below, occasionally counterclockwise above; theca
0.5-1.0(-1.5) mm in length, yellow-brown to brown, ovoid to
elliptical, exothecial cells rectangular, walls thin, stomates
phaneropore, at base of theca, annulus of two rows of weakly
vesiculose cells; peristome absent. Operculum long-rostrate,
0.4-0.6(-1.8) mm in length, sometimes longer than the theca,
inclined, cells in straight rows. Calyptra cucullate, smooth,
1.2-1.5(-2.0) mm in length. Spores 9-12(-19) jlm in diameter,
light brown, weakly to strongly papillose. Lamina/ KOH color
reaction yellow to yellow-orange. Reported chromosome number n = 13.
A rather large but singularly homogeneous group distributed
mainly in tropical, arctic or mountane areas of the world.
Anoectangium is unusual in that the distinctions between the
many species are mainly in characters considered variable in
other genera. A close study of the Middle American representation (Zander 1977c) resulted in extensive synonymy with only
MERCEYOIDEAE
137
Plate 38 Anoectangium. 1-9. A. afrocompactum. l. Habit. 2-4. Three leaves. 5. Leaf apex. 6. Basal cells. 7. Leaf section. 8. Papillae. 9.
Perigoniate branch. 10-13. A. aestivum. 10. Leaf. 11. Leaf section. 12. Leaf apex. 13. Papillae. 14-15. A. angustifolium. 14. Leaf. 15. Leaf
apex. 16-17. A. clarum. 16. Leaf. 17. Leaf apex. 18-20. A. mafatense. 18. Leaf. 19. Leaf apex. 20. Leaf section. 21-22. A. magnirete. 21. Leaf.
22. Leaf apex. 23-25. A. strachyanum. 23. Leaf. 24. Leaf apex. 25. Perichaetialleaf. 26-28. A. wilmsianum. 26. Leaf. 27. Leaf apex. 28. Leaf
section.
138
GENERA OF THE POTTIACEAE
one species, A. aestivum (Pl. 38, f. 10-13), recognized for the
area. The most common expression of the species in the area was
found to have small, superficially bulging !aminal cells with
multifid papillae centered over the lumens. Considered weakly
distinguishable were certain uncommon, mostly local variants
with various leaf shapes and usually superficially flat, larger
upper lamina! cells having thickened walls, often transversely or
longitudinally elongate medially, the lamina! papillae low, simple,
seldom multifid. Some collections, however, showed independent
segregation of these features.
Review of a series of exotic taxa for this study indicate that
the Middle American variation is repeated on a larger and possibly more distinctly stepwise scale, with taxonomic distinctions
dependent on combinations of leaf shape, and details of the
areolation and papillae morphology. A Mexican morphological
variant with a tendency for upper !aminal cells to have massive,
centered, capituliform papillae, and for the cells to protrude superficially on one side of the leaf or the other in spaced rows or
patches (these often bistratose) was found to occur also in South
Africa (as A. wi/msianum, Natal, Cathedral Peak Forest Station,
Magill 5532, PRE). Whether this represents a vicariance event or
polytopic differentiation is unknown; however, it may be pointed
out that another South African taxon, Tortula ammonsiana, also
occurs rarely in eastern North America, which is a disjunction of
equal geographic magnitude.
Anoectangium is generally distinguishable from a similar
genus with very short sporophyte-bearing branches arranged laterally on the axis, Molendoa, by its constant lack of a ventral costal
stereid band (Pl. 38, f. 7, 11, 20, 28), but see the discussion of
Molendoa for additional comments. Norris and Koponen (1989)
suggested that "the apparent lateral perichaetia (Pl. 38, f. 1) may
be interpreted as resulting from innovation below the still very
immature perichaetia." The narrow leaves appear to distinguish
Anoectangium from Pottioideae taxa with single stereid bands,
and the rather strongly differentiated stem sclerodermis is clearly
that of the Merceyoideae. The single stereid band, however, cannot be attributed to small plant size alone (as is the case in certain
single-stereid banded collections of species of Didymodon and
Gymnostomum, and other genera of Merceyoideae). Syntrichia
abruptinervis is similar to Anoectangium species in the lanceolate
leaves with a deep, narrow groove along the costa, but differs significantly in the inflated basal cells, costa arcuate in section,
hydroid strand present, and the red KOH reaction. Anoectangium
shares the distinctive deep groove running up the ventral side of
the leaf at the costa with Barbula species, along with similar
papillae and KOH color reaction, among other characters. Molendoa, on the other hand, is similar to Didymodon in many characters, though placed in the Hyophileae of Cladograms 14-16.
Additional literature: Geissler (1985), Malta (1921), Newton
(1983), Rashid (1970), Saxena and Gill (1986), Saxena and
Rashid (1980), Zander and Vitt (1979).
Number of accepted species: 47.
Species examined: A. abyssinicum (BM), A. aestivum, A.
afrocompactum (NY), A. angustifolium (NY), A. bicolor (NY), A.
borbonense (FH), A. clarum (BUF), A. eukilimandscharicum
(BM), A. hobsonii (NY), A. humblotii (FH), A. impressum (BM,
FH), A. keniae (PC), A. lineare (NY), A. mafatense (FH), A.
magnirete (FH), A. raphidostegium (NY), A. rufoviride (DUKE),
A. shepherdae (PC), A. strachyanum (BUF, NY), A. thompsonii
(DUKE, NY), A. walkeri (DUKE), A. wilmsianum (NY).
New combinations: Anoectangium keniae (P. de Ia Yarde)
Zand., comb. nov. (Gymnostomum keniae P. de Ia Yarde, Rev.
Bryol. Lichenol. 22: 10, 1953). Anoectangium shepherdae
(Card. & Dix.) Zand., comb. nov. (Hymenostylium shepherdae
Card. & Dix., J. Bot. 48: 307, 1910).
New synonymy: Anoectangium compactum var. alaskanum
Card. & Ther. Ami = Anoectangium aestivum (Hedw.) Mitt.
Anoectangium sordidum Mitt. = Hymenostylium recurvirostrum
(Hedw.) Dix.
Plate 39.
30. GYROWEISIA
Gyroweisia Schimp., Syn. Muse. Eur., ed. 2. 38, 1876, nom.
cons. Lectotype: Gyroweisia tenuis (Hedw.) Schimp.
Weisiodon Schimp., Coroll. 9, 1856, nom. rejic. Type:
Weisiodon rejlexus (Brid.) Schimp.
Gyroveisia Schimp. ex Luis., Broteria ser. Bot. 8: 36, 1909,
orthogr. var.
Gymnostomum subg. Gymnoweisia B.&S. in BSG, Bryol. Eur.
1: 78, 1846 (fasc. 33-36 Mon. 4). Lectotype: Gyroweisia
tenue Hedw.
Weissia subg. Weisiopsis BSG, Bryol. Eur. 1: 5. 1851 (fasc.
46-47. Consp. 1: VII), nom. illeg. Type: Weissia rejlexa
Brid.
Trichostomum sect. Weisiodon (Schimp.) Lindh., Oefv. K.
Vet. Ak. Foerh. 21: 213, 1864, as "Weissiodon." Type:
Trichostomum rejlexum (Brid.) Lindh.
Weissia sect. Spathulidium C. Miill., Linnaea 40: 298, 1876.
Type: Weissia tophicola C. Miill.
From gyrus, circle + o + Weis[s]ia, a genus; a Weissia-like
moss with a well-developed persistent annulus.
Plants low, gregarious or forming a thin turf, green above,
tan below. Stems short, branching occasionally, to 0.4 em in
length, transverse section rounded-pentagonal, central strand
present or absent, sclerodermis present (substereid), hyaloderrnis absent; axillary hairs of ca. 5(-10) cells, basal 1-3
brownish. Leaves appressed-incurved when dry, weakly spreading, strict to reflexed when moist, narrowly ligulate to longovate, ca. 0.7-1.4 mm in length, upper lamina shallow-grooved
along costa or flat, margins plane to weakly recurved, entire,
occasionally bistratose throughout; apex rounded to rounded
acute or obtuse, sometimes acuminate or apiculate by a sharp
cell; base scarcely differentiated to ovate, occasionally sheathing; costa percurrent or ending ca. 4-8 cells below apex, superficial cells elongate ventrally and dorsally, 2-8 rows of cells
across costa ventrally at midleaf, costal transverse section semicircular to rounded, ventral stereid band absent or weak, often
superficially exposed, dorsal band present and semicircular in
section when well developed, epidermis usually present ventrally, often present dorsally, guide cells 2(-6) in 1 layer,
hydroid strand absent; upper /aminal cells quadrate or shortrectangular, ca. 8-11 Jlm in width, 1-2: 1, walls thin to evenly
thickened, superficially flat to convex; papillae hollow, simple
to indistinctly bifid, scattered, ca. 4 per lumen, occasionally
absent; basal cells differentiated across leaf base in lower 1/4 to
1/2 of leaf, rectangular, commonly enlarged or inflated and
hyaline, usually little wider than upper cells, 3-5:1, walls thin to
evenly thickened. Propagula often present, oval to spindleshaped, of several cells, borne on basal rhizoids, brown.
MERCEYOIDEAE
139
Plate 39. Gyroweisill. 1-13. G. tenuis. 1. Sporangiate plant. 2. Sterile plant. 3. Section of stem. 4--6. Three leaves. 7. Areolation. 8-9. Two
sections at midleaf. 10. Papillae. 11. Propagula. 12-13. Two perichaetialleaves. 14-21. G. reflexa. 14. Sporangiate plant. 15. Perigoniate plant.
16. Sterile plant. 17-18. Two leaves. 19. Areolation. 20. Perichaetialleaf. 21. Peristome. 22-30. G. yuenannensis. 22. Habit. 23-24. Two
leaves. 25. Leaf apex. 26. Leaf base. 27-28. Two sections at midleaf. 29. Perichaetialleaf. 30. Peristome.
140
GENERA OF THE POTTIACEAE
Dioicous or autoicous, occasionally heteroicous. Perichaetia terminal, inner leaves lanceolate, usually well differentiated, to 1.5
mm in length, often strongly sheathing the seta, lower cells rectangular to long rhomboidal. Perigonia terminal, gemmate, on
somewhat smaller plants, or as buds at base of perichaetiate plant.
Seta ca. 1.5--6.0 mm in length, 1 per perichaetium, yellowish
brown, twisted clockwise; theca ca. 0.8-1.5 mm in length, yellowish brown, oval to short-cylindrical, neck sometimes well
differentiated, exothecial cells short-rectangular to rhomboidal,
thin-walled, stomates phaneropore, occasionally somewhat
enlarged, at base of capsule, annulus of 2-3 rows of highly
vesiculose cells, usually revoluble but often merely persistent;
peristome teeth absent or ca. 16, rudimentary, short, ligulate or
oblong and much perforate, lightly papillose to closely spiculose,
ca. 30-80 jlm in length, with ca. 3-4 articulations, straight, basal
membrane low. Operculum short-conic to narrowly rostrate, ca.
0.2-0.6 mm in length, cells straight. Calyptra cucullate, smooth,
ca. 0.7-1.4 mm in length. Spores ca. 8-14 j..lm in diameter, light
brown, smooth to papillose. Lamina! KOH color reaction yellow
or orange. Reported chromosome number n = 13.
Found on thin soil over calcareous rock, in widely scattered
localities across North America, Europe, the Middle East, Africa
and China.
Gyroweisia has long been a "wastebasket" genus (Zander
1977c) wherein several small, feature-poor species have been set
aside. Generally, these taxa have ligulate leaves with subpercurrent costae and enlarged, hyaline basal cells (Pl. 39, f. 7, 19,
25-26), a vesiculose annulus, and a rudimentary peristome or
sometimes none at all (Pl. 39, f. 21, 30). Little attention has been
given to characters of the areolation and anatomy, which may be
used to better place the species. Past work (Zander 1977c; Hill
1981) and the present study has assigned many of these taxa to
more appropriate genera, and even further reduction in the size of
the genus is probable.
Gyroweisia may be viewed (see also Zander & Hermann
1986) as part of a complex evolutionary series, also including
Gymnostomum, Leptobarbula and Barbula sect. Convolutae,
involving morphological reduction of plant size and expression of
the peristome. Gyroweisia differs from a morphologically similar
genus, Gymnostomum, by the occasional presence of a peristome
(albeit rudimentary), the large annulus, the sterile plants (Pl. 39, f.
2) distinctly smaller than the sporophyte-bearing gametophytes,
basal leaf cells differentiated higher up the leaf (lower l/4 to 1/2)
(Pl. 39, f. 7, 19, 26), the more common presence ofpropagula (Pl.
39, f. 11-see also Sergio 1984), and the perichaetialleaves much
larger than the cauline (Pl. 39, f. 12, 13, 20, 29). The transformation series conceived above is, however, not supported by the
cladistic analysis, probably because the analysis utilizes more
than just the most obvious characteristics of the taxa involved.
Additional literature: Andrews (1922c), Conard (1945b),
Crundwell (1981), Egunyomi and Olarinmoye (1978), Geheeb
(1906b), Sergio (1972b), Steere (1939c), Theriot (1923).
Number of accepted species: 6.
Species examined: G. monterreia (BUF), G. reflexa (BUF,
DUKE, NY), G. tenuis (BUF, DUKE, MICH, NY), G.
yuenannensis (H).
New homotypic synonymy: Gyroweisia lindigii (Hampe)
Broth.= Didymodon lindigii (Hampe) Zand. mixed with Didymodon tophaceus (Brid.) Lisa and Didymodon australasiae (Hook.
& Grev.) Zand., judging from the several apparent isotypes that
are present at FH and NY.
Plate40.
31. BELLIBARBULA
Bellibarbula Chen, Type: Bellibarbula kurziana Chen,
Hedwigia 80: 223, 1941, India, "Sikkirn, Phalloot Top,"
Kurz 2026, BM, lectotyp. nov.; NY, isotype.
From bel/us, beautiful + i + Barbula, a genus.
Plants forming a low turf, green to red-orange above, blackish red-brown below. Stems branching occasionally, to 3.0 em
in length, transverse section rounded-pentagonal, central strand
present, sclerodermis thick, hyalodermis absent; axillary hairs
short, 3-4 cells in length, basal 1(-2) cells yellow; sparsely
radiculose. Leaves appressed, incurved when dry, weakly
spreading when moist, short-lanceolate, often concave at
midleaf, ca. 0.7-1.5(-2.0) mm in length, upper lamina broadly
and deeply channeled along costa, margins strongly recurved in
lower 3/4 or to near apex, entire; apex rather narrowly blunt to
acute and sometimes apiculate; base ovate; costa ending 2-4
cells below apex or percurrent, sinuose above midleaf, superficial cells elongate or quadrate near apex ventrally, elongate dorsally, ca. 3-4 rows of cells across costa ventrally at midleaf,
costal transverse section semicircular or circular, two stereid
bands present, epidermis present on both sides of costa, guide
cells 4 in 1 layer, hydroid strand absent; upper /aminal cells
rounded-quadrate to short-rectangular, occasionally transversely elongated in patches, ca. 10-13 IJ.m in width, 1-2:1(-2),
walls thin to incrassate, lumens angular to oval or elliptical,
superficially flat to weakly convex; papillae small, simple to
bifid, solid or hollow, 4-6 per lumen; basal cells not
differentiated or weakly developed medially, quadrate to rectangular, not wider than upper cells, 1(-4):1, walls thin- to
thick-walled . Dioicous. Perichaetia terminal, highly
differentiated, inner leaves long-rectangular, apiculate, to 2.0
mm in length, convolute-sheathing, cells long-rhomboidal to
near apex. Perigonia terminal, gemmate, often in series from
subterminal innovations. Seta ca. 4--6 mm in length, 1 per
perichaetium, reddish brown, twisted clockwise; theca 0.8-1.4
mm in length, reddish brown, cylindrical, exothecial cells thinwalled, rhomboidal, stomates numerous at base of theca, annulus of highly vesiculose cells, revoluble; peristome absent.
Operculum short-conic, ca. 0.4-0.5 mm in length, cells straight.
Calyptra cucullate, smooth, ca. 2.0 mm in length. Spores 15-18
IJ.m in diameter, brownish, lightly papillose. Lamina/ KOH
color reaction red.
Found on rock in montane situations in the Himalayas of
India, China (Yunnan), the Appalachians of the U.S.A., and
mountain ranges in Mexico.
Bellibarbula was proposed by Chen (1941) for two species
distinguished from other Barbuleae by strongly convolutesheathing perichaetial leaves (Pl. 40, f. 10, 11, 16), peristome
absent, ovate-lanceolate leaves, papillose upper !aminal cells
(Pl. 40, f. 9) and little-differentiated leaf bases (Pl. 40, f. 7). The
genus proves to be similar to Bryoerythrophyllum in the red coloration and small, bifid !aminal papillae. Bryoerythrophyllum
shows a tendency towards Bellibarbula in that some of its species likewise have an eperistomate capsule with short-conic
operculum. Bellibarbula is easily distinguished, however, by
the costa usually sinuose above (Pl. 40, f. 4, 5, 13, 14), rather
MERCEYOIDEAE
141
Plate 40. Bellibarbula. 1-12. B. kuniana. l. Sporangiate plant. 2. Perigoniate plant. 3. Section of stem. 4-5. Two leaves. 6. Leaf apex. 7. Basal
cells. 8. Transverse section at midleaf. 9. Upper !aminal papillae. 10-11. Two perichaetialleaves. 12. Fragment of revoluble annulus.13-16. B.
recurva. 13-14. Two leaves. 15. Leaf apex. 16. Perichaetialleaf.
142
GENERA OF THE POTTIACEAE
concave upper lamina (at least in collections with incrassate cell
walls), and the highly differentiated, convolute-sheathing inner
perichaetial leaves with long, rhomboidal cells extending to the
apiculate apex. Didymodon nigrescens and related species are
similar to Bellibarbula in the red coloration, incrassate upper
laminal cells, generally elongate ventral costal cells and short
basal cells, but differ in their very thin costae and lack of papillae
or papillae low and massive.
Judging from study of a range of specimens and the types at
BM and PC, the two species of Bellibarbula are only weakly distinguishable by the features mentioned in Chen •s ( 1941) key and
as discussed (in Chinese) by Chen (1963). The type of B. kurziana
includes some plants with much the same morphology as the second species. Bellibarbula obtusicuspis of China (type at PC!) is
clearly the same as what has been known as Bryoerythrophyllum
recurvum, and a new combination is necessary along with extension of known morphological variation (description by Zander
1978g) and geographic range of the species (United States and
Mexico).
Number of accepted species: 2.
Species examined: B. kurziana (BM, NY), B. recurva (BUF,
DUKE, MICH, PC, TENN).
New heterotypic synonymy: Bellibarbula obtusicuspis
(Besch.) Chen= Bellibarbula recurva (Griff.) Zand.
New combinations: Bellibarbula recurva (Griff.) Zand.,
comb. nov. (Gymnostomum recurvum Griff., Calcutta J. Nat. Hist.
2: 482, 1842, see discussion of nomenclature by Robinson 1968;
Didymodon recurvus (Griff.) Broth; Bryoerythrophyllum recurvum (Griff.) Saito).
32. STREPTOPOGON
Plates 41-42.
Streptopogon Wils. in Mitt., Kew J. Bot. 3: 51, 1851. Type:
Streptopogon erythrodontus (Tayl.) Wils.
Striptopogon Hampe, Flora 45: 450, 1862, orthogr. var.
Sect. Streptopogon
Sect. Streptopogon (Wils.) Mitt., Phil. Trans. R. Soc. London
168: 33, 1879, nom. superjl.
Streptopogon sect. Eustreptopogon C. Mtill., Gen. Muse. Fr.
423, 1900, nom. illeg.
Sect. Calymperella C. Mtill., Hedwigia 33: 128, 1894.
Sect. Streptopogonella Demar. & P. Yarde, Bull. Jard. Bot.
Bruxelles 26: 270, 1956. Type: Streptopogon calymperoides
Demar. & P. Yarde.
From O'tp£7t'to<;. twisted + o + 1tc6yolv, -rovo<;, beard.
Plants growing in tufts or mats, green above, reddish brown
below. Stems commonly branching, ca. 0.5-3.0 em in length,
transverse section rounded-pentagonal to elliptical, central strand
absent (inner cells of central cylinder occasionally crushed),
sclerodermis weakly developed to distinct, hyalodermis absent;
axillary hairs of ca. 7-10 cells, basal 1-3 cells often brownish;
often red-tomentose below. Leaves appressed-incurved to lax and
weakly contorted when dry, widely spreading and recurved when
moist, ovate-lanceolate, 2.0-6.5 mm in length, upper lamina
keeled to broadly channeled, margins recurved to revolute at leaf
base or to near apex, entire to denticulate or serrulate above,
sometimes bordered with elongate cells below or throughout;
apex acute to acuminate, occasionally rounded and cucullate; base
scarcely differentiated to rectangular, somewhat decurrent; costa
usually strongly bulging dorsally, percurrent to excurrent as a
weakly denticulate awn, occasionally subpercurrent by several
cells, superficial cells on both sides elongate and smooth, 2
rows of cells across costa ventrally at midleaf, costal transverse
section round, stereid band single, usually strong and round to
semicircular, ventral and dorsal epidermis present or occasionally absent, guide cells 2 in 1 layer, hydroid strand absent;
upper /aminal cells rectangular to rhombic, occasionally fusiform near apex leaf, 15-20 jlm in width, 2(-3):1, walls thin to
thick, occasionally porose, superficially equally convex on both
sides of lamina; papillae absent; basal cells only weakly
differentiated from the upper cells, rectangular, to 25 jlm in
width, 3-5:1, walls thin to weakly thickened and porose. Uni- or
multiseriate (with internal longitudinal walls) elliptical propagula borne on dorsal leaf apex, apex of costa or upper leaf
margins, seldom remaining on mature leaves, clavate or cylindrical, of 8-16 cells. Dioicous, paroicous, synoicous or
autoicous. Perichaetia terminal, leaves somewhat enlarged,
inner leaves long-elliptical to oblong, to 9.0 mm in length, not
sheathing, lower cells thin-walled and rectangular in lower 3/4.
Perigonia in dioicous species terminal on branches of muchbranching smaller plants. Seta usually short, 0.5-9.0 mm in
length, 1(-2) per perichaetium, yellowish, occasionally twisted
clockwise; theca 3.0--3.5 mm in length, yellowish brown, elliptical, often with a short, rugose neck, mouth of theca occasionally flaring or forming a narrow ring, columella may protrude
past mouth of theca after removal of operculum, exothecial cells
rectangular, walls thin to weakly thickened, stomates
phaneropore, at base of theca, annulus of ca. 4 rows of weakly
vesiculose cells, persistent; peristome teeth 16-32 (reportedly
absent in one species), often coming off with operculum, often
strongly spreading when wet, twisted when dry, linear, filamentous, variously cleft, densely spiculose, ca. 1000--1400 jlm,
with several articulations, twisted counterclockwise ca. once,
basal membrane 70--800 jlm in height, densely spiculose. Operculum conic, ca. 0.9-2.0 mm in length, cells twisted counterclockwise ca. once. Calyptra conic-mitrate, lobed below,
strongly papillose (prorulose) with upward pointing simple
papillae (but reportedly smooth in three species), 2.0--3.0 mm in
length. Spores ca. 13-15(-30) j.l.m in diameter, light brown,
densely papillose. Lamina/ KOH color reaction usually red,
occasionally yellow to yellowish orange. Reported chromosome
number n = 20.
Found on rocks and tree branches and trunks; Latin America, central and southern Africa, Madagascar and Hawaii.
Important characters for Streptopogon are the plant's reddish color below, Jack of a central strand in the stem (Pl. 41, f.
2, 11), ovate-Janceolate leaves, costa with one stereid band and
no hydroids (Pl. 41, f. 8; 42, f. 5, 14, 22), upper !aminal cells
smooth, basal cells little differentiated from the upper, clavate
propagula often present on leaves or costa (Pl. 41, f. 14, 16, 19;
42, f. 17), perichaetial leaves not or little sheathing, seta often
short (Pl. 41 I), calyptra usually scabrous-prorulose (Pl. 41, f.
9-10; 42, f. 9-as in Hypodontium), and laminal KOH reaction
usually red. The peristome (Pl. 42, f. 7, 15) often is only weakly
attached basally and may be removed with the operculum
resulting in what appear to be gymnostomous capsules; however, Griffin (1979b) reported, from a collection with a single
sporophyte with semidetached operculum and calyptra, what
appears to be a true gymnostomous condition in S. calymperes.
MERCEYOIDEAE
143
Plate 41. Streptopogon. 1-10. S. erythrodontus. I. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8.
Transverse section at midleaf. 9. Calyptra. 10. Detail of calyptra. 11-16. S. calymperes. 11. Transverse section of stem. 12-13. Two leaves. 14.
Leaf apex. 15. Transverse section at midleaf. 16. Propagula. 17-19. S. cavifolius. 17-18. Two leaves. 19. Leaf apex.
144
GENERA OF THE POTTIACEAE
Plate 42. Streptopogon. 1-9. S. clavipes. 1-3. Three leaves. 4. Leaf apex. 5. Section at midleaf. 6. Perichaetialleaf. 7. Peristome. 8. Calyptra. 9.
Detail of calyptra. 10-15. S. heterophyllus. 10-12. Three leaves. 13. Leaf apex. 14. Section at midleaf. 15. Peristome. 16-17. S. matudianus.
16. Leaf. 17. Leaf apex. 18-22. S. stenophyllus. 18-20. Three leaves. 21. Leaf apex. 22. Section at midleaf.
MERCEYOIDEAE
Crum (1952b) found both the synoicous and paroicous condition
in a specimen of S. heterophyllus from southern Peru; the type of
this species at W! is synoicous and autoicous. Streptopogon
stenophyllus (Pl. 42, f. 18-22) and S. ca/ymperoides are unique in
their yellow to yellowish orange reaction to KOH; the former is,
however, clearly a member of the genus, possessing characteristic
weak costa, lack of a stem central strand, papillose long-mitrate
calyptra, seta only slightly longer than the 2 mm-long capsule,
and little-differentiated perichaetial leaves, while the latter,
though known only from extremely fragmentary, sterile material,
is much like S. calymperes. "Streptopogon peruvianus" (L!),
although cited as a correct name in Index Muscorum (van der
Wijk et al. 1959-1969) lacks a description, and is thus a nomen
nudum.
Additional literature: Brown ( l898a), Griffin ( l986b ), Salmon
(1903), Sharp (1986).
Number of accepted species: 14.
Species examined: S. ca/ymperes (NY), S. ca/ymperoides
(FH), S. cavifolius (TENN), S. clavipes (BUF), S. erythrodontus
(NY), S. heterophy//us (L), S. matudianus (TENN), S.
stenophyllus (PC).
New heterotypic synonymy: Streptopogon australis Mitt. =
"Daltonia angustifolia? Dozy & Molk." fide H. Robinson's annotation of the type (NY); the type is definitely assignable to that
widely distributed species; also note the strong yellow KOH reaction.
33.BARBULA
Plates 43-45.
Barbu/a Hedw., Spec. Muse. 115, 1801, nom. cons. non Loureiro,
1790. Lectotype: Barbu/a unguicu/ata Hedw.
Barbiferus Poir., Enc. Meth. Bot. Suppl. 1(2): 587, 1811, nom.
illeg.
Mallia Schrank. ex Lindb., Utkast Nat. Grupp. Eur. Bladmoss.
38, 1878, hom. illeg. incl. gen. prior. non Mallia Gmell.,
1791, nee Mallia, 1806, nee Mallia Mart., 1826, nom. cons.
(p .p. Barbula Hedw. et p.p. Tortula Hedw.).
Sect. Barbula
Tortula subg. Barbula (Hedw.) De Not., Mem. R. Ace. Sc.
Torino 40: 287, 1838.
Barbula subg. Barbula Schimp., Coroll. 31, 1856.
Barbula subg. Helicopogon (Mitt.) Lindb., Musci Scand. 22,
1879, nom. illeg. incl. typ. gen.
Barbu/a subg. Eubarbu/a (C. Mtill.) Kindb., Eur. N. Amer.
Bryin. 2: 246, 1897, nom. il/eg.
Barhu/a subg. Tortobarhula Szafr., Fl. Polska Mchy 1: 213,
1957 1958] nom. in val. descr. polon.
Bryum sect. Barbu/a (Hedw.) Relh., Fl. Cantabr. ed. 2: 426,
1802.
Barbula sect. Barbu/a Rebent., Prod. Fl. Neomarch. 257, 1804,
nom . il/eg.
Barbu/a sect. Cau/escentes Hi.ib., Muse. Germ. 317, 1833, nom.
nud. incl. typ . gen.
Tortula sect. Unguiculatae De Not., Mem. R. Ace. Sc. Torino
40: 287, 1838. Type: Barbula unguicu/ata Hedw.
Barbula sect. Unguicu/atae BSG, Bryol. Eur. 2: 80, 1842 (fasc.
13-15 Mon. 18), nom . il/eg. incl. typ. gen.
Barbula sect. Senophyllum C. Mi.ill., Syn. l: 606, 1849 [Not an
error for "steno-", see protologue of Pottia sect. Senophyllaria
C. Mi.ill.)
r
145
Barbula sect. Eubarbu/a C. Mi.ill., Syn. 1: 623, 1849, nom.
il/eg. exc/. typ. gen . cons.
Tortula sect. Barbula (Hedw.) Mitt., J. Linn. Soc. Bot. 12:
144, 158, 1869.
Tortula sect. Helicopogon Mitt., J. Linn. Soc. Bot. 12: 142,
150, 1869.
Barhu/a sect. Fa/ax Lazaro e lbiza, Bot. Oeser. Comp. Fl.
Esp. 1: 586, 1869, nom. illeg. incl. typ . gen. cons.
Barbula sect. Eubarbula Lindb. ex Braithw., Brit. Moss Fl. 1:
261, 1887, nom. il/eg. excl. typ . gen . cons.
Barhula sect. Helicopogon (Mitt.) Braithw., Brit. Moss. Fl. 1:
274, 1887.
Barbula sect. Senophyllum C. Mi.ill. ex Podp., Consp. Muse.
Eur. 200, 1954.
Sect. Hyophilade/phus C. Mi.ill., Syn. l: 604, 1849.
Barbula subg. Hyophilade/phus (C. Mi.ill.) Zand., Phytologia
44: 201, 1979. Lectotype: Barbu/a agraria Hedw. fide
Zander, Phytologia 44, 201, 1979.
Tortula sect. Hyophilade/phus (C. Mi.ill.) Broth., Nat. Pfl.
1(3): 429, 1902.
Barbu/a sect. Agrariae Steere in Grout, Moss Fl. N. Amer.
1(3): 173, 1938, nom. illeg. Type: Barbu/a agraria Hedw.
Sect. Bulhibarbula C. Mtill., Flora 62: 379, 1879.
Barbu/a sect. Rhystobarbu/a C. Mi.ill., Gen. Muse. Fr. 463,
1900, nom . il/eg. incl. sect. prior.
Tortu/a sect. Rhystobarbula Dix., J. Bot. 80:41, 1941.
Tortula sect. Bulbibarbu/a (C. Mi.ill.) Wijk & Marg., Taxon 7:
290, 1958. Type: Tortula eubryum C. Mi.ill.
Sect. Convolllfae B.&S. in BSG, Bryol. Eur. 2: 91, 1842 (fasc.
13-15 Mon. 29).
Streblotrichum P. Beauv., Mag. Enc. 5: 31'7, 1804. Lectotype:
Streb/otrichum convo/utum (Hedw.) P. Beauv. fide Saito, J.
Hattori Bot. Lab. 39: 499, 1975.
Tortula sect. Convo/utae De Not., Mem. R. Ace. Sc. Torino
40: 287, 1838. Type: Tortu/a convoluta (Hedw.) Gaertn.,
Meyer & Scherb.
Tortu/a sect. Leptopogon Mitt., J. Linn. Soc. Bot. 12: 143,
156, 1869. Type: Tortula ca/yculosa Mitt., /ectotyp. nov.
Tortula subg. Streb/otrichum (P. Beauv.) Chev., Fl. Gen. Env.
Paris 2: 51, 1827.
Barbula subg. Odontophylla Saito, J. Hattori Bot. Lab. 39:
499, 1975. Type: Barbula hiroshii Saito.
Barbu/a sect. Leptopogon (Mitt.) Lindb., Muse. Scand. 22,
1879.
Barbula subg. Streblotrichum (P. Beauv.) Limpr., Laubm.
Deutsch!. 1: 626, 1888.
Sect. Hydrogonium (C. Mi.ill.) Saito, J. Hattori Bot. Lab. 39:
492, 1975.
Hydrogonium (C. Mtill.) Jaeg., Ber. St. Gall. Naturw. Ges.
1877-78: 405, 1880 (Ad. 2: 669). Lectotype: Hydrogonium
ehrenbergii (Lor.) Jaeg. fide Saito, J. Hattori Bot. Lab. 39:
492, 1975.
Didymodon subg. Hydrogonium (C. Mi.ill.) Kindb., Eur. N.
Amer. Bryin. 2: 273, 1897.
Barbula subg. Hydrogonium (C. Mi.ill.) Fleisch., Musci Fl.
Buitenzorg 1: 352, 1904.
Trichostomum sect. Hydrogonium C. Mi.ill., Linnaea 40: 297,
1876.
Semibarhu/a Herz. ex Hilp., Beih. Bot. Zentralbl. 50: 626,
1933. Type : Semibarbula indica (Hook.) Hilp.
146
GENERA OF THE POTTIACEAE
Hydrogonium sect. Barbuliella Chen, Hedwigia 80: 233, 1941.
Hydrogonium sect. Euhydrogonium Chen, Hedwigia 80: 233,
1941, nom. illeg.
Sect. Pachynoma (Mitt.) Par., Ind. Bryol. ed. 2, 3: 348, 1905.
Tortula sect. Pachynoma Mitt., J. Linn. Soc. Bot. 12: 143, 151,
1869.
Sect. Pseudocrossidiella Ther. in Felipp., Rev. Bryol. n. ser. 2:
216, 1930. Type: Barbula subgrimmiacea Ther.
Subsect. Purpureaeformes Kindb., Eur. N. Amer. Bryin. 2: 246,
1897. Type: Barbula purpurea C. Mtill.
From barba, beard + -uta, diminutive; little beard, referring to the
peristome of 32 hair-like divisions.
Plants loosely caespitose or forming cushions, yellowish
brown, brown or blackish above, yellowish brown to reddish
brown below. Stems branching irregularly, ca. 0.2-3.0 em in
length, transverse section rounded-pentagonal or irregular, central
strand present or seldom absent, sclerodermis usually present,
hyalodermis occasionally present; axillary hairs 2-10 cells in
length, usually all hyaline, occasionally basal l-2 cells firmwalled; rhizoids sparse to common. Leaves appressed-incurved to
weakly spreading, often contorted or twisted about stem and occasionally catenulate when dry, spreading when moist, spathulate,
ligulate or more usually broadly lanceolate to long-triangular,
usually ca. 1-3 mm in length, upper lamina usually deeply and
narrowly grooved ventrally along costa, occasionally only
broadly concave, lamina unistratose or seldom bistratose in
patches, margins usually recurved in lower 112-213 of leaf, occasionally plane, entire or occasionally denticulate near apex or
above midleaf, seldom dentate, margins occasionally thickwalled, seldom bi- or multistratose; apex rounded to obtusely
acute, usually mucronate, occasionally entire or short-apiculate;
base usually weakly differentiated to ovate, sometimes broadened
and sheathing, sometimes narrowly decurrent; costa percurrent to
short-excurrent as a sharp mucro, seldom short-awned, occasionally ending a few cells below the apex, superficial cells elongate
or occasionally quadrate to short-rectangular ventrally, usually
elongate but sometimes quadrate to short-rectangular dorsally,
2-3(-5) rows of cells across costa ventrally at midleaf, costal
transverse section oval to semicircular, two stereid bands usually
present, usually small but distinct ventrally, usually strong dorsally and crescent-shaped but sometimes nearly semicircular,
epidermis usually differentiated ventrally, usually present but
weakly differentiated dorsally, guide cells usually 2-4 in 1 layer,
hydroid strand occasionally present; upper /aminal cells quadrate
to short-rectangular, 6-13 Jlm in width, usually 1:1, walls thin to
evenly thickened, usually superficially bulging on both sides,
occasionally ventrally bulging and dorsally flat or nearly so;
papillae hollow or solid, multifid or bifid, 2-3 per lumen, seldom
simple or absent, usually obscuring the lumens, occasionally
mamillose ventrally; basal cells usually differentiated, reaching
across leaf or reaching higher medially or occasionally marginally, rectangular, usually little wider than upper lamina[ cells,
3-5: l, walls thin to evenly thickened. Propagula when present
borne on basal rhizoids or axillary on stalks, often rather large,
30-300 Jlm in length, of 1-50 cells, clavate to ovate, occasionally
armed, multicellular, green to reddish. Dioicous, possibly occasionally rhizautoicous. Perichaetia terminal, inner leaves little
differentiated or ovate to long-lanceolate, sometimes strongly
sheathing, lower cells long rhomboidal in lower l/2 of leaf, occa-
sionally to 3/4 or more. Perigonia gemmate, often prominent.
Seta 0.5-1.5 em in length, usually l per perichaetium, yellowish
to reddish brown, twisted clockwise below, often counterclockwise above; theca usually 0.8-1.5 mm in length, yellowish to
reddish brown, ovate to long-cylindrical, exothecial cells rec- ·
tangular or occasionally rhomboidal, walls thin to evenly thickened, stomates phaneropore, at base of theca, annulus weakly
differentiated to strong, of l-3 rows of vesiculose cells, usually
persistent, occasionally revoluble or deciduous in pieces; peristome teeth of 32 narrow rami, seldom short or rudimentary,
filamentous to narrowly triangular, usually densely spiculose,
with many articulations, usually strongly twisted 1.5 to 2 times
counterclockwise, occasionally straight, to 1200 Jlm in length,
basal membrane low but distinct, ca. 40 Jlm in height, granulate
to spiculose. Operculum usually long-conic, ca. 0.8-1.7 mm in
length, cells twisted counterclockwise. Calyptra cucullate,
smooth, usually 1.5-3.0 mm in length. Spores usually 9-16 Jlm
in diameter, light brown, weakly papillose. Lamina/ KOH color
reaction yellow, occasionally yellowish orange. Reported chromosome number n = 10, lO+m, 11, l2+m, 13, l3+m, 13+2m,
14, l4+2m, 16, 24, 26.
A cosmopolitan genus found in a wide variety of habitats,
mainly on soil and acid or calcareous rock.
Saito's (1975a) distinctions between the genera Barbula and
Didymodon are to a large extent followed in this treatment.
Axillary hair characters, however, are considered here to be
more variable than indicated by Saito. In Barbula, the hairs are
often but not always entirely much elongate and hyaline (Pl. 44,
f. 21-as opposed to the short, basally brown-celled and otherwise hyaline hairs of Didymodon). Some species of Barbu/a
have elongate, entirely hyaline hairs (e.g. B. hiroshii and B.
unguiculata), some have firm-walled hair basal cells (e.g. B.
riograndensis), and others may be variable within the same species (e.g. B. indica). (Completely hyaline hairs may be interpreted as all firm-walled rather than all thin-walled. Firmwalled basal cells are either hyaline or somewhat more yellowish than the distal cells. The cells of the completely hyaline
hairs in Barbula are, in fact, rather firm-walled.) In Didymodon,
however, the brownish, firm walls of the basal cells of the
axillary hairs appear to develop earlier than they do in those
species of Barbula that have them. Small, dark rectangles are
generally visible under the microscope through the transparent
bases of the leaves of the extreme stem apex of Didymodon species, while they are seldom so evident in Barbula. Careful stripping of the subapical leaves will uncover the young hairs.
Saito's (l975a) use of the shape of the superficial cells of
the ventral surface of the costa as a distinction (Barbula with
elongate cells-Pl. 44, f. 9, 14, 20, Didymodon sect. Didymodon
with short or quadrate cells) is variable for Barbula world wide,
though it apparently holds for the Japanese species he revised;
Eddy (1990) also considered the elongate ventral costal cells to
be taxonomically important in the Malesia area. The two genera, however, can usually be distinguished quickly by the morphology of the lamina! papillae and the leaf apex. In Barbula,
the upper !aminal papillae are rough, knobby, obscuring the
lumens, and protuberant along the upper !aminal margins (Pl.
43, f. 5), while in Didymodon these are generally low, difficult
to distinguish, and little evident along the upper margins. There
are, of course, exceptions, notably Barbula sect. Hydrogonium
and Didymodon sect. Vineales. Bryoerythrophyllum is quite like
MERCEYOIDEAE
147
Plate 43. Barbula. 1-8. B. costesii. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse section
at, midleaf. 8. Peristome. 9-14. B. agraria. 9. Habit. 10--11. Two leaves. 12. Leaf apex. 13. Transverse section at midleaf. 14. Peristome. 15-20.
B. ·amplexifolia. 15--17. Three leaves. 18. Leaf apex. 19. Transverse section at midleaf. 20. Propagula.
148
GENERA OF THE POTTIACEAE
Barbula in papillae morphology but reacts red, not yellow, to
KOH. Barbula is unlike Didymodon in that its leaf apex usually
ends in an apiculus of one or a few clear cells or in a rather
strong, sharp mucro; in Didymodon, the costa ends before or in
the apex, or, if excurrent, is rather blunt and opaque, seldom
apiculate by one or more clear cells. As Saito (l975a) pointed out,
in Barbula the propagula (Pl. 43, f. 20; 44, f. 5, 10) range from
small and ovate to large and irregular in shape, while Didymodon
has only small ovate propagula. Both Barbula and Didymodon,
however, remain rather heterogeneous, and segregate genera will
surely be described in the future to include, for instance, such
oddities as B. integrifolia and B. eubryum. Compounding this are
species of Didymodon still remaining as combinations in Barbula;
transfer of some of these is done here, but much remains to be
accomplished by revisionists. Barbula may be hypothesized as the
start of a reduction series continuing through Barbula sect.
Convolutae, Leptobarbula, Gyroweisia and Gymnostomum. Gyroweisia is distinct in the combination of enlarged basal cells,
vesiculose and often revoluble annulus, and peristome absent or
reduced. It may well be, however, that Barbula will be broken up
in the future into segregate genera consisting of narrowly conceived reduction series of species crossing presently recognized
limits of these similar genera.
Barbula sect. Convolutae is distinctive in the mostly plane
leaf margins, usual presence of a stem hyalodermis, commonly
strongly differentiated perichaetial leaves, and generally a yellow
seta. It is problematically quite similar to Trichostomum, except
for the presence of a twisted peristome, and may be related. However, in many species of sect. Convolutae the basal cells extend
farthest up the leaf medially (in others merely straight across) or
reaching highest at the midpoint between the costa and the margins on both sides). What is apparently the type of sect.
Convolutae, B. convoluta, has a stem section like that of the
Merceyoideae (central strand present, comparatively large inner
cylinder cells, thin cortex of abruptly smaller stereid cells, hyalodermis little or not differentiated) rather than the Trichostomoideae (central strand variably present, inner cylinder cells often relatively small, cortex commonly of substereid cells, hyalodermis
often well differentiated and sometimes in more than one layer).
This is also true of B. indica, which, although having plane leaf
margins, is clearly a Barhula by its stem anatomy, though probably best placed in Hydrogonium since the generitype of
Hydrogonium is clearly a related but derived species. The type of
B. cancellata (a name recently much in use in North America) at
NY is B. indica var. indica; it bears small propagula. Section
Convolutae characteristically has rather large propagula; axillary
or rhizoidal propagula are typical of many genera of Barbuleae
but are uncommon in genera of Pottieae or Trichostomoideae. It is
quite possible, in any case, that upon revision at least certain species of sect. Convolutae (such as B. amplexifolia, Pl. 43, f. 15-20,
which has the stem anatomy of the Trichostomoideae) will be
seen to better belong in or near Trichostomum, or even Tortella.
Norris and Koponen (1989), on the other hand, in their treatment
of the bryophytes of an area in New Guinea, indicated that planemargined Barhula species lack a hyalodermis and have little taper
to the costae, while Trichostomum species with which they might
be confused have a hyalodermis and tapering costae. Further
investigation is necessary at the revision level. They also pointed
out that sterile specimens of Hydrogotzium can be distinguished
from Dicranella (Dicranaceae) by the former's quadrate cells of
the ventral surface of the costa and a regular arrangement of the
!aminal cells in rows.
The genus Tetrapterum is similar to sect. Convolutae in its
gametophyte morphology, but differs significantly in the sporophyte. Barhula calycina, which may belong with Tetrapterum,
is unusual in its basal cells differentiated into two groups of
about equal size, of hyaline, thin-walled, smooth cells towards
the margins, and yellow, thick-walled, papillose cells medially.
This is reminiscent of a corresponding morphology in Pseudosymblepharis species. Barbula subcalycina has very lax basal
cells reminiscent of those of Tortella humilis. Stone (1991) provided good distinctions between B. calycina and B. suhcalycina.
Barbula calyculosa, B. fendleri and B. fide/is are probably the
same as B. convoluta.
Many species of sect. Hydrogonium (the type of which is
Barbula ehrenbergii) may have evolved from ancestors quite
like B. indica towards a hygric habitat and large size. Barbula
leucodontoides (Pl. 44, f. 18-21-isotype, NY!) has a dorsally
prorulose costa, and is surely very closely related to B. indica.
Other species (e.g. B. zambesiaca) have rather flaccid leaves
with reduced papillae and the dorsal surface of the costa is
merely bumpy. Barbula javanica, with its ventrally bulging and
dorsally smooth (or nearly so) upper lamina, may share ancestors with the Hyophileae. Note that cladistic analysis does not
support a close relationship between Hyophila and Barbula, and
perhaps the section Hydrogonium should be recognized at the
generic level. Barbula agraria (Pl. 43, f. 9-14), with its broad
leaves and ventrally bulging upper !aminal cells may also
belong with the Hyophileae. Both sect. Hydrogonium and sect.
Hyophiladelphus have the essentially tropical lowland distribution characteristic of the Hyophileae. Cladistic analysis at the
species level might detail support for this possible dismemberment of Barbula along these lines.
Barhula marginatula (Pl. 44, f. 22-25) is at an end point in
the evolution of sect. Hydrogonium with the additional character of a cartilaginous, denticulate leaf border of elongate cells.
Gangulee (1972) indicated that B. marginatula may represent a
"new genus of barbuloid mosses" but if such were recognized
the genus should include the types of Hydrogonium and Semibarbula, which clearly belong to the lineage. Barbula
pachyloma (refered to Cinclidotus by Hilpert 1933 but recognized as a good species of Barhula by Eddy 1990 and Norris
and Koponen 1989) may well belong here too, having multistratose upper lamina! borders of stereid cells covered with parenchymatous cells (much like those of Calymperaceae species),
but prorulae are not evident; it is much like Dialytrichia in
appearance and moist habitat but differs in the stereid cells of
the !aminal margin, widely channeled (non-keeled) leaves, and
quadrate ventral superficial cells of the costa.
Species previously referred to Barbula but which have longawned, marginally highly revolute leaves with strongly flattened dorsal stereid band and the ventral band often absent, and
the perichaetialleaves usually enlarged and convolute-sheathing
are better recognized in Pseudocrossidium ((q.v. ).
Certain species of Ditrichum, namely D. tortipes (Mitt.) Par.
and D. amhiguum Best (cf Crum & Anderson 1981, Grout 1927
and Robinson 1968-these two species probably conspecific),
have once-twisted, densely spiculose peristome teeth and longtriangular, apically rough, setaceous leaves with rectanguh;1,
smooth upper !aminal cells much like the leaves of forms/of
/
MERCEYOIDEAE
149
Plate 44. Barbu/a. 1-'6. B. comosa. l-2. Two leaves. 3. Leaf apex. 4. Basal cells. 5. Propagula. 6. Perichaetialleaf. 7-10. B. hiroshii. 7-8. Two
leaves. 9. Leaf apex. 10. Propagula. 11-15. B. hispaniolensis. 11-13. Three leaves. 14. Leaf apeX. 15. Transverse section at midleaf. 16-17. B.
integrifolia. 16. Leaf. 17. Leaf apex. 18-21. B. /eucodontoides. 18-19. Two leaves. 20. Leaf apex. 21. Axillary hairs. 22-25. B. marginatula.
22. Transverse section of stem. 23-24. Two leaves. 25. Leaf apex, lateral view.
150
GENERA OF THE POTTIACEAE
Barbula arcuata. These Ditrichum species may actually belong
inBarbula. Future investigation, at least of the B. arcuata complex, should deal with this question. Barbula arcuata differs significantly in the red rather than yellow, more strongly twisted
peristome, and the leaves only weakly denticulate and margins
unistratose. Its costal section, however, is strikingly like that of
the Ditrichum species, especially in the strongly differentiated
dorsal epidermal cells.
A quite unusual species is Barbula hispaniolensis (Pl. 44, f.
11-15), which is similar to Bryoeyrthrophyllum ferruginascens,
except that the leaves are bright yellow in KOH solution and the
costa has a narrow ventral groove (characteristic of Barbu/a).
This species is retained here pending further study; one might
hypothesize a chemical treatment used during collecting that
might have changed the color reaction, but casual experiments
show that neither boiling in formaldehyde solution or in ethyl
alcohol causes irreversible changes to the red KOH reaction of
true B. ferruginascens.
Additional literature: Abramova et a!. ( 1967}, Coats and
Mahler (1985}, Conard (1945a, 1951a), Crum (1956, 1965d,
1967a), Crundwell (1976), Dhingra-Babbar (1988), DhingraBabbar and Chopra (1985), Field (1990), Hoffmann (1957},
Maheu (1908), Roorda van Eysinga (1972), Saito (1971a,
Sharma and Chopra (1987), Sharma and Sharma (1980, 1986},
1975a}, Steere (1938a, 1939b), Takio (1989), Takio et a!.
(1986}, Weber (1972), Zander (1979f, 1981a). Most bibliographic references to Barbula also discuss species of Didymodon sensu Saito (1975a) and the present treatment.
Number of accepted species: 205, plus 8 remaining in
Hydrogonium, 1 in Semibarbula, and 3 in Streblotrichum.
Species examined: B. afrofontana (NY), B. agraria, B.
arcuata (BM, BUF, FH, NY, TENN), B. amplexifolia (BM,
BUF, MICH, NY), B. bicolor (BUF), B. calycina (NY}, B.
calyculosa (NY), B. clavicostata (PC}, B. comosa (NY), B.
convoluta, B. costesii (NY), B. crocea (BUF), B. ehrenbergii, B.
enderesii (BUF}, B. eubryum (BUF, H, US), B. eustegia (NY},
B. fendleri (NY}, B. fide/is (NY}, B. hiroshii (TNS}, B.
hispaniolensis (NY), B. indica, B. integrifolia (US}, B. isoindica
(NY), B. javanica (NY), B . lambarenensis (NY), B.
leucodontoides (NY}, B. microcalycina (NY}, B. munyensis
Plate 45. Barbula. 1-4. B. munyensis. 1-3. Three leaves. 4. Leaf apex. 5-9. B. pseudoehrenbergii. 5-7. Three leaves. 8. Leaf apex, dorsal
view. 9. Transverse section at midleaf. 10-12. B. riograndensis. 10. Leaf. II. Leaf apex. 12. Transverse section at midleaf. 13-15. B. williamsii. 13-14. Two leaves. 15. Capsule with calyptra.
MERCEYOIDEAE
(NY), B. occidentalis (NY), B. orizabensis (BM, BUF, FH,
TENN), B. pachyloma (BUF), B. peruviana (NY)-near B.
indica, B. pseudoehrenbergii (NY), B. rechingeri (H), B.
riograndensis (BUF, FH), B. semirosulata (PC), B. spathulifolia
(BM), B. subcalycina (NY), B. wil/iamsii (NY), B. unguiculata,
B. zambesiaca (NY).
New heterotypic synonymy: Barbula tonkinensis (Besch.)
Broth. = Barbula indica (Hook.) Spreng.
New combinations and new names: Barbula clavicostata
(Ren. & Hilp.) Zand., comb. nov. (Hyophila clavicostata Ren. &
Card., Act. Soc. Linn. Bordeaux 53: 21, 1898 and Prodr. Fl.
Bryol. Madag. 123, 1898; Hydrogonium clavicostatum (Ren. &
Card.) Hilp.). Barbula isoindica Zand., nom. nov. (Pottia
papil/inervis Lor., Moostud. 160, 1864; Barbula papillinervis
(Lor.) Broth., hom. il/eg., Nat. Pfl. 1(3): 408, 1902). Barbula
semirosulata Zand., nom. nov. (Gymnostomiella rosulata P.
Yarde, Rev. Bryol. Lichenol. 26: 1, 1957). Barbula (sect.
Hydrogonium) spathulifolia (Dix. & P. Yarde) Zand., comb. nov.
(Merceyopsis spathulifolia Dix. & P. Yarde, Arch. Bot. 1(8-9):
164, 1927).
34. GYMNOSTOMUM
Plate 46.
Gymnostomum Nees & Hornsch., Bryol. Germ. 1: 112, 1823,
nom. cons., non Gymnostomum Hedw., 1801, nom. rejic.
Lectotype: Gymnostomum calcareum Nees & Hornsch.
Sect. Gymnostomum
Gymnostomum subg. Eugymnostomum Schimp., Syn. 39, 1860,
nom. il/eg.
Gymnostomum subg. Gymnostomum Kindb., Eur. N. Amer.
Bryin. 2:284,288, 1897, nom. il/eg.
Bryum sect. Gymnostomum (Hedw.) Relh., A. Cantabr. ed. 2:
424, 1802.
Gymnostomum sect. Holomitria Wallr., Fl. Crypt. Germ. 1: 94,
1831, p.p.
Trichostomum sect. Pycnocaulus Lindh. ex Milde, Bryol. Siles.
106, 1869.
Sect. Diastoma Griff., Calcutta J. Nat. Hist. 2: 481, 1842.
From "(Uj.lV6~, naked + o +<Tt6j.lCX, -cx'to~. mouth; the peristome is
absent.
Plants growing in turf or cushions, light to dark or olive green
above, light to dark brown below. Stems branching often, to 2.7
em in length, transverse section rounded-pentagonal to occasionally rounded-triangular, central strand usually present, weak,
sclerodermis absent or occasionally present, hyalodermis present
or absent; axillary hairs 3-10 cells in length, the basall-2 usually
brownish; sparsely radiculose or occasionally red tomentose.
Leaves appressed to appressed-incurved when dry, weakly
spreading to spreading-recurved when moist, usually ligulate,
occasionally ovate to circular, short, to 1.1(-1.8) mrn in length,
upper lamina flat to broadly convex, occasionally keeled, margins
plane or occasionally recurved below midleaf, entire or minutely
crenulate by projecting papillae, sometimes bistratose marginally
above midleaf; apex rounded obtuse to broadly acute, often
apiculate; base scarcely differentiated or seldom ovaterectangular, sometimes denticulate marginally; costa ending 2-5
cells below apex, only occasionally percurrent, occasionally swollen at or above midleaf, ventrally usually bulging, ventral superficial cells quadrate or short-rectangular, occasionally elongate,
151
dorsally elongate or occasionally short-rectangular to quadrate
above midleaf, both sides of costa papillose, 2-4(-6) rows of
cells across costa ventrally at midleaf, costal transverse section
ovate to semicircular, stereid bands weak or lacking ventrally,
present but often weak dorsally, ventral epidermis present, dorsal occasionally little differentiated, guide cells 2(-4) in 1 layer,
hydroid strand absent; upper /aminal cells subquadrate, 7-11
j.lm in width, 1: 1, walls thin to weakly evenly thickened, seldom
irregularly thickened and lumens angular, homogeneous, superficially flat to convex on both sides, often appearing wrinkled in
section because of hollow papillae, cells of apex often in rows
that "criss-cross" at right angles just below the apex; papillae
simple to bifid, low, small, scattered, generally crowded, 3-5
per lumen, hollow or solid; basal cells differentiated across leaf
or rising higher medially, rectangular, little wider than upper
cells, 2-4:1, walls thin. Propagula occasionally present, spherical to obovoid or spindle-shaped, of 5-10 usually multiseriate
cells, borne on branching stalks in leaf axils. Dioicous. Perichaetia terminal, inner leaves ovate-lanceolate, to 1.5 mrn in
length, sheathing below midleaf, sometimes marginally
serrulate, cells rectangular and occasionally bulging below
midleaf. Perigonia gemmate, terminal. Seta generally 0.3-0.6
em in length, 1 per perichaetium, yellowish to reddish brown,
twisted clockwise; theca 0.5-0.8 mrn in length, yellowish to
reddish brown, ovoid to elliptical, occasionally with a weak but
high circumstomal ring, exothecial cells quadrate to rectangular,
walls thin to somewhat thickened, stomates phaneropore, at
base of theca, annulus of 1-3 rows of smaller, transversely
elongated, occasionally vesiculose cells; peristome teeth absent.
Operculum rostrate to conic-rostrate, ca. 0.4-0.5 mm in length,
cells in straight rows. Calyptra cucullate, smooth, 0.5-1.2 mm
in length. Spores 9-15 j.lm in diameter, brownish, essentially
smooth to clearly papillose. Laminal KOH color reaction usually yellow to yellow-orange, occasionally red in patches.
Reported chromosome number n = 13.
Found on rock (usually calcareous, occasionally acid) in
very moist areas; widely distributed on most continents.
Gymnostomum might be viewed as a much reduced,
hygrophilic segregate of Barbula, an end-member of a phyletic
series beginning with Barbula sect. Barbula, and extending
through B. sect. Convolutae, Leptobarbula and Gyroweisia.
Cladistic analysis (Cladograms 11 and 14), however, does not
support more than a close relationship with Barbula. The generally well developed papillae (Pl. 46, f. 9) of Gymnostomum,
especially as they roughen the upper laminal margins, are like
those of Barbu/a, but the firm-walled basal cells of the axillary
hairs and the costa ending below the apex are more characteristic of Didymodon.
Large plants of Gymnostomum aeruginosum (e.g. U.S.A.,
Flowers, Utah, 331, US, and Czechoslovakia, Pilous 857,
DUKE) have a tendency toward a leaf base differentiated in
shape (becoming rectangular) but the apex of such large plants
is more like that of Hymenostylium, the costa being percurrent
or vaguely excurrent as a broadly triangular mucro, or
percurrent and the apex rounded-acute, as is commonly the case
in Didymodon sect. Didymodon, not excurrent and clearly distinct from the tissue of the lamina as is the case with Barbula.
Hilpert's (1933) combination Barbula mosis (Lor.) Hilp. shows
a sensitivity on his part to the Barbu/a relationship, as does K.
Saito's annotation of a combination in Barbula on the type of
152
GENERA OF THE POTTIACEAE
Plate 46. Gymnostomum. 1-9. G. aeruginosum. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5-6. Two leaf apices. 7. Basal cells.
8. Transverse section at midleaf. 9. Papillae. 10-13. G. bewsii. 10-11. Two leaves. 12. Leaf apex. 13. Transverse section at midleaf. 14-19. G.
viridulum. 14. Habit. 15-17. Three leaves. 18. Leaf areolation. 19. Propagula.
153
MERCEYOIDEAE
G. hymenostylioides (as Merceyopsis) at H, with the comment:
"Though the species lacks peristome teeth, it seems to be on the
same evolutionary (or reductive) line with Barbula indica. I think
it may be more natural to treat it [as] a member of the genus
Barbula."
Large specimens of Gymnostomum aeruginosum may also
have distinctly serrulate lower leaf margins, much as is the case in
Molendoa hornschuchiana, a large end-member of a stature gradient extending through M. sendtneriana, and the serrulate perichaetial leaves of some large specimens of G. aeruginosum and
those of G. bewsii are quite similar to those of M. sendtneriana.
Evidence of apparent phenocopy phenomena (including presence
of propagula in small forms of M. sendtneriana) between the two
genera at the small end of both of their stature gradients (Zander
1977c, p. 261) supports this relationship. Euc/adium, however,
also has denticulate lower !aminal margins and is not closely
related to Gymnostomum or Molendoa, and it is possible that this
characteristic is merely typical of hygrophilic species. Although
Molendoa when sterile can generally be distinguished from Gymnostomum by the percurrent or excurrent costa and more massive,
scab-like !aminal papillae, this is not always the case. Additional
discussion on the morphological similarities of Gymnostomum
and Molendoa has been given by Newton (1983), who favored a
more analytic approach to species distinctions. Cladistic analysis
gives differing results as to the relationship of Gymnostomum and
Molendoa, but see discussion of Cladograms 14-16.
Within Gymnostomum itself, there is a clear reduction series
based on leaf shape (ligulate to ovate) from G. aeruginosum (Pl.
46, f. 3-4) to G. mosis, with elaborations such as propagula in G.
virdulum (Pl. 46, f. 19, and rarely in G. aeruginosum), thickened
costae in G. hymenostylioides, and bistratose upper lamina! margins in G. bewsii (Pl. 46, f. 13-often present in G. aeruginosum
from the southeastern U.S.A.). Gymnostomum viridulum (Pl. 46,
f. 14-19; cf Sergio 1984, also Whitehouse & Crundwell 1991,
1992) is doubtfully different from G. mosis, while G.
hymenostylioides is very similar to both taxa in its leaf being widest at the middle but the latter has a longer leaf and very stout
costa. Although I have synonymized G. ca/careum with G.
aeruginosum (Zander 1977c), because these appear to intergrade
in the New World, the two are apparently distinct in Europe (see
e.g. Whitehouse & Crundwell 1991, 1992) and, for this treatment
to be of maximum service, both names are recognized here.
Khanna (1976) has pointed out that "G. recurvirostrum [=
Hymenostylium recurvirostrum] .. . , G. calcareum ... and G.
aeruginosum cannot be called clearly marked species because of
the presence of intergrading forms among them, at least in the
area under investigation [the.Himalayas]. However, they are morphologically distinct from each other and statistical distances
among them can be defined."
Eventually, certain of the names presently accepted as combinations in Gymnostomum will probably be assigned to other genera, such as Hymenostylium or Anoectangium. For instance, G.
chenii is surely the same as H. recurvirostrum var. cylindricum, if
one may judge from Saito's (1973c) detailed illustrations.
Additional literature: Andrews ( 1922b), Brown ( 1894a),
Crum & Anderson (1956), Crundwell (1981), Geheeb (1906a),
Khanna (1976), Nyholm and Hedenas (1986), Pierrot (1973,
1989), Stirling ( 1968).
Number of accepted species: 24.
Species examined: G. aeruginosum, G. bewsii (PRE), G.
ca/careum (BUF), G. hymenostylioides (BM, H), G. luisieri
(LISU), G. mosis (BUF, MO).
New combination: Gymnostomum hymenostylioides (Broth.
& Dix.) Zand., comb. nov. (Merceyopsis hymenostylioides
Broth. & Dix., J. Bot. 48: 302, 1910).
Plate47.
35. SCOPELOPHILA
Scope/ophila (Mitt.) Lindb., Acta Soc. Sci. Fenn. 10: 269, 1872.
Type: Scopelophila ligulata (Spruce) Spruce.
Merceya Schimp., Syn. Muse. ed. 2: 852, 1876. Type:
Merceya ligulata (Spruce) Schimp.
Merceyopsis Broth. & Dix. ex Dix., J. Bot. 48: 301, 1910.
Type: Merceyopsis pellucida Broth. & Dix., J. Bot. 48: 301,
1910, India, W. Ghats, Panchgani, Sedgwick 35, H,
/ectotyp. nov.
Weissia sect. Scopelophila Mitt., J. Linn. Soc. Bot. 12: 134,
1869.
Pottia sect. Orthotrichella C. Miill., Gen. Muse. Fr. 392,
1900. Type: Pottia gedeana Lac.
lookout place, headland, a crag, high rock + o
love, dear, beloved; referring to the characteristic
rocky habitat.
Plants forming a thin or thick turf, greenish yellow to brown
above, blackish green or weakly iridescent metallic tan or yellowish brown below. Stems seldom branching and then irregularly, to 4 em in length, transverse section rounded-pentagonal,
central strand absent, sc/erodermis absent, hyalodermis absent;
axillary hairs 3-5 cells in length, basal cell usually brownish;
sparsely radiculose or occasionally with a thick, reddish brown
tomentum. Leaves often crowded, incurved to spreading, contorted, usually carinate, occasionally with undulate upper margins when dry, spreading when moist, Ungulate to ligulate or
ob/anceo/ate, widest at or above mid/eaf, to 2.5 mm in length,
upper lamina narrowly grooved along costa or sometimes
broadly channeled, margins plane or somewhat recurved below,
entire to minutely crenulate or denticulate above, often bordered
by a few rows of thicker walled cells above; apex broadly acute
or obtuse, often with a broad apiculus, occasionally rounded;
base scarcely differentiated in shape to long-elliptical, occasionally wasp-waisted, occasionally slightly decurrent; costa slender, percurrent or ending up to 2-8 cells below the apex, occasionally excurrent as a short mucro, superficial cells quadrate to
rectangular ventrally, short-rectangular to elongate dorsally, 2-4
rows of cells across costa ventrally at midleaf, costal transverse
section semicircular to round, one stereid band present, generally well distinguished from the superficial parenchymatous layers, epidermis absent or present ventrally, present dorsally,
guide cells 2(-4) in 1 layer, hydroid strand absent; upper
/aminal cells rounded-quadrate to hexagonal or shortrectangular, ca. 8-14 jlm, often heterogeneous in size and
shape, 1(-2): 1, walls thin to evenly thickened or weakly
collenchymatous, thicker near margins, superficially flat or
somewhat bulging on ventral surface; papillae lacking, occasionally low-verrucose; basal cells differentiated across leaf,
extending higher medially, rectangular, occasionally inflated,
scarcely wider than the upper cells to inflated, 2-3:1, walls
hyaline or deep brown, bordered on margins by one or more
rows of narrow rectangular cells. Propagula rare, on stalks from
From
+
aK61tEAO~.
<p{A.o~.
154
GENERA OF THE POTTIACEAE
Plate 47. Scopelophila. 1-7. S. ligulata. 1. Habit. 2. Transverse section of stem. 3. Leaf. 4. Leaf apex. 5. Basal cells. 6. Transverse section at
midleaf. 7. Propagula. 8-12. S. cataractae. 8. Habit. 9-10. Two leaves. 11. Leaf apex. 12. Transverse section at midleaf.
MERCEYOIDEAE
155
Plate 48. Gymnostomiella. 1-4. G. vernicosa. 1-2. Two habits. 3. Leaf. 4. Propagula. S. G. longinervis. 5. Leaf. 6-11. G. monodii. 6. Habit. 7.
Transverse section of stem. 8. Leaf. 9-11. Transverse sections near midleaf. 12-16. G. orcuUii. 12-14. Habits of sterile and propaguliferous
plants. 15. Perigoniate plant. 16. Propagula.
156
GENERA OF THE POTTIACEAE
the stem, greenish brown, clavate to ellipsoidal or filamentous and
branching, ca. 12 J.l.m in diameter, of 2 or more rounded cells.
Dioicous. Perichaetia terminal, inner leaves little differentiated
from the cauline. Perigonia terminal, outer leaves loosely sheathing, inner deltoid. Seta 2-6 mm in length, 1(-2) per perichaetium,
brown to yellowish brown, twisted clockwise below; theca
0.6--2.2 mrn in length, brown to yellowish brown or blackish,
short-elliptical to short-cylindrical, exothecial cells quadratehexagonal to rectangular, occasionally bulging, 2-3:1, walls thin,
stomates present and commonly difficult to distinguish or often
apparently absent, phaneropore, at base of theca; annulus weakly
differentiated or of 1-4 rows of vesiculose cells, deciduous in
fragments; peristome absent. Operculum conic-rostrate, erect or
oblique, ca. 0.7-1.5 mm in length, cells in straight rows. Calyptra
cucullate, smooth, ca. 0.8 mm in length. Spores 8-13j.l.m in diameter, light brown, finely papillose or smooth. Lamina/ KOH color
reaction yellow to yellowish orange. Reported chromosome number n = 13.
Scopelophila is a small genus usually associated with highly
mineralized soils, found in mountainous areas of North, Central
and South America, Europe, central Africa, Asia and Oceania
(Hawaii).
Distinguishing characters of Scopelophila include the stem
section of nearly homogenous, wide-lumened cells (Pl. 47, f. 2),
lacking sclerodermis, hyalodermis, or central strand, the usually
spathulate, oblanceolate leaves often with a distinctive broad
apiculus (Pl. 4 7, f. 4 ), upper !aminal cells lacking papillae and
often heterogeneous in size and shape, costal section with a single
stereid band imbedded in parenchymatous tissue (Pl. 47, f. 6, 12),
and the capsule lacking a peristome. The leaves are often blackish
in the lower parts of the plant, and this may be associated with
iron ions in the substrate as the leaves of many mosses react with
a black coloration to ferric chloride solution.
There is an interesting body of literature (reviewed by Persson
1948, among others, and most recently by Shaw and Anderson
1988) on Scopelophila as a "copper moss," meaning that it is one
of a small number of taxa characteristically found growing in
association with copper, zinc and iron ores.
The genus Merceyopsis was created (Dixon 1910) with seven
species, apparently as an attempt to group certain eperistomate
species (except in the case of M. spathulifolia, the type of which
had only fragmentary sporophytes) that are seen here (see also
Noguchi 1956) as members of Barbu/a, Gymnostomum, Hymenostylium, Scopelophi/a, Trichostomum or Tuerckheimia (see
Merceyopsis in the list of recognized taxa for dispositions).
Dixon's (1910, p. 298) preliminary discussion of his treatment
indicates that the lectotype of Merceyopsis should be M. pellucida
Broth. & Dix. ex Dix. (= Scope/ophila cataractae (Mitt.) Broth.
cf Noguchi 1956; Zander 1967).
Additional literature: Arts (1988), Bartram (1924b), Corley
and Perry (1985), Crundwell (1986), Frahm (1990b), Hoe (1973),
Jones (1961), Lampton (1966), Lecointe and Schumacker (1988),
MArtensson and Berggren (1954), Melick (1975), Nagano and
Schimizu (1973), Nagano et al. (1969), Noguchi and Furuta
(1956), Noguchi and Ochi (1956), Reese (1989), Rumsey and
Newton (1989), Satake (1990, 1991), Satake et al. (1988, 1990),
Schatz (1955), Schumacker and Brugues (1991), Schumacker et
al. (1989), Shaw (1987a,b), Shaw and Anderson (1989), Shaw
and Beer (1989), Sotiaux et al. (1987), Takenaka and Satake
(1991), Thyssen and Poelt (1958), Zander (1967, 1986b).
Number of accepted species: 3 plus 1 remaining in
Merceya.
Species examined: S. cataractae (BUF, DUKE, FH, H, L,
MEX, TENN, US), S. infericola (BUF), S. ligu/ata.
New heterotypic synonymy: Desmatodon africanus P.
Yarde (PC)= Scope/ophila cataractae (Mitt.) Broth.
36. GYMNOSTOMIELLA
Plate 48.
Gymnostomiella Fleisch., Musci Fl. Buitenzorg 1: 309, 1904.
Type: Gymnostomiella vernicosa (Hook.) Fleisch.
Gymnomitriella Sak., Bot. Mag. Tokyo 56: 221, 1942, err.
pro. Gymnostomiella Fleisch.
Pottia sect. Splachnobryella C. Miill., Gen. Muse. Fr. 389,
1900. Type: Pottia vernicosa (Hook.) Hampe.
From Gymnostomum, a genus + i + -ella, diminutive; resembling the genus Gymnostomum.
Delicate plants in dense tufts or mats, light green to blackish green above, brown below. Stems short, often branching,
1.5-6.0 mm in length, transverse section rounded-pentagonal,
central strand present, comparatively large, sclerodermis
absent or weak, hyalodermis absent; axillary hairs ca. 3 cells in
length, moniliform, basal cell with thicker walls than the distal
cells; rhizoids present below. Leaves erect or laxly spreading
when dry, erect to spreading, lax when moist, obovate to
oblong-obovate, larger above, 0.3--0.4 mm in length, upper lamina flat, margins plane, crenulate above by bulging cell walls,
occasionally serrulate, occasionally with 1 or 2 additional large
teeth above; apex rounded or very broadly acute, occasionally
apiculate by a sharp, conical cell; base not differentiated in
shape; costa slender and ending at midleaf or ending near apex,
seldom percurrent, superficial cells on both sides elongate, ca. 2
rows of cells across costa ventrally at midleaf, costal transverse
section elliptical, stereid band single, small, present centrally,
often absent and costal section appearing homogeneous, epidermis absent ventrally and present dorsally, guide cells 2 in 1
layer, hydroid strand absent; upper lamina/ cells quadrate to
hexagonal, occasionally longer than broad, 12-18 J.l.m in width,
1(-2): 1, walls thin, delicate, superficially convex on both sides;
1-3 small, simple, hollow, conical papillae per lumen, scattered
over lamina; basal cells weakly differentiated across lamina,
rectangular, little wider than upper cells, 2-3:1, walls thin. Propagula elliptical to clavate, to 280 J.l.m in length, with ca. 6
transverse and 6 inner longitudinal walls, consisting of about 14
cells, borne in leaf axils or on ventral surface of leaves, sometimes 1-3 in a cup-like terminal rosette of leaves. Dioicous.
Perichaetia terminal, inner leaves ovate, to 0.7 mrn in length,
sheathing, smooth, lower cells laxly rectangular, paraphyses
moniliform. Perigonia terminal, paraphyses absent, plants often
small, nearly stemless. Seta 3-6 mm in length, 1 per
perichaetium, yellowish brown, twisted clockwise; theca
0.6--0.8 mm in length, black or brown, often shiny as if varnished, ovate to short-elliptical, exothecial cells large, roundedhexagonal, 35-55 J.l.m in diameter, mostly thin-walled, stomates
phaneropore, at base of theca but often absent, annulus of about
2 rows of little-differentiated cells; peristome teeth absent
Operculum relatively large, obliquely rostrate from a low-conic
base, 0.8-1.0 mrn in length, cells weakly twisted clockwise.
Calyptra cucullate, smooth, ca. 1.3 mrn in length. Spores 11-15
MERCEYOIDEAE
Jlm in diameter, light brown, finely papillose. Laminal KOH color
reaction yellow, or negative to black, or pink to deep purple.
Reported chromosome number n = 13.
This is a small genus of southern and eastern Asia, Australia,
northern and central Africa, subtropical and tropical North America, Central America, the West Indies and Brazil; found on limy
rock usually in association with cyanobacteria (as is the case with
Luisierella barbula).
In the absence of a peristome, this genus is placed in the Pottiaceae largely because of its papillose, obovate or spathulate
leaves (Pl. 48, f. 3, 5, 8). Although without obvious close relatives, it has some similarity to the genus Chenia and Hennediella
(Pottioideae) through the large hyaline laminal cells, simple papillae, and serrulate upper margins. Cladistic analysis indicates that a
better phylogenetic hypothesis would be placement in the Barbuleae. Unusual characters are the central location of the single
(sub)stereid band (Pl. 48, f. 9-11), the moniliform axillary hairs
(approached in shape by the weakly bulging cells of the hairs of
Molendoa species), the often nearly spherical capsule with unusually large exothecial cells and stomates often absent, and odd variation in KOH color reaction of the lamina, including black and
purple. The possible relationships of Gymnostomiella with the
Splachnobryaceae are discussed by Andrews (1949) and Crum
(1949) and in an overview of the Splachnobryaceae by A.
Koponen (1981). A convenient key to the species of Gymnostomiella was constructed from descriptions by Sloover (1977).
The propagula of G. vernicosa (Pl. 48, f. 4) are borne in leaf
axils and in clusters of 1-3 in a somewhat swollen cup-like terminal rosette of ovate leaves with the general appearance of a
perigonium. The propagula, with their clavate shape, internal
transverse and longitudinal cross walls, are somewhat similar to
antheridia. Thus, the propaguliferous sterile plants may easily be
mistaken for perigoniate plants. Also, Aeischer (1902-22, Vol. 1,
p. 31 0) correctly indicated that the perigonia of actual perigoniate
plants of G. vernicosa, which differ from the propaguliferous sterile plants by being nearly stemless, lack the paraphyses usually
expected in perigonia.
Of some significance is the relatively tiny size of the leaves in
respect to that of the stem, sporophyte and propagula, in conjunction with the leaves' simple anatomy and serrulate upper margins.
The leaves are apparently heterochronically paedomorphic (cf
discussion of Mishler 1986a of this phenonemon in Tortula) in
that they may not have developed much beyond the "scale leaf'
stage of very young shoots. It would be interesting to attempt to
"force" the mature stage by modifying developmental processes
(cf Basile & Basile 1984).
Redfearn (1991) recently synonymized the American G.
orcuttii (Pl. 48, f. 12-16) with the Asian G. vernicosa. These two
taxa are here conservatively retained as separate species as the
former often has multipapillose upper lamina! cells and the latter
unipapillose cells. Gymnostomiella orcuttii is, however, much like
the Asian G. burmensis, and the variation thus may not be geographical. In any case, a revision of all species of the genus is
sorely needed, and considerable synonymy might be expected, in
all probability supporting Redfearn's evaluation.
Additional literature: Eckel (l985b ), Potier de la Varde
(1953), Schomherst (1944), Seki and Miyagi (1980), Stone
(1985), Vital (1984).
Number of accepted species: 6.
Species examined: B. burmensis (FH), G. longinervis (NY),
157
G. monodii (PC), G. orcuttii, G. vernicosa (BM).
Plates 49-50.
37. DIDYMODON
Didymodon Hedw., Sp. Muse. 104, 1801. Lectotype: Didymodon rigidulus Hedw .,fide Grout, Moss Fl. N. Amer. 1: 186,
1939.
Pottia sect. Gomphoneuron C. Miill., Linnaea 42: 310, 1879.
Type: Pottia lorentzii C. Miill. (= Didymodon lorentzianus
(C. Miill.) Broth. fide van der Wijk et al., Ind. Muse. 4: 540,
1967).
Pottia sect. Senophyllaria C. Miill., Linnaea 42: 311, 1879.
Sect. Didymodon
Didymodum Hedw. ex P. Beauv., Mag. Enc. 5: 309, 1804,
nom. illeg. incl. gen . prior.
Didimodon P. Beauv., Mem. Soc. Linn. Paris (fasc. planch.):
3 f. 5, 1822, nom. illeg. orthogr. var.
Dydimodon Hedw. ex Amott, Mem. Soc. Linn. Paris 5: 263,
1827, nom . illeg. orthogr. var.
Trichostomum subg. Didymodon (Hedw.) Tum., Muse. Hib.
Spic. 34, 1804.
Didymodon subg. Didymodon (Hedw.) Boul., A. Crypt. Est
Muscin. 504, 1872.
Didymodon subg. Eudidymodon Kindb., Eur. N. Amer. Bryin.
2: 273, 1897, nom. illeg.
Barbula sect. Luridae Moenk., Laubm. Eur. 281, 1927.
Lectotype nov.: Didymodon rigidulus Hedw.
Barbula sect. Acutae Steere in Grout, Moss Fl. N. Amer. 1(3):
174, 1938.
Barbula sect. Didymodon (Hedw.) Giac., Atti !st. Bot. Univ.
Lab. Critt. Pavia ser. 5, 4: 208, 1947.
Barbula subsect. Acutiformes Kindb., Eur. N. Amer. Bryin. 2:
246, 1897. Type: Barbula acuta (Brid.) Brid.
Barbula subsect. Rigidulae Chen, Hedwigia 80: 193, 1941.
Sect. Asteriscium (C. Miill.) Zand., Cryptogamie, Bryol.
Lichenol. 2: 383, 1981 [1982]. Type: Barbula umbrosa C.
Miill.
Husnotiella Card., Rev. Bryol. 36: 71, 1909. Type: Husnotiella revoluta Card.
Trichostomopsis Card., Rev. Bryol. 36: 73, 1909. Type:
Trichostomopsis. crispifolia Card.
Asteriscium (C. Miill.) Hilp., Beih. Bot. Centralbl. 50(2): 618,
1933, hom. illeg. non Cbam. & Schlecht., 1826.
Barbula sect. Asteriscium C. Miill., Linnaea 42: 342, 1879.
Didymodon sect. Craspedophyllon Card., Rev. Bryol. 36: 81,
1909.
Sect. Fallaces (De Not.) Zand., Phytologia 44: 209, 1979. Type:
Barbulafallax Hedw.
Geheebia Schimp., Syn. ed. 2: 233, 1876. Type: Geheebia
cataractarum Schimp.
Dactylhymenium Card., Rev. Bryol. 36: 72, 1909. Type:
Dactylhymenium pringlei Card.
Limneria Stirt., Trans. Bot. Soc. Edinburgh 26: 428, 1915.
Type: Limneria viridula Stirt.
Prionidium Hilp., Beih. Bot. Centralbl. 50(2): 640, 1933.
Type: Prionidium setschwanicum (Broth.) Hilp.
Trichostomum subg. Zygotrichodon Schimp., Syn. ed. 2: 169,
1876. Type: Trichostomum tophaceum Brid.
Barbula subg. Geheebia (Schimp.) Szafr.;A. Po1ska Mchy l:
213, 1957 [1958].
158
GENERA OF THE POTTIACEAE
Tortula sect. Fallaces De Not., Mem. Roy. Ace. Sci. Torino 40:
287, 1838. Type: Tortulafallax (Hedw.) Tum.
Barbula sect. Graciles Milde, Bryol. Siles. 117, 1869.
Lectotype: Barbula rigidicaulis C. Miill. fide Saito, J. Hattori
Bot. Lab. 39: 601, 1975.
Barbula sect. Pseudodidymodon Kindb., Eur. N. Amer. Bryin.
2: 246, 1897, nom. illeg. incl. sect. prior.
Barbula sect. Rejlexae Monk., Laubm. Eur. 280, 1927, nom.
illeg. incl. sect. prior.
Barbula sect. Fallaces (De Not.) Steere in Grout, Moss Fl. N.
Amer. 1: 174, 1938.
Didymodon sect. Graciles (Milde) Saito, J. Hattori Bot. Lab. 39:
501, 1975, see Zander, Phytologia 41: 24, 1978.
Barbula subsect. Fallaciformes Kindb., Eur. N. Amer. Bryin. 2:
246, 1897. Type: Barbulafallax Hedw.
Barbula subsect. Rejlexae (Monk.) Chen, Hedwigia 80: 203,
1941, nom. illeg. incl. sect. prior.
Sect. Rufidulus (Chen) Zand. (a comb. nov. made below).
Barbula sect. Rufidula Chen, Hedwigia 80: 210, 1941.
Sect. Vineales (Steere) Zand., Phytologia 41: 24, 1978. Lectotype
nov.: Didymodon vinealis (Brid.) Zand.
Barbula sect. Vineales Steere in Grout, Moss Fl. N. Amer. 1:
174, 1938.
Barbula sect. Rubiginosae Steere in Grout, Moss Fl. N. Amer.
1: 174, 1938. Type: Barbula rubiginosa Mitt.
Barbula subsect. Vinealiformes Kindb., Eur. N. Amer. Bryin. 2:
246, 1897. Type: Barbula vinealis Brid.
From a!a'Cf.l.o~. double, twin + oaou~ (o&ov), oaov'to~. tooth;
the twin divisions of the peristome teeth.
Plants gregarious or more usually forming turfs or cushions,
light to blackish, olive or reddish green above, brown to reddish
brown or tan below. Stems branching seldom to often, to 2(-9) em
in length, transverse section rounded-pentagonal, occasionally
rounded-triangular, central strand usually distinct, seldom absent,
sclerodermis usually present, hyalodermis only occasionally
present; axillary hairs of ca. 5 cells, basal 1-2 cells brownish;
indumentum usually absent. Leaves often crowded, appressedincurved and occasionally twisted or curled when dry, spreading
when moist, ovate or more usually lanceolate or long-lanceolate
to occasionally long-triangular, ca. 0.8-3.0(-{).0) mm in length,
upper lamina usually broadly concave, occasionally narrowly
channeled or keeled, margins seldom plane or more usually
recurved or occasionally revolute, entire or occasionally weakly
dentate or crenulate, occasionally bistratose in patches or entirely
so; apex narrowly acute to rounded, seldom cucullate; base
weakly differentiated to ovate, occasionally oblong and halfsheathing the stem, occasionally decurrent, shoulders rarely
present; costa ending several cells below apex to excurrent as a
blunt awn, superficial cells quadrate to elongate ventrally, usually
short-rectangular dorsally above midleaf, smooth or occasionally
papillose, 2-4(-8) rows of cells across costa ventrally at midleaf,
costal transverse section ovate, semicircular or reniform, stereid
bands usually weak, occasionally absent ventrally, ventral epidermis present or seldom absent, dorsal present but usually weak,
guide cells 2-6(-8) in 1(-2) layers, hydroid strand seldom
present; upper !aminal cells subquadrate to hexagonal or rounded
angular, occasionally short-rectangular or rhomboidal, usually
8-13 Jlm in width, 1: l, occasionally bistratose in patches or
entirely, walls thin to thickened, lumens sometimes angular, occa-
sionally trigonous, superficially weakly to strongly convex on
both surfaces; papillae usually low, simple to bifid, usually
solid, not obscuring the lumens, occasionally absent or multiplex; basal cells usually weakly differentiated to occasionally
strongly differentiated across leaf or extending higher medially,
occasionally little different from upper cells, quadrate to rectangular, seldom bulging, usually little wider than the upper, ca.
2-4: l, walls usually rather thin, occasionally porose, smooth to
papillose. Propagula occasionally present, green, usually comparatively small, spherical to elliptical, of 1-10 cells, usually
borne in leaf axi/s, occasionally on ventral surface of costa or
on basal rhizoids; occasionally the leaf apex swollen and fragile. Dioicous (occasionally possibly rhizautoicous). Perichaetia
terminal, inner leaves ovate to long-lanceolate, occasionally
enlarged, not or occasionally sheathing in lower l/2, seldom
convolute-sheathing, lower cells rhomboidal-rectangular in
lower l/2. Perigonia terminal or occasionally as small buds on
protonema near archegoniophore. Seta mostly 0.5-2.0 em in
length, l(-2) per perichaetium, yellowish to reddish brown,
twisted clockwise below, occasionally counterclockwise above;
theca ca. l-3 mm in length, yellowish to reddish brown, elliptical to cylindrical, exothecial cells rectangular, thin-walled,
stomates phaneropore, at base of theca, annulus of l-3 rows of
hexagonal, often vesiculose cells, often deciduous in pieces or
revoluble; peristome teeth 16, or 32 and grouped in pairs, occasionally rudimentary or rarely absent, oblong to linear or longtriangular, often perforate or cleft medially, papillose to
spiculose or spirally striate, to 700(-1300) j.!m, often of many
articulations, usually straight or weakly twisted counterclockwise, but sometimes strongly twisted, basal membrane absent
or low, occasionally to 70 j.!m in height, papillose or spiculose.
Operculum short- to long-conic or conic-rostrate, ca. 0.5-1.2
mm in length, cells in straight rows or weakly twisted counterclockwise, occasionally to twice twisted. Calyptra cucullate,
smooth, ca. 2.0-2.5 mm in length. Spores ca. 7-15 j.!m in diameter, light brown, smooth to papillose. Lamina! KOH color reaction yellow or red. Reported chromosome number n = 12,
l2+m, 13, 14.
Found on a variety of substrates, mostly rock or soil; a cosmopolitan genus widely diversified in temperate and montane
regions.
Didymodon and Barbula, although often treated as one
genus, are here distinguished along the lines set out by Saito
(1975a) and summarized by Zander (1978e; see also treatment
of Barbula). In addition, Didymodon as presented here remains
an apparent potpourri of phyletic lines; Steere (1947) pointed
out that Didymodon "is a synthetic genus composed of discordant elements originating in several genera of the Pottiaceae,
agreeing among themselves only in their (supposedly!) [sic]
untwisted peristome teeth." Just as Barbula and Bryoerythrophyllum are closely related groups now separated as fairly
homogeneous genera by a variety of characters, chief among
them KOH color reactions, Didymodon, which is suspiciously
heterogeneous in KOH color reactions, may prove to be profitably split by recognizing various sections as genera. Obvious
candidates are described below.
Didymodon sect. Vineales
This section is characterized by the leaves spreading to widely
spreading and occasionally recurved when moist, concave
MERCEYOIDEAE
159
Plate 49. Didymodon. 1-14. D. rigidulus. I. Habit of sporophyte-bearing plant. 2. Perigoniate plant. 3. Transverse section of stem. 4--6. Three
leaves. 7. Leaf apex. 8-10. Three transverse sections at midleaf. 11. Leaf base. 12. Propagula. 13-14. Two peristomes. 15-22. D.
amblyophyllus. 15. Habit. 16-19. Four leaves. 20. Leaf apex. 21. Operculum. 22-23. Ca1yptra. 24-28. D. bartramii. 24-26. Three leaves. 27.
Leaf apex. 28. Transverse section at midleaf.
160
GENERA OF THE POTTIACEAE
across the leaf to keeled and narrowly channeled along the adaxial
surface of the costa, margins weakly decurrent to strongly so in
robust plants, weakly recurved below to recurved or revolute to
near the apex, often apiculate by a conical cell, the costa usually
percurrent to short-excurrent in a broad mucro, the upper !aminal
cells occasionally bistratose along the leaf margins, epapillose to
papillae simple or irregular to more often spiculose-multiplex,
1-4 over each lumen, the adaxial superficial cells of the costa
quadrate in the upper half of the leaf, the guide cells occasionally
in two layers, and the adaxial stereid band often absent (often
replaced by substereid cells); the peristome is absent or rudimentary to well developed and twisted up to 2.5 turns; spores released
usually in spring, also summer; KOH color reaction usually red to
red-orange (high magnification might be needed to ascertain the
exact hue of the internal upper !aminal cell walls). A "marker"
character, not always present but otherwise distinctive, is the
absence of the quadrate ventral costal cells at the extreme leaf
apex, resulting in a short, boat-shaped groove bottomed by
epapillose elongate cells. Some characteristic species of sect.
Vineales are D. brachyphyllus, D. cordatus, D. herzogii, D.
luehmanii (Pl. 50, f. 8-11}, D. luridus, D. nicholsonii, D.
occidentalis, D. reedii, D. sinuosus, D. tectorum and D. vinealis.
Didymodon sect. Fa/laces
This section is distinguished by the leaves spreading to often
strongly recurved when moist, concave to keeled, margins weakly
to strongly decurrent, plane to recurved in lower 2/3, not
apiculate, the costa ending below the apex to short-excurrent, the
upper !aminal cells unistratose, epapillose to papillae simple,
hemispherical or occasionally conic-apiculate, usually 1-2 over
each lumen, the adaxial superficial cells of the costa shortrectangular to elongate in the upper half of the leaf (except D.
asperifolius) and the adaxial stereid band usually present; peristome rudimentary to well developed and twisted up to 2 turns;
spores mature usually in winter or spring; KOH color reaction
usually reddish orange. The correct name at the generic level
would be Geheebia. This section is morphologically similar to
Hymenostylium by the commonly exposed ventral stereid band
and often angular upper medial cell lumens.
Although Sollman (1983) has synonymized D. constrictus
with D. (sect. Vineales) vinealis, the type of the former at NY
(" 170") is a mixed collection but largely composed of plants identical with other NY specimens ("Walanchoon" and "Lachen")
labeled in Mitten's hand as "Barbula constricta." These are not
D. vinealis but have clearly elongate ventral cells; the Asian D.
constrictus is near to or the same as the dark-red colored D. (sect.
Fa/laces) laevigatus of the Andes. Specimens identified in various herbaria as D. constrictus but with quadrate ventral costal
cells are usually D. (sect. Didymdon) rigidulus var. icmadophilus,
which has a similar very short leaf base filled with quadrate cells
and a very long-acuminate upper leaf.
.
Although Syed and Crundwell (1973) indicated that Didymodon maxima (as Barbula maxima) lacked a central strand, it usually has one, albeit weakly developed, even in European plants
(e.g. Ireland: Crundwell & Warburg 1962, NY). Trigones, similar
to those of Didymodon gigante~s. are present in the leaves of
most specimens. Hill (1981) and H. Crum and D. Hall (in press)
have pointed <?Ut that Didymodon ferrugineus (Schimp. ex Besch.)
Hill is the correct name for D. fa/lax var. reflexus at the species
level. Some characteristic species of sect. Fa/laces are D.
asperifolius, D. calycinus (Pl. 50, f. 1-3}, D. ceratodonteus (Pl.
50, f. 4-7), D. constrictus, D. fa/lax, D. ferrugineus, D.
erosodenticulatus, D. giganteus, D. hastatus, D. inundatus, D.
johansenii, D. laevigatus, D. maxima; D. michiganensis, D.
nigrescens, D. spadiceus (Pl. 50, f. 12-16), D. tomaculosus (differs from D.fallax only in the presence of rhizoidal propagula},
D. tophaceus and D. waymouthii (Pl. 50, f. 17-21 ).
Didymodon sect. Asteriscium
This section is characterized by the stem occasionally with a
hyalodermis, leaves spreading to spreading-recurved when
moist, occasionally squarrose from a shortly sheathing base,
broadly channeled, leaf margins not decurrent, plane to broadly
recurved throughout, the costa ending 1--6 cells below apex to
short-excurrent, often rather broad at midleaf, adaxial surface
convex, upper lamina! cells unistratose to bistratose evenly or in
patches along the leaf margins, papillae absent to large, low,
simple to bifid 1(-4) per cell lumen, adaxial superficial cells of
costa quadrate to elongate, adaxial stereid band absent or very
small, hydroid groups often present; KOH color reaction usually
yellow or yellowish orange. If recognized at the genus level, the
correct name would be Husnotiella or Trichostomopsis, both
published at the same time. This section represents at least
superficially a morphological intermediate, as noted by Hilpert
(1933), between Didymodon and Erythrophyllopsis in that the
upper lamina of D. challaense is bistratose in patches medially
or completely. Some characteristic species are D. australasiae,
D. bartramii (Pl. 49, f. 24-26), D. challaense, D. revolutus and
D. umbrosus. Probably also belonging here is D . marginatum.
Hydroid strands are of scattered occurrence in the genus Didymodon, having been seen in D. australasiae, D. ceratodonteus,
D. /uehmanii, D. revolutus and D. xanthocarpus. This feature is
evidently of little taxonomic utility at the present time.
Didymodon tophaceus, a relatively common species of wet
habitats, may lack a peristome in some specimens, which occasionally appear as taxonomic types of certain synonyms (e.g.
Gymnostomum knightii Schimp. in Knight, BM) originally
described as members of other, characteristically eperistomate
genera.
The type of Didymodon occidentalis (NY) lacks a peristome
entirely (it is not retained in the operculum on dehiscence). The
sect. Vineales is similar to Bryoerythrophyllum in red coloration
in KOH but is distinguishable from the latter by the usually
sharply acute leaves, smaller upper !aminal papillae, and often
more than one layer of guide cells in the costa, which also often
lacks a ventral stereid band.
Didymodon lindigii of the Andes has bistratose upper margins and apex, and rudimentary peristome, and is apparently
close to Didymodon rigidulus (Pl. 49, f. 1-14). The several
apparent isotypes at NY consist of considerable admixture with
D. tophaceus and D. australasiae, while the isotype at FH is
solely of the last two; lectotypification is sorely needed, with
attention to possible confusion with Astomum lindigii (cf. Mitten's 1859 treatment of this last species).
Weissia waymouthii (Pl. 50, f. 17-21) is here transferred to
Didymodon (sect. Fa/laces) reflecting its recurved lower leaf
margins, elongate ventral cells of costa, lack of a differentiated
ventral costal epidermis, upper !aminal cells pellucid, thickwalled, with rounded lumens and medially longitudinally
MERCEYOIDEAE
161
Plate SO. Didymodon. 1-3. D. calycinus. 1-2. Two leaves. 3. Leaf apex. 4-7. D. ceratodonteus. 4-5. Two leaves. 6. Leaf apex. 7. Transverse
section at midleaf. 8-11. D. luehmanii. 8-9. Two leaves. 10. Leaf apex, dorsal view. 11. Transverse section at midleaf. 12-16. D. spadiceus.
12-13. Two leaves. 14. Leaf apex. 15. Transverse section at midleaf. 16. Peristome. 17-21. D. waymouthii. 17-19. Three leaves. 20. Leaf apex.
21. Transverse section at midleaf. 22-26. D. xanthocarpus. 22-23. Two leaves. 24. Leaf apex. 25. Peristome. 26. Operculum.
162
GENERA OF THE POTTIACEAE
elongate, and papillae low and simple. The absence of a central
strand in D. waymouthii is unusual in the genus, and might indicate a relationship with Hymenostylium, but the species is retained
at least tentatively in Didymodon because of the presence of a
peristome and costa ending a few cells below the apex. The KOH
color reaction is reddish orange, not the characteristic yellow of
Weissia species.
Additional, selected literature: Abramova et al. (1987), Allen
(1992), Andrews (1941), Bartram (1926a,c), Conard (1945a,
1951a), Corley et al (1987), Crum (1965b, 1969a), Crundwell
(1976), Crundwell and Nyholm (1965), Crundwell and
Whitehouse (1978), Dismier (1905), Diill (1984), Diill-Hermanns
(1984), Diill-Hermanns and Diill (1985), Eckel (l986a), Ellis and
Smith (1983), Guerra and Ros (1987), Herzog (1905), Hill (1978),
Hillier (1931), Lal and Parihar (1980), Matteri (l988a), Mitten
(1867), Philibert (1885), Preston and Whitehouse (1985), Robinson (1968, 1970), Savicz-Ljubitzkaja (1965b), Sollman (1983),
Steere (1938b), Vashistha and Chopra (1984), Werner (1982),
Williams (1913), Zander (1978b,h, 1981c), Zander and Delgadillo
(1984).
Number of accepted species: 122, plus 1 in Husnotiel/a, and 1
in Trichostomopsis.
Species examined: D. a/ticaulis (DUKE, TENN), D.
amblyophyllus (NY), D. anserinocapitatus (NY), D. asperifolius,
D. austra/asiae, D. bartramii (F, US), D. brachyphyllus (BUF),
D. calycinus (BM, NY), D. cardotii, D. ceratodonteus (NY, PC),
D. challaense (H), D. constrictus (BUF, NY), D. cordatus (BUF,
F), D. crassicostatus (FH), D. deciduus (NY), D. fal/ax, D.
ferrugineus, D. giganteus, D. herzogii (JE, L), D. hampei (BM,
BUF, FH, TENN), D. hastatus (NY), D. humidus (NY), D.
imperfectus (H), D. incrassatolimbatus (BM, FH, NY, TENN), D.
inundatus (NY), D. japonicus (H), D. johansenii (ALA, MICH),
D. /aevigatus (BUF, NY), D.lamyanus (F), D. /eskeoides (ALTA,
BUF, NY, UBC), D. lindigii (NY), D. lingu/atus (NY, BM), D.
luehmanii (BUF), D. luridus, D. jackvancei (NY), D. marginatum
(NY), D. maxima (NY), D. michiganensis (DUKE, NY, TENN),
D. minusculus (NY), D. nicholsonii (BUF, HSC, UBC, US), D.
nigrescens (COLO, DUKE, FH, NY, US), D. occidentalis (NY),
D. patagonicus (NY), D. perobtusus (H, NY), D. pruinosus (NY,
BUF), D. reedii (US), D. revolutus, D. riEJ.idulus, D. rivicola
(BUF), D. rufidulus (BUF), D. sinuosus (BUF, NY), D.
spadiceus, D. stewartii (NY), D. subandreaeoides (ALA, CANM,
NY), D. subtorquatus (US), D. taylori (NY), D. tectorum (H), D.
tomaculosus (BUF), D. tophaceopsis (BUF, NY, US), D.
tophaceus, D. torquatus (HSC), D. umbrosus, D. uruguayensis
(US), D. vinea/is, D. waymouthii (NY), D. xanthocarpus (NY).
New heterotypic synonymy: Gyroweisia boliviano Williams =
Didymodon tophaceus (Brid.) Lisa. Husnotiella baueri Bartr. in
Bauer = Didymodon tophaceus (Brid.) Lisa. Gymnostomum
knightii Schimp. in Knight= Didymodon tophaceus (Brid.) Lisa.
New combinations and names:
Didymodon sect. Ruftdulus (Chen) Zand., comb. nov. (Barbula
sect. Ruftdula Chen, Hedwigia 80: 210, 1941).
Didymodon anserinocapitatus (X.-j. Li) Zand., comb. nov. (Barbula anserinocapitata X.-j. Li, Acta Bot. Yunnan. 3: 103,
1981).
Didymodon bartramii Zand., nom. nov. (Didymodon angustifolius
Bartr. non Wamst. nee Hen., Fieldiana Bot. 28: 4, 1951).
Didymodon cardotii (Dus.) Zand., comb. nov. (Barbula cardotii
Dus., Bot. Not. 1905: 299, 1905).
Didymodon challaense (Broth. in Herz.) Zand., comb. nov.
(Trichostomum chal/aense Broth. in Herz., Biblioth. Bot.
87: 30, 1916; Erythrophyllopsis challaensis (Broth. in
Herz.) Hilp.).
Didymodon crassicostatus (Bartr.) Zand., comb. nov. (Barbula
crassicostata Bartr., Bryologist49: 114, 1946).
Didymodon deciduus Zand., nom. nov. (Barbula decidua C.
Miill., Linnaea 43: 425, 1882, hom. illeg .; Barbula
uruguensis Par., nom. nov., Ind. Bryol. 101, 1894).
Didymodon herzogii Zand., nom. nov. (Trichostomum
ferrugineum Hen., Biblioth. Bot. 87: 31, 1916; Gertrudiella
ferruginea (Herz.) Hilp.; non Didymodon ferrugineus
(Schimp. ex Besch.) Hill).
Didymodon hampei Zand., nom . nov. (Trichostomum
obtusifolium Hampe, Bot. Zeit. 28: 4~. 1870; Gyroweisia
obtusifolia Broth.; non Didymodon obtusifolius Schkuhr).
Didymodon hastatus (Mitt.) Zand., comb. nov. (Barbula hastata
Mitt., J. Linn. Soc. Bot. Suppl. 1: 34, 1859).
Didymodon humidus (Mitt.) Zand., comb. nov. (Tortula humida
Mitt., J. Linn. Soc. Bot. 12: 162, 1869).
Didymodon imperfectus (C. Miill.) Zand., comb. nov. (Trichostomum imperfectum C. Miill., Linnaea 42: 214, 1879;
Barbula imperfecta (C. Miill.) Broth.).
Didymodon lindigii (Hampe) Zand., comb. nov. (Hyophi/a
lindigii Hampe Ann. Sc. Nat. Bot. ser. 5, 3: 343, 1865;
Barbula lindigii Hampe, Bot. Zeit. 27: 867, 1869).
Didymodon jackvancei Zand., nom. nov. (Husnotiella plicata
Magill, A. S. Afr. I. Bryophyta 1: 222, 1981; non Didymodon plicatus (C. Miill.) Mont.).
Didymodon marginatum (Robins.) Zand., comb. nov. (Trichostomum marginatum Robins., Phytologia 12: 389, 1971).
Didymodon minusculus (Williams) Zand., comb. nov. (Tortula
minuscula Williams, Bull. Torrey Bot. Cl. 42: 399, 1915).
Didymodon pruinosus (Mitt.) Zand., comb. nov. (Tortula
pruinosa Mitt., I. Linn. Soc. Bot. 12: 152, 1869; Barbula
pruinosa (Mitt.) Jaeg.).
Didymodon rigidulus var. ditrichoides (Broth.) Zand., comb. et
stat. nov. (Barbula ditrichoides Broth., Sitzungsber. Ak.
Wiss. Wien Math. Nat. Kl. 133: 566, 1924).
Didymodon stewartii (Bartr.) Zand., comb. nov. (Barbula
stewartii Bartr., Bull. Torr. Bot. Cl. 82: 23, 1955).
Didymodon taylorii Zand., nom. nov. (Tortula campylocarpa
Tayl., London I. Bot. 7: 187, 1848; Barbula campylocarpa
(Tayl.) C. Miill.; non Didymodon campylocarpus (C. Miill.)
Broth.).
Didymodon tophaceopsis Zand., nom. nov. (Gyroweisia latifolia
Dix., S. Afr. J. Sci. 18: 309, 1922; Husnotiella /atifolia
(Dix.) Zand. & Magill; non Didymodon /atifolius Wahlenb.
in Web. & Mohr).
Didymodon uruguayensis Zand., nom. nov. (Barbula
uruguayensis Broth. is an hom. illeg. of Barbula uruguensis
Par. fide Margadant in litt., itself a nom. nov. for Barbula
decidua C. Miill. a quite different species here recognised as
Didymodon deciduus (C. Miill.) Zand., Bib. K. Svensk. Vet.
Ak. Foerh. 36 afd. 3(7): 18, 1900).
Didymodon waymouthii (R. Br. ter) Zand., comb. nov. (Weissia
waymouthii R. Br. ter, Trans. New Zealand lnst. Bull. 31:
439, 1899; Weissia lancifolia var. waymouthii (R. Br. ter)
Wijk & Marg.).
Subfamily POTTIOIDEAE
Pottioideae (Limpr.) Broth., Nat. Pfl. 1(3): 381, 1902 "Pottieae."
Hyophilaceae Hampe, Linnaea 20:68, 1847, nom. nud., nom. rejic. Type: Hyophila Brid.
Astomaceae Schimp., Coroll. Bryol. Eur. 7, 1855 [1856]. Type: Astomum Hampe.
Phascaceae Schimp., Coroll. Bryol. Eur. 4, 1855 [1856]. Type: Phascum Hedw.
Weissiaceae Schimp., Coroll. Bryol. Eur. 7, 1855 [1856] "Weisiaceae." Type: Weissia Hedw.
Eupottiaceae Hampe, Flora 50: 67, 1867, nom. illeg.
Euweisiaceae Hampe, Flora 50: 67, 1867, nom. illeg.
Astomataceae Magill, Taxon 26: 597, 1977, orth. err. pro Astomaceae Schimp., 1855 [1856].
Tortuloideae Visotska, Citol. Genet. (Kiev) 1(4): 38, 1963, nom. inval. typ. non. cit.
This subclade is distinguished from subclades lower in the tree (Timmielloideae, Erythrophyllopsoideae, Gertrudielloideae, Chionolomoideae and Trichostomoideae) by the advanced characters of stem hyalodermis absent and perichaetial leaves sheathing (cf
Cladogram 15). The features of the immediate ancestral node are: stem sclerodermis not or little differentiated, leaf base little
differentiated in shape, and seta nearly absent to short, less than 1 em in length.
The Pottioideae is clearly discernible, both without including the Weissia group in Cladograms 8-10 and 14 and with the Weissia group as a basal part of the lineage in Cladograms 2, 3 and 4. Apparently the more highly evolved of the Pottioideae (those with
only one stereid band, !aminal margins mostly recurved, and upper !aminal cells mostly equally bulging on both surfaces of the lamina) were ultimately derived from ancestors of Leptobarbula and Weissia (with two stereid bands and lanceolate leaves) through the
loss of the ventral stereid band (beginning with the shared ancestor of Luisierella and Weisiopsis). Thus, from a side branch with
spathulate leaves and plane or incurved leaf margins, with ventrally bulging lamina! cells and mostly two costal stereid bands (e.g.
ancestors of Hyophila and its relatives), the somewhat reduced group of Stegonia, Crossidium, Pterygoneurum, Globulinella,
Aloina and Aloinella was derived.
The following salient traits have developed in the more highly evolved genera of this subclade: spathulate leaves, loss of the
ventral stereid band, and development of distinct infrageneric sporophyte reduction series; stem central strand and leaf hydroid
strand usually present; stem-borne stalked clavate propagula very rare; the leaf margins usually recurved below, plane or recurved
above; the upper lamina! cells generally rather large and equally convex on both superficial faces; elaborations of specialized
photosynthetic tissue not uncommon; and the basal membrane of the peristome occasionally very high.
The gametophytic characters of many species of this subfamily, especially of Tortula sect. Pottia, are strikingly like those of
species of Bryaceae. A separate derivation of these species from the Bryaceae is improbable because the twisted peristome would
have had to have been derived in separate families. Note, however, that a twisted peristome is present, albeit rarely, in the Ditrichaceae. An examination of various Bryum species shows they generally have a stout, little twisted seta. The seta of Ditrichaceae species, on the other hand is like that of the Pottiaceae in being relatively thin, generally twisted clockwise above and counterclockwise
below.
The presence of a small ventral stereid band in various species (e.g. Tortula rubra and Globulinella benoistii) of genera characteristically having only one stereid band (the dorsal) indicates that the traditional combination of characters for this subfamily was
never entirely consistent. This, plus (1) the inclusion of the many genera with two stereid bands in the Pottioideae (see Cladograms
14-16) at the base of the Pottieae subclade, and (2) certain single stereid-banded genera commonly are removed from the Pottieae
upon cladistic analysis (e.g., Calyptopogon placed near Tortella, and Gymnostomiella and Scopelophila near Didymodon in Cladogram 14) demonstrates that the number of stereid bands is as easily influenced by evolutionary pressures as many other characters
in the Pottiaceae. The traditional Pottioideae (these genera here treated as the more highly evolved members of the Pottieae and of
the Hyophileae) is a paraphyletic group.
Tribe HYOPHILEAE
Hyophileae Chen, Hedwigia 80: 179, 1941.
Pleuroweisieae Limpr., Laubm. Deutsch!. 1: 240, 1888 (rank not given, inoper. in priority).
Pleuroweisioideae (Limpr.) Broth., Nat. Pfl. ed. 2, 10: 243, 1924.
Pleuroweisieae (Limpr.) Chen, Hedwigia 80:41, 1941.
This tribe is distinguished at the immediate ancestral node by the features: leaves tubulose .when dry, leaf margins incurved or
involute, and upper lamina! cell walls ventrally bulging and dorsally nearly flat. Molendoa, which includes as a synonym
Pleuroweisia, the type of the tribe Pleuroweisieae, is included in this group. Although the Pleuroweisieae has been often recognized
164
GENERA OF THE POTTIACEAE
at the tribal level in the past (see review by Saito 1975a), the name Hyophileae (typified by Hyophila), which was conveniently published at the same time, is chosen here (I.C.B.N., Art. 57.2) as tribal name because Molendoa is placed with the Barbuleae in several
cladograms with various outgroups and must be considered of doubtful position. The distribution of the tribe Hyophileae is essentially worldwide, but with a concentration of significant taxa in tropical areas (cf Cladogram 14). Genera with single stereid bands
in their costa apparently have also lost the ventrally bulging and dorsally flat !aminal cell feature, though these genera may each
have one or more species with some degree of this one-sided bulging.
Plate 51.
38. HYPODONTIUM
Hypodontium C. Mtill., Hedwigia 38: 96, 1899. Type: Hypodontium dregei (Homsch.) C. Miill. (lectotype by Magilll981).
From U7to, under, beneath, less than + doou<; (oorov),
006V't0<;, tooth + -ium, characteristic; referring to the poorly
developed peristome teeth.
Plants robust, growing in a dense turf, light or dark green,
often glaucous above, light tan to blackish brown below. Stems
branching often, to 7 em in length, transverse section roundedpentagonal, central strand present, sclerodermis weak, hyalodermis present, often weak; axillary hairs long and narrow, ca. 5
cells in length, basal 1-2 cells thick-walled; radiculose or redtomentose. Leaves tubulose, incurved, often contorted when dry,
widely spreading from a sheathing base when moist, lanceo/ate,
3.5-6.0 mm in length, upper lamina broadly channeled, margins
plane to broadly incurved or involute, occasionally tubulose near
apex, entire above but sometimes denticulate along basal margins,
bordered by 1-5 rows of narrow, thick-walled cells to midleaf or
near apex; apex rounded-acute, occasionally broadly mucronate;
base sheathing in the lower 114 of leaf, with distinct shoulders;
costa percurrent or ending in a short, broad mucro, occasionally
ventrally strongly bulging, ventral superficial cells quadrate, dorsally elongate, papillose to spinose, 7-12 rows of cells across
costa ventrally at midleaf, costal transverse section ovate or semicircular, 2 stereid bands present, the ventral occasionally larger,
dorsally lunate, epidermis usually strongly differentiated ventrally, absent or weakly differentiated dorsally, guide cells 4-6 in 1
layer, hydroid strand absent; upper /aminal cells roundedquadrate, 8-11 j..Lm in width, 1: 1, walls evenly thickened, superficially ventrally bulging, dorsally weakly convex; papillae thick,
columnar, to 20 j..Lm in height, one per lumen on each side of the
lamina, apex of papilla low-spiculose, papillae larger medially
and ventrally on the leaf; basal cells differentiated across leaf, rising higher marginally in a vee, rectangular, 15-25 j..Lm in width,
2-4: 1, wider medially, walls hyaline, with distinct pores or these
grading into transverse slits. Dioicous. Perichaetia terminal, inner
leaves /inear-lanceolate, long-awned, fragile, (r-7 mm in length,
tubulose-sheathing in lower 1/2-2/3, lower cells rectangular and
hyaline except at base of awn. Perigonia consisting of flattened
axillary buds in clusters near apex of perigoniate plants similar in
stature to the perichaetiate. Seta 5-8 mm in length, 1 per
perichaetium, yellowish or reddish brown, twisted counterclockwise; theca fleshy, 1.7-2.0 mm in length, yellowish or reddish
brown, ovate to cylindrical, exothecial cells short-rectangular,
30-40 j..Lm in width, 1-3:1, very thick-walled, stomates
phaneropore, at base of capsule, annulus weakly differentiated, of
smaller, transversely elongated cells; peristome teeth 16, flat and
long-triangular or broadly lanceolate, smooth, 150-220 j..Lm, with
of 8-10 articulations, straight, strongly incurved, basal membrane
absent or very low, smooth; fragmentary remains of an exostome
remaining attached to the inside of the operculum or as scattered
transverse, narrowly elliptical plates attached externally to the
joints of the peristome teeth. Operculum long-rostrate, ca. 1 mm
in length, cells straight. Calyptra cucu/late, rough in upper 1/2
with forward-pointing papillae, 2.7-3.0 mm in length. Spores
large, ca. 30-40 j..Lm in diameter, brown, finely to very coarsely
papillose. Lamina! KOH color reaction yellow.
The genus is endemic to southern Africa, where it is found
on soil, rock or tree trunks.
Magill's (1981) treatment of the genus is detailed and up to
date. Reese et al. (1986), in an instructive review of the genera
of Calymperaceae, where Hypodontium has heretofore been put,
suggested this genus might be pottiaceous. Edwards (1980a)
accepted Hypodontium as Calyrnperaceae, and did not discuss it
extensively. Edwards (1979, p. 328) pointed out the fragmentary remains of 32 narrow exostome teeth that remain attached
to the inside of the operculum of Hypodontium and fall with it;
scattered, narrowly elliptical, transversely oriented plates
attached to the external faces of joints of the peristome were
seen in this study. An examination of a series of collections of
H. dregei (Pl. 51, f. 1-12) and H. pomiformis (Pl. 51, f. 13-16)
from PRE indicates that Hypodontium lacks many of the central
characters of genera of Calymperaceae, e.g. the campanulate
calyptra of Calymperes, a stem central strand, the intramarginal
bands of elongated cells, and the enlarged but short-rectangular
to quadrate basal cells of Ca/ymperes, Mitthyridium and
Syrrhopodon . The elongate, hyaline basal cells of Hypodontium
are similar to those of many of the genera of Pottiaceae. Characters of Hypodontium most characteristic of the Calymperaceae
are the peristome teeth (Pl. 51, f. 12) being short and incurved,
16, deep orange, triangular, smooth or weakly verrucose (compare those of Syrrhopodon, or even Octoblepharum cf Magill
1981 ); upper basal cells of at least the outer perichaetial leaves
sharply differentiated and inflated-quadrate like the cancellinae
of Calymperes; cauline leaves lanceolate, bordered by muchelongate cells with thick walls, apex broadly acute to rounded
and margins plane to incurved and often spinose, base with high
shoulders; the upper !aminal cells usually bulging ventrally and
weakly convex dorsally; and upper !aminal and dorsal costal
papillae very thick (Pl. 51, f. 7, 16); none of these characters is,
however, unique to the Calymperaceae. The peristome teeth, in
being red, flat and triangular, are quite like those of Oreoweisia
(Dicranaceae), a genus, which, however, has a rather dissimilar
gametophyte morphology.
Unusual in both the Pottiaceae and Calymperaceae are the
inner perichaetial leaves (Pl. 51, f. 9) sheathing below but narrowly subulate or awned apically (as in Bryobartramia of the
Encalyptaceae, or species of Diphyscium, Diphysciaceae, and
approached in the awned perichaetial leaves of some species of
Pseudocrossidium). Calymperes has a persistent calyptra, opening by slits, which is unique to Calymperaceae, while the
cucullate calyptra of Syrrhopodon often has a distinct collar on
the lower margin that opens late. The calyptra of Hypodontium
(Pl. 51, f. 11) is comparatively small, cucullate, and perched on
the rostrum of the operculum, and is more typical of Pottiaceae,
POITIOIDEAE
165
Plate 51. Hypodontium. 1-12. H. dregei. I. Habit. 2. Transverse section at midleaf. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Upper laminal papillae. 9. Perichaetialleaf. 10. Perigoniate bud. 11. Sporophyte with calyptra. 12. Vertical view of peristome teeth. 13-16. H. pomiformis. 13-14. Two leaves. 15. Leaf apex. 16. Transverse section at midleaf.
166
GENERA OF THE POTTIACEAE
although also found in Mitthyridium, Calymperaceae (Nowak
1980).
Except for the peristome, Hypodontium has the same capsule
morphology of fleshy appearance; large, thick-walled exothecial
cells; annulus largely undifferentiated; and spores relatively large,
as has Tridontium (here removed to the Grimmiaceae near
Scouleria) and Tetracoscinodon (Merceyoideae), the last with
similarly sheathing, distally narrowly subulate (but not quite
awned) perichaetial leaves. The peristome teeth of these three
genera are also somewhat alike, being broadly triangularlanceolate, while Hypodontium and Tetracoscinodon both have
smooth teeth, but the gametophytes are disparate in appearance. If
there were some distinctive autapomorphy or compelling combination of unusual characters, Hypodontium might be placed in a
family of its own.
Number of accepted species: 2.
Species examined: H. dregei (PRE), H. pomiformis (PRE).
39. HYMENOSTYLIELLA
Plate 52.
Hymenostyliella Bartr., Philippine J. Sci. 68: 108, 1939. Type:
Hymenostyliella involuta (Card. & Ther.) Bartr.
From Hymenostylium, a genus + i + -ella, diminutive; resembling the genus Hymenostylium.
Plants in cushions, yellow-green above, yellow-brown
below. Stems branching irregularly, to 3 em in length, transverse section rounded-pentagonal, central strand strong, sclerodermis present, hyalodermis absent; axillary hairs of 7-10 cells,
basal 1-2 yellowish brown; red-tomentose. Leaves involute,
Plate 52. Hymenostyliella. 1-8; H. 1/anosii. 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7.
Perichaetialleaf. 8. Perigoniate branchlet.
167
POTTIOIDEAE
incurved, curled when dry, spreading-recurved when moist, longlanceolate, widest just below midleaf, ca. 4.5 mm in length, upper
lamina broadly channeled across leaf, margins involute in upper
415, plane below, entire or more commonly distantly dentate in
upper 114; apex narrowly acute; base not differentiated in shape,
somewhat decurrent; costa short-excurrent as a mucro, occasionally flexuose or dentate, superficial cells ventrally quadrate to
short-rectangular, bulging, once prorulose at distal ends, dorsally
elongate, 4-5 rows of cells across costa ventrally at midleaf,
costal transverse section semicircular, 2 stereid bands present,
epidermis present ventrally, weakly developed dorsally, guide
cells 4 in 1 layer, hydroid strand absent; upper /aminal cells
rounded-hexagonal or somewhat longitudinally elongate, 10-13
jlm in width, 1(-2): 1, walls thick, trigonous, superficially strongly
bulging ventrally, flat dorsally; papillae absent; basal cells
weakly differentiated across leaf or somewhat higher along margins, rectangular, similar to upper cells in width, 2-4:1, walls
moderately thick-walled, somewhat porose. Dioicous. Perichaetia
on short lateral branchlets, inner leaves lanceolate, to 3.0 mm in
length, strongly sheathing in lower half, cells long-rhomboidal in
lower 4/5. Perigonia lateral, small, gemmate, occurring singly or
in clusters. Seta ca. 5-6 mm in length, 1 per perichaetium, reddish
brown, twisted clockwise; theca ca. 1.2-1.5 mm in length, reddish brown, obovoid to elliptical, exothecial cells rhomboidal,
thick-walled, stomates phaneropore, at base of theca, annulus of
2-3 rows of dark, weakly vesiculose cells; peristome absent.
Operculum long-rostrate, longer than the theca , to 1.5 mm in
length, cells in straight rows. Calyptra cucullate, smooth, ca. 1.7
mm in length. Spores ca. 10-13 jlm in diameter, light brown,
essentially smooth. Lamina! KOH color reaction yellow. [This
description is based on H. llanosii collections at PC and US. Robinson (1971a) recently transferred Timmie/la alata Herz. to this
genus (without examination of authentic material but probably
correctly), saying this species is closely related but is distinguished from H. llanosii by a dorsal costal surface winged with
two ridges up to 12 cells in height and a cucullate leaf apex.]
A rare taxon found on rock in wet areas in the Philippines,
India and Brazil.
This genus bears an immediate resemblance to Timmie/la by
the elongate, involute leaf and ventrally bulging-mamillose upper
!aminal cells (Pl. 52, f. 6), but as Chen (1941) pointed out, the latter genus has a peristome, as well as a bistratose upper lamina.
Unusual characters in Hymenostyliella are the strongly involute
leaf margins, ventrally bulging-mamillose and trigonous (Pl. 52, f.
4) upper !aminal cells, and sporophytes borne laterally on short
branches (Pl. 52, f. 1). Saito (l975a) noted the monopodia!
branching of Hymenostyliella in his discussion of H. japonica (as
a synonym of Didymodon japonicus). The long-lanceolate leaf
shape and distant teeth of this genus are reminiscent of Tuerckheimia, which differs by the distinctive massive !aminal papillae
and terminal perichaetium. Robinson (l971a) indicated a similarity to Hyophila (Cladogram 15 bears out a relationship) through
the ventrally bulging-mamillose upper lamina! cells. Hyophila
differs, however, in its spathulate leaves, generally has a hydroid
strand, and is acrocarpous; Ganguleea, with laterally borne
sporophytes is seemingly more closely related, but see Cladogram
15. The characteristic trigonous areolation is found to greater or
lesser degree in genera of other subfamilies, such as Hymenostylium, Leptodontium, Reimersia, Trichostomum and Tuerckheimia.
Number of accepted species: 3.
Species examined: H. llanosii (PC, US).
40. MOLENDOA
Plate 53.
Molendoa Lindh., Utkast Nat. Grupp. Eur. Bladmoss. 29, 1878.
Type: Molendoa hornschuchiana (Hook.) Lindh. ex Limpr.
Pleuroweisia Limpr. ex Schlieph., Flora 68: 359, 1885. Type:
Pleuroweisia schliephackei Limpr. ex Schlieph.
Ozobryum Smith Merrill, Novon 2: 255, 1992. Type: Molendoa ogalalensis (Smith Merrill) Zand.
Anoectangium subg. Molendoa (Lindh.) Kindb., Eur. N.
Amer. Bryin. 2: 317, 1897.
Named for Ludwig Molendo, 1833-1902, a German
muscologist.
Plants in a compact turf, dark to light green, occasionally
glaucous above, brown, often tan below. Stems branching seldom to often, to 4.0 em in length, transverse section roundedtriangular to pentagonal, central strand present, usually strong,
sclerodermis present, usually weak, hyalodermis absent or occasionally weakly developed; axillary hairs of 3-15 cells, usually
all hyaline or occasionally basal 1-2 cells thick-walled; light
brown to reddish tomentum sometimes present. Leaves usually
crowded, appressed incurved to weakly spreading, usually
twisted, occasionally tubulose when dry, spreading to
spreading-recurved when moist, variously oval, ligulate, longoblong, linear, ovate- to linear-lanceolate, (0.3-)1.0-4.0 mm in
length, upper lamina flat to broadly or occasionally narrowly
channeled, margins plane to weakly recurved in lower 112,
entire or occasionally sinuolate above, occasionally denticulate
at shoulders of the base (when dilated), upper margins often
bistratose entirely or in patches; apex broadly to narrowly
acute, occasionally broadly rounded and somewhat cucullate;
base scarcely differentiated to elliptical and sheathing; costa
ending 1-3(-6) cells below apex, percurrent, or excurrent as a
stout mucro, superficial cells quadrate to elongate on both sides,
2-7 rows of cells across costa ventrally at midleaf, costal transverse section circular, semicircular to flattened reniform, stereid
bands absent to weak ventrally, present dorsally and flattened
in section, epidermis present ventrally, usually present but weak
dorsally, guide cells 2-4(-7) in 1 layer, hydroid strand absent;
upper /aminal cells often irregular in shape, rounded-quadrate,
oval, rounded-triangular, ca. 8-10 Jlm in width, 1: 1, walls
evenly thickened to weakly trigonous, superficially flat to bulging; papillae usually crowded, low, irregularly scab/ike, occasionally simple to bifid or massively multifid; basal cells
differentiated across leaf or reaching higher along costa or margins, rectangular, little wider than the upper cells, (l-)2-5:1,
walls thin, evenly thickened to porose. Propagula rarely
present, obovoid to spindle-shaped, ca. 35-50 jlm long,
multicellular, borne in leaf axils. Dioicous. Perichaetia terminal
on short lateral branches, inner leaves ovate-lanceolate, often
marginally serrulate, to 3 mm in length, sheathing the seta,
lower cells rhomboidal in lower 1/2 to throughout. Perigonia
lateral. Seta 0.2-0.7 em in length, I per perichaetium, yellow to
brown, twisted clockwise below; theca 0.6-1.5 mm in length,
yellow-brown, ovoid to cylindrical, often macrostomous, exothecial cells thin-walled, stomates phaneropore, at base of theca,
annulus of 2-3 rows of transversely elongated, hexagonal cells,
weakly vesiculose; peristome teeth absent. Operculum very
168
GENERA OF THE POTTIACEAE
Plate 53. Molendoa. 1-9. M. duthei. I. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7-8. Perichaetial
leaves. 9. Sporophyte on short lateral branch. 10-15. M. platyphyllum. 10. Habit. 11-13. Three leaves. 14. Leaf apex. 15. Perichaetium. 16-18.
M. warburgii. 16-17. Two leaves. 18. Leaf apex. 19-22. M. seravschanica. 19-20. Two leaves. 21. Leaf apex. 22. Transverse section at
midleaf. 23-26. M. schliephackei. 23-25. Three leaves. 26. Leaf apex.
POTIIOIDEAE
long-rostrate, oblique, ca. 0.8-1.5 mm in length, cells straight.
Calyptra cucullate, smooth, 1.0-2.5 mm in length. Spores ca.
9-12 J.lm in diameter, brown, essentially smooth or weakly to
weakly papillose. Lamina/ KOH reaction light yellow. Reported
chromosome number n = 13.
This genus is found on all continents except Australia and
Antarctica, in montane or arctic areas, growing mainly on soil or
rock (often calcareous) in wet or seasonally moist places.
Diagnostic characters are few and somewhat variable. For this
reason species of Molendoa are often confused with Anoectangium species, which also are eperistomate and pleurocarpous,
and also often have triangular stem sections. Molendoa differs
from the latter principally by the presence of a ventral costal
stereid band (Pl. 53, f. 6, 22), which, unfortunately, is often absent
in Molendoa specimens of small size. Other characters that are
found in some but not all species of Molendoa (or which occur
only in well-developed populations of certain species), but which
are not found in Anoectangium, may be used to refer specimens to
the correct genus. These occasional indicator features of Molendoa include glaucous color of the upper leaves, becoming light
tan in lower parts of the plant; leaves linear-lanceolate or not
keeled; apex rounded, not apiculate; upper margins bistratose
(Anoectangium may be bistratose in transverse spaced rows
across upper leaf); leaf base much dilated and shoulders sometimes denticulate; superficial cells of the costa quadrate; costa
ending before the apex or very broad (up to 7 rows of cells across
costa ventrally at midleaf and up to 7 guide cells in the costa);
costal transverse section flattened reniform, or showing two
stereid bands; !aminal cells heterogeneous in shape (Pl. 53, f. 14),
or superficial walls (as seen in section) flat and distinctly thicker
than the contiguous, anticlinal walls (Pl. 53 6, 22); papillae low,
scab-like, crowded; KOH reaction pale yellow (versus yelloworange); theca macrostomous (Pl. 53, f. 9).
The leaf shape, leaf section, and propagula are similar to those
of Didymodon (sensu Saito l975a), but Molendoa differs from
that genus by the rather distinctive scab-like papillae (in most species) and, of course, the lateral gametoecia. Molendoa clavuligera
(not seen) is reported to have clavate leaf apices similar to those
of Didymodon johansenii. Molendoa is, on analysis, more closely
related to Didymodon than to Anoectangium in some cladograms
and is placed in the Hyophileae of the Pottioideae in Cladograms
14-16..
Pleuroweisia is here treated as a synonym of Molendoa (following the suggestion of Hilpert 1933) in that the key characters
of the former are also found in Molendoa . The recognition of the
genus Pleuroweisia by several modern authors (Brotherus
1924-25; Hilpert 1933; Chen 1941; Savicz-Ljubiskaja &
Smimova 1970) is probably due to acceptance of technical characters advanced by Limpricht (1890) that in fact have no substance. The calyptrae, alleged (Limpricht 1890; Chen 1941) to
cover only the rostrum of the operculum, are also found perched
on the rostrum in Molendoa sendtneriana in both microstomous
(e.g. Mexico: Muller 1834, MICH. type of Anoectangium
glaucescens) and macrostomous (e.g. Mexico: Le Sueur E6a, FH)
collections. The type specimen (leaves figured in Pl. 53, f. 23-26
from an isotype, Switzerland, Pontresina, Roseg-Gletschers, Graf,
9-7-1883, H) of Molendoa schliephackei (= Pleuroweisia
schliephackei) has calyptrae present only on immature capsules,
and, in fact, these partly cover the thecae as well as the opercula.
Other characters, such as the recurved leaf margins, rounded
169
apex, rather thick-walled laminal cells, indistinct stem central
strand, ovoid perichaetialleaves that sheath the seta, ovoid capsules, and somewhat vesiculose annular cells are also found in
various combinations in Molendoa, specifically the highly variable (Pilous 1958; Zander l977c) M. sendtneriana. The chromosome number of M. schliephackei has been reported (Fritsch
1972) to be the same as that of Molendoa sendtneriana, n = 13.
Characters that may prove of significance for M. schliephackei
at the species level are the strongly recurved !aminal margins
and the rather large leaf cells (l2-14J.lm in diameter), although
these are features seen so far only in the type specimen. Other
specimens examined that were identified as this species are
within the range of variation accepted (Zander 1977c) for M.
sendtneriana in the New World, and none match the type of the
species in all significant respects. Perhaps Geissler (1985) has a
better solution to the problem of variation among these taxa; she
viewed M. sendtneriana, M. schliephackei, M. tenuinerve and
M. taeniatifolia as synonyms of M. hornschuchiana (as Anoectangium hornschuchianum).
Previously placed in Anoectangium, Molendoa platyphyllum
(Pl. 53, f. 10-15) of the Peruvian Andes and M. warburgii (Pl.
53, f. 16-18) of the United Kingdom are similar in the small
plant size, much-branched stems, leaf shape (ovate-triangular
with a constricted apex) and costa thin (the former has two
stereid bands, but none have yet been detected in the latter). The
two differ in papillae shape (the former has low, scattered scabs,
while the latter has massive simple or less thickened bifid papillae), but these two species are evidently closely related. Long
(1982c) found propagula in the latter species, and he rightly
suggested these indicate a closer relationship to Molendoa than
to Anoectangium. The mid-North American M . ogalalensis,
type of Ozobryum, is similar to these but differs in the lack of a
stem sclerodermis, the broader leaf apex, single massive papilla
over each bulging lumen, and superficial walls of upper laminal
cells not distinctly thicker than the internal walls; this species is
distinctive but, in my opinion, not at the generic level. Molendoa ogalalensis is similar in stem section, leaf shape and papillae morphology to Quaesticula navicularis, which differs, however, in the incurved upper leaf margins and terminal
perichaetium.
The Bryotheca E. Levier specimen of Molendoa roylei at
NY is Hymenostylium recurvirostrum and other distributed
duplicates may also be this species. Hymenostylium characteristically (but not always) lacks a central strand. This and the rectangular, often irregularly thickened median leaf cells are good
distinguishing characters for sterile specimens.
Additional literature: Castelli (1966, 1968), Gyorffy (1910,
1912, 1914, 1946), Herzog (1943), Iwatsuki & Sharp
(1958-this paper actually refers to Tuerckheimia svihlae
(Bartr.) Zand. as noted by Saito 1972b and Zander 1979b, both
as Tuerckheimia angustifolia).
Number of accepted species: 15.
Species examined: M. duthei (NY), M. hornschuchiana, M.
ogalalensis (BUF), M. platyphyllum (F, NY), M. schliephackei
(BP, H, NY, PC), M. seravschanica (H), M. sendtneriana, M.
warburgii (BUF).
New heterotypic synonymy: Hymenostylium secundum C.
Mtill. = Molendoa sendtneriana (BSG) Limpr. Hymenostylium
validinerve Dix. & P. Yarde = Molendoa sendtneriana (BSG)
Limpr.
170
GENERA OF THE POTTIACEAE
New combinations: Molendoa ogalalensis (Smith Merrill)
Zand., comb. nov. (Ozobryum ogalalense Smith Merrill, Novon 2:
255, 1992). Molendoa platyphyllum (Williams) Zand., comb. nov.
(Anoectangium platyphyllum Williams, Field Mus. Nat. Hist.
4(5): 130, 1927. Molendoa schliephackei (Limpr. ex Schlieph.)
Zand., comb. nov. (Pleuroweisia schliephackei Limpr. ex
Schlieph., Flora 68: 359, 1885 [Pleuroweissia schliephackei
Limpr., Jahresber. Schles. Ges. Vaterl. Cult. 61: 224, 1884, nom.
inval. sin. descr.]). Molendoa warburgii (Crundw. & Hill) Zand.,
comb. nov. (Anoectangium warburgii Crundw. & Hill, J. Bryol. 9:
435,1977 [1978]).
Plates 54-56.
41. HYOPHILA
Hyophila Brid., Bryol. Univ. l(Suppl.): 760, 1827. Lectotype:
Hyophila javanica (Nees & Blume) Brid. fide Hampe in Bot.
Zeit. 4: 266, 1846.
Rottleria Brid., Bryol. Univ. 1: 105, 1826, hom. illeg. non
Wild., 1797.
Hygrophila Syd., Bot. Jahresber. 39(1): 99, 1912, nom. inval.
error pro Hyophila Brid.
Hyophila subg. Gymnohyophila Card. in Grand., Hist. Madag.
39: 208, 1915, nom. illeg. incl. type. gen.
Gymnostomum sect. Hyophila (Brid.) Reichenb., Consp. Regn.
Veg. 1: 33, 1828.
Pottia sect. Hyophila (Brid.) C. Miill., Syn. 1: 558, 1849.
Weissia sect. Hyophila (Brid.) Mitt., J. Linn. Soc. Bot. 12: 135,
1869.
From 00>, to wet, rain + <pO..o~. love, dear, beloved; referring to a
preference for wet habitats.
Plants turf-forming or occasionally loosely caespitose, green
above, red to reddish brown or dark green below. Stems branching
occasionally, to 1.0 em in length, transverse section roundedpentagonal or triangular, central strand usually strong, occasionally absent or central portion of stem hollow, central cylinder
often of rather thick-walled cells grading into a sc/erodermis,
sclerodermis usually present, often strong, in several layers,
hyalodermis absent or present (often only weakly differentiated);
axillary hairs 6-10 cells in length, these all hyaline; radiculose.
Leaves often rosulate, tubulose-twisted, incurved and occasionally contorted when dry, spreading, occasionally rather fragile
when moist, commonly spathulate or ligulate, often ovate, oblong
or elliptical, usually narrowed to the base, to 2.5 mm in length,
upper lamina broadly channeled, shallowly grooved along costa,
occasionally concave, margins plane to broadly incurved, occasionally narrowly recurved in lower 2/3, entire or denticulate to
dentate in upper 1/4 or just at apex, apex broadly acute to
rounded, rarely cucullate or emarginate; base not differentiated in
shape or half-sheathing, occasionally weakly auricled; costa
subpercurrent or percurrent, ending in an apiculus or occasionally
a mucro, superficial cells ventrally quadrate or more commonly
short-rectangular, often bulging, dorsally elongate, 2-6 rows of
cells across costa ventrally at midleaf, costal transverse section
semicircular, 2 stereid bands present, ventral and dorsal epidermis
present, guide cells 4(-6) in 1 layer, hydroid strand occasionally
present; upper /aminal cells rounded-quadrate or hexagonal, usually small, 6-13 J.lm in width, 1:1, walls thin to evenly thickened,
either superficially ventrally strongly bulging and dorsally weakly
convex, or bulging equally on both sides; papillae absent or sim-
pie, solid, often only on dorsal surface of lamina, occasionally
weakly bifid; basal cells differentiated across leaf or medially,
usually restricted to small area near insertion, shortrectangular, 10-20 J.lm in width, 1-4:1, walls thin to evenly
thickened. Propagula often present, clavate, stellate or dentateelliptical, often to 300 )lm in length, borne on stout, branching
stalks in leaf axils. Dioicous or monoicous (autoicous,
paroicous). Perichaetia terminal and inner leaves little different
from or smaller than the cauline, half-sheathing, lower cells
thin-walled and rhomboidal in lower half. Perigonia as lateral
buds on perichaetiate plants or terminal on perigoniate plants,
inner leaves little differentiated. Seta 0.4-0. 7 em in length,
1(-2) per perichaetium, reddish to yellowish brown, twisted
clockwise; theca 1.0-2.3 mm in length, reddish to yellowish
brown, long-ovoid to cylindrical, exothecial cells quadrate to
rectangular, 20-45 )lm in width, 1-5:1, walls thin or much
thickened both on exposed and contiguous sides, stomates
phaneropore, at base of theca, annulus 1-3 rows of vesiculose
cells, deciduous in pieces or persistent on theca or operculum;
peristome teeth absent. OpercuJum conic to long-conic or rostrate, 0.5-0.8 mm in length, cells not twisted. Calyptra cucullate,
often twisted about the seta when mature, smooth, 2-3 mm in
length. Spores 7-24 J.lm in diameter, light brown, papillose.
Lamina/ KOH color reaction yellow, occasionally with a red
blush medially above midleaf, occasionally cells near leaf insertion red. Reported chromosome number n =7, 13, 13+m.
Found on rock, soil and trees, generally in moist or wet
areas, throughout the tropic and temperate zones.
Abundant synonymy and combinations in other genera will
certainly be necessary on revision for presently accepted correct
names in Hyophila. Recent synonymy by various authors
includes several broad-leaved sterile or eperistomate taxa in
Hyophila transferred to or synonymized with species in such
genera as Barbula, Didymodon, Bryoerythrophyllum, Gymnostomum, Gyroweisia, Plaubelia, Scopelophila, Tortula, Trichostomum and Weissia. Many correct names in Hyophila provisionally accepted during this study are based on type or authentic specimens expected, on revision, to prove to be H. involuta
(Pl. 54, f. 1-11); this species requires comprehensive evaluation, however, and extensive synonymy within the genus was
not attempted. The species illustrated here are mostly those
fairly different from H. involuta.
Many species of the genus are morphologically similar to
Trichostomum; these seem to intergrade with Trichostomum
species through such similar, intermediate taxa as Plaubelia,
Hyophila nymaniana (Pl. 55, f. 18-25) and T. planifolium.
Trichostomum itself certainly shows tendencies towards the
Hyophila morphotype (e.g. Magill 1981, p. 262 refers Hyophila
zeyheri of South Africa to the synonymy of Trichostomum
brachydontium because of peristome variation from rudimentary to absent, see also Sergio 1985). The upper laminal cells
ventrally bulging and dorsally flattened are developed only
medially in H. bartramiana and H . subcucul/ata (Pl. 56, f.
6-10), two closely related monoicous species that also have
!aminal papillae. Ventrally bulging upper lamina! cells (usually
also ornamented with papillae) are also found in many species
of Weissia, but Weissia is apparently not closely related to
Trichostomum (see Cladogram 14 and others) on analysis using
the full character set, although it is related to Hyophila. Hyophila appears to be a weakly segregated end point of a parallel
POITIOIDEAE
171
Plate 54. Hyophila. 1-11. H. involuta. I. Habit of sporangiate plant. 2. Perigoniate plant. 3. Transverse section of stem. 4. Two leaves. 5. Leaf
apex. 6. Basal cells. 7. Transverse section at midleaf. 8. Propagula. 9. Thecal mouth. 10. Portion of transverse section of thecal wall. 11.
Calyptra. twisted about operculum. 12-17. H. acutifolia. 12-14. Three leaves. 15. Leaf apex. 16. Transverse section at midleaf. 17. Thecal
mouth.
172
GENERA OF THE POTTIACEAE
Plate 55. Hyophihl. 1-5. H. apicuhlta. 1-3. Three leaves. 4. Leaf apex. 5. Portion of thecal wall in section. 6-11. H . beruensis. 6. Section of
stem. 7-9. Three leaves. 10. Leaf apex. 11. Section at midleaf. 12-17. H. propagulifera. 12. Section of stem. 13-14. Two leaves. 15. Leaf apex.
16. Section at midleaf. 17. Propagula.18-25. H. nymaniana. 18. Section of stem. 19-20. Two leaves. 21. Leaf apex. 22. Section at midleaf. 23.
Propagula. 24. Mouth of theca. 25. Section of theca.
POTTIOIDEAE
line of evolution towards Trichostomum species (sect.
Laminanchium) with ventrally bulging upper !aminal cells, or else
the latter is better seen as a separate taxon at the genus level in the
Hyophileae or even transferred to Hyophila.
Some species of Hyophila, including the generitype, H.
involuta, have certain of the characters of Barbu/a (Norris and
Koponen 1989 felt that Hyophila is closely related to Barbula)
including recurved lower lamina! margins, medially differentiated
basal lamina! cells, ventrally colliculate but dorsally smooth upper
lamina (e.g. B. javanica) and armed, often caltrop-shaped propagula (as in Barbula indica and Barbula sect. Hydrogonium).
Also, the capsules of some species (e.g., H. involuta and H.
nymaniana) are large, dark brown and thick-walled, reminiscent
of capsules of Barbula or Tortula, while others (e.g., H.
acutifolia) are short, yellowish and thin-walled, like those of
Trichostomum or Weissia; intermediates are few. These features
indicate that Hyophila as presently recognized is polyphyletic. Of
some significance is that H . siamensis (Pl. 56, f. 1-5) could be
placed with Barbula by its narrowly recurved marginal !aminal
cells and other characters, except that the spathulate leaf shape
and lack of a peristome at the present time militate against it; the
same is true for H . nymaniana, which differs widely from other
Hyophila species in the presence of punctate papillae (curiously
like those of certain species of Rhachitheciaceae) on the upper
!aminal cells, which bulge on both sides of the leaf, the absence of
a stem central strand, and the bright red KOH color reaction of the
basal lamina! cells at the leaf insertion (the last two characters are
173
unusual but duplicated in Leptodontium viticulosoides and Tortuta cuneifolia var. blissii). On the other hand, H . apiculata (Pl.
55, f. 1-5) appears to be a phenocopy of B. agraria in the shortovate leaf shape, acute apex and sharply excurrent costa. Additional study is needed for satisfactory disposition of these Hyophila species.
Distinctive characters often but not always found in Hyophila species in various combinations and which must serve to
distinguish Hyophila from Trichostomum and Barbula include: ·
leaves ligulate, narrowed to the insertion, broadly concave in
transverse section; upper !aminal cell surfaces epapillose, ventrally bulging and dorsally weakly convex (Pl. 54, f. 7); basal
!aminal cells short and poorly differentiated or differentiated
cells restricted to a small area near the insertion; propagula
present, armed (Pl. 54, f. 8; 55, f. 23); monoicous; capsule
eperistomate, and exothecial cells thick-walled (Pl. 55, f. 25).
No species of Hyophila has all these characters and few have
most. Hyophila differs from Plaubelia mainly in being eperistomate and the ventral surface of the costa usually consisting
of elongate cells. The leaf hydroid strand is also often absent in
Hyophila , but these characters may ultimately prove insufficient
to separate the two taxa at the generic level. Two species previously placed in Hyophila probably because of their lingulate
leaves and eperistomate capsules are here referred to Tisserantiella of the Rhachitheciaceae (one as a synonym, the second as a
comb. nov., see Excluded Taxa) because their costa mostly end
below the leaf apex and the upper lamina! papillae are
Plate 56. Hyophila. 1-5. H. siamensis. 1-2. Two leaves. 3. Leaf apex. 4. Basal cells. 5. Transverse section at midleaf. 6-10. H. subcucu/lata.
6-8. Three leaves. 9. Leaf apex, lateral view. 10. Transverse section at midleaf.
174
GENERA OF THE POTTIACEAE
extremely small, solid, punctifonn to short-spiculose, with the
appearance of tiny specks of light under the compound lens (these
characters are unique only in combination).
Additional literature: Andrews and Redfearn (1965), Deguchi
et al. (1991), Gao et al. (1991), Long (1978), Mehra (1984, 1988),
Nawawi and Mohamed (1989), Olarinmoye (1981), Rahbar and
Chopra (1980), Sakurai (1954), Sharma and Chopra (1986), Sharp
(1955), Smith and Whitehouse (1974), Yang (1965).
Number of accepted species: 88.
Species examined: H. acutifolia (TNS), H. apiculata (FH), H.
bartramiana (BUF, MICH, TENN), H. beruensis (H), H. blanda
(NY), H. grossidens (H), H. involuta, H. /atifolia (H), H.
mattogrossensis (H), H. nymaniana (BUF, NY, TENN), H.
potierii (FH), H. propagulifera (H, NY, TNS), H. siamensis
(BM), H. spathulata (NY), H. subcucullata (NY), H. subjlaccida
(NY), H. usambarica (H), H. viridula (BUF).
New heterotypic synonymy: Gymnostomum denticulatum
Griff. (NY)= Hyophila involuta (Hook.) Jaeg.
42. PLAUBELIA
Plate 57.
P/aubelia Brid., Bryol. Univ. 1: 522, 1826 (nom. rejic. vs. Trichostomum Bruch, 1829). Type: P/aubelia tortuosa Brid.
Hyophilopsis Crum, Bryologist 68: 69, 1965, non Card. & Dix.
1911, hom. il/eg.
Neohyophila Crum, Bryologist 68: 470, 1965, nom. nov. for
Hyophilopsis Crum, hom. il/eg. Type: Neohyophila sprengelii
(Schwaegr.) Crum.
Tortula sect. Plaubelia (Brid.) Mitt., J. Linn. Soc. Bot. 143,
154, 1869.
Named for a mycologist surnamed Plaubel; a specialist of the
1820's in Puccinia Pers. and other Uredinales. Major mycological
references provide no further infonnation on this person.
Plants fonning turf or loosely caespitose, green above, green
or sometimes brown below. Stems often branching, to 4 mm in
length, in transverse section rounded-pentagonal, central strand
strong, sclerodennis present, often weak, hyalodennis absent;
axillary hairs of up to 5 uniseriate cells, the basal 1-2 yellow;
rhizoids sparse. Leaves incurved to spreading, often tubulose
when dry, widely spreading when moist, usually rosulate,
spathulate to oblong, to 3.2 mm in length, ventral surface flat to
broadly concave across leaf; margins incurved, involute, or occasionally plane, sometimes broadly involute at the apex, entire to
distantly denticulate above; apex rounded acute to broadly
obtuse, apiculate or occasionally entire; base little differentiated
in shape; costa percurrent or ending up to 4 cells below the apex,
adaxial surface cells bulging, rounded-hexagonal, ca. 4 rows of
cells across ventral surface of costa at midleaf, dorsal cells elongate, costa in transverse section rounded, showing one or two
stereid bands, the dorsal semicircular in section except for a
ventral indentation at the hydroid strand, ventral epidennis
differentiated, 2-4 guide cells in 1 layer, dorsal epidennis slightly
differentiated, hydroid strand present (this occasionally difficult
to demonstrate in small plants); upper lamina/ cells roundedhexagonal, 8-10 J..lm in width, I: I, walls evenly thickened, ventrally strongly convex and dorsally nearly flat; upper lamina! papillae often absent, when present solid, small, simple, 1-2 per lumen
dorsally or occasionally present on both sides of lamina; basal
cells not differentiated or often fonning a small group medially or
weakly differentiated across the leaf, quadrate to shortrectangular, to 15 J..lm wide, 2-3:1, hyaline to somewhat yellowish; 1-2 rows of inflated, hyaline cells across insertion sometimes also present, occasionally fonning small auricles. Propagula in leaf axils, clavate, often multi-branched at the wider
end, mostly 100-300 J..lm in length, with occasional internal
walls. Dioicous. Perichaetia terminal, inner leaves ovatelanceolate to ligulate, slightly larger or somewhat shorter than
outer leaves, weakly or strongly sheathing, cells longrhomboidal or rectangular in lower half. Perigonia tenninal,
inner leaves ovate, outer leaves large. Seta to 0.8 em in length,
yellow- to red-brown, twisted clockwise below, often counterclockwise above, 1 per perichaetium; theca ca. 1.0-1.5 mm in
length, red- or yellow-brown, ellipsoidal or oblong, exothecial
cells somewhat bulging, rectangular, ca. 15-35 J..lm in width,
2-5: 1, walls thin to weakly porose and thickened, stomates
phaneropore, present at base of capsule above a comparatively
well developed neck; annulus of strongly vesiculose cells, persistent on the capsule mouth, detaching in pieces or revoluble;
peristome of 16 spiculose, lanceolate to long-linear teeth, cleft
to near base or variously cleft or perforate, often only perforate
at base and entire, to 180 J..lm in length, up to 9 articulations,
straight, low-spiculose, basal membrane absent or to 25 J..lm in
height, low-spiculose. Operculum rostrate, to 0.9 mm in length,
cells in straight rows. Calyptra cucullate, smooth, 0.~2.0 mm
in length. Spores yellow, weakly papillose, small, ca. 8-10 J..lm
in diameter. Lamina/ KOH color reaction yellow in upper
leaves, often orange-brown in lower.
Found on rock, soil, especially calcareous substrates, in
extreme southwestern and southeastern U.S.A., Mexico, Central
America, the West Indies, Venezuela, Brazil, South Africa and
Bunna.
Bride! (182~27) established Plaubelia to include the single
species Plaubelia tortuosa Brid. (= Desmatodon sprengelii
(Schwaegr.) Williams, a taxonomic synonym fide Williams
1919). The name Plaubelia fell from use after being rejected
against the later (1829) generic name Trichostomum Bruch; M.
Crosby (in litt.) noted that Plaubelia is acceptable when it is not
a taxonomic synonym of Trichostomum Bruch. Crum (1965a)
created Hyophilopsis (and later the nomen novum Neohyophila,
a heterotypic synonymn of Plaubelia) as a segregate of Desmatodon, citing, as a unique combination of features, the
spathulate, rosulate leaves (Pl. 57, f. 1, 9, 10) with erect or
incurved margins, !aminal cells ventrally bulging, and the
ventral epidermis of bulging cells (Pl. 57, f. 5). He further
referred the genus to the Pottioideae, citing a single stereid
band, but I agree with Delgadillo and Cardenas (1982) that this
character is variable, as one or two bands may be present in different leaves of the same plant. Actually Plaubelia is apparently
most closely related to Hyophila, from which it differs most
saliently in the presence of a peristome (Pl. 57, f. 8, 18). The
strongly involute margins are reminiscent of Weissia. The presence of a hydroid strand (Pl. 57, f. 7, 16) and of roundedquadrate ventral epidennal cells (as seen from above) on the
costa are also good characters distinguishing Plaubelia from the
similar and very commonly distributed Hyophila involuta, as
well as several other Hyophila species but not all (Hyophila
greatly needs revision). Globulinella is very similar in many
respects, but is distinguishable by the costa not bulging dorsally
and the usually cucullate leaf apex. Crum and Anderson (1981)
POTIIOIDEAE
175
Plate 57. Ploubelia. 1-8. P. sprengelii. 1. Habit. 2-4. lbree leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse section at midleaf. 8. Peristome.
9-18. P. involuta. 9. Habit of sporangiate plant. 10. Perigoniate plant. 11. Transverse section of stem. 12-14. lbree leaves. 15. Leaf apex. 16.
Transverse section at midleaf. 17. Propagula. 18. Peristome. 19. P. perinvoluta. 19. Habit.
176
GENERA OF THE POTTIACEAE
placed the species Merceyopsis angulosa with Plaubelia (without
actually making a combination), but that species is better viewed
as the monotypic genus, Ganguleea ((q.v.).
Two new combinations are here added to Plaubelia from the
Old World, where they were previously recognized in Desmatodon and Weisiopsis. These additional species differ from the
American P. sprengelii in their strongly involute upper !aminal
margins. They are otherwise little different from each other and
may prove synonymous.
Additional literature: Crum (l965c), Delgadillo & Zander
(1985), Saito (l973d), Zander (l983c).
Number of accepted species: 3.
Species examined: P. involuta (NY), P. perinvoluta (FH), P.
sprengelii.
New combinations: Plaubelia involuta (Magill) Zand., comb.
nov. (Weisiopsis involuta Magill, A. S. Afr. Mosses 1: 225, 1981
[1982]). Plaubelia perinvoluta Zand., nom. nov. (Desmatodon
involutus Bartr., hom. illeg., Farlowia 1: 175, 1943). Plaubelia
sprengelii (Schwaegr.) Zand., comb. nov. (Barbula sprengelii
Schwaegr., Spec. Muse. Suppl. 2(1): 64, 1823; Desmatodon
sprengelii (Schwaegr.) Williams; Trichostomum sprengelii
(Schwaegr.) Lindh.). Plaubelia sprengelii var. stomatodonta ·
(Card.) Zand., comb. nov. (Hyophila stomatodonta Card., Rev.
Bryol. 36: 76, 1909; Desmatodon stomatodontus (Card.) Williams; Neohyophila stomatodonta (Card.) Crum).
Plate 58.
43. TENIOLOPHORA
Teniolophora Reese, Bryologist 65: 67, 1962. Type:
Teniolophorajluviatile (Williams) Reese.
Teniola Reese, Bryoloigst 62: 221, 1959 [1960], nom. inval.
From teniola, a band of intramarginal differentiated cells + o +
qx5po<;, that which is brought in, from <papero, to bear or carry. to
possess; referring to the differentiated border of elongated cells
otherwise characteristic of the Calymperaceae.
Plants growing in a thick turf, dark green above, brown
below. Stems to 3 em in length, often branching, in transverse
section pentagonal, central strand strong, sclerodermis present,
hyalodermis absent, axillary hairs ca. 10 cells in length, basafl-2
brown. Leaves oppressed, incurved and tubulose when dry,
spreading when moist, elliptical, to 2.5 mm in length, broadly
channeled across ventral surface; margins plane to incurved,
entire to denticulate near apex, with a bistratose inframarginal
border of 5-6 rows of thick-walled, rectangular or rhomboidal
cells extending from leaf base to near apex; apex rounded to
broadly acute, often apiculate or sharply reflexed; base scarcely
differentiated in shape; costa percurrent, Cr10 cells across ventral
surface at midleaf, superficial cells quadrate to short-rectangular
ventrally, elongate dorsally, transverse section elliptical, two
strong stereid bands present, epidermal cells differentiated on
both surfaces, guide cells 4-8 in 1 layer, hydroids absent; upper
lamina/ cells short-rectangular to hexagonal, small, 7-9 Jlm in
width, 1:1, walls evenly thickened, ventrally bulging, dorsally
nearly flat, homogeneous; papillae apparently absent or low and
broad, evident only in leaf section and only present on dorsal surface, simple, one per lumen; basal lamina! cells differentiated
across leaf, rectangular,1ittle wider than upper !aminal cells, walls
evenly thickened, 2-3: l. Propagula borne on stout stalks in upper
leaf axils, multicellular, large, to 350 Jlm long, obovoid to ellipti-
cal. Apparently dioicous, perichaetia terminal, inner leaves not
differentiated. Perigonia and sporophyte not known. Lamina/
KOH color reaction deep yellow in upper leaves, deep yelloworange in lower leaves.
Found on moist rocks near streams or other moist situations;
presently only known from the West Indies in Puerto Rico and
Haiti.
This genus was established largely on the basis of the
differentiated !aminal intramarginal border (Pl. 58, f. 5, 8), similar to the teniolae of Calymperes. Excepting this band of thickwalled cells and firm-walled basal cells of the axillary hairs,
characters of Teniolophora are much the same as those of Hyophila involuta. Specimens of this latter taxon (e.g. Brazil, Rio
de Janeiro, Glaziou 9228, NY) may be found with similar large,
obovoid, unarmed propagula (Pl. 58, f. 9, 10) rather than the
more usual armed propagula.
·
Literature: Reese (1959, 1962).
Number of accepted species: 1.
Species examined: T.fluviatile (NY).
Plates 59-62.
44. WEISSIA
Weissia Hedw., Sp. Muse. 168, 1801. Lectotype: Weissia
controversa Hedw.fide Mitten, Kew J. Bot. 8: 258, 1856.
Cavanillea Borkh., Disp. Pl. 251, 1809, hom. illeg. non
Medikus, 1787, non Desf. in Lam., 1792.
Subg. Weissia
Simophyllum Lindh., Act. Soc. Sci. Fenn. 10: 74, 1871, nom.
illeg. incl. gen. prior.
Rechingerella Frohl., Ann. Naturhist. Mus. Wien 66: 36, 1962
[1963], hom. illeg. non Petrak. Type: Rechingerella
macedonica J. Frohl.
Weissia subg. Weisia BSG, Bryol. Eur. 1: 5, 1851 (fasc. 47.
Consp. vol. 1: VII), nom. illeg.
Weissia subg. Microweisia BSG, Bryol. Eur. 1: 5, 1851 (fasc.
4Cr47 Consp. vol. 1: VII).
Weissia subg. Euweisia Schimp., Syn. 54, 1860, nom. illeg.
Hymenostomum subg. Weisia (Hedw.) Andrews, Bryologist
23: 31, 1920, nom. illeg. prior. ut gen.
Tortella subg. Nanotortella C. Jens., Danm. Moss. 2: 318,
1923, nom. illeg. incl. typ. gen. prior., p .p.
Bryum sect. Weissia (Hedw.) Relh., Fl. Cantabr. 425, 1802.
Type: Bryum virens Relh.
Weissia sect. Controversae Nees & Homsch., Bryol. Germ.
2(2): 25, 31, 1831.
Weissia sect. Viridulae BSG, Bryol. Eur. 1: 66, 1846 (fasc.
33-36 Mon. 6).
Weissia sect. Euweisia C. Miill., Syn. 1: 651, 1849, nom. illeg.
Didymodon sect. Pusilli Kindb., Eur. N. Amer. Bryin. 2: 272,
1897. Type: Didymodon triumphans (De Not.) Kindb.
Barbuto sect. Edentella C. Miill., Gen. Muse. Fr. 453, 1900.
Type: Barbuto schweinfurthiana C. Miill.
Weissia subsect. Sphalerostomae Nees & Homsch., Bryol.
Germ. 2(2): 24, 25, 1831, nom. illeg.
Subg. Astomum (Hampe) Kindb., Eur. N. Amer. Bryin. 2: 283,
1897.
Astomum Hampe, Flora 20: 285, 1837. Lectotype: Astomum
crispum (Hedw.) Hampe fide Margadant, Acta Bot. Neerl.
8: 274, 1959.
Sphaerangium Schimp., Syn. 12, 1860, nom. illeg. incl. gen.
POTIIOIDEAE
177
Plate 58. Teniolophora. 1-10. T. fluviatile. 1. Habit. 2-4. lbree leaves. 5. Leaf apex. 6. Upper marginal cells, showing intramarginal border. 7.
Basal cells. 8. Transverse section at midleaf. 9. Stalked propagula on stem. 10. Propagulum.
178
GENERA OF THE POTTIACEAE
prior.
Systegium Schimp., Syn. 30, 1860. Lectotype: Systegium
crispum (Hedw.) Schimp.
Astomum subg. Euastomum (C. Miill.) Broth., Nat. Pfl. 1(3):
384, 1901, nom. illeg.
Hymenostomum subg. Astomum (Hampe) Andrews, Bryologist
23 : 31, 1920.
Tortella subg. Systegium C. Jens., Danm. Moss. 2: 317, 1923,
hom. illeg. Type: Tortella crispa (Hedw.) C. Jens.
Phascum sect. Astomum (Hampe) Angstr. in Fries, Summ. Veg.
Scand. 1: 97, 1846.
Weissia sect. Astomum (C. Miill.) C. Miill., Bot. Zeit. 5: 98,
1847, as autonym.
Weissia sect. Systegium Lindh. , Ofv. K. Vet. Ak. Forh. 21: 230,
1864. Lectotype: Weissia crispa (Hedw.) Mitt.
Acaulon sect. Macrobryum C. Miill., Linnaea 43: 353, 1882.
Type: Acaulon lorentzii C. Miill.
Phascum sect. Systegium C. Mtill., Hedwigia 37: 76, 1898, nom.
il/eg. incl. sect. prior.
Subg. Hymenostomum (R. Br.) Limpr., Laubm. Deutschl. 1: 225,
1886, as autonym.
Hymenostomum R. Br., Trans. Linn. Soc. London 12(2): 572,
1818. Type: Hymenostomum microstomum (Hedw.) R. Br.
Gymnostomum subg. Hymenostomum (R. Br.) Schimp., Syn. 33,
1860.
Mollia subg. Hymenostomum (R. Br.) Lindh., Musci Scand. 21,
1879.
Weissia subg. Hymenostomum Limpr., Laubm. Deutschl. 1:
225, 1886, nom. illeg.
Hymenostomum subg. Kleioweisia Limpr., Laubm. Deutsch). 1:
224, 1886. Type: Hymenostomum rostellatum (Brid.) Schimp.
Weissia subg. Hymenostomum (R. Br.) Kindb., Eur. N. Amer.
Bryin. 2: 283, 1897, nom . illeg.
Hymenostomum subg. Euhymenostomum Andrews, Bryologist
23: 31 , 1920, nom. il/eg.
Weissia sect. Hymenostomum (R. Br.) BSG, 1: 56, 1846 (fasc.
33-36 Mon. 6), as autonym.
Hymenostomum sect. Microstoma B.&S . in BSG, Bryol. Eur. 1:
56, 1846 (fasc. 33-36 Mon. 6), nom. illeg. incl. typ. gen.
Type: Hymenostomum microstomum (Hedw.) R. Br.
Hymenostomum sect. Tortilia B.&S. in BSG, Bryol. Eur. 1: 56,
1846 (fasc. 33-36 Mon. 6). Type: Hymenostomum tortile
(Schwaegr.) B.&S .
Weissia sect. Hymenostomum (R. Br.) C. Miill., Syn. Muse. 1:
660, 1849.
Mol/ia sect. Hymenostomum (R. Br.) Braithw., Brit. Moss A. 1:
230, 1885.
Subg. Phasconica (C. Miill.) Zand. see below.
Phasconica C. Miill., Linnaea 43: 438, 1882. Type: Phasconica
lorentzii C. Miill., lectotyp. nov.
Subg. Pseudopottia Kindb., Eur. N. Amer. Bryin. 2: 283, 284,
1897.
Sect. Gymnostomum Mitt. , J. Linn. Soc. Bot. 12: 41, 129, 131,
1869.
Sect. Tortularia Mitt., J. Linn. Soc. Bot. 12: 15, 130, 139, 1869.
Subsect. Hapalostomae Nees & Hornsch., Bryol. Germ. 2(2): 24,
25, 1831.
Named for Friedrich Wilhelm Weiss (1744-1826) of Gottingen,
Germany, a lichenologist.
Plants in low cushions or turfs or loosely caespitose, green
above, brown to tan or yellow below. Stems branching irregularly, to 1.0 em in length, transverse section roundedpentagonal, central strand present, occasionally hollow or very
thick, sclerodermis weakly differentiated in 1-2 layers, hyalodermis weakly differentiated to distinct, seldom absent; axillary
hairs to 10 cells in length, basal 1-2 cells thicker walled;
sparsely radiculose. Leaves incurved, tubulose, often contorted
or spiralled when dry, spreading and occasionally sharply
reflexed above a sheathing base when moist, long-ligulate,
oblong or triangular to lanceolate, 1.5-2.5(-4.0) mm in length,
upper lamina broadly channeled across leaf, seldom narrowly
channeled along costa, margins sharply incurved (or occasionally tightly involute) above the leaf base, seldom merely erectincurved near apex, entire, occasionally fragile and breaking off
in large rectangles; apex sharply acute to subulate, occasionally
broadly acute, obtuse, or weakly cucullate; base scarcely
differentiated to ovate or rectangular, occasionally halfsheathing; costa shortly and sharply mucronate, seldom
subpercurrent, superficial cells quadrate or occasionally shortrectangular to elongate ventrally, elongate dorsally , 4-8(-10)
rows of cells across costa ventrally at midleaf, costal transverse
section ovate, occasionally circular or semicircular, two stereid
bands present, usually of about equal size, differentiated epidermis present ventrally, absent or occasionally weakly
differentiated dorsally, guide cells 4-6(-8) in 1 layer, hydroid
strand absent or occasionally present; upper laminal cells
subquadrate to hexagonal, 7- 13 ~min width, 1:1, walls thin to
evenly thickened, superficially strongly bulging on both
exposed surfaces or more strongly protuberant on ventral surface; papillae bifid, 2-6 per lumen, occasionally fused into a
capitulate papilla covering the lumen, occasionally spiculosebranching, seldom absent; basal cells differentiated across leaf,
occasionally rising higher along margins in a vee, rectangular,
occasionally rhomboidal, 2-5:1, walls thin to evenly thickened.
Rhizoid-borne tuber-like propagula reported for W. controversa.
Monoicous, occasionally dioicous. Perichaetia terminal, inner
leaves little different from cauline leaves or somewhat larger,
occasionally weakly sheathing the seta, lower cells longrhomboidal to midleaf. Perigonia appearing as stalked lateral
buds on perichaetiate plants (but variably present) or terminal
on usually smaller perigoniate plants. Seta 0.1-1.3 em in length,
1(-2) per perichaetium, yellowish brown, twisted clockwise;
theca 1.0-2.2 mm in length, occasionally inclined, yellowish to
reddish brown, elliptical to ovate or cylindrical, occasionally
nearly spherical or urceolate, exothecial cells rectangular, walls
thin or becoming rather thick, rarely superficially bulging,
stomates phaneropore at base of capsule, annulus of 2-6 rows
of persistent, vesiculose cells, often on a differentiated collar, or
annulus not differentiated; c/eistocarpous or stegocarpous, peristome teeth 16, short or rudimentary or absent, occasionally
lifts with operculum, oblong-truncate to long-triangular, often
irregularly cleft into two rami or variously perforate, papillose
to spiculose or spirally ridged, to 125(-250) ~m . with up to 10
articulations, straight or twisted weakly counterclockwise, basal
membrane absent or low,low-spiculose, mouth of capsule occasionally closed by a hymenium, this internal to the peristome
(when peristome present), seldom macrostomous. Operculum
conic to rostrate, 0.3-0.8(-1.5) mm in length, cells straight or
rarely somewhat twisted counterclockwise.
POTIIOIDEAE
179
Plate 59. Weissia. 1-11. W. controversa. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Upper lamina! papillae. 9-10. Two peristomes. 11. Transverse section of capsule wall. 12-17. W. artocosana. 12-14.
'Three leaves. 15-16. Axillary hairs. 17. Transverse section at midleaf.18-22. W. brachycarpa. 18. Habit. 19-21. 'Three leaves. 22. Leaf apex.
180
GENERA OF THE POTTIACEAE
Plate 60. Weissia. 1-5. W. breutelii. 1-3. Three leaves. 4. Leaf apex. 5. Transverse section at midleaf. 6-9. W. crispa. 6-8. Three leaves. 9.
Leaf apex. 10-15. W. edentula. 10-11. Two leaves. 12. Leaf apex. 13. Perichaetialleaf. 14. Capsule with operculum. 15. Transverse section of
capsule wall. 16-20. W. jamaicensis. 16-17. Two leaves. 18. Leaf apex. 19. Transverse section at midleaf. 20. Peristome teeth. 21-24. W.
longidens. 21-22. Two leaves. 23. Leaf apex. 24. Peristome teeth.
POTIIOIDEAE
Calyptra cucullate (to short-triangular on cleistocarpous capsules),
smooth, 0.4-2.5 mm in length. Spores 14-28 j.lm in diameter,
brown to yellowish brown, low-papillose or occasionally highly
papillose. Lamina/ KOH color reaction yellow, usually pale yellow. Reported chromosome number n = 13, 13+m, 13+0-4m, 14,
26 (subg. Weissia); 13, 13+m, 26 (subg. Astomum); 8, 13, 14, 26
(subg. Hymenostomum).
A large genus found on all continents except Antarctica,
mostly growing on soil.
Weissia is superficially not well distinguished from Trichostomum. Andrews (1945) felt that the Weissia complex should be
"extended to cover Trichostomum." In Weissia, the central strand
nearly universally present (Pl. 59, f. 2; 62, f. 7) at least in the
specimens examined), upper laminal margins are usually narrowly and sharply incurved (Pl. 59, f. 7; 60, f. 5) as is the case
with Chionoloma and Weissiodicranum (those of Weissia
termitidarum and certain others are loosely involute), the upper
!aminal cells are often more strongly bulging ventrally ,than dorsally (although usually overlain with papillae), the hydroid strand
is usually absent (but present in W. controversa), the sexual condition is commonly monoicous (position of antheridia often varying between or within collections of a particular species), perichaetial leaves are often larger than the cauline, peristomes are
generally short with the teeth more or less flattened (Pl. 59, f.
9-10) or absent, and spores are rather large, 14-28 j.lm in diameter. When present, the ventrally colliculate condition, i.e. bulging
upper !aminal cells associated with slightly thicker, flattened dorsal superficial walls, has not been emphasized by previous
authors, but is common among the species examined, being
present to various degrees in, e.g. W. artocosana (Pl. 59, f. 17),
W. breutelii (Pl. 60, f. 5), W. canaliculata, W. condensa, W.
controversa (Pl. 59, f. 7), W. ghatensis, W. glaziouii, W.
latiuscula, W. ligulaefolia, W. macrospora, W. occidentalis, W.
striata (Pl. 61, f. 19), W. termitidarum, W. triumphans, W.
veviridis and W. wimmeriana.
The best-developed species of Weissia, W. jamaicensis (Pl.
60, f. 16-20), includes a potpourri of features characteristic of
other genera. It has the basal vee of hyaline cells of Tortella , the
much-enlarged ventral stereid band and dilated leaf . base of
Pseudosymblepharis, and a peristome of 16 straight, linearlanceolate teeth cleft to a basal membrane into two spiculose rami
quite like that of Trichostomum crispulum (which, itself, occasionally develops a basal vee and margins incurved near the apex,
but lacks the swollen ventral stereid band). In this case, the species are distinctive, but the generic definitions break down.
Trichostomum crispulum and T. perligulatum have leaf margins
that are often incurved near the apex but generally have plane or
broadly incurved margins throughout most of the leaf. Some species, such as W. crispa (Pl. 60, f. 6-9) and the doubtfully distinct
W. muhlenbergiana (Pl. 61, f. 1-6, see Crum & Anderson's 1981
discussion), have perichaetial leaves with plane to erect-incurved
margins but cauline leaves (and those on sterile stems) with
weakly but narrowly incurved upper margins, and are perhaps
ultimately better assigned .to Trichostomum in the T. crispu/um
relationship. Weissia rutilans (Pl. 61, f. 7-11) has upper lamina!
margins not easily assigned to either plane or narrowly incurved
status. In any case, to provide an interim classification that better
reflects perceived relationships, all Weissia taxa with clearly
plane upper cauline leaf margins examined in the course of this
study are here transferred to Trichostomum.
181
The fact that peristome teeth may occasionally be represented in some species only by very much reduced, truncate and
colorless fragments, these often hidden below the rim of the
capsule mouth, indicates that past generic level distinctions
between peristomate and gymnostomous species of Weissia
with similar leaves may not be worthwhile. There is some correlation between thick capsule walls and lack of a peristome
(reflected in distinctions between Weissia taxa of my treatments
in the Moss Flora of Mexico, in press), but this must be investigated in greater detail. Stoneburner (1985) demonstrated significant variation in peristome expression in species of Weissia
from the southwestern United States, and even found rudimentary teeth in a species (W. ligulaefolia) hitherto considered
gymnostomous. She further discussed the "evanescent membrane" of hymenostomous species as a poor taxonomic character, explaining it as the upper surface of a somewhat persistent
columella in both eperistomate and peristomate taxa. The
present study confirms this in that a "hymenium" can often be
seen below the peristome teeth of recently dehisced capsules of,
for instance, W. controversa. This is an old problem. Referring
to Brown's (1818) description of the then new genus Hymenostomum, Greville and Arnott (1824b) wrote: "In Dr. Hooker's
excellent figure of Gymnostomum xanthocarpum in the Musci
Exotici; the membrane which closes the sporular bag is admirably delineated.... It ought to be observed, that Palisot has
described this membrane in the Mosses without a peristome, but
denied it to the others" and (1824a) again asserted: "Now, every
moss possesses this 'membranula indivisa,' arising from the lining of the theca; but in all the species of Gymnostomum, it is
peculiarly evident at some particular stage." However, Greville
and Arnott (l824b) admitted that: "The membrane to which we
have already alluded, as closing the mouth of the sporular bag,
there is every reason to suppose, is stronger and more durable in
all such as have a naked peristome."
A further taxonomic complication is that the peristome teeth
of occasional specimens, e.g. of W. triumphans, are somewhat
twisted counterclockwise, possibly an indication of a relationship of these species to Tortella.
Regarding the synonymy of Weissia and Astomum, a review
of the problem and new evidence (existence of sporophytes of
intermediate morphology) was offered by Stoneburner (1985).
Many combinations at the genus level in Astomum and Hymenostomum remain as "correct" names in the list of species and
infraspecies given at the end of this treatment. A mass transfer
of these names to Weissia was considered but rejected because
many names, on examination of authentic material, will prove
to be more appropriately placed in Trichostomum (as emended
here). Recognition here of subgenera of Weissia that are based
largely on sporophyte morphology reflects recent studies by
other authors (excepting subg. Phasconica). Quite probably,
intraspecific lines of evolution involving sporophytic features of
species with essentially identical gametopbytes will be found
that cut across presently recognized subgeneric limits. Unusual
characters of sporophytes, such as the thick thecal walls of species of subg. Hymenostomum or the strongly bulging exothecial
cells of W. macrospora, however, should also be evaluated and
taken into account.
A monotypic genus now in the . Viridivelleraceae, Viridivellus Stone, may actually represent reduction of the Weissia
gametophyte to an extreme, leaving only the stem and
182
GENERA OF THE POTTIACEAE
Plate 61. Weissia. 1-6. W. muhlenbergiana. I. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Sporophyte. 6. Calyptra. 7-11. W. rutilans. 1-9. Three
leaves. 10. Leaf apex. 11. Peristome, rudimentary, vertical view. 12-15. W. semidiaphana. 12-13. Two leaves. 14. Leaf apex. 15. Capsule.
16-20. W. striatll. 16--17. Two leaves. 18. Leaf apex. 19. Transverse section at midleaf. 20. Capsule.
POTTIOIDEAE
perichaetialleaves on a persistent protonema; Stone (l980b) indicated that this genus may be better placed in the Pottiaceae. It
should be examined during future revisions of the genus Weissia.
Brotherus (1924-25) placed Phasconica between Tetrapterum
and Hymenostomum, among genera he felt intermediate between
Astomum and Weissia . Hilpert (1933), who saw material of the
generitype, Phasconia lorentzii, noted the closely inrolled upper
laminal margins and suggested a relationship with Astomum.
Crundwell and Nyholm (1972a, 1974) included Phasconica in
Weissia subg. Astomum. Stone (1980b) carefully described and
illustrated what is surely Phasconica balansae from Australia,
showing the strongly incurved upper larninal margins characteristic of Weissia; however, she indicated a reluctance to follow
Crundwell and Nyholm (l972a, 1974) in assigning the genus to
Weissia subg. Astomum because of its macrostomous capsule with
dehiscent flattened operculum, stout columella, and ventral surface of laminal cells distinctly mamillose and strongly papillose,
among other characters. Magill (1981) suggested that because of
similarities of gametophyte and dissimilarities of sporophyte (ex
descr.) this genus "is reminiscent of the relationship between
Weissia and the segregate genera Astomum Hampe and Hymenostomum R. Br." Saito (1975, p. 417) reduced Phasconica to synonymy with Weissia, with no further explanation. Material of
183
Phasconica C. Miill. s. str., seen during this study was found
mixed in the type of Trachycarpidium verrucosum (P.
tisserantii, PRE!, is referred to Trichostomum as a synonym of
T. unguiculatum see Crundwell & Nyholm 1974), and the genus
Phasconica is here given a new status as a subgenus of Weissia
differing from other subgenera mainly by the macrostomous,
eperistomate capsule with a short seta and a well differentiated,
dehiscent operculum. Because of the unusual sporophyte morphology, subg. Phasconia may not be part of a reduction series
in Weissia, but may instead be a remnant of a more primitive
and now largely extinct lineage; further analysis should be done
at the species level. Weissia platystegia (illustrated and discussed as Astomum platystegium by Eddy (1990) and Norris and
Koponen (1989) belongs in subg. Phasconica.
The singular Pseudosymblepharis socotrdna, including the
synonym Barbula (sect. Edentella) schweinfurthiana (isotype at
BM) is transferred to Weissia (as W. artocosana, a nom. nov.)
on account of the weakly papillose, ventrally bulging upper
laminal cells, ventral stereid band smaller than the dorsal (Pl.
59, f. 17), and the lack of a peristome, all characters found in
Weissia. The strongly sheathing leaf bases are found in both
genera, and are variably demonstrated among species of
Pseudosymblepharis. This species is gametophytically rather
Plate 62. Weissia. l-6. W. triumphans. 1-2. Two leaves. 3. Leaf apex. 4. Theca. 5. Peristome. 6. Operculum showing slightly twisted cells.
7-11. W. veviridis. 7. Transverse section of stem. 8-10. Three leaves. 11. Leaf apex.
184
GENERA OF THE POTTIACEAE
similar to Rhamphidium but differs in the entire leaves and lack
of a hydroid strand, as well as absence of a peristome.
Hybrids have not only been reported between species of different subgenera of Weissia, but also between Weissia crispa and
Torte/la flavovirens (Nicholson 1910). The sporophytes in this
last case apparently had short setae; the thecae were ovoid, partially cleistocarpous, rudimentarily peristomate, and covered by
very large calyptrae (characteristic of Torte/la flavovirens). The
spores were also reported as remaining adherent in tetrads.
Additional literature: Anderson and Lemmon (1972, 1973,
1974), Andrews (1920, 1922a, 1924, 1933), Bryan (1956),
Crundwell (197lb), Dietert (1979), Eckel (1986b), Flowers
(1973b), Grout (1900), Khanna (1960, 1964), Lemmon (1968),
Lewinski (1983), Mizushima (1957), Nicholson (1905a, 1906),
Reese (1988, 1991), Reese and Lemmon (1965), Robinson
(1966), Shaw (1987a), Stoneburner (1981, 1986), Stoneburner
and Wyatt (1985), Williams (1966b), Zander (1985a), Zuttere et
al. (1984).
Number of accepted species: 97, plus 11 in Astomum and 16
in Hymenostomum.
Species examined: W. abbreviata (NY), W. andersoniana
(BUF, TENN), W. argentinica (NY), W. artocosana (BM, NY),
W. balansae (PC), W. balansaeana (MO), W. bizotii (PC), W.
brachycarpa (BUF, NY), W. breutelii (BUF, NY), W.
canalicu/ata (NY), W. condensa (BUF, TENN, NY), W.
controversa, W. crispa (BUF, DUKE), W. diffidentia (US), W.
edentula (MICH. NY), W. fa/lax (NY), W. fe/ipponei (PC), W.
ghatensis (BM), W. glaziouii (NY), W. jamesonii (TENN), W.
jamaicensis (BUF, NY, TENN), W. /atiuscula (BM), W.
ligulaefolia (BUF, FH, NY), W. lineaefolia (NY), W. longidens
(DUKE, NY), W. /ongifolia (NY), W. ludoviciana (DUKE), W.
macrospora (NY), W. muhlenbergianum (DUKE), W. newcomeri
(FH), W. occidentalis (COLO), W. opaca (NY), W. ova/is (NY),
W. platystegia (BM), W. riograndensis (H), W. rutilans (BUF,
NY), W. semidiaphana (BUF, FH), W. sharpii (BUF, DUKE), W.
subacaulis (NY), W. submicacea (NY), W. termitidarum (NY),
W. triumphans (NY), W. unguiculata (NY), W. veviridis (H), W.
wimmeriana (NY).
New heterotypic synonymy: Barbula (sect. Edentella)
schweinfurthiana C. Miill. = Weissia artocosana (Mitt.) Zand.
Trichostomum bermudanum (Mitt.) Par. = Weissia jamaicensis
(Mitt.) Grout. Trichostomum ekmanii Ther. = Weissia jamaicensis
(Mitt.) Grout. Trichostomum linealifolium C. Miill. = Weissia
jamaicensis (Mitt.) Grout.
New names, statuses and combinations:
Weissia subg. Phasconica (C. Miill.) Zand., comb. et stat. nov.
(Phasconica C. Miill., Linnaea 43: 438, 1882).
Weissia abbreviata (Thwait. & Mitt.) Zand., comb. nov. (Systegium abbreviatum Thwait. & Mitt., J. Linn. Soc. Bot. 13: 299,
1873; Astomum abbreviatum (Thwait. & Mitt.) Fleisch.).
Weissia artocosana (Mitt.) Zand., nom. nov. (Symblepharis
socotrana Mitt., Trans. R. Soc. Edinburgh 31: 331, 1888;
Pseudosymblepharis socotrana (Mitt.) Ther.).
Weissia balansae (C. Miill.) Zand., comb. nov. (Phasconica
balansae C. Miill., Linnaea 43: 438, 1882).
Weissia (subg. Phasconica) bizotii Zand ., nom . nov .
(Kleioweisiopsis involuta Biz., Rev. Bryol. Lichenol. 40: 119,
1974 invalid, single element not cited, fide Crosby et al.,
1992). Lectotype: Herb. Bizot; paratype: Herb. Pocs.
Weissia diffidentia Zand., nom. nov. (Phascum recurvirostum C.
Miill., Flora 71: 5, 1888; Tetrapterum recurvirostrum (C.
Miill.) Broth.).
Weissia glaziouii Zand., nom . nov. (Hymenostomum striatum
Geh. & Hampe, Vid. Medd. Naturh. For. Kjoebenh. ser. 4,1:
84, 1879).
Weissia (subg. Phasconica) /orentzii (C. Miill.) Zand., comb.
nov. (Phasconica /orentzii C. Miill., Linnaea 43: 438, 1882).
Weissia neocaledonica (Ther.) Zand., comb. nov. (Aschisma
neocaledonicum Ther., Diagn. Esp. Var. Nouv. Mouss. 8: 4,
1910; Astomum neocaledonicum (Ther.) Andrews).
Weissia riograndensis (Broth.) Zand., comb. nov.
(Hymenostomum riograndense Broth., Bih. K. Svensk. Vet.
Ak. Handl. 26 Afd. 3(7): 19, 1900).
Weissia triumphans var. monspe/iensis (Schimp.) Zand., comb.
nov. (Trichostomum monspeliense Schirnp., Syn. ed. 2, 175,
1876; Trichostomum triumphans var. monspeliense
(Schimp.) Husn.).
Weissia veviridis Zand., nom. nov. (Trichostomum perviride
Broth., Dansk. Bot. Ark. 2(9): 3, 1918).
Plate 63.
45. WEISSIODICRANUM
Weissiodicranum Reese in Reese & Buck, Bryologist 94: 308,
1991. Type: Weissiodicranum insularum Reese in Reese &
Buck.
From Weissia, a genus+ o + Dicranum, a genus (Dicranaceae);
this name refers to genera of two different families, characters
of which are shared by Weissiodicranum.
Plants forming low cushions or turfs, green above, tan
below. Stems branching occasionally, ca.0.5(-1.0) em in length,
transverse section rounded-pentagonal, central strand distinct
and rather large, sclerodermis absent, hyalodermis absent;
axillary hairs 4-10 cells in length, hyaline or weakly thickwalled basally; rhizoids sparse. Leaves incurved, tubulose, contorted when dry, spreading when moist, longly ligulate to
lanceolate, ca. 2.5 mm in length, upper lamina broadly channeled across leaf, margins sharply incurved to involute, entire,
apex narrowly acute; base scarcely differentiated or rectangular;
costa shortly and sharply mucronate, superficial cells quadrate,
elongate and smooth dorsally, with 4 rows of cells across costa
ventrally at midleaf, costal transverse section nearly round to
elliptical or ovate, 2 stereid bands present, differentiated epidermis present both ventrally and dorsally, guide cells (2-)4 in 1
layer, hydroid strand absent; upper /aminal cells subquadrate,
ca. 8-10 Jlm in width, 1: 1, walls evenly thickened, superficially
strongly bulging ventrally, nearly flat dorsally; low molariform
papillae present qn upper part of costa ventrally; basal cells
differentiated across leaf, rectangular, 10-13 Jlm in width,
2-4:1, walls thin to evenly thickened, several "alar" cells at
insertion inflated in 1-3 rows, extending from margin about 2/3
way to costa, decurrent as a pad on stem. Probably dioicous,
only archegonia known. Perichaetia terminal, inner leaves little
different. Perigonia and sporophytes unknown. Lamina/ color
reaction to KOH yellow.
A monotypic genus found on soil at low to medium elevations 'in the West Indies (Puerto Rico and Jamaica) and the
Galapagos Islands of Ecuador.
As Reese and Buck (1991) indicated, this genus is distinc-
185
POTIIOIDEAE
tive in its inflated alar cells (Pl. 63, f. 8), similar to those of the
genus Dicranum Hedw. (Dicranaceae), differing as well from a
very similar species, Weissia breutelii, by the absence of a dorsal
costal epidermis and by the bulging dorsal surface of the costa
(Pl. 63, f. 9). The alar cells are enlarged and banana-shaped, grading below into a unique pad of inflated cells that are 1-2 cells in
thickness on the stem. This is apparently an extreme development
of a tendency already present in Weissia (a synapomorphy not
used in the Data Set), in which leaf insertions consist of one or
two rows of swollen cells that occasionally partially detach from
the stem laterally when the leaf is removed for examination and
giving the appearance of differentiated alar cells. Examined species of Weissia with small groups of distinctly differentiated but
inconspicuous alar cells include: W. abbreviata, W. balsansae,
W. crispum, W. rutilans, and W. veviridis; there are presumably
others. It is almost certain, because of the complex and characteristic costal anatomy, and because somewhat differentiated
alar cells are found in many species of Weissia, that Weissiodicranum is derived from ancestors of pottiaceous stock, not of
Dicranaceae.
Number of accepted species: l.
Species examined: Weissiodicranum insulanum (BUF).
3
Plate 63. Weissiodicranum. 1-9. W. insu/anum. 1. Habit. 2. Transverse section of stem. 3-6. Four leaves. 7. Leaf apex. 8. Basal cells. 9.
Transverse section at midleaf.
186
GENERA OF THE POTTIACEAE
Plate 64.
46. QUAESTICULA
Quaesticula Zand., gen. nov.
Type: Cuba, "moist rocky ledges," Wright, Cuban Mosses 29 (as
"Barbula linearis Swartz"), NY, lectotype of Weissia
navicularis Mitt.
Plantae interdum in regione superna glaucae. Caules filum
centra/em validum evolventes, sclerodermide carentes,
hyalodermide praediti. Folia in statu sicco incurva tubulosaque ,
ligulatolanceolata, marginibus incurvis vel/ate involutis, integris,
apice rotundato, base vix in forma distincto, costa subpercurrenti,
in regione mediana cellulis superficialibus ventra/iter valde
tumescentibus, stratis stereidarum plerumque duobus, cellulis
laminalibus supernis rotundatoquadratis vel hexagonis,
parietibus crassitie aequis evolventibus , in superficie
tumescentibus, papillis magnis, plerumque /uminem obtegentibus,
simplicibus vel bifidis, plerumque per luminem unicis praedita.
The name refers to the long and careful evaluation required
for placing the new genus in proper taxonomic perspective, and is
patterned after the hepatic name Pinguicula.
Plants forming a low, dense turf, dark to light green,
occasionally glaucous above, tan below. Stems branching occasionally, to 0.6 mm in length, transverse section roundedpentagonal, central strand strong, sclerodermis absent, hyalodermis absent; axillary hairs conspicuous, elongate, of 6-8
hyaline cells; sparsely radiculose. Leaves contorted, incurved and
tubulose when dry, spreading when moist, ligulate-lanceo/ate, ca.
1.5-2.0 mm in length, upper lamina broadly channeled across
leaf, margins incurved to broadly involute, entire (crenulate by
projecting cell walls); apex rounded, sometimes weakly cucullate;
base scarcely differentiated; costa strong, often up to 1/3 leaf
width at base, subpercurrent, ending 2-4 cells below apex, costa
with lamina inserted nearly laterally, superficial cells roundedquadrate, strongly bulging ventrally , dorsally elongate and
smooth, 3--6 rows of cells across costa ventrally at midleaf, costal
transverse section circular or ovate, stereid bands ventrally distinct or occasionally absent, dorsally strong and semicircular in
shape, ventral and dorsal epidermises well differentiated, guide
cells (2-)4 in 1 layer, hydroid strand absent; upper /aminal cells
somewhat bulliform, rounded-quadrate or hexagonal, 10-14 1.1m
in width, 1: 1, walls evenly thickened, superficially strongly bulging ventrally, weakly convex dorsally to almost as strongly convex
as ventrally, papillae massive, usually covering the lumens, simple to occasionally bifid, usually one over each lumen ; basal cells
differentiated across lower 1/5 of leaf, rectangular, 10-14 1.1m, the
marginal cells narrower in width, 3-4:1, walls thin. Dioicous.
Perichaetia terminal, inner leaves long-ligulate, to 3 mm in length,
sheathing in lower 1/4, lower cells long-rectangular. Perigonia
terminal, budlike. Seta 0.9-1.2 em in length, 1 per perichaetium,
brown, twisted clockwise; theca 0.7-1.3 mm in length, brown,
ellipsoidal, exothecial cells rectangular, 20-25 1.1m in width,
3-4: 1, thin-walled, stomates phaneropore, at base of capsule,
annulus of 3 rows of strongly vesiculose, persistent cells; peristome teeth 32, longly filamentous, to 700 1.1m in length, spiculose,
twisted almost once counterclockwise, basal membrane 10 to 35
1.1m, papillose. Operculum conic, 0.7-1.0 mm in length, cells
twisted counterclockwise nearly once. Calyptra long, cucullate,
smooth, 2.5-3.0 mm in length. Spores 10-13 1.1m in diameter,
light brown, essentially smooth. Lamina/ KOH color reaction yellow.
Found on rocks (and once on a palm tree trunk) near mountain streams and waterfalls in Mexico and the West Indies.
The incurved upper !aminal margins and the upper !aminal
cells (lumens examined in transverse section) bulging more
strongly ventrally than dorsally (Pl. 64, f. 8-9) are also characteristic traits of Weissia, Plaubelia and Hyophila, but, in addition to the twisted peristome (Pl. 64, f. 10), the narrowly
rounded leaf apices distinguish Q. navicularis from the first, the
lack of a costal hydroid strand from the second, and the leaves
are far too narrow for inclusion of this species in the third. The
significant gametophytic morphology of Quaesticu/a is, in fact,
nearly that of Trichostomum crispulum, especially the ligulate
leaves, apex often cucullate, margins broadly incurved, costa
subpercurrent and protuberant dorsally, but the upper lamina!
cells differ in being evenly thick-walled and otherwise
quasibulliform, somewhat more strongly convex dorsally than
ventrally, and papillae massive, apparently one broad lens-like
papilla over each lumen in the type and most known collections,
but commonly two over each lumen in one collection (Mexico:
Eckel 8511141 BUF); thus approaching the papilla morphology
of Tuerckheimia. The distinctly twisted peristome is unlike that
known for Tuerckheimia and Trichostomum. One might hypothesize that, of genera with superficially strongly bulging upper
lamina! cells and massive papillae, Tuerckheimia is derived
from ancestors of Trichostomum subg. Oxystegus, while
Quaesticula is derived from ancestors of Trichostomum subg.
Crispu/iformes (see subgeneric descriptions as given in the
treatment of Trichostomum) . Luisierella may have been derived
from ancestors of Quaesticula through reduction of sporophyte
(weakly differentiated peristome) and gametophyte (loss of
ventral costal stereid band), and elaboration of sexual condition
and the strongly differentiated vee-shaped basal cell region.
Some of this speculation is supported by the consensus tree of
Cladogram 14; perhaps if Trichostomum subg. Crispuliformes
were treated as a separate genus, it might be referred to the
Hyophileae. Analysis at the species level is needed.
Only one specimen of Weissia navicularis Mitt. is referred
to in the protologue; of the variously distributed isotypes of
Wright 29, the lower left specimen, from Mitten's herbarium,
on the NY sheet is here designated as lectotype of Weissia
navicularis Mitt. and has been marked as such.
Literature: Brotherus (1910), Iwatsuki & Sharp (1958),
Zander (1978£), Zander & Eckel (1987).
Number of accepted species: 1.
Species examined: Quaesticula navicularis (BUF, NY).
New heterotypic synonymy: Tuerckheimia ca/cu/osa Zand.
& Eckel= Quaesticula navicularis (Mitt.) Zand.
New combination: Quaesticula navicularis (Mitt.) Zand.,
comb. nov. (Weissia navicularis Mitt., J. Linn. Soc. Bot. 12:
139, 1869; Tortula navicularis (Mitt.) Broth.).
47. GANGULEEA
Plate 65.
Ganguleea Zand., Phytologia 65 : 426, 1989. Type: Gangu/eea
angulosa (Broth. & Dix.) Zand.
Named for H. C. Gangulee, 1957-, author of "Mosses of Eastern India and Adjacent Regions" (1969-1980).
Plants in loose turf, green above, brown to blackish brown
POTTIOIDEAE
187
i \ :'
.i.........) •• • . ;'
4
5
Plate 64. Quaesticula. 1-11. Q. navicularis. I. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8-9.
Transverse sections of leaves. 10. Peristome teeth. II. Operculum.
188
GENERA OF THE POTTIACEAE
Plate 65. Ganguleea. 1-14. G. angulosa. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse
section at midleaf. 9. Cluster of sporophyte-bearing perichaetia. 10. Capsule. 11. Exothecial cells with plicae. 12. Operculum. 13. Calyptra. 14.
Calyptra inserted on operculum.
POTTIOIDEAE
below, generally rosulate. Stems branching rather often, to 0.3 em
in length, transverse section rounded elliptical to triangular, central strand absent, sclerodermis absent or present (variable in
same stem), hyalodermis absent; axillary hairs 4-6 cells in length,
the basal cell fum-walled; rhizoids sparse. Leaves incurved and
tubulose when dry, spreading when moist, spathulate to oblongelliptical, 1.5-2.0 mrn in length, upper lamina broadly channeled,
margins incurved, entire; apex broadly rounded and apicu/ate;
base very narrow, constricted above the insertion; costa
subpercurrent to short-excurrent as a sharp mucro, costa with lamina inserted ventrally but divergent at 180', superficial cells
rounded-quadrate, bulging ventrally, dorsally elongate, 2-4 rows
of cells across costa ventrally at midleaf, costal transverse section
circular, stereid band single, strong and circular in shape, ventral
and dorsal epidermis present, cells flattened, guide cells 2,
flattened-elliptical, in 1 layer, hydroid strand absent; upper
/aminal cells rounded-quadrate to hexagonal, 8-12 jlm in width
but rather heterogeneous in size, 1:1, lumens usually rounded,
walls usually evenly thickened, superficially strongly bulging
ventrally, nearly flat dorsally; papillae absent; basal cells
differentiated in a small, triangular, juxtacostal area near the
insertion, bulging-rectangular, ca. 18 jlm in width, 2-3:1, walls
thin to thickened and trigonous. Autoicous, paroicous, occasionally probably rhizautoicous. Pleurocarpous. Perichaetia borne on
a short lateral branch, inner perichaetiate leaves lanceolate,
entire to weakly serrulate, short, to 0.8 mm in length, not sheathing, lower cells short-rectangular to rhomboidal, walls thin. Perigonia borne in clusters just below perichaetia, occasionally terminal on a separate plant. Seta 2.5-5.5 mm in length, 1 per
perichaetium, brown, twisted clockwise; theca 0.5-0.8 mm in
length, brown, often glistening, ovate, macrostomous, with 8
plicae, exothecial cells thin-walled, quadrate to short-rectangular,
ca. 18 jlm in width, 8 longitudinal rows of thick-walled cells in
two ranks each along ridges of plicae, stomates phaneropore, at
base of capsule, annulus of 2 rows of thin-walled, weakly
vesiculose cells; peristome teeth absent. Operculum long-conic to
rostrate, erect, ca. 0.7 mm in length, cells straight. Calyptra longconic, sometimes not cleft, inserted on operculum, smooth to
somewhat rough apically with projecting cell walls, ca. 1 mm in
length. Spores 10-13 jlm in diameter, light yellow, essentially
smooth. Lamina/ KOH color reaction yellow.
A rarely collected monotypic genus found on soil over rock,
Himalayas of India and Nepal and mountains above Rio de
Janeiro, southeastern Brazil (Schafer-Verwimp 8403, BUF).
Like Anoectangium, Molendoa and Pleurochaete, this genus
bears its sporophytes laterally on the main stem axis, at the ends
of very short branches bearing highly modified perichaetialleaves
(Pl. 65, f. 1, 9). Ganguleea differs from these genera, however, in
being monoicous. There is considerable resemblance to Weisiopsis, in which it was placed by Hilpert (1933), especially in the
plicate capsule (Pl. 65, f. 10-11), leaves with narrowed base (Pl.
65, f. 3-5, 7), colliculate ventral surface, margins incurved, and
costal section with a usually very strong and rounded stereid band
(Pl. 65, f. 8). Ganguleea may be derived from ancestors of that
genus through the loss of the stem central strand (Pl. 65, f. 2), further narrowing of the leaf base, loss of peristome, and development of pleurocarpy. An isotype at H has the sporophytes fallen
with the short perichaetial branchlets (with their much different
leaf morphology) attached and giving the appearance of two different species in the packet. The Brazilian specimen mentioned
189
above differs from that from Nepal (Norkett 6118, BM) in having a long-rostrate, slightly inclined operculum with a cleft
calyptra, while the Nepal specimen has an erect-conic operculum and long-conic, uncleft calyptra; it may be possible to
distinguish the two at the specific level on this basis, but more
specimens need to be examined.
Additional literature: Gangulee (1972), Vohra and Wadhwa
(1966).
Number of accepted species: l.
Species examined: Ganguleea angulosa (BM, BUF, H).
48. WEISIOPSIS
Plate 66-67.
Weisiopsis Broth., Ofv. Finsk. Vet. Soc. Forh. 62A(9): 7, 1921.
Lectotype: Weisiopsis anomala (Broth. &. Par.) Broth. fide
Saito, J. Hattori Bot. Lab. 39: 525, 1975.
Hyophila subg. Hyophilodonta Card. in Grand., Hist. Madag.
39: 214, 1915. Type: Hyophi/a subplicata Ren. & Card.
From Weis[s]ia, a genus + lf'ljf<n~, -E~, appearance; resembling the genus Weissia.
Plants small, in a low turf, light green above, brown below.
Stems branching occasionally, to 3 mm in length, transverse
section rounded-pentagonal, central strand present, scleradermis absent or of substereid or occasionally stereid cells,
hyalodermis absent; axillary hairs of ca. 3-5 cells, the basal 1
thick-walled; radiculose below. Leaves usually tubulose and
erect-incurved when dry, tubu/ose and spreading when moist,
long-ligulate to spathulate, to 2.2 mrn in length, upper lamina
flat or narrowly grooved along costa, margins plane or broadly
and weakly incurved, entire or minutely crenulate by projecting
cell walls, occasionally serrulate at extreme apex; apex rounded
to broadly acute, often apiculate; base oblong or elliptical; costa
tapering, ending 3-6 cells below apex or percurrent or shortexcurrent into an apiculus, superficial cells elongate and smooth
or rounded-quadrate and bulging ventrally, elongate and smooth
dorsally, 2(-3) rows of cells across costa ventrally at midleaf,
costal transverse section nearly round, stereid band one, usually round in section, ventral epidermis often absent, dorsal epidermis present, guide cells 2(-4) in 1 layer, often flattened and
lenticular in section, hydroid strand absent; upper /aminal cells
rounded-hexagonal to quadrate, 6-13 jlm in width, 1:1, walls
evenly thickened to somewhat trigonous, superficially ventrally
strongly bulging, weakly convex dorsally; papillae absent; basal
cells differentiated across leaf or larger medially up to l/3 leaf
length, often abruptly enlarged and sharply differentiated from
the upper !aminal cells, rectangular, often bulging, 20-25 jlm in
width, 3-5:1, walls thin and hyaline or yellowish brown and
thick-walled, marginal cells narrower. Monoicous (autoicous or
occasionally heteroicous or rhizautoicous). Perichaetia leaves
little differentiated or smaller, terminal. Perigonia lateral and
budlike, occasionally terminal on equal-sized, entirely
perigoniate plants. Seta 4-6 mm in length, 1 per perichaetium,
yellowish brown, twisted clockwise; theca 0.6-1.0 mrn in
length, yellowish brown, ovoid to cylindrical, occasionally
plicate with 8 low ridges (best seen in transverse section) with 2
rows of somewhat thick-walled exothecial cells on the crests,
other exothecial cells rectangular to rhomboid, 25-35 jlm in
width, 2-4:1, thin-walled, rarely superficially bulging, stomates
phaneropore at base of capsule, annulus of 2-3 rows of strongly
190
GENERA OF THE POTTIACEAE
Plate 66. Weisiopsis. 1-8. W. anomala. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Peristome. 9-16. W. nigeriana. 9. Habit. 10-12. Three leaves. 13. Leaf apex. 14. Basal cells. 15. Transverse section at
midleaf. 16. Capsule.
POTIIOIDEAE
191
Plate 67. Weisiopsis. 1-11. W. norrisii. 1-2. Two leaves. 3. Leaf apex. 4. Basal cells. 5. Transverse section at midleaf. 6-11. W. oblonga. 6.
Habit. 7-9. lbree leaves. 10. Transverse section at midleaf. 11. Peristome. 12-21. W. plicata. 12. Habit of sporophyte-bearing plant. 13.
Entirely perigoniate plant. 14. Autoicous bud. 15-17. Three leaves. 18. Leaf apex. 19. Transverse section at midleaf. 20. Transverse section of
theca. 21. Peristome.
192
GENERA OF THE POTTIACEAE
vesiculose cells, occasionally smaller and not vesiculose; peristome teeth absent or more usually 16, inserted below the mouth
of the theca, not contiguous at base, linear-subulate, entire or
occasionally branching above or perforate at base, finely
spiculose to smooth, to 200 11m. with 5-7 articulations, straight,
basal membrane absent. Operculum long-conic to rostrate from a
low-conic base, 0.3-0.7 mm in length, cells straight. Calyptra
cucullate, 1.{}-1.5 mm in length. Spores 8-13 jlm in diameter, yellow, essentially smooth. Lamina! KOH color reaction yellow.
A small genus of southern and eastern Asia, Mexico, Central
America, Brazil, central and southern Africa, and Madagascar;
growing on rocks and soil.
Important characters for Weisiopsis are the broad, tubulose
leaves with plane or incurved margins; colliculate ventral surface
of the lamina (only weakly developed in W. nigeriana); guide
cells often lenticular in section (Pl. 67, f. 5, 10); single, cylindrical stereid band (Pl. 66, f. 7); occasionally plicate thecae (Pl. 67,
f. 20), theca occasionally with bulging exothecial cells (Pl. 66, f.
16-W. nigeriana); basal lamina! cells abruptly enlarged and
rather thick-walled above (Pl. 66, f. 6, 14; 67, f. 4); somewhat
distant peristome teeth (Pl. 67, f. 11, 21) when present; and
essentially monoicous sexuality. Ganguleea is its closest relative. Weisiopsis has much the same costal section and !aminal
basal cells as species of Gyroweisia but differs in the ventrally
bulging upper lamina! cells and the spaced peristome teeth.
Weisiopsis is possibly composed of species from different lines
of reduction, but ultimate relationships cannot be distinguished
without rigorous analysis at the species level.
Weisiopsis plicata (type NY!) has thecae with plicae almost
as strongly developed as those of Ganguleea ((q.v.), and it may
Plate 68. Luisierella. 1-11. L. barbula. l. Habit. 2-6. Five leaves, one enlarged. 7. Leaf apex, ventral view. 8. Basal cells. 9. Transverse section
at midleaf. 10-11. Two peristomes, one comparatively rudimentary.
POTTIOIDEAE
be a transitional species between Gangu/eea and Weisiopsis. In
addition, W. plicata has entirely archegoniate plants, autoicous
plants (perigonia as a bud just below the perichaetium), and
entirely perigoniate plants (perigonia terminal) of the same size as
the sporophyte-bearing gametophytes. Much patience is required
to adequately determine the sexual condition of certain species of
Pottiaceae.
A specimen at NY labelled "Weisiopsis bahiensis ...Burchell,
Catalogus Geographicus Plantarum Brasiliae Tropicae, No.
7737-2" is not the type of that species; the specimen is
taxonomically Trichostomum brachydontium Bruch.
Additional literature: Hilpert (1933), Noguchi (1951).
Number of accepted species: 7.
Species examined: W. anomala (DUKE, NY), W. nigeriana
(NY, UIH), W. norissii (BUF), W. oblonga (BUF, PC, TENN), W.
plicata (NY).
New heterotypic synonymy: Gyroweisia pocsii Biz. = Weisiopsis nigeriana Egun. & Olar.
New combinations: Weisiopsis cucullatifolia (Gao, Jia & Cao)
Zand., comb. nov. (Hyophila cucullatifolia Gao, Jia & Cao, Bull.
Bot. Res. (Harbin) 11(2): 29. 1991), ex descr. et icon. Weisiopsis
nigeriana (Egun. & Olar.) Zand., comb. nov. (Gyroweisia
nigeriana Egun. & Olar., Bryologist 81: 443, 1978). Weisiopsis
norrisii (Zand.) Zand., comb. nov. (Scope/ophila norrisii Zand.,
Bryologist 88: 353, 1985 [1986]).
49. LUISIERELLA
Plate 68.
Luisierel/a Ther. & P. Yarde, Bull. Soc. Bot. France 83: 73, 1936.
Type: Luisierel/a pusilla Ther. & P. Yarde.
Named for Rev. Pere Alphonse Luisier, 1872-1957, a French
muscologist +-ella, diminutive.
Plants dull or blackish green, scattered or gregarious, often
growing in a thin crust associated with cyanobacteria. Stems
occasionally branching, to 2 mm in length, in transverse section
rounded-pentagonal, central strand absent, outer cortex not
differentiated, hyalodermis apparently absent; axillary hairs ca. 5
cells in length, basal cell mostly yellowish. Leaves tubulose, contorted when dry, sometimes rosulate, spreading-recurved from the
base when moist, ligulate-lanceolate to long-elliptical, ca. 1.0--2.0
mm in length; margins plane to erect-incurved, crenulate above
by projecting cells; apex rounded to obtusely acute; base elliptical; costa thin above, rather wide and much flattened below, ending ca. 4-5 cells below the apex, ventral superficial cells similar
to the /aminal cells and 4-6 cells across at midleaf, the dorsal
elongate, costal transverse section showing a single stereid band,
this crescent-shaped, 2-4 guide cells in one layer, epidermis
present only ventrally, hydroids lacking; upper /aminal cells with
a persistent green color, irregularly rounded-hexagonal, 8-12jlm
in width, 1: I (-2), often somewhat transversely elongate at the leaf
margins, superficially ventrally bulging and dorsally nearly flat,
lumens rounded; papillae absent or perhaps represented by
weakly developed, broad, low, flat thickenings above the lumens;
basal lamina/ cells sharply differentiated in a vee, inflated,
rectangular, 20--28 jlm in width, ca. 3: I, walls hyaline, thin.
Gynodioicous; plants synoicous, or different gametoecia on a single plant synoicous or only archegoniate, or some plants entirely
archegoniate but not producing sporophytes. Perichaetia terminal,
enclosing a highly swollen vaginula, inner perichaetial leaves
...
193
smaller than the cauline. Seta 1(-2) per perichaetium, ca. 4-5
mm in length, red-orange, twisted clockwise above; theca 1-2
mm in length, narrowly cylindrical, occasionally slightly
curved, exothecial cells ca. 20 11m in width, 3-4:1, thick-walled,
stomates phaneropore, at base of capsule; annulus of 2 rows of
strongly vesiculose cells, persistent; peristome extremely variable in development, absent, rudimentary or to 500 jlm in
length, when elongate often broken off with the operculum,
teeth 16, linear to Janceolate, cleft to near base and often perforate below, straight, red when well developed, finely papillose
to densely spiculose, basal membrane absent or weakly developed, weakly spiculose or papillose. Operculum long-conic, to I
mm in length, cells straight. Calyptra cucullate, smooth, ca. 2
mm in length. Spores light brown, essentially smooth, small, ca.
8 jlm in diameter. Lamina! KOH color reaction light yellow to
orange, occasionally negative.
Growing on calcareous rock in southeastern U.S.A., Mexico, Central America, the West Indies, Brazil, Japan and Java.
This monotypic genus is easily recognized by the blackish
coloration of the ligulate, tubulose (when dry) leaves with
crenulate upper margins, upper !aminal cells bulging ventrally
and flat dorsally, and ventral surface of the costa covered with
bulging, rounded-hexagonal cells (Pl. 68, f. 7). The basal cells
are inflated and arranged in a sharply differentiated vee (Pl. 68,
f. 8), similar to those of Torte/la, a genus differing in its generally acute leaf apices with costa short-excurrent, and two stereid
bands in the costa, among other characters. Eperistomate specimens might be confused with Hyophi/a, Gyroweisia or Weisiopsis species, but none of these has the basal vee of inflated
hyaline cells. Javan specimens labeled as the synonym .Gyroweisia brevicaulis differ weakly from American collections in
being somewhat more robust and less commonly associated
with the thin turf of cyanobacteria almost universal in New
World collections.
Most descriptions of this taxon refer to a dioicous sexual
condition. A thorough examination of collections indicates that
sporophyte-bearing plants in any particular collection Jack
antheridia, but other plants are variously entirely archegoniate,
synoicous or both archegoniate and synoicous in different
gametoecia on the same plant. This difference is probably the
reason for past incorrect ascription of dioicous sexual condition.
Using a modified form of Wyatt's (1985) nomenclature for sexuality, Luisierella is actually gynodioicous (with both
monoicous and gynoicous plants), and individual plants are
either perigoniate or synoicous or gynomonoicous (with both
perigoniate and, in this case, synoicous gametoecia). The Jack
of sporophytes on synoicous plants may be due to dichogamy,
but this needs further investigation. Deguchi (1987) recently
described a monoicous (synoicous and cryptoicous) and
dioicous or rhizautoicous condition for L. barbula in Asia.
Luisierella may be confused with species of Calymperaceae, which may share the bulging, lens-like upper !aminal
cells, but the vee-shaped area of enlarged basal cells is diagnostic.
Additional literature: Crum & Anderson (1961), Harada and
Deguchi (1990), Potier de Ia Yarde (1936), Steere (1945).
Number of species recognized: 1.
Species examined: L. barbula (BP, BM-as Gyroweisia
brevicaulis, BUF, DUKE, FH, NY, TENN).
New heterotypic synonymy: Trichostomum brevicau/e
194
GENERA OF THE POTTIACEAE
Plate 69. StegoniiJ. 1-13. S. lotifoliiJ. 1-2. Habits. 3. Transverse section of stem. 4-6. lbree leaves. 7. Leaf apex. 8. Basal cells. 9. Ventral
surface of costa. 10-11. Transverse sections near midleaf. 12. Theca. 13. Peristome. 14-18. S. hyalinotricha. 14-15. Two leaves. 16. Leaf apex.
17-18. Capsules.
POTTIOIDEAE
Hampe ex C. Miill. (Gyroweisia brevicaulis (Hampe ex C. Mtill.)
Broth.) = Luisierella barbula (Schwaegr.) Steere. Weisiopsis
hollandii Bartr. = Luisierella barbula (Schwaegr.) Steere.
Plate 69.
50. STEGONIA
Stegonia Vent., Rev. Bryol. 10: 96, 1883. Type: Stegonia /atifo/ia
(Schwaegr.) Vent.
Hyalophyllum Warnst., Hedwigia 53: 284, 1913, nom. illeg.
incl. gen. prior.
Didymodon subg. Stegonia (Vent.) Kindb., Eur. N. Amer.
Bryin. 2: 272, 1897.
Pottia sect. Stegonia (Vent.) C. Miill., Gen. Muse. Fr. 386,
1900.
From O'ttyro, to cover closely, so as to keep out wet, to cover,
protect, to contain; a reference to the concave, protecting leaves.
Plants gregarious, bu/biform, white to whitish green or occasionally just green above, brown below, usually unbranched,
short, to 3 mrn in length, transverse section rounded, central
strand present, sclerodermis absent, hyalodermis absent; axillary
hairs ca. 3-6 cells in length, the basal 2 cells firm-walled; rhizoids
scarce. Leaves appressed when dry, appressed or loosely involute
above and weakly spreading below when moist, nearly circular to
very broadly ovate, sometimes elliptical, ca. 1.5-2.0 mrn in
length, upper lamina deeply concave, margins broadly recurved
to revolute, occasionally plane and seemingly broadly incurved
because of broad leaf concavity, commonly serrulate near apex,
occasionally entire, upper 1/3 to 1/4 of leaf usually hyaline, of
rhomboidal thick-walled cells; apex broadly acute or occasionally
rounded; base not differentiated in shape; costa thin, excurrent as
a flexuose hyaline awn or occasionally percurrent or
subpercurrent, costa with lamina inserted laterally, superficial
cells ventrally short-rectangular, smooth and bulliform, dorsally
rectangular and smooth, ca. 3-4 rows of cells across costa ventrally at midleaf, costal transverse section round, stereid band
present dorsally (occasionally of substereids), ventral epidermis
present, strongly bulging, dorsal epidermis absent above midleaf
but present below, guide cells 2 in I layer (often as two thickwalled cells larger than stereid cells but smaller than ventral
epidermal cells) but often absent above, hydroid strand present
and large (occasionally absent above), ventral epidermal cells
forming a bulging ridge on leaf; upper /aminal cells rhomboidal
or hexagonal and more elongate towards the leaf apex, often
hyaline in upper 1/3 to 1/4 of leaf, 13-22 Jlm in width, 2-3:1,
walls usually thin medially but thick-walled near apex and dorsally superficially thick-walled and ventrally thin-walled, about
equally convex on both free surfaces; papillae absent; basal cells
differentiated across leaf, rectangular, occasionally bulging, little
differentiated or to 30 jlm in width and 4:1, walls often somewhat
thicker than those of upper cells. Monoicous (paroicous or
autoicous). Perichaetia terminal, inner leaves little differentiated.
Setae various in length, very short (7~100 jlm in length) or 0.2 to
1.2 em in length, 1(-2) per perichaetium, brown, straight or
twisted clockwise below and occasionally counterclockwise
above; capsule cleistocarpous, immersed and ca. 0.7 mm in
length, ovate and very shortly apiculate, or stegocarpous and
195
theca exerted and 1-2 mm in length, brown, shiny, elliptical to
cylindrical and somewhat curved, exothecial cells 18-30 jlm in
width, 1-2: 1, thin-walled, stomates phaneropore, at base of capsule, annulus when present of 2-3 rows of persistent, vesiculose
cells; peristome teeth absent in stegocarpous capsules or rudimentary or 16 cleft to near base, occasionally perforate,
ligulate, spiculose, to 125 jlm in length, with up to 5 articulations, straight, basal membrane absent. Operculum when
differentiated short-rostrate, ca. 0.7 mm in length, cells straight
or twisted very weakly counterclockwise. Calyptra not seen,
reported as cucullate and smooth. Spores 20-45 Jlm in diameter,
light to dark brown, essentially smooth to papillose. Lamina/
KOH color reaction yellow. Reported chromosome number n =
26.
Found on soil in Europe, Asia, North Africa, western North
America, Mexico.
The two species, Stegonia /atifolia and S. hyalinotrichum,
have essentially identical gametophytes. Characters of importance are the bulbiform habit; deeply concave leaves with bases
not differentiated in shape; upper !aminal cells smooth and
rhomboidal to hexagonal, cells of the upper 1/3-1/4 of leaf (in
most specimens) hyaline and thick-walled (Pl. 69, f. 7, 16), dorsallaminal cell walls thick-walled (as seen in section or in optical sections of folded portions of the leaf) and the ventral thinwalled in the upper portion of the leaf (Pl. 69, f. 1~11); costa
thin, in section dorsally flattened, the single stereid band represented by stereid, substereid or thin-walled cells in various
specimens or even in parts of the same leaf (the lower portions
of leaf with less thickened cell walls), and the two guide cells
not strongly differentiated but, unlike Tortula sect.
Schizophascum, are not absent.
This genus, like Tortu/a, shows a reduction series in sporophyte characters, S. /atifolia having a peristome (Pl. 69, f.
13-this poorly developed and often apparently absent), S.
mourettii (not seen) said to be eperistomate, and S.
hyalinotrichum cleistocarpous (Pl. 69, f. 17-18). The character
of dorsally thickened !aminal walls is present in Hilpertia
(which has much the same costal section and hyaline leaf apex)
and some species of Acaulon, and may be absent in Stegonia
latifolia in the lower part of the lamina (flowers 1973a) or
almost throughout the leaf (Mosses N. Amer. 484, BUF, has
cell walls little thickened on both exposed sides except in the
extreme apex), and dorsal wall thickening in leaves of S.
hyalinotrichum is only present at the medial apical cells. Of pottiaceous genera, Hilpertia rs most similar in general morphology, but is apparently only distantly related (see Cladograms
9-10, 12-16). Hilpertia differs saliently in the revolute and
strongly photosynthetic leaf margins, differentiated perichaetial
leaves, and red lamina! KOH color reaction. One specimen
labeled "Phascum hyalinotrichum" at MICH (Steere 17728) is a
ttematode-infested Bryum sp.
Number of accepted species: 3.
Species examined: S. hyalinotrichum (MICH), S. /atifolia.
New combinations: Stegonia hyalinotrichum (Card. &
Ther.) Zand., comb. nov. (Phascum hyalinotrichum Card. &
Ther., Bot. Gaz. 37: 363, 1904).
196
GENERA OF THE POTTIACEAE
51. CROSSIDIUM
Plates 70-71.
Crossidium Jur., Laubmoosfl. Oest.-Ungam 127, 1882, nom.
cons. Type: Crossidium squamiferum (Viv.) Jur.
Chloronotus Vent., Fauna A. Venet. 1(3): 124, 1868, nom.
rejic.
Pseudaloina Delg., Bryologist 85: 401, 1982 [1983]. Type:
Pseudaloina woodii Delg.
Barbula sect. Chloronotae BSG, Bryol. Eur. 2: 74, 1842 (fasc.
13-15 Mon. 12).
Barbula sect. Argyrobarbula C. Mull., Syn. 1: 597, 1849.
Tortula sect. Chloronotae (BSG) Spruce, Ann. Mag. Nat. Hist.
ser. 2, 3: 374, 1849.
Tortula sect. Squamigerae Lindh., Oefv. K. Vet. Ak. Foerh. 21:
214, 1864.
Sect. Pseudocrossidium Holz. & Bartr., Bryologist 26: 69, 1923.
Type: Crossidium aberrans Holz. & Bartr., /ectotyp. nov.
Crossidium sect. Paenecrossidium Holz. & Bartr., Bryologist
27: 54, 1924, nom. illeg. incl. sect. prior.
From Kpoaooc;, fringe, tassel + -tOtOV, diminutive; the "tasslelike" fringe on the ventral surface of the costa.
Plants low, gregarious or forming a thin turf, green above,
reddish brown to tan below. Stems mostly buried in soil, branching occasionally, to 4 mm in length, transverse section roundedpentagonal to elliptical, central strand present, occasionally hollow, sclerodermis absent, hyalodermis absent; axillary hairs of
2-6 cells, all clear or basal 1-2 thicker walled; rhizoids sparse
above, usually pale and densely clothing the buried portions of
the stem. Leaves appressed-incurved, occasionally spiralled
about the stem when dry, weakly spreading when moist, ovate to
short-triangular, occasionally spathulate, 0.7-1.5 mm in length,
upper lamina broadly concave, margins recurved to revolute
below or to near apex (but occasionally incurved above to
infolded, often also narrowly recurved), entire or occasionally
denticulate or serrulate near apex or above midleaf, marginal cells
often less papillose than medial or forming a smooth or hyaline
membranous border; apex broadly acute to rounded, usually concave, occasionally emarginate or cucullate; base short-ovate,
occasionally rectangular; costa usually ending in an apiculus or a
short or long (to 300 ~-tm), smooth or denticulate awn, seldom
merely percurrent, superficial cells of upper ventral half of costa
forming a pad of separate filaments, these often branching, 1-9
cells in length, apical cell smooth or papillose, costal superficial
cells elongate dorsally, 3-6 rows of cells across costa ventrally at
midleaf, costal transverse section round to elliptical, a single
stereid band present dorsally, mostly oval, in section,
differentiated epidermis absent to weakly discernible dorsally,
guide cells 2-5 in 1(-2) layers, hydroid strand present, strong,
occasionally encircled by stereid cells; upper /aminal cells
subquadrate to hexagonal or rhomboidal, ca. 13-19 ~-tm in width,
1(-2):1, walls thin or evenly thickened, occasionally somewhat
trigonous and lumens rounded, superficially equally weakly convex or strongly convex ventrally and nearly flat dorsally, dorsal
superficial walls occasionally thickened; epapillose or with hollow, simple "cee-shaped" papillae, usually occurring medially on
the leaf, also often mostly dorsally, 1-4 papillae per lumen; basal
cells differentiated across leaf, rising higher medially, rectangular, occasionally quadrate, to 25~-tm in width, ca. (1-)2-3:1, walls
thin, hyaline, transverse walls often thickened. Dioicous (possibly
occasionally rhizautoicous) or monoicous. Perichaetial terminal,
leaves similar to the cauline or broadly ovate, occasionally rudimentary, inner leaves triangular to broadly ovate or occasionally
spathulate, little or distinctly sheathing below, lower cells occasionally hyaline, rectangular to rhomboidal throughout. Perigonia terminal, sometimes cladautoicous or appearing as
autoicous buds in upper leaf axils. Seta elongate, ca. 0.4-1.6 em
in length, 1 per perichaetium, orange to reddish brown, occasionally yellow, twisted clockwise below, counterclockwise
above; theca ca. 0.8-1.5 mm in length, reddish brown, longellipsoidal to cylindrical, exothecial cells rectangular, ca. 20-30
~-tm in width, 2-4:1, thin-walled, stomates phaneropore, at base
of theca, annulus of 1-2 layers of somewhat vesiculose cells;
peristome teeth 16, cleft to near base into 32 paired rami, longsubulate to linear, spiculose, to 700 ~-tm, with several articulations, twisted weakly counterclockwise, basal membrane low or
to ca. 35 ~-tm in height, spiculose. Operculum conic, occasionally retaining fragments of peristome, ca. 500-900 ~-tm in
length, cells weakly twisted counterclockwise. Calyptra
cucullate, smooth, ca. 1.5-3.0 mm in length. Spores ca. 10-15
~-tm in diameter, yellowish brown, essentially smooth to weakly
papillose. Lamina/ KOH color reaction yellow, orange in older
leaves, occasionally with red patches medially above. Reported
chromosome number n =24.
Found in arid regions of most continents, on soil or rock.
The genus is easily distinguished by its dry habitat; leaves
concave, ovate to spathulate, upper margins recurved or plane
(occasionally broadly incurved but then also marginally
recurved-Pl. 71, f. 4, 14); costa with a ventral cushion of
multicellular, branching filaments of photosynthetic tissue
inserted only on the costa, with one stereid band and a prominent hydroid strand (this often encircled by stereid cells, Pl. 70,
f. 6, 12, 71, f. 14, 18); laminal cells unistratose; seta elongate;
capsule peristomate (Pl. 70, f. 7); and KOH color reaction yellow (occasionally lamina medially orange, especially dorsally at
thickened superficial walls). It differs from Pseudocrossidium
most consistently by the oval (rather than reniform) section of
the dorsal stereid band, upper laminal margins never
differentiated as photosynthetic tissue (rather than occasionally
so in Pseudocrossidium), medial upper leaf cells equally weakly
convex or strongly convex ventrally and nearly flat dorsally (not
strongly bulging . on both sides as is usually the case with
Pseudocrossidium), and the absence of a ventral stereid band
(occasionally present in Pseudocrossidium).
Crossidium squamiferum and related taxa of sect. Crossidium have a characteristic areolation (Pl. 71, f. 13) similar to that
of Aloina (upper cells rhomboidal) and the upper margins are
membranous and incurved over the pad of filaments, but the
lamina is entirely unistratose, the filaments are inserted only on
the costa (not also on a portion of the lamina), and the reaction
to KOH is yellow. Crossidium woodii (Pl. 71, f. 15-18), previouslfrecognizec;l in the monotypic genus Pseudaloina, belongs
here and is very similar to C. apiculatum. Species of Crossidium sect. Pseudocrossidium, such as C. aberrans (Pl. 70, f.
8-13), C. rosei, C. geheebii (Pl. 71, f. 5-7) and C. seriatum (Pl.
71, f. 8-10), may be more closely related to Tortula sect. Tortula (through such species as T. atherodes and T. protobryoides
with their flask-shaped protuberant ventral costal cells) or Tortula sect. Tortula (through Tortula atrovirens and T. revolvens
with their ventral costal pads of vertically elongate cells) than to
Crossidium s. str., lacking the distinctive cylindrical terminal
POITIOIDEAE
197
Plate 70. Crossidium. 1-1. C. spiralifolium. l. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7. Peristome. 8-13. C. aberrans. 8. Transverse section of stem. 9-10. Three leaves. 11. Leaf apex. 12. Transverse section at midleaf. 13. Perichaetial
leaf.
198
GENERA OF THE POTTIACEAE
Plate 71. Crossidium. 1-4. C. crassinerve. I. Transverse section of stem. 2-3. Two leaves. 4. Leaf apex. 5-7. C. geheebii. 5-6. Two leaves. 7.
Leaf apex. 8-10. C. seriotum. 8-9. Two leaves. 10. Leaf apex. 11-14. C. squamiferum. 11-12. Two leaves. 13. Leaf apex. 14. Transverse
section at midleaf. 15-18. C. woodii. 15-16. Two leaves. 17. Leaf apex. 18. Transverse section at midleaf.
POITIOIDEAE
cell of the costal filaments and strongly differentiated, epapillose
upper !aminal cells of species of sect. Crossidium. Although
Crossidium is placed with the Hyophileae in Cladograms 14-16,
perhaps sect. Pseudocrossidium would be referred to the Pottieae
if recognized as a separate genus.
Crossidium is clearly related to Pterygoneurum (see Cladogram 14 and others); the photosynthetic tissue of the former, however, is probably derived from progressive elaboration of a ventral
costal pad of cells (such as that of Tortula muralis) into filaments,
while the latter may have been derived through elaboration of
ridges of ventral costal cells like those in T. atherodes. The filaments may be considered merely deeply laciniate lamellae, but on
dissection, there is no immediate evidence of this in Crossidium
but see discussion of Pterygoneurum. Stark and Castetter (1987)
mention both Crossidium and Pterygoneurum as probably perennial in arid habitats.
Additional literature: Cano et al. (1992), Delgadillo
(l973a,b,c, 1975a, 1982), Frey and Ktirschner (1984, 1988b,
1991), Fuertes-Lasala (1983), Holzinger and Bartram (1923a,b),
Mcintosh (1989).
Number of accepted species: 13.
Species examined: C. aberrans (BUF, DUKE), C. apicu/atum
(BUF, PRE), C. crassinerve (BUF, DUKE, NY, TENN) C.
geheebii (H), C. seriatum (BUF, DUKE), C. spiralifolium (PRE),
C. squamiferum (BUF, DUKE), C. woodii (E, MEXU).
New combinations: Crossidium woodii (Delg.) Zand., comb.
nov. (Pseudaloina woodii Delg., Bryologist 85: 401, 1982
[1983]).
52. PTERYGONEURUM
Plate 72-73.
Pterygoneurum Jur., Laubm. Oest. Ungarn 95, 1882, nom. et orth.
cons., as "Pterigoneurum". Lectotype: Pterygoneurum cavifolium Jur., nom. il/eg.
Fiedleria Rabenh., Flora 31: 252, 1848, hom. illeg. non
Reichenb., 1844. Type: Fiedleria subsessilis (Brid.) Rabenh.
Pharomitrium Schimp., Syn. 120, 1860, nom. rejic.
Pterigoneurum Jur., Hedwigia 21: 154, 1882, nom . il/eg.
Pterogoneuron Jur. in Kindb., Eur. N. Amer. Bryin. 2: 272,
1897, orthogr. var.
Portia subg. Pterygoneurum (Jur.) Boul., Muscin. France 469,
1884.
Barbula sect. Lamel/igerae Schimp., Syn. ed. 2: 193, 1876.
Type: Barbula cavifolia Schimp.
Tortula sect. Pterygoneuron (Jur.) Dix., Stud. Handb. Brit.
Moss.176, 1896.
Pottia sect. Lamellipottia C. Mtill., Gen. Muse. Fr. 386, 1900.
Type: Pottia cavifolia Ftirnr.
Tortula subsect. Pterygoneuron (Jur.) Braithw., Brit. Moss. Fl.
1: 206, 1885.
From 1tt4lu~. -"Oyo~. wing + o + VE~ov, nerve, sinew, tendon;
referring to the ridge-like lamellae of the dorsal surface of the
costa.
Plants short or bulbiform, gregarious or forming a thin turf,
green above, light brown below. Stems branching occasionally,
0.5-5.0 mm in length, transverse section rounded-pentagonal,
central strand present, distinct, sc/erodermis absent, hya/odermis
absent; axillary hairs ca. 7 cells in length, all hyaline or basal 1-2
cells brown; buried stem densely clothed with pale, brown
199
rhizoids. Leaves appressed when dry, weakly spreading when
moist, ovate to ligulate, lamina 0.5-1.5 mm in length, awn to
2.0 mm in length, upper lamina broadly concave, weakly
cucul/ate, margins weakly to broadly incurved to near apex,
plane below, occasionally broadly recurved above, entire or
denticulate above, occasionally dentate to erose near costa at
apex; apex obtuse to rounded; base not differentiated in shape
or somewhat widened; costa long-excurrent as a smooth or
denticulate hyaline awn, but awn occasionally absent in lower
cauline leaves or innermost perichaetial bracts, superficial cells
of costa in medial portion of lamina differentiated as longitudinal lamellae ventrally, elongate dorsally, 4-6 dorsal rows of
cells across costa dorsally at midleaf, costal transverse section
circular, one stereid band present dorsally but this often weak,
differentiated epidermis usually present dorsally, guide cells 2
in 1 layer, hydroid strand present, usually large, (2-)3 lamellae
longitudinally inserted on ventral surface of costa, ca. 12 cells
in height, occasionally lobed, smooth or papillose, cells bulging
superficially; upper /aminal cells quadrate to short-rectangular,
often transversely elongated, 10-13(-20) Jlm in width,
(2-)1:1(-2), walls thin to weakly thickened and often weakly
trigonous, superficially flat or somewhat bulging on both sides
or only ventrally; papillae absent or small, simple, solid or hollow, present dorsally in medial portion of leaf, 2-3 per lumen;
basal cells differentiated across leaf base, occasionally rising
higher medially, rectangular, little wider than upper cells or
weakly inflated, ca. 2-5:1, walls thin. Autoicous, paroicous or
c/adautoicous. Perichaetia terminal, inner leaves elliptical, little
differentiated, to 1.5 mm in length, not sheathing, lower cells
hyaline, rhomboidal-rectangular in lower 1/2. Perigonia occurring as small buds in upper leaf axils or terminal on short
branches from base of perichaetiate plant. Seta 0.4-3.5 mm in
length, 1 per perichaetium, dark reddish to yellowish brown,
twisted counterclockwise; capsule usually stegocarpous but
cleistocarpous in P. kozlovii (not seen); theca 0.5-1.5 mm in
length, dark reddish brown to yellowish brown, shortcylindrical, often macrostomous, variously sulcate or ridged
when dry, exothecial cells rectangular, outer exposed walls
occasionally much thickened (as seen in transverse section),
stomates phaneropore, at base of theca, annulus of ca. 1-3 rows
of weakly vesiculose cells; peristome teeth absent or seldom
present, usually removed with the dehiscence of the operculum,
rudimentary, consisting of a latticework of elliptical fragments
borne on a conical hyaline membrane, weakly papillose, to 300
Jlm in length, with several articulations, weakly twisted counterclockwise, basal membrane low, weakly papillose. Operculum
rostrate to conic-rostrate, ca. 0.6-1.4 mm in length, cells
straight or twisted counterclockwise. Calyptra cucul/ate or
mitrate and lobed, smooth, 0.9-2.0 mm in length. Spores large,
25-38 Jlm in diameter, brown, occasionally hyaline, papillose,
occasionally smooth. Lamina/ KOH color reaction yellow,
occasionally with red or orange patches medially near apex or
on leaf base. Reported chromosome number n = 26, 52.
Found largely in areas of dry climate on soil or occasionally
rock, on most continents.
Pterygoneurum is easily identified by the short, often
bulbiform habit (Pl. 72, f. 1) with concave, ovate to spathulate
leaves with lamellae on the ventral surface of the costa (Pl. 72,
f. 7) and capsule stegocarpous. Species with short setae and
short capsules have mitrate calyptrae, those with elongate setae
GENERA OF THE POTTIACEAE
200
16
Plate 72. Pterygoneurum. 1-11. P. ovatum. 1. Habit. 2. Transverse section of stem. 3-6. Four leaves. 7. Leaf apex. 8. Basal cells. 9. Transverse
section at midleaf. 10. Perichaetialleaf. 11. Capsule. 12-17. P. lame/tatum. 12-14. Three leaves. 15. Leaf apex. 16. Sporophyte. 17. Peristome
(rudimentary).
POTIIOIDEAE
and capsules have cucullate calyptrae. The species of Pterygoneurum examined have extraordinarily similar gametophytes but a
range of sporophyte morphology ranging from long-exerted, elliptical, peristomate capsules (Pl. 72, f. 16-17) to short-cylindrical or
obovate, eperistomate capsules on short setae (Pl. 73, f. 7).
Although one might expect that other species with lamellate
leaves and cleistocarpous capsules might also belong here, the
candidate taxon Acaulon subg. Alaticosta truly belongs with
Acau/on (Pottieae) because of the characteristically nearly circular, more deeply concave leaves; larger, rectangular to rhomboid
mediallaminal cells (20 11m in width, 2-3:1); calyptra shorter (ca.
201
0.4 mm); and leaves with an orange to red KOH color reaction
throughout the leaf blade. Pterygoneurum is quite like Tortula
sect. Pottia, and is approached in most characters by T.
cuneifolia, T. grandiretis and T. ca/ifornica. It is surprising that
Pterygoneurum appears to be rather distantly related to Tortula
in Cladograms 14-16, which indicate a derivation from Hyophila-like ancestors. Cladograms 9, 10 and 13 show, on the
other hand, a close relationship; these hypotheses, of course,
must be tested through additional study, preferably at the species level.
The neatly arranged trellis-like peristome illustrated for P.
Plate 73. Pterygoneurum. l-6. P. macleanum. 1. Transverse section of stem. 2~ . Three leaves. 5. Leaf apex. 6. Transverse section at midleaf.
7-13. P. subsessile. 7. Habit. 8-10. Three upper cauline leaves. 11. Lower cauline leaf. 12. Leaf apex. 13. Transverse section at midleaf.
202
GENERA OF THE POTTIACEAE
lamel/atum by Flowers (l973a) and others was not seen in the
course of this study; a more rudimentary arrangement of small
plates over a membrane was seen instead (Pl. 72, f. 17). The
calyptra of P. ca/ifornicum is actually not cucullate as was indicated in the original description, but is more accurately described
as "long-mitrate," with one long split and 2-4 smaller clefts basally forming small lobes. Both Flowers (l973a) and Wareham
(l939c) note that P. subsessile has a mitrate calyptra that may at
times appear cucullate through uneven splitting. Mcintosh (1989)
reviewed the possibility (Boros 1953, Corley et al. 1981) that P.
kozlovii might represent be a hybrid of Tortula atherodes (Pl. 84,
f. 16-22, discussed as the synonym Phascum cuspidatum) and
another species of Pterygoneurum; the problem, apparently, is the
reduced capsule of P. kozlovii, which is here accepted as a common and natural outcome of evolution in arid land mosses and
requires no postulation of hybridization to explain it away.
Stegonia is similar to Pterygoneurum in the strongly concave
ovate leaves, piliferous leaf apex at the base of which the leaf
margins are commonly erose, !aminal papillae absent, and inflated
bulging ventral costal cells, but lacks the ventral costal
photosynthetic lamellae. Frey et al. ( 1990) reviewed the literature
on the phylogenetic derivation of Pterygoneurum, which indicates
a derivation from (ancestors of) Crossidium. The presence on the
ventral costal lamellae of filaments similar to those of Crossidium
was cited as important evidence of this. The cladistic analysis of
the present study supports such a derivation.
Additional literature: Abramova et al. (1973), PospBil (1975),
Smith (1985).
Number of accepted species: 12.
Species examined: P. californicum (MICH), P. /amellatum
(BUF, DUKE, NY), P. mac/eanum (NY), P. ovatum (BUF,
DUKE, MICH, NY).
53. GLOBULINELLA
Plate 74.
Globulinella Steere, J. Washington Acad. Sci. 36: 221, 1946.
Lectotype: Globulinella globifera (Hampe) Steere fide van der
Wijk et al. (1959-69).
Seligeria subg. Globulina C. Mtill., Gen. Muse. Fr. 306, 1800.
Type: G/obulinella globifera (Hampe) Steere.
From globuline, this from globulus, diminutive of globus, ball +
-ella, diminutive; the bead-like shape of these tiny plants.
Plants small, scattered, gregarious or forming a turf, green
above, tan below. Stems branching seldom, then by
subperichaetial innovation, ca. 0.4-0.6 em in length, transverse
section rounded-pentagonal, central strand strong, sclerodermis
absent, hyalodermis absent or weakly differentiated; axillary hairs
clavate, short, of 2-3 cells, the basal 1 often yellowish. Leaves
appressed when dry, weakly spreading when moist, spathulate or
ovate to orbicular, 0.7-1.3 mm in length, upper lamina concave
to broadly channeled, margins plane or incurved, entire; apex
broadly rounded, usually cucullate; base not differentiated in
shape; costa ending 4-6 cells below apex, often laterally spurred,
ventrally protuberant, dorsally flat or bulging, superficial cells
ventrally quadrate to short-rectangular, bulging, dorsally elongate,
2-6 rows of cells across costa ventrally at midleaf, costal transverse section round to elliptical, stereid band usually absent ventrally, dorsally present and round or flat in shape, ventral and dorsal epidermises present, the dorsal sometimes only present later-
ally, guide cells 2-4 in 1 layer, hydroid strand indistinct, but
apparently present; upper /aminal cells subquadrate to shortrectangular or rhomboidal, lumens rounded, 9-13 )lm in width,
l-2: l, walls irregularly thickened, superficially weakly convex
on both sides or occasionally somewhat more bulging ventrally;
papillae absent; basal cells weakly differentiated across leaf,
short-rectangular, ca. 10-13 Jlffi in width, 2-3:1, walls thin to
evenly thickened or indistinctly porose. Propagula rare, small,
of several cells, short-clavate to spherical, ca. 40 Jlm in length,
borne in leaf axils. Dioicous. Perichaetia terminal, inner leaves
little differentiated, ovate, less cucullate, somewhat enlarged,
usually sheathing the seta, cells elliptical to rhomboidal in lower
l/4-3/4. Perigonia terminal. Seta 0.4-0.8 em in length, 1 per
perichaetium, yellow to brown, twisted clockwise above or
straight; theca 0.6-1.6 mm in length, brown, ovoid to elliptical,
exothecial cells rectangular, walls thin to evenly thickened,
stomates phaneropore, at base of theca, annulus of 2 rows of
vesiculose cells; peristome teeth 16, irregularly cleft or
anastomosing, ligulate to lanceolate, densely spiculose, 65-115
Jlm, with 3-6 articulations, straight, basal membrane absent or
low, spiculose. Operculum rostrate, inclined, 0.6-0.7 mm in
length, cells straight. Calyptra cucullate, smooth, ca. 1.0 mm in
length. Spores 8-13 Jlm in diameter, brown, essentially smooth
to weakly papillose. Lamina/ KOH color reaction yellow.
A small genus known from southwestern U.S.A. south
through Mexico and Central America to Ecuador, found on soil
in dry, montane areas.
Globulinella (see also discussion of Globulina in treatment
of Bryoerythrophyllum) can be recognized by the following
combination of distinctive character states: small, julaceous
plants growing on soil, leaves usually broadly rounded and
deeply concave, often cucullate (Pl. 74, f. 2-6), upper laminal
cell walls incrassate, costa bulging ventrally (Pl. 74, f. 7, 10),
seta elongate, and peristome teeth 16, irregularly cleft and
densely spiculose (Pl. 74, f. 8). Saitoella peruviana, known only
from sterile material, has previously been recognized in Globulinella (sharing the features of rounded, cucullate leaves with
upper cells mainly rhomboidal and thick-walled, and costa ending before the apex) and, in the light of Cladogram 14, may represent convergent evolution. Saitoella differs in its upper
!aminal cells being papillose only medially in the leaf, the complete lack of costal guide cells (the single layer of ventrally
exposed costal parenchyma is papillose, and therefore probably
best interpreted as epidermal tissue), and its red reaction to
KOH solution. Naked, lateral archegonia borne singly on the
stems of both G. g/obifera and S. peruviana were seen, but this
is also the case in certain species of other gertera of Pottiaceae.
Propagula were found in only one specimen (of G. globifera
from Mexico: Zander 4900, BUF). The two stereid bands seen
in sections of the costae of a recent collection of G. benoistii
(Pl. 74, f. 10--Ecuador, Steere E35, NY) indicate .a possible
relationship of this species (and the similar, but singly stereidbanded G. g/obifera) with Plaubelia, with which it shares a
short leaf with incurved upper margins, rounded leaf apex,
smooth (in majority of Plaubelia specimens) upper !aminal
cells, general lack of a ventral stereid band, and 16 irregularly
cleft peristome teeth. Plaubelia differs in its oblong leaves, ventrally colliculate areolation, and non-spurred costa.
Literature: Magill ( 1977 a), Steere & Chapman (1946),
Zander (1976).
POTTIOIDEAE
Number of accepted species: 2.
Species examined: Globulinella benoistii (NY, PC), Globulinella globifera (BUF, FH, TENN).
Plates 75-76.
54.ALOINA
Aloina Kindb., Bih. K. Svensk. Vet. Ak. Hand!. 6(19): 22, 1882,
nom. cons. Lectotype: Aloina aloides (Schultz) Kindb.
Aloidella Vent., Comm. Fauna A. Venet. 1(3): 124, 1868, nom.
rejic.
Barbula subg. Tortula BSG, Bryol. Eur. 2: 1, 1851 (fasc. 45-47
Consp. 2: III).
203
Barbula subg. Aloidella (Vent.) Schimp., Syn. ed. 2: 188,
1876.
Tortula subg. Aloidella (Vent.) C. Jens., Medd. Groen1and 3:
379, 1887.
Barbula sect. Aloideae Fiirnr., Flora 12: 598, 1829. Type:
Tortula aloides (Schultz) De Not.
Tortula sect. Aloideae (Fiirnr.) De Not., Mem. R. Ace. Sc.
Torino 40: 287, 1838.
Barbula sect. Aloina C. Miill., Syn. 1: 596, 1849, nom. illeg.
incl. sect. prior.
Tortula sect. Aloidella De Not., Muse. Ita!. 1: 3, 14, 1862,
nom. illeg. incl. sect. prior.
Plate 74. Globulinella. 1-8. G. globifera. 1. Habit. 2-5. Four leaves. 6. Leaf areolation. 7. Transverse section at midleaf. 8. Peristome. 9-10.
G. benoistii. 9. Leaf. 10. Transverse section at midleaf.
204
GENERA OF THE POTTIACEAE
Plate 75. Aloina. 1-14. A. bifrons. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 6. Basal cells. 8. Transverse section
of upper lamina. 9. Theca. 10-14. A. aloides. 10. Perigoniate plant, some bracts removed. 11-12. Two leaves. 13. Leaf apex, lateral view. 14.
Transverse section at midleaf. 15-18. A. brevirostris. 15-16. Two leaves. 17. Leaf apex and marginal cells with membranous border. 18.
Perichaetial1eaf.
POITIOIDEAE
Tortula sect. Aloina Mitt., J. Linn. Soc. Bot. 12: 143, 157, 1869,
nom. illeg. incl. sect. prior.
Barbula sect. Aloidella (Vent.) Lesq. & James, Man. N. Amer.
Moss. 115, 1884, nom . illeg. incl. sect. prior.
Tortula subsect. Aloideae (Fiirnr.) Braithw., Brit. Moss Fl. 1:
208, 1855.
From Aloe + -ina, characteristic of; fleshy leaved like Aloe
(Liliaceae).
Plants gregarious or in a thin turf, deep green above, brown
below. Stems branching irregularly, fleshy, buried in soil, ca. 0.5
mm exposed above soil surface, transverse section roundedpentagonal, cells thin-walled, central strand absent or present and
distinct, sclerodermis and hyalodermis absent; axillary hairs ca. 6
205
cells in length, all clear; buried stem densely covered with pale
rhizoids. Leaves incurved and somewhat tubulose when dry,
spreading-incurved when moist, obovate or ovate to ligulate, ca.
3.0-3.5 mm in length, upper lamina broadly and deeply channeled, margins strongly infolded, entire to weakly sinuose or
denticulate above, marginal cells often longitudinally elongate
or forming a membranous border; apex rounded, cucullate;
base broadly rectangular, sheathing, hyaline, often with distinct
shoulders; costa brownish, subpercurrent to percurrent or
mucronate or long-excurrent as a smooth or serrate awn, ventrally covered from midleaf to near /aminal apex with
photosynthetic filaments, which are also inserted on the medial
portion of lamina, dorsally with elongate superficial cells or
these occasionally quadrate to short-rectangular near
Plate 76. Aloina. 1-4. A. hamulus. l. Transverse section of stem. 2-3. Two leaves. 4. Leaf apex, dorsal view. S-8. A. rigida. 5-6. Two leaves.
7. Leaf apex. 8. Peristome. 9-11. A. roseae. 9-10. Two leaves. 11. Leaf apex, dorsal view.
206
GENERA OF THE POTTIACEAE
apex, 10-20 rows of cells across costa dorsally at midleaf, costal
transverse section broadly reniform, stereid band present only
dorsally, crescent-shaped, occasionally weak or absent, dorsal
epidermis weakly differentiated, guide cells ca. 6-12 in 1(-2) layers, hydroid strand present, ventral epidermis differentiated as a
series of uniseriate filaments ca. 3-4 cells in length, extending
over bistratose portion of lamina, occasionally branching, apical
cell of filament occasionally thickened and papillose; upper
/aminal cells transversely elongate, rectangular to rhomboidal,
medially bistratose, ca. 8-13 Jlm in width, 1-2:1, walls thickened,
lumens rounded, superficially flat to weakly convex, thickened
dorsally; papillae absent (cells near apex occasionally rough
when awn is rough); basal cells differentiated across leaf, medially bistratose, rectangular, ca. 25 Jlm in width, 3-5:1, walls thin,
occasionally transversely slightly thickened. Dioicous or
monoicous. Perichaetia terminal, inner leaves usually little different from cauline leaves, occasionally with margins plane and not
infolded, innermost leaves serrulate on margins and cells lax
throughout. Perigonia terminal, clustered, inner leaves triangular,
paraphyses filamentous, uniseriate. Seta ca. 0.8-1.5 em in length,
1(-2) per perichaetium, brown to reddish brown, twisted clockwise below, counterclockwise above; theca ca. 2-3 mm in length,
reddish brown, cylindrical, exothecial cells rectangular, ca. 25-30
Jlm in width, 2-3:1, thin-walled, stomates phaneropore at base of
capsule, annulus of ca. 2 rows of vesiculose cells, persistent; peristome teeth 16, linear, variously cleft above or more usually to
basal membrane, spiculose, to ca. 1000 Jlm in length, with several
articulations, twisted 0.5-1.0 times counterclockwise, basal membrane absent or present, to 60 J.lm in height, spiculose. Operculum
conic to rostrate, ca. 0.5-1.0 mrn in length, cells spiralling counterclockwise. Calyptra cucullate, smooth, ca. 2.5 mm in length.
Spores ca. 10-15 J.lm in diameter, light brown, smooth or papillose. Lamina/ KOH color reaction red. Reported chromosome
number n =23+m, 25+2m, 24, 26, 28, 48.
A widespread genus found in North, Central and South America, Europe, Asia, northern and southern Africa, Australia and
New Zealand, growing on soil, occasionally walls and rock.
Delgadillo (1973b, 1975a) recently produced a useful revision
of this genus, supplying workable distinctions between Aloina and
genera with similar morphology, and discussing his conclusions
about their phylogenetic relationships. Delgadillo (1975a) and
Brotherus ( 1924-25) both used the absence of a central strand in
stem sections to assist in separating Aloina from related genera;
however, this may be misleading in that some species of Aloina
(e.g. A. brevirostris, Pl. 75, f. 15-18, and A. hamulus, Pl. 76, f.
1-4) may in fact have a distinct central strand. These species,
however, lack the stem satellite hydroid strands of Aloinella. The
taxonomically most important features of Aloina are the fleshy,
buried stem (like that of Crossidium), the upper !aminal margins
strongly folded over the pad of photosynthetic filaments, which
itself covers the upper costa and the bistratose medial portion of
the lamina (Pl. 75, f. 8), and the KOH reaction red (Aloinella is
golden yellow in KOH). The pad of filaments may not be homologous in derivation with those of Aloinella and Crossidium,
because they arise from a portion of the lamina. The genus Pseudaloina is treated here as a synonym of Crossidium ((q. v.).
Additional literature: Baczurina (1972), Barker (1904 ), Craig
(1939), Delgadillo (1973c, 1982), Koponen and Oittinen q967),
Mitten (1874), Persson (1944), Steere (1950), Weber (1979).
Number of accepted species: 12.
Species examined: A . aloides (BUF, NY), A. bifrons (NY),
A. brevirostris (BUF), A. hamulus (BUF, NY), A. rigida (BUF),
A. rosei (NY).
Plate 77.
55. ALOINELLA
Aloinella Card., Rev. Bryol. 36: 76, 1909. Type: Aloinella
catenula Card.
Barbula sect. Climacocaulon C. MUll., Linnaea 42: 329,
1879. Type: Aloinella galeata (C. MUll.) Broth. see van der
Wijk et al. (1959-69, 5: 301).
From Aloina, a genus + -ella, diminutive; resembling the genus
Aloina.
Plants growing in a turf or somewhat mixed with other species, yellowish green above, tan to dark or reddish brown
below. Stems branching often and irregularly, to 2 em in length,
transverse section rounded-pentagonal, cells of central cylinder
often large, to 25 Jlm in diameter, thin-walled, central strand
strong, occasionally additional small hydroid strands embedded
in outer cortex; sclerodermis absent, outer cortex of substereid
or occasionally stereid cells, hyalodermis absent; axillary hairs
to 9 cells in length, basal 1-3 cells thick-walled; sparsely
radiculose. Leaves appressed to incurved-catenulate when dry,
oppressed to weakly spreading when moist, short- to long-ovate
or obovate, occasionally circular or oblong, 0.4-1.0 mrn in
length, upper lamina deeply concave, margins weakly and
broadly incurved, entire or weakly denticulate at apex or
weakly serrulate above or throughout, occasionally bordered by
2-3 rows of thin-walled, hyaline cells; apex rounded, deeply
cucullate; base not differentiated . in shape; costa broad, flat,
ending a few to several cells below the apex, superficial cells
elongate dorsally, ca. 12 rows of cells across costa dorsally at
midleaf, costa often strongly and broadly decurrent at leaf base,
costal transverse section broadly reniform, stereid band single,
crescent-shaped, epidermis modified ventrally as a pad of
crowded filaments restricted to the costa, each filament uniseriate and ca. 2-6 cells in length, erect or slanted distally, occasionally hollow-papillose, epidermis not differentiated or
weakly differentiated dorsally, guide cells 4-8 in 1 layer, often
differentiated only medially on the costa, hydroid strand
present; upper /aminal cells rectangular to rhomboidal, 9-15
Jlm in width, 1(-2):1, walls evenly thickened, lumens angular,
superficially weakly convex on both exposed surfaces, occasionally bulging slightly more ventrally than dorsally, dorsal
superficial walls often thickened; papillae absent or lamina
papillose medially with 3-6 bifid, scattered papillae per lumen,
often papillose only dorsally; basal cells weakly differentiated
across base, rectangular, to l8Jlm in width, 1-4:1, walls thin to
evenly thickened. Dioicous. Perichaetia terminal, inner leaves
ovate to short-elliptical, not cucullate, to 1.9 mm in length,
sheathing, lower cells lax, thin-walled, rectangular-rhomboidal
throughout. Perigonia terminal, swollen-gemmate, paraphyses
occasionally of cells with internal longitudinal walls. Seta
0.5-1.1 em in length, 1 per perichaetium, yellowish to reddish
brown, twisted clockwise; theca 0.9-2.0 mm in length, reddish
brown, long-ellipsoidal to cylindrical, exothecial cells rectangular, thin-walled, ca. 18-28 J.lm in width, 2-3:1, stomates
phaneropore, at base of theca, annulus of 1-3 rows of
vesiculose cells; peristome teeth 16, short, entire or cleft into 2
POTIIOIDEAE
207
Plate 77. Aloinella. 1-7. A. catenula. 1. Habit. 2. Transverse section of stern showing subepidermal satellite hydroid strands. 3-5. Three leaves.
6. Leaf areolation. 7. Transverse section at rnidleaf. 8-14. A. boliviana. 8. Habit. 9-10. Two leaves. 11. Upper rnarginallarninal cells. 12-13.
Perichaetialleaves. 14. Peristorne. 15-17. A. cucullifera. 15-17. Three leaves. 18-21. A. venezuelana. 13-19. Two leaves. 20. Transverse
section at rnidleaf. 21. Upper !aminal papillae.
208
GENERA OF THE POTTIACEAE
rami, occasionally anastomosing, 110--165 ~m in length, with
several articulations, straight or nearly so, densely spiculose, basal
membrane lacking. Operculum conic-rostrate, ca. 0.4-1.0 mm in
length, cells in straight rows. Calyptra cucullate, smooth, ca. 1
mm in length. Spores 11-22 ~min diameter, brown, lightly papillose. Lamina/ KOH color reaction golden yellow.
Restricted to South America and Mexico, growing on soil at
high elevations.
This genus is relatively homogeneous (the species differ by
few but probably not insignificant characters) and has been
revised recently (by Delgadillo l975a). The striking features of
Aloinella are the appressed, deeply cucullate leaves (Pl. 77, f. 3-5,
15-17, 9-10) with ventral photosynthetic filaments arising from
and restricted to the broad, flat costa (Pl. 77, f. 7, 20). The genus
is unusual for the multiseriate-celled paraphyses of the perigonia and the small satellite hydroid strands in the stem (Pl. 77,
f. 2), these latter probably associated with the long decurrency
of the costa. Satellite strands have also been found in the stem
of Timmia (Mastracci 1993).
Additional literature: Brotherus (1911), Delgadillo
(1973b,c), Griffin (1975).
Number of accepted species: 7.
Species examined: A. boliviano (BUF, NY), A. catenula
(BUF, PC, TENN), A. cucullifera (NY), A. venezuelana
(FLAS).
Tribe POTTIEAE
Pottieae (Limpr.) Dix., Stud. Handb. Brit. Moss. 174, 1924.
Pottieae Limpr., Laubm. Deutsch!. 1: 518, 1888, rank not given.
This subclade is distinguished at the immediate ancestral node by the character states: stem short, less than 1 em in length and peristome of 32 similar rami, distinctly twisted. There is a clear evolutionary transformation from taxa with lanceolate leaves and two
stereid bands in the costa to spathulate leaves and one stereid band in the costa, paralleling that in the Hyophileae. The Pottieae is
the only suprageneric group with many genera having the seta twisted counterclockwise. The distribution of species is generally
worldwide, with a concentration in South America.
56. LEPTOBARBULA
Plate 78.
Leptobarbula Schimp., Rev. Bryol. 2: 17, 1875. Lectotype:
Leptobarbula meridionalis Schimp. fide Philibert, Rev.
Bryol. 9: 18, 1882.
Barbula sect. Leptobarbula (Schimp.) Kindb., Eur. N. Amer.
Bryin. 2: 246, 1897.
From f..£1tt6~, peeled, fine, small, thin, delicate + o + Barbula,
a genus; a small Barbula-like moss.
Plants loosely caespitose, in a thin turf, yellow-green
above, brown below. Stems seldom branching, to 3 mm in
length, rounded-pentagonal in transverse section, central strand
strong, sclerodermis absent or weakly developed, hyalodermis
absent; axillary hairs of ca. 6-7 cells, the basal 1-2 brown.
Cauline leaves much different from the more obvious perichaetial leaves, cauline leaves small, spreading-recurved to incurved
and twisted when dry, spreading-recurved when wet, ligulate,
ca. 0.7 mm in length; margins plane, entire; apex acute to
obtuse; base scarcely differentiated in shape; costa percurrent
or ending 1-2 cells below the apex, 2-3 rows of cells in width
at midleaf, superficial cells papillose, ventrally quadrate, dorsally short-rectangular, costal transverse section semicircular to
ovate, stereid bands two, the ventral often weak or absent, epidermis present on both sides, guide cells 2--4 in l layer, hydroid
strand absent; upper /aminal cells quadrate, ca. 10 ~min diameter, 1:1, walls thin, superficially flat to weakly convex; papillae bifid to multiplex, 2-4 per lumen, hollow; basal cells
differentiated across leaf, reaching higher medially, rectangular, ca. 10 ~min width, 2-3:1, walls moderately thickened to
porose. Dioicous. Perichaetia terminal, inner leaves longligulate to long-lanceolate, convolute-sheathing, to 1.5 mm in
length, with distinct, often denticulate shoulders, basal cells
long rhomboidal and thick-walled, filling most of leaf. Perigonia terminal, gemmate, inner leaves ovate-triangular. Seta
short, 0.6 to 0.9 em in length, yellow-brown, occasionally red
below, straight or twisted clockwise above, 1 per
perichaetium; theca 0.7-2.0 mm in length, red-brown, elliptical to cylindrical; exothecial cells rhomboidal, 40--45 ~m in
width, 2--4:1, walls thin; stomates phaneropore, at base of
theca; annulus strongly vesiculose, persistent or revoluble;
peristome of 32 filamentous , yellow, spiculose teeth, to ca.
300 ~m in length, of several articulations, twisted counterclockwise about once, basal membrane low, 25-50 ~m in
height, sometimes hidden by a persistent annulus, lowspiculose. Operculum long-conic to rostrate, 0.5-0.7 mm in
length, cells in counterclockwise twisted rows. Calyptra
cucullate, smooth, ca. 1.6 mm in length. Spores ca. 8 ~m in
diameter, light brown, essentially smooth. Lamina/ KOH
color reaction yellow.
The genus is of local occurrence in the Mediterranean
region of Europe, the Middle East and north Africa.
Schimper at first took this taxon to be a Seligeria by reason of its tiny size and generally slender habit (Pl. 78, f. 1--4),
but its large, filamentous, somewhat twisted peristome (Pl. 78,
f. 13) shows a clear relationship to the Pottiaceae. Characters
diagnostic of Leptobarbula are the small size of the plants,
twisted peristome teeth, convolute-sheathing perichaetial
leaves (Pl. 78, f. 12), ligulate cauline leaves with plane margins, little differentiated base, two costal stereid bands (Pl. 78,
f. 9-10), small quadrate upper lamina! cells with bifid to
multifid papillae (Pl. 78, f. 11), basal cells differentiated
mostly medially, and !aminal color in KOH yellow. Plants of
POTIIOIDEAE
Leptobarbula grow in close mixtures of perichaetiate,
perigoniate and sterile stems, which gave Casares-Gil (1932)
cause to postulate possible rhizautoicy.
Hilpert (1933) recognized a relationship with Barbula when
he made the combination Streblotrichum bericum (De Not.)
Hilp. ("in niichste Niihe von Str. bicolor"). It is indeed similar
to Barbula sect. Convolutae by the yellow setae (at least
above); twisted, spiculose peristome; the highly differentiated
perichaetial leaves; reflexed, plane-margined cauline leaves
with upper !aminal cells having thin walls and bifid to multifid
papillae; basal !aminal cells differentiated more highly in the
leaf medially than marginally; and yellow color in KOH.
Appleyard et al. (1985), however, felt that Leptobarbula was
"allied to Gymnostomum and Gyroweisia, differing apparently
only in the revoluble annulus and well-developed peristome"
(Gyroweisia usually has a revoluble annulus). The cladistic
analysis indicates that Leptobarbula is probably a relict of a
now much depleted lineage inserted near the base of the Pottioideae.
Additional literature: Corbiere (1890), Dglevskaja (1963),
Loeske (1916), Philibert (1882a,c), Pilous (1952), Whitehouse
209
and During (1987), Zander and Hermann (1986).
Number of accepted species: 1.
Species examined: L. berica (BUF, CU, DUKE, Fl, NY,
WTU).
Plate 79.
57. TETRAPTERUM
Tetrapterum Hampe ex Jaeg., Ber. S. Gall. Naturw. Ges.
1868-1869: 85, 1869. Type: Tetrapterum australe
Hampe.
From 'tt'tpd-, four+ 7t'ttp6v, feather, wing, fin, leaf; referring to the strongly ridged capsules.
Plants in low cushions or turfs, green above, brown
below. Stems branching occasionally, to 4.5 mm in length,
transverse section rounded-pentagonal, central strand very
strong, sc/erodermis absent or rarely weakly developed,
hyalodermis absent or seldom weakly differentiated; axillary
hairs rather long, of 8-10 cells, the basal 2 thicker walled.
Leaves incurved when dry, spreading when moist, oblonglanceolate to long-lanceolate, occasionally ligulate, ca.
1
Plate 78. Leptobarbula. 1-13. L. berica. 1. Habit of sporophyte-bearing plant. 2-. Perigoniate plants. 4. Sterile plant. 5-7. Three leaves. 8. Leaf
areolation. 9-10. Transverse sections near midleaf. 11. Papillae. 12. Perichaetialleaf. 13. Peristome.
210
GENERA OF THE POTTIACEAE
Plate 79. Tetrapterum. 1-11. T. tetragonum. l. Habit. 2. Transverse section of stem. 3--1. Two leaves. 5. Leaf apex. 6. Basal cells. 7.
Transverse section at midleaf. 8. Upper lamina! papillae. 9. Sporophyte. 10. Transverse section of capsule. 11. Calyptra. 12-19. T. cylindricum.
12. Habit. 13-15. lbree leaves. 16. Leaf apex. 17-18. Transverse sections near midleaf. 19. Perichaetialleaf. 20-23. T. sullivanii. 20-21. Two
leaves. 22. Leaf apex. 23. Transverse section at midleaf. 24-26. T. weymouthii. 24-25. Two leaves. 26. Leaf apex.
POITIOIDEAE
1.5-2.0 mm in length, upper lamina broadly chrumeled, margins
plane, entire; apex broadly to narrowly acute; base oblong to
broadly elliptical; costa excurrent as a sharp, smooth mucro,
usually rather broad below midleaf, superficial cells ventrally
quadrate, papillose, dorsally elongate below grading to shortrectangular above, papillose, 6-12 rows of cells across costa
ventrally at midleaf, costal transverse section semicircular to
reniform, two stereid bands present, ventral and dorsal epidermises present, the dorsal often only weakly developed, guide
cells 4-6 in 1 layer, hydroid strand present, often multiple and
occasionally on both sides of guide cells; upper /aminal cells
quadrate, occasionally hexagonal or rectangular, 8-13 J..Lm in
width, 1(-2):1, walls evenly thickened, lumens q~adrate, superficially flat to convex; papillae hollow or solid, crowded, low,
4-6 per lumen, bi-(tri-)fid; basal cells differentiated across leaf,
rectangular, slightly wider than upper cells, 3-5:1, walls thin.
Monoicous (autoicous). Perichaetia terminal, inner leaves longlanceolate from a high, often shouldered base, to 2.5 mm in
length, strongly sheathing in lower half, lower cells longrectangular to long-rhomboidal. Perigonia gemmate, as lateral
buds or terminal on a branch. Seta short, 0.5-1.5 mm in length,
1 per perichaetium, pale yellowish brown, straight, distal end of
seta with a narrow, pale band of cells forming an abscission
layer; capsule c/eistocarpous, 1.5-1.9 mm in length, yellowish
translucent brown, ovate to elliptical, nearly cylindrical in
transverse section or with 4(-8) flat sides and then often with
sharp longitudinal folds or wings when dry, exothecial cells
rectangular, ca. 20-24 J..Lm in width, 2-3: I, thin-walled
anticlinally but superficially thick-walled, spore sac easily visible and surrounded by a large air space, stomates phaneropore,
at base of theca, annulus absent, operculum and peristome
undifferentiated. Calyptra cucul/ate, smooth, 1.4-1.6 mm in
length. Spores rather large, ca. 25-28 J..Lm in diameter, yellowish, strongly spiculose to verrucose. Lamina/ KOH color reaction deep yellow to orange.
Found on bare soil in dry areas of South America, Australia,
Tasmania and South Africa.
Tetrapterum is similar in many gametophytic characters to
Barbula sect. Convolutae, particularly those species with broad
costae having quadrate ventral cells, but Tetrapterum may be
distinguished by the short seta with a distal abscission layer in a
pale band just below the capsule (Pl. 79, f. 9), and the capsule
cleistocarpous, apically blunt, often with several flat sides, and
the spore sac surrounded with a large air space. Like Aschisma
and Tortula sect. Schizophascum, the capsule tends to rupture
along distinct encircling lines of apparent weakness at the transverse exothecial cell walls. It is possible that Barbula calycina,
B. microcalycina and B. subcalycina belong with Tetrapterum
in that the gametophytes are nearly identical and no propagula
typical of Barbula sect. Convolutae are known for these. Final
disposition requires a revision of Barbula.
Andrews (1945) discussed Tetrapterum, recognizing only T.
tetragonum. He was of the opinion that of the 11 species recognized by Brotherus (1924-25), those with cylindrical capsules
are not Tetrapterum but instead belong to Astomum. I disagree
with this (which in any case would have excluded the
generitype) in part and emphasize other characters (above), thus
211
retaining T. cylindricum among other species. Catcheside
(1980) likewise recognized T. cylindricum. Andrews (1945)
felt that Tetrapterum was ultimately related to Trichostomum,
but cladistic study indicates a somewhat more distant relationship. The yellow KOH reaction does not support Hilpert's
(1933) suggestion of a relationship with Bryoerythrophyllum
(as Erythrophyllum), although it is true that the costal sections
are quite alike. The large superficial cells that are lateral to the
costa and just dorsal of the !aminal insertion (Pl. 79, f. 7, 17,
18, 23) are reminiscent of those of Pseudocrossidium.
The distal end of the capsule (Pl. 79, f. 1, 9) of all species
of Tetrapterum recognized here is broadly rounded-acute or
very bluntly apiculate, lacking the distinct rostrum or apiculus
of capsules of the reduced, cleistocarpous Trichostomum species placed in Tetrapterum by Brotherus (1924-25). Moistened capsules, even when quite mature, contain a substantial
air space around the spore sac. After abscission of the capsule
at the distal end of the seta, the capsule apparently disperses
as an integral unit possibly by floating in seasonal floods; free
capsules with a seed-like appearance are commonly found in
the loose debris within collection packets. This condition is,
however, not unique to Tetrapterum, but is most highly developed in this genus. Brotherus (1924-25) made several combinations in Tetrapterum reflecting his emendation of the genus
as including all those species of Trichostomoideae with undifferentiated opercula and long-elliptical capsules. It is apparent
that most (Tetrapterum lilliputanum is here transferred to Tortel/a) of Brotherus' combinations were of species better
viewed as species of Trichostomum at the ends of one or more
evolutionary series involving reduction of the sporophyte,
though this needs detailed evaluation at the species level.
Tetrapterum species may be distinguished from cleistocarpous Trichostomum species by a hydroid strand being visible in costal sections. The capsules of T. tetragonum are actually 4-8 sided when dry, and the corners may form longitudi;
nal, flattened ridges or low wings (Pl. 79, f. 10).
In light of the discussion above, the genus may well be
represented by only a few taxa, since the Australian species
seen during this study are rather similar, but synonymy, if
any, should await revision.
Number of accepted species: 8.
Species examined: T. cylindricum (NY), T. sullivanii (H),
T. tetragonum (BM, NY), T. weymouthii (H).
58. TRACHYCARPIDIUM
Plate 80.
Trachycarpidium Broth., Nat. Pfl. 1(3): 383, 1901. Type:
Trachycarpidium verrucosum (Besch.) Broth.
Astomum subg. Pycnocaulon (C. Mtill.) Broth., Nat. Pfl. 1(3):
384, 1901.
Acaulon sect. Pycnocaulon C. Miill., Linnaea 37: 144, 1872.
Type: Acaulon brisbanicum C. Miill.
From 'tpdxuc;, rough + Kapm5c;, fruit + -\~>tov, diminutive +
-ium, characteristic of.
Plants gregarious or forming a thin turf, light green
above, green or brownish below. Stems sometimes branchirig,
212
GENERA OF THE POTTIACEAE
Plate 80. Trachycarpidium. 1-10. T. brisbanicum. 1. Habit. 2-4. Three leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse section at midleaf. 8.
Lamina! papillae. 9. Sporophyte. 10. Calyptra. 11. T. echinatum. 11. Sporophyte and perichaetialleaf. 12-16. T. tisserantii. 12-13. Two leaves.
14. Leaf apex. 15. Transverse section at midleaf. 16. Mature capsule fallen from seta.
POTTIOIDEAE
ca. 0.3 mm in length, transverse section rounded-pentagonal,
central strand present but weak, sclerodermis absent, hyalodermis absent; axillary hairs ca. 3 cells in length, basal cell yellowish. Leaves erect-spreading, weakly incurved and occasionally somewhat twisted when dry, erect-spreading and weakly
recurved when moist, long-lanceolate, 1.5-2.0 mm in length,
upper lamina shallowly but narrowly grooved along costa, this
sometimes masked when costa is stout, margins plane, entire
(but minutely crenulate by projecting papillae), sometimes marginal cells rectangular and epapillose near apex or throughout;
apex narrowly acute; base short-ovate to oblong; costa stout,
excurrent as a thick, sharp, smooth mucro or short awn,
superficial cells elongate ventrally and elongate dorsally, ca.
4-6 rows of cells across costa ventrally at midleaf, costal transverse section semicircular to ovate, with two stereid bands,
weak or strong ventrally, strong dorsally, epidermis present and
weakly developed ventrally, absent or present and weakly
developed dorsally, guide cells 2-4 in 1 layer, hydroid strand
variably absent or present, small; upper laminal cells hexagonal
to subquadrate, occasionally short-rectangular, ca. 7-10 J.!m in
width, 1(-2): 1, walls thin to weakly evenly thickened, superficially weakly convex or strongly bulging on both sides; papillae
bifid, 2-3 per lumen, solid or hollow, crowded or somewhat
centered over lumens; basal cells differentiated across leaf base
and rising distinctly higher along margins, inflated-rectangular,
ca. 15-20 Jlm in width, 2-6:1, walls thin, hyaline. Autoicous or
rhizautoicous. Perichaetial leaves little different than the
cauline, lower cells rhomboidal to long-rectangular in lower
1/4. Perigonia terminal on small buds or plantlets at the base of
perichaetiate plants. Seta very short, ca. 75 Jlm in length, 1 per
perichaetium, hyaline, straight; theca cleistocarpous,falling off
distal end of seta, ovate to short-elliptical, 300-400 Jlm in
length plus a 150-200 J.!m sh\lfP apiculus (theca occasionally
very broad at base), with protuberances basally or throughout,
walls very light yellowish brown, translucent, columella apparently absent in mature capsules, exothecial cells with superficial
walls considerably and evenly thickened, cells hexagonal to
'short-rectangular, strongly projecting-mamillose at base or
throughout (except the apiculus), stomates phaneropore, at base
of theca, annulus absent. Calyptra campanulate to short-coniccucullate, smooth to slightly rough, 350-400 Jlm in length.
Spores ca. 25 Jlm in diameter, brown, low-spiculose. Laminal
KOH color reaction yellow.
Found on bare soil; Brazil, central Africa, Australia, New
Caledonia, New Guinea.
Trachycarpidium is a genus of pygmy mosses (Pl. 80, f. 1)
characterized by long-lanceolate, plane-margined, entire leaves
with a stout costa ending in a short awn, the basal cells
differentiated in a vee up the margins (Pl. 80, f. 6) as in Tortel/a, and bulging, strongly protuberant cells of the body (not
the apiculus) of the immersed, cleistocarpous capsule (Pl. 80, f.
9, 11, 16). It is similar to Tortella eckendorffii and Tortel/a
fruchartii, and may be viewed as an apparent reduction product
with further elaboration of the capsule; on detailed analysis,
however, the cladograms do not reflect this hypothesis. Like
Bryoceuthospora and Uleobryum, the exothecial walls of the
capsule are mamillose. They are only very light yellow in color,
213
easily transmitting the darker brown color of the spores, but
are not definitely hyaline as in Uleobryum. Trachycarpidium
also differs from Bryoceuthospora and Uleobryum in the even
thickening of the superficial walls, that is, these are without a
central lens-like spot. Superficially bulging exothecial cells
were also observed in the operculate taxa Weisiopsis
nigeriana and Weissia macrospora, apparently as a convergent development. The capsule is, in Trachycarpidium, easily
broken off the end of the short, weak seta (Pl. 80, f. 16).
Bryobartramia Sainsb. (Encalyptaceae, see Excluded
Taxa) is similar in its nearly spherical capsule, hyaline exothecial cells, large spores, short seta, papillose calyptra, and
yellow KOH reaction, but differs most significantly in the
calyptra not detaching from the vaginula and inflated as an
epigonium (Sainsbury 1948; Stone 1977a), the brown seta, the
cauline leaves with only one stereid band, and the perichaetial
leaves subulate, often consisting only of a costa.
Type material of Trachycarpidium verrucosum loaned by
PC was Weissia (subg. Phasconica) balsanae; this material,
however, was not in the original packet and was evidently
segregated from the true type material, presumably still at PC,
which is a Trachycarpidium from the illustration by Brotherus
(1924-25).
Additional literature: Dixon (1942b), Potier de la Yarde
(1928), Stone (1975).
Number of accepted species: 5.
Species examined: T. brisbanicum (BUF, HSC, NY,
MICH), T. echinatum (BM), T. lonchophyllum (NY), T.
tisserantii (BM, US).
New combination: Trachycarpidium lonchophyllum
(Roth) Zand., comb. nov. (Astomum lonchophyllum Roth,
Aussereur. Laubm. 182, 1910).
59. ASCHISMA
Plate 81.
Aschisma Lindh., Utkast Nat. Grupp. Eur. Bladm. 28, 1878.
Type: Aschisma carniolicum (Web. & Mohr) Lindh.
Phascum subg. Aschisma (Lindh.) Kindb., Eur. N. Amer.
Bryin. 2: 403, 1897.
From the alpha privative + crxfcrJ.!CX, -a·to<;, a split; referring
to the capsule lacking an operculum.
Plants very small, gregarious or forming a thin green turf.
Stems seldom branching, ca. 0.2 mm in length, transverse section rounded-pentagonal, central strand present, weak, sclerodermis not well differentiated, hyalodermis absent; axillary
hairs ca. 3 cells in length, basal cell thicker walled. Leaves
incurved and tubulose when dry, weakly spreading when
moist, oblong or triangular to short-lanceolate, often falcate
and plicate on one side, 0.6--1.0 mm in length, upper lamina
broadly channeled, margins plane to occasionally weakly
incurved, entire to sharply serrulate above midleaf, sometimes
bordered above, below or throughout by 1-3 rows of weakly
papillose, elongate cells; apex broadly or rounded-acute; base
little differentiated in shape; costa excurrent as a sharp
mucro, this occasionally rough or denticulate, superficial
cells elongate both ventrally and dorsally, 2-4 rows of cells
214
GENERA OF THE POTTIACEAE
Plate 81. Aschisma. 1-11. A. carniolicum. 1. Habit. 2-4. Three leaves. 5. Leaf apex, dorsal view. 6. Basal cells. 7. Transverse section through
leaf base. 8. Transverse section through upper leaf. 9. Sporophyte. 10. Exothecial cells. 11. Calyptra. 12-17. A. kansanum. 12. Habit. 13-15.
Three leaves. 16. Leaf apex. 17. Transverse section at midleaf.
POITIOIDEAE
across costa ventrally at midleaf, costal transverse section circular to elliptical, two stereid bands present, mostly substereid
ventrally, often also substereid dorsally, epidermis not ·
differentiated, guide cells 2-4 in 1 layer, hydroid strand apparently absent or occasionally present; upper !aminal cells quadrate to occasionally short-rectangular, 9-12 ~m in width,
1:1 (-2), walls evenly thickened, superficially flat or bulging
only ventrally or on both exposed surfaces; papillae stout, bifid,
crowded, scattered or centered over lumens, ca. 4 per lumen;
basal cells strongly differentiated, rising higher along margins
in a weak vee, rectangular, little wider than upper cells, 3-5:1,
walls very thin to evenly thickened. Monoicous. Perichaetia terminal, inner leaves long-elliptical, to 1.5 mm in length, concave
or not sheathing, lower cells oblong. Antheridia paroicous in
axils of bracts or in stalked autoicous buds at base of the
perichaetiate plant. Seta very short, ca. 50 ~m in length, 1 per
perichaetium, hyaline, straight; capsule cleistocarpous, 0.3 mm
in diameter, yellowish brown, spherical, with a small blunt
apiculus (ca. 30 ~m in length), exothecial cells rectangular,
215
mostly ca. 13-18 ~min width, 4-5:1, circling capsule longitudinally in several even bands with the appearance of a palisade, stomates absent, annulus absent; peristome teeth absent.
Calyptra conic, split once or twice, rough, 0.2-0.3 mm in
length. Spores large, 18-24 ~min diameter, yellowish brown,
essentially smooth to spiculose-papillose. Lamina/ KOH color
reaction yellow.
A genus widely distributed in the Mediterranean area and
-disjunctive to central North America; growing in dry climates
on soil, occasionally under translucent rocks in exposed situations.
The most prominent traits of this distinctive genus are the
dry habitat; small size of the plants; spherical, cleistocarpous
capsules with yellow, rectangular cells arranged in neat encircling bands, lacking stomates (Pl. 81, f. 9-10); the weak,
hyaline seta; the falcate leaves usually with a distinct border
of epapillose cells (Pl. 81, f. 5, 6, 16) and the leaf base not
differentiated in shape; and the costa with usually merely
substereid cells in two bands (Pl. 81, f. 7, 8, 17). As is the
Plate 82. Bryoceuthospora. 1-13. B. mexicana. I. Habit of sporophyte-bearing plant. 2. Perigoniate plant. 3-7. Five leaves, one enlarged. 8.
Leaf apex. 9. Upper marginal cells. 10. Basal cells. II. Transverse section at midleaf. 12. Papillae. 13. Calyptra.
216
GENERA OF THE POTTIACEAE
case with Tetrapterum and Tortula sect. Schizophascum, both
Pottioideae but rather distantly related, the capsule tends to rupture along the transverse walls at the butt ends of the exothecial
cells, which are superficially rather thick-walled.
Aschisma carniolicum and A. kansanum are fairly distinct in
the material seen. The latter (Pl. 81, f. 12-17) has more strongly
bordered and serrulate margins, more strongly bulging upper
!aminal cells with centered (not scattered) papillae, a larger,
more strongly denticulate mucro, and spores less ornamented
and toward the small end of the size range for the genus. The
protonema of A. kansanum survives arid conditions under translucent pebbles, ultimately producing leafy axes and sporophytes
peripherally (Cridland 1959); this scenario is also the case with
Bruchia brevifolia (Bruchiaceae) (cf Rushing 1989) and, at
least occasionally, Syntrichia inermis (Weger & During 1989).
Additional literature: Andrews (1915), Cardenas (1988),
Sergio (1972a).
Number of accepted species: 2.
Species examined: A. carniolicum (BUF, NY), A. kansanum
(NY).
60. BRYOCEUTHOSPORA
Plate 82.
Bryoceuthospora Crum & Anders., Bryologist 62: 66, 1959.
Type: Bryoceuthospora mexicana (Bartr.) Crum & Anders.
Ceuthospora Crum & Anders., J. Elisha Mitchell Sci. Soc. 74:
31, 1958, hom. illeg. non Fries, 1825.
From ~puov, a moss + KEOOo~. hidden + o + cmopd, seed,
spore; referring to the immersed capsules.
Plants small, gregarious, often with a persistent protonema,
yellow-green. Stems not branching, 0.5-4.0 mm in length,
transverse section rounded-pentagonal, central strand absent or
weak, outer cortex cells similar to those of central cylinder,
hyalodermis absent; axillary hairs ca. 4 cells in length, the basal
1 yellowish. Leaves few (to 12), crowded and larger above,
widely spreading and weakly contorted when dry, patent and
recurved above midleaf when moist, oblong-lanceolate to longligulate, grading to the long-lanceolate perichaetial leaves, to
1.5 mm in length, upper lamina broadly channeled above, margins broadly incurved to plane, irregularly dentate to closely
serrate above midleaf, sometimes entire except near the apex,
somewhat undulate; apex acute; base scarcely differentiated in
shape; costa strong, to 40-45 Jlm in width at midleaf, ending
2-4 cells below apex, superficial cells long-rectangular and
smooth on both sides, with several rows of cells across costa at
midleaf, costal transverse section circular to ovoid, stereid
bands two, strong, epidermis absent or weakly differentiated on
both sides, guide cells 2-4 in 1 layer, hydroid strand present but
often weak or apparently absent; upper /aminal cells quadrate
to hexagonal or short-rectangular, heterogeneous in shape,
(7-)9-12 Jlm in width, 1(-2):1, walls thin, superficially weakly
bulging; papillae large, simple, 1(-2) over each lumen, hollow
to solid, often weak or present on only a few of the medial
upper lamina! cells, basal cells differentiated across leaf base or
rising higher marginally in a vee, rectangular, mostly 9-12 Jlm
in width, 2-4:1, walls thin, somewhat bulging. Dioicous (or
possibly rhizautoicous) or monoicous. Perichaetia terminal,
inner leaves about twice the length of the cauline leaves.
Paroicous or apparently rhizautoicous and perigoniate plants
smaller than the perichaetiate, associated on the protonema.
Seta very short, 35-40 Jlm in length, 1(-2) per perichaetium,
yellow-green; theca 230-360 Jlm (apiculus 45-90 Jlm) in
length, translucent yellow to yellowish brown, the brown
spore mass evident, cleistocarpous, spherical and conicapiculate, surface smooth to co/lieu/ate, exothecial cells hexagonal to rhomboidal, thin-walled but with superficial, medial
lens-like thickenings, stomates at base of capsule, annulus not
differentiated. Calyptra conic-campanulate and distinctly 4lobed, cells smooth to bulging conic-mamil/ose, especially on
the lobes, ca. 255 Jlm in length. Spores large, oval to elliptical, 25-30(-35) Jlm in longest diameter, dark brown, densely
spiculose. Lamina! KOH color reaction yellow.
This genus is known only from Mexico and Angola,
growing on soil.
Bryoceuthospora is quite similar in general appearance to
Trachycarpidium of Africa and the Australasian region (see
Stone 1975) and to Aschisma of central Europe and North
American prairie states. It differs gametophytically from both
in the longer leaves (Pl. 82, f. 8) and low, unipapillose upper
!aminal cells (Pl. 82, f. 12). Trachycarpidium has similarly
mamillose cells of the calyptra (Pl. 82, f. 13), but the cells of
the exothecium of that genus are more strongly protuberant.
Important features of Bryoceuthospora (holotype: Mexico,
Sinaloa, Bartram 505, FH) are the dentate, broadly channeled
leaves, margins nearly plane (not strongly and narrowly
incurved as in most Weissia species), the strong costa, the
cleistocarpous capsule with convex exothecial cells, and the
(often) rough calyptra. The rather translucent exothecial cell
walls relate it to Uleobryum, which has entirely colorless cell
walls but a rather different gametophyte. Ephemerum species
(both species of Bryoceuthospora were originally described in
Ephemerum) look much like Bryoceuthospora, and may even
have similarly unipapillose leaves (e.g. E. cohaerens (Hedw.)
Hampe) with two (faint) stereid bands, but the leaves of that
genus are more strongly serrate, the costa is rather thin, the
exothecial cells are not bulging, and the calyptrae are smooth,
distinctly campanulate rather than conic-campanulate. Bryoceuthospora aethiopica, recently reported from Mexico
(Cardenas 1988), is distinguishable from B. mexicana by its
paroicous sexual condition, weak central strand, cauline
leaves about half the length of the perichaetial and more
strongly papillose (pluripapillose in the lower leaves), and
smooth calyptrae; in spite of the species' name, the type (BM)
was apparently collected near Golungo Alto, now in Angola.
Additional literature: Crum & Anderson (1958b).
Number of accepted species: 2.
Species examined: B. aethiopica (BM), B. mexicana (FH).
New combination: Bryoceuthospora aethiopica (Welw. &
Dub.) Zand., comb. nov. (Ephemerum aethiopicum Welw. &
Dub., Mem. Soc. Phys. Hist. Nat. Geneve 21: 443, 1871;
Aschisma aethiopica (Welw. & Dub.) Lindb.).
POTfiOIDEAE
217
Plate 83. Uleobryum. 1-10. U. peruvianum. 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Basal cells. 6. Transverse section at midleaf. 7. Laminal
papillae. 8. Sporophyte. 9. Exothecial cells. 10. Calyptra. 11-19. U. curtisii. 11. Habit. 12-15. Four leaves. 16. Leaf apex. 17. Basal cells. 18.
Transverse section at midleaf. 19. Sporophyte.
218
GENERA OF THE POTTIACEAE
Plate 83.
61. ULEOBRYUM
Uleobryum Broth., Hedwigia 45: 271, 1906. Type: Uleobryum
peruvianum Broth.
Named for Ernst H. G. Ule, 1854-1915, a German collector of
South American plants + o + ~puov, a moss.
Plants very short, gregarious or forming a thin turf, green
above, brown below. Stems branching occasionally, 1-2 mm in
length, transverse section rounded-pentagonal, central strand
present, sclerodermis absent, hya1odermis absent; axillary hairs
ca. 4 cells, basal 1 brownish. Leaves appressed-incurved when
dry, spreading when moist, spathulate to obovate or elliptical,
ca. 1 mm in length, upper lamina narrowly grooved along
costa, margins plane, minutely crenulate-serrulate by projecting papillae and cell walls, marginal row of cells often smooth
and rectangular; apex acute to rounded, mucronate; base
scarcely differentiated in shape; costa percurrent to shortexcurrent, occasionally ending 1-2 cells below apex, superficial
cells elongate ventrally and dorsally, 2-6 rows of cells across
costa ventrally at midleaf, costal transverse section round to
semicircular, with two stereid bands (the ventral occasionally
absent), epidermis variably present or absent ventrally, absent
dorsally, guide cells 2-4 in 1 layer, hydroid strand usually
present, often small; upper !aminal cells hexagonal or
subquadrate to short-rectangular, 8-13 flm in width, 1(-2):1,
walls thin to evenly thickened, superficially convex on both
sides, sometimes more convex ventrally; papillae simple or
bifid, 2-4 per cell, occasionally indistinct, mostly hollow; basal
cells differentiated across leaf base, somewhat higher along
margins, rectangular, slightly inflated, to 20 flm in width,
mostly 3-4:1, walls thin, hyaline. Autoicous. Perichaetia terminal, inner leaves with base often oblong, otherwise little different from the cauline leaves, not or little sheathing, lower cells
little different from those of the cauline leaves. Perigonia
present as small axillary buds on perichaetiate plants. Seta very
short, ca. 30--40 flm in length, 1(-2) per perichaetiate plant,
hyaline, straight; theca spherical with a strong, conical
apiculus, ca. 350--400 flm (plus 110-150 flm for apiculus) in
diameter, c/eistocarpous, glistening, transmitting the brown
color of the contained spores, exothecial cells hexagonal,
hyaline (entirely transparent, colorless), mamillose, superficial
walls centrally thickened and lens-/ike, columella apparently
absent at maturity, stomates ca. 4, phaneropore, at base of theca,
annulus absent, operculum not differentiated. Calyptra coniccampanulate, split once or twice, cells slightly rough to strongly
mamillose, ca. 250-300 fJ.m in length. Spores large, ca. 25-33
flm in diameter, brown, closely low-spiculose. Lamina/ KOH
color reaction yellow.
Found on bare soil; West Indies, Mexico, Peru, Brazil and
Australia.
Uleobryum is quite like Bryoceuthospora in its small size,
spherical capsules with hyaline setae (Pl. 83, f. 8, 19), the transparent capsule walls that are thickened on the surface to form
bulging lenses (Pl. 83, f. 9) and which transmit the color of the
large spores, the conic-campanulate calyptrae that are usually
rough or pustulate apically (Pl. 83, f. 10), the leaves serrulate
(very weakly so in Uleobryum), and the costa with two stereid
bands (Pl. 83, f. 6, 18) and elongate superficial cells ventrally.
It differs significantly, albeit not altogether satisfactorily,
from Bryoceuthospora in the shorter, lingulate to elliptical
leaves with pluripapillose leaf cells (Pl. 83, f. 7). The two
genera are quite close; see also Cladogram 14. Trachycarpidium has very lightly yellow-brown colored exothecial cells
and the gametophyte is similar in areolation to that of U.
curtisii (Pl. 83, f. 11-19), but it differs in a teardrop-shaped
capsule that is strongly pustulate, and the exothecial cells are
evenly thickened superficially, not just medially as in U/eobryum. Uleobryum occultum of Brazil differs little from the
Australian U. curtisii, being slightly more strongly serrulate
on the leaf margins by projecting cell walls.
Additional literature: Cardenas (1988), Stone (1984,
1985).
Number of accepted species: 3.
Species examined: U. curtisii (MELU), U. occultum
(SPA), U. peruvianum (H, NY).
New heterotypic synonymy: Phascum brittoniae Crum &
Steere (nom. nov. for Phascum sessile E. Britt.) = Uleobryum
peruvianum Broth.
New combination: Uleobryum occultum (Roth) Zand.,
comb. nov. (Aschisma occultum Roth, Aussereur. Laubm.
173, 1911).
62. TORTULA
Plates 84-89.
Tortula Hedw., Sp. Muse. 122, 1801, nom. cons. non
Roxburgh, 1800. Lectotype: Tortula subulata Hedw.
Beccaria C. Mtill., Nuov. Giorn. Bot. Ital. 4: 11, 1872.
Bauriella Wamst., Hedwigia 57: 88, 1915, nom. inval. prov.
Type: Tortula polyseta (C. Mtill.) Warnst.
Tortula sect.? Piliferae De Not., Mem. R. Ace. Sc. Torino
40: 287, 1838, rank not indicated; inoperative in piority
I.C.B.N. Art. 35.2.
Barbula sect. Amphidiopsis C. Mtill., Linnaea 42: 332,
1879. Type: Barbula amphidiifolia C. Mtill.
Barbula sect. Pilifera Lazaro e lbiza, Bot. Descr. Comp. Fl.
Esp. 1: 586, 1896.
Barbula sect. Orthopodiae Kindb., Eur. N. Amer. Bryin. 2:
245, 1897.
Barbula sect. Catillaria C. Mtill., Gen. Muse. Fr. 425, 1900.
Type: Barbula pel/ata Schimp.
Pottia sect. Beccaria (C. Mtill.) C. Mtill., Gen. Muse. Fr.
389, 1900.
Portia subsect. Acutae C. Jens., Skand. Bladmfl. 203, 1939,
nom. inval. descr. suec.
See sectional synonymy for additional nomenclature.
From tortus, twisted + -uta, diminutive; referring to the
twisted peristome teeth.
Plants forming cushions or turfs, green or occasionally
blackish green above, yellow-brown to dark brown below.
Stems branching occasionally, to 2 em in length, transverse
section rounded-pentagonal, central strand present or very
rarely absent, sclerodermis absent, hyalodermis absent;
axillary hairs ca. 5-8 cells in length, basal 1-3 cells thicker
POTIIOIDEAE
walled; rhizoids often dense. Leaves appressed-incurved to lax
when dry, weakly to widely spreading when moist, usually
obovate to spathulate, occasionally ovate to elliptical or
ligulate, 1-4(-6) mm in length, upper lamina nearly flat to concave, broadly channeled, occasionally grooved along costa,
margins recurved below or rarely plane, entire or occasionally
weakly serrulate near apex, marginal 1-4 rows of cells often
less papillose and smaller than the medial or walls thicker,
occasionally marginal cells elongate, rarely bistratose; apex
broadly acute to rounded; base scarcely differentiated in shape
to elliptical, rarely weakly auricled; costa short- to longexcurrent as an awn, occasionally percurrent or subpercurrent,
costa with lamina inserted laterally or to 45', superficial cells
quadrate or occasionally short-rectangular and papillose or
smooth ventrally, dorsally short-rectangular to elongate and
papillose or smooth, 3-4(-5) rows of cells across costa ventrally at midleaf, costal transverse section circular to semicircular, ventral stereid band absent or occasionally small and represented by a few cells, dorsally present and round, elliptical or
semicircular in shape, epidermis present ventrally and dorsally
or occasionally only laterally on the dorsal side, rarely absent
dorsally, guide cells 2(-3) in 1(-2) layers or rarely absent,
hydroid strand usually present, often large, very rarely absent;
rarely an elliptical pad of cells bulging from ventral surface of
the costa; upper /aminal cells rounded-quadrate to hexagonal,
occasionally rhomboidal, ca. 15-19 jlm in width, 1-2:1, walls
thin or seldom evenly thickened, superficially convex; papillae
usually hollow, simple or bifid, 4-6 per lumen, occasionally on
a conical salient, rarely absent; basal cells differentiated across
leaf or higher medially, rectangular, often rather lax, 18-25 jlm
in width, 2-5: 1, walls thin, hyaline, rarely little differentiated.
Propagula absent. Dioicous or monoicous (commonly
autoicous or paroicous). Perichaetia terminal, inner leaves little
differentiated or somewhat larger than the cauline. Perigonia
terminal or as autoicous buds in subperichaetial or lower leaf
axils. Seta very short or to 2.5 em in length, I (very rarely 2)
per perichaetium, yellowish brown to brown, twisted counterclockwise, clockwise or not twisted; theca stegocarpous or else
cleistocarpous, 0.5-3.0(-7.0) mm in length, yellowish brown to
dark brown, spherical, ovate, elliptical or cylindrical, occasionally inclined, occasionally macrostomous, exothecial cells rectangular, 25-30 IJ.m, ca. 2-3:1, rarely 4-5:1, walls thin or
evenly thickened, stomates present at base of theca,
phaneropore, annulus of 1-2 rows of vesiculose cells, persistent
or very rarely revoluble, occasionally absent or rarely with up to
8 circumferential weak lines of dehiscence; peristome teeth of
32 filaments or 16 triangular teeth or rudimentary or absent,
long or shortly triangular, cleft to near base, spiculose, up to
2000 jlm in length, with many articulations, straight to twisted
counterclockwise, basal membrane absent or low or up to 1000
jlm in height, tessellated and spiculose. Operculum when
differentiated long-conic, occasionally shortly rostrate, 0.5-2.5
mm in length, cells twisted counterclockwise. Calyptra
cucullate, smooth, 2.5-6.0 mm in length. Spores 13-30(-50)
jlm in diameter, light brown, papillose, rarely densely spiculose.
Lamina/ KOH color reaction usually yellow, occasionally red
medially, occasionally negative, rarely reddish orange.
219
Reported chromosome numbers: Sect. Tortula: n = 13+m, 14,
24, 26, 27, 28, 30, 39, 40, 48, 48+m, 50, 52, 60, 66. Sect.
Pottia: n = 12, 13, 15, 20, 21, 24, 25, 26, 26+m, 27,
28+1-2acc, 30, 32, 42, 52. The most often reported number
for both sections is n = 26.
Found on most continents in various habitats, mainly soil.
With the segregation of various genera (Zander 1989:
Chenia, Dolotortula, Hennediella, Hilpertia, Sagenotortula,
Stonea, Syntrichia), Tortula becomes a fairly homogeneous
group with very similar gametophytes and a characteristic tendency to reduction in the sporopltyte. Thus, the correlations of
season of sporophyte maturation (Zander 1979d) with genus
(Pottia having spring and winter sporophyte maturation dates
and Tortula in the traditional sense having dates mainly in the
spring and summer) is probably a reflection of "life strategy"
(cf During 1979) rather than phylogeny, though this needs to
be tested. Major characters of Tortula as presented here
include presence of stem central strand (rarely absent or
present in different stems of same collection, e.g. T.
brevissima) and absence of sclerodermis and hyalodermis (Pl.
84, f. 2, 17); leaves usually obovate to spathulate, margins
usually narrowly recurved below and entire; costal stereid
band usually semicircular to rounded in section, hydroid
strand present, dorsal epidermis usually present (Pl. 84, f. 7, 8,
21); upper !aminal cells usually rather large and clear (i.e.
walls usually relatively unobscured by the papillae, Pl. 84, f.
9); propagula absent; upper !aminal KOH reaction usually
yellow.
Tortula is distinguishable from Syntrichia by the semicircular to rounded stereid band (not crescent-shaped) and yellow KOH reaction of the upper !aminal cells (not red). Unlike
Syntrichia, a dorsal costal epidermis is usually differentiated,
either completely or occasionally only laterally (Pl. 89, f.
15-as is also the case in Hennediella) . The yellow KOH
reaction is usually present, but some few taxa have no color
reaction (e.g. T. entosthodontacea), or the ba~al cells may be
brick red (e.g. T. raucopapil/osa), or the leaves may blush red
medially in the upper part of the leaf (e.g. T. atrovirens), or
the older leaves may be red and the younger yellow (e.g. T.
lingulata), or all leaves may have a reddish orange cast (e.g.
T. nevadensis). But the characters of these taxa otherwise are
those of Tortula as here emended. Hennediella is distin"
guished by its red KOH reaction, commonly dentate or serrate
and plane upper lamina! margins, and superficially flattened
upper !aminal cells. Papillae may be absent in some Tortula
species or variously expressed in different specimens of the
same species.
Chamberlain (1978) recognized Pottia caespitosa, but the
leaves have plane margins, the upper lamina! cells are rather
small and thick-walled, and costal sections show two stereid
bands. This small-statured species is actually a Trichostomum
(as witness the combination Trichostomum caespitosum
(Bruch ex Brid.) Jur.), differing somewhat from other species
of that genus by the broadly sheathing perichaetialleaves and
quite neckless capsule.
Upon examination of the sporophytes of a considerable
range of species of Pottieae, there was found to be no sharp
220
GENERA OF THE POTIIACEAE
Plate 84. Tortula. 1-15. T. subulato. 1. Habit, with sporophyte. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7.
Transverse section at midleaf. 8. Transverse section near leaf apex. 9. Upper lamina! papillae. 10. Perichaetialleaf. 11. Autoicous bud. 12.
Theca. 13. Portion of basal membrane and peristome. 14. Operculum. 15. Calyptra. 16-22. T. atherodes. 16. Habit. 17. Transverse section of
stem. 18-19. Two leaves. 20. Leaf apex, lateral view. 21. Transverse section at midleaf. 22. Sporophyte.
POITIOIDEAE
221
Plate 85. Tortula. 1-4. T. altipes. 1-2. Two leaves. 3. Transverse section at midleaf. 4. Portion of peristome. S-8. T. atrovirens. 5-6. Two
leaves. 7. Leaf apex. 8. Transverse section at midleaf. 9-14. T. canescens. 9. Habit. 10-11. Two leaves. 12. Leaf apex. 13. Transverse section at
midleaf. 14. Peristome. 15-20. T. cernua. 15. Leaf. 16. Leaf apex. 17. Transverse section at midleaf. 88. Theca with peristome. 19. Portion of
peristome. 20. Calyptra. 21-25. T. cuneifolia var. blissii. 21. Habit. 22-23. Two leaves. 24. Transverse section at midleaf. 25. Peristome.
26-29. T. deciduidentata. 26-27. Two leaves. 28. Leaf apex. 29. Transverse section at midleaf.
222
GENERA OF THE POTTIACEAE
difference between traditional "Tortula" peristomes with 32
similar rami (Pl. 84, f. 12-13) and "Desmatodon" peristomes
with 16 teeth cleft to near the base or to a basal membrane (i.e.,
32 paired rami, Pl. 86, f. 16; 87, f. 9-10). Even if some difference was statistically demonstrable, it would cut across
observed (and as taxonomically recognized here) clearly
defmed generic groupings based on several gametophytic characters. It is simpler to entertain convergence of one character
(short, probably reduced peristomes having teeth paired) than
convergent evolution of several distinctive gametophyte
morphotypes two or more times. The flattened basal portion of
"Desmatodon" peristome teeth may be explained as a cleft portion of the basal membrane. The proximally flat teeth of Desmatodon species are also different from the almost terete filaments of Tortula with long, twisted peristomes simply because
the long filaments of Tortula have distal regions little wider
than their thickness. A cladistic evaluation at the species level
may clarify this.
Visotska's (1967) proposal of a subfamily Tortuloideae (no
type cited), based on a chromosome number of x = 12 and
intended to contain Tortula, Aloina and Crossidium, was criticized on cytological grounds by Nyholm and Wigh (1973)
because several species of Tortula have a basic chromosome
number of x = 13. Evaluation of chromosome counts given by
Fritsch (1982, 1991) gave both 12 and 13 as basic numbers for
both Tortula s. str. and Syntrichia as conceived in the present
study; however, Newton (1972) found n = 7 for S. robusta and
Ramsay (1974) found n = 6+m for S. papil/osa.
An electron microscopical study by Lewinsky (1974) of
spore ornamentation in 10 European species of Tortula s. lat.
showed differences between the spores of the specimens studied
(only one or two collections were examined for each species
although 20-30 spores from two to five capsules were studied
in each species), which probably represent differences between
the species, but she found no evidence of differences between
traditional sections of the genus.
Mishler's (1986a) cladogram of postulated phylogenetic
relationships of several species of Tortula s. lat. recognized
Tortula s. str. (as recognized here) as a primitive group (he
listed T. subulata, T. mucronifolia and T. muralis) distinct from
several other species (all recognized here as Syntrichia) by the
upper laminal cells not strongly mammilose.
Corley et al. (1981) gave an up-to-date presentation of the
sections of the genus as represented in Europe. Their apprehension of Desmatodon as a rather small assemblage of the type
species and closely related taxa presages the present study.
They stated that Desmatodon "is not defmed by sound technical
characters, and there has been much confusion about which species should be assigned to it. As with Didymodon there has been
too much emphasis placed on the peristome, which is not a conservative character in Pottiaceae."
Tortula muralis, T. leucostoma and T. altipes occasionally
have a small ventral (sub)stereid band (Pl. 85, f. 3). The costa
is, however, rounded in section and generally unlike that of
Barbula.
Selected literature on Tortula s. str. (here including Phascum, Desmatodon and Portia) : Arts (1987a,b, 1988), Bachelot
(1813), Bennett (1965), Brown (1894b,c), Bryan (1956),
Carri6n et al. (1990), Chamerlain (1978), Crundwell (1953,
1955, 1956), Dixon (1927b), Favali and Gianni (1973),
Guerra et al. (1988, 1991, 1992), Hausler (1984), Hernnstadt
and Heyn (1989), Holzinger (1925), Hughes (1969, 1979,
1982), Hughes and Wiggin (1969), Jimenez et al. (1990),
Kanda (1981), Lazarenko (1969, 1974), Lazarenko and
Lesnyak (1972), Lazarenko et al. (1961), Lewinsky (1974),
Lightowlers (1984, 1985a,c, 1986a,b,c), Lobachevskaya et al.
(1986), Matteri (1977a,b), Mishler (1985b, 1986a, 1990),
Mishler and Newton (1988), Ripetskij (1978, 1979),
Ripetskij et al. (1983), Risse (1985), Rumsey (1992), Rungby
(1957), Sainsbury (1936), Saito (1973a), Savicz-Ljubitzkaya
and Smirnova (1963b, 1965), Sergio (1972a, 1978), Springer
(1935), Steere (1939a, 1940), Stone (1989), Toth (1987),
Ulycna (1977), Wareham (1939a), Wareham and Whitney
(1939), Warnstorf (1912, 1916), Zander (1989).
Number of accepted species: 163, none remaining in Desmatodon, plus 10 as yet undistributed in Phascum, plus 30
undistributed in Portia.
Species examined: see below.
New heterotypic synonymy: Didymodon schimperi
(Mont.) Broth.= Tortula atrovirens (Sm.) Lindh.
New taxa, combinations, statuses and names:
Tortula altipes (Broth.) Zand., comb. nov. (Desmatodon
altipes Broth., Act. Hort. Bot. Ac. Sc. U.R.S.S. 42: 154,
1931).
Tortula argentinica (Broth.) Zand., comb. nov. (Desmatodon
argentinicus Broth., Ark. Bot. 15(6): 5, 1918), not seen.
Tortula atherodes Zand., nom. nov. (Phascum cuspidatum
Schreb. ex Hedw., Spec. Muse. 22, 1801).
Tortula atherodes var. arcuata (Herrnstadt & Heyn) Zand.,
comb. nov. (Phascum cuspidatum var. arcuatum
Herrnstadt & Heyn, Bryologist 94: 175, 1991), not seen.
Tortula atherodes var. affinis (Nees & Hornsch.) Zand.,
comb. nov. (Phascum affine Nees & Hornsch., Bryol.
Germ. 1: 74, 1823; Phascum cuspidatum var. affine (Nees
& Hornsch.) Hampe), not seen.
Tortula atherodes var. curviseta (Dicks.) Zand., comb. nov.
(Phascum curvisetum Dicks., Pl. Crypt. Brit. 4: 2, 1801;
Phascum cuspidatum var. curvisetum (Dicks.) Nees &
Hornsch.), not seen.
Tortula atherodes var. diaphora (Hag.) Zand., comb. nov.
(Phascum acaulon var. diaphorum Hag., K. Norsk. Vid.
Selsk. Skrift. 1928(3): 19, 1929; Phascum cuspidatum var.
diaphorum (Hag.) C. Jens.), not seen.
Tortula atherodes var. elata (Brid.) Zand., comb. nov. (Phascum elatum Brid., J. Bot. (Schrader) 1800(1): 269, 1801;
Phascum cuspidatum var. e/atum (Brid.) Drumm.), not
seen.
Tortula atherodes var. intertexta (Brid.) Zand., comb. nov.
(Phascum intertextum Brid., Mant. Muse. 8, 1819; Phascum cuspidatum var. intertextum (Brid.) Brid., not seen.)
Tortula atherodes var. marginata (Hernnstadt & Heyn) Zand.,
comb. nov. (Phascum cuspidatum var. marginatum
Hernnstadt & Heyn, Bryologist 94: 175, 1991), not seen . .
POITIOIDEAE
Tortula atherodes var. mitraeformis (Limpr.) Zand., comb. nov.
(Phascum cuspidatum var. mitraeforme Limpr., Laubm.
Deutsch!. 1: 187, 1885), not seen.
Tortula atherodes var. papillosa (Lindb.) Zand., comb. nov.
(Phascum papillosum Lindb., Oefv. K. Vet. Ak. Foerh. 21:
217, 1864; Phascum cuspidatum var. papillosum (Lindb.)
Roth); Phascum cuspidatum ssp. papillosum Guerra & Ros
in Guerra, Jimenez, Ros & Carri6n), not seen.
Tortula atherodes var. pilifera (Hedw.) Zand., comb. nov.
(Phascum piliferum Scherb. ex Hedw., Spec. Muse. 20,
1801; Phascum cuspidatum var. piliferum Scherb. ex
Hedw.) Hook. & Tayl.).
Tortula atherodes var. retortifolia (Guerra & Ros in Guerra,
Jimenez, Ros & Carri6n) Zand., comb. nov. (Phascum
cuspidatum var. retortifolium Guerra & Ros in Guerra,
Jimenez, Ros & Carri6n, Cryptogamie, Bryol. Lichenol. 12:
390, 1991),'not seen.
Tortula atherodes var. schreberiana (Dicks.) Zand., comb. nov.
(Phascum schreberianum Dicks., Pl. Crypt. Brit. 4: 2, 1801;
Phascum cuspidatum var. schreberianum (Dicks.) Brid.),
not seen.
Tortula atrovirens var. leucodonta (Corb.) Zand., comb. nov.
(Barbula atrovirens var. leucodonta Corb., Mem. Soc. Sc.
Nat. Cherbourg 26: 244, 1889; Desmatodon convolutus var.
/eucodontus (Corb.) Wijk & Marg.), not seen.
Tortula bogosica (C. Mtill.) Zand., comb. nov. (Desmatodon
bogosicus C. Mtill., Nuov. Giom. Bot. Ital. 4: 12, 1872).
Tortula capillaris (Chen) Zand., comb. nov. (Desmatodon
capil/aris Chen, Hedwigia 80: 287, 1941), not seen.
Tortula cernua var. xanthopus (Kindb.) Zand., comb. nov.
(Desmatodon cernuus var. xanthopus Kindb., Ottawa Natural. 4: 61, 1890), not seen.
Tortula cuneifolia var. blissii Zand., var. nov. (Pl. 85, f. 21-25.)
A varietate typica gametophytis subatris, in solutione KOH
colorem profundiorem evolventibus, foliis rigide appressis,
seta curta, crassa, longitudine 4-5 mm, latitudine 0.2-0.3
mm dijfert.
Differs from the typical variety by the blackish gametophytes, these more strongly colored in KOH; leaves stiffy
appressed; seta short, ca. 4-5 in length, stout, 0.2-0.3 mm in
width. Type: Canada, Northwest Territories, Cornwallis
Island, Resolute Bay area, L. C. Bliss, 1977, holotype, BUF;
isotype, ALTA. The leaves of var. blissii are identical in
morphology to those of muticous-leaved forms of the European species T. cuneifolia, but show a very strong color
reaction to KOH: bright yellow upper lamina and deep
brick-red basal cells, colors which are pale in European
specimens at BUF. The short, thick seta is apparently
unique to this arctic variety; European specimens have setae
0.7-1.5 mm in length and ca. 0.15 mm in width. Like European specimens, the capsule is variably macro- and
microstomous, and the operculum is broadly to narrowly
conic. This specimen was incorrectly reported (Vitt &
Zander 1978) as a second known station for Crumia
deciduidentata (here treated as a species of Tortula near T.
cuneifolia), which has a similar short, thick seta and
smooth, weakly bordered leaves that are bright yellow in
223
KOH except for brick-red basal cells, but which differs in
its capsule about twice as long (2.7-3.3 mm vs. 1.3-1.6
mm) and proportionately thicker; operculum pushed off
by the elongating columella; spores larger (ca. 18 jlm vs.
ca. 13 jlm); leaves ovate to clearly spathulate (vs. short- to
long-ovate); constricted leaf apex; and much enlarged
basal cells. A section across the leaf base of var. blissii
shows what appear to be two stereid cell groups of equal
size separated by a group of hydroid cells (also seen in
sections of the basal portion of the costa of Tortula
muralis, and see discussion of Phascopsis).
Tortula entosthodontacea (Card. & Dix.) Zand., comb. nov.
(Hyophilopsis entosthodontacea Card. & Dix., J. Bot. 49:
137, 1911.).
Tortula euryphylla Zand., nom. nov. (Dicranum latifolium
Hedw., Spec. Muse. 140, 1801; Desmatodon /atifolius
(Hedw.) Brid.).
Tortula euryphy/la ssp. brevifolia (Kindb.) Zand., comb. nov.
(Tortula latifolia (Hedw.) Lindb. ssp. br.evifolia Kindb.,
Bib. K. Svensk. Vet. Ak. Handl. 7(9): 135, 1883; Desmatodon latifolius ssp. brevifolius (Kindb.) Kindb.).
Tortula euryphy/la var. eucalyptrata (Lindb.) Zand., comb.
nov. (Tortula eucalyptrata Lindb., Bot. Not. 1886: 100,
1886; Desmatodon latifolius varr eucalyptratus (Lindb.)
Kaur.).
Tortula euryphy/la var. jlavescens (Brid.) Zand., comb. nov.
(Dicranum latifolium var. jlavescens Brid., Spec. Muse.
140, 1801; Desmatodon latifolius var.jlavescens).
Tortula euryphy/la var. spelaea (Amann) Zand., comb. nov.
(Desamtodon spelaeus Amann, Bull. Murithienne 40: 46,
1919; Desmatodon latifolius var. spelaeus (Amann)
Podp.).
Tortula euryphy/la var. subobliqua (Lindb.) Zand., comb. nov.
(Desmatodon latifolius var. suboliquus Lindb., Oefv. K.
Vet. Ak. Foerh. 23: d553, 1867).
Tortula chungtienia Zand. , nom. nov. (Desmatodon
yuennanensis Broth., Symb. Sin. 4: 44, 1929).
Tortula deciduidentata (Sharp & Iwats.) Zand., comb. nov.
(Crumia deciduidentata Sharp & Iwats., J. Hattori Bot.
Lab. 32: 95, 1969).
Tortula kabir-khanii (Broth.) Zand., comb. nov. (Desmatodon
kabir-khanii Broth., Mitteil. lnst. Allg. Bot. Hamburg 8:
400, 1931), not seen.
Tortula lanceola Zand., nom. nov. (Encalypta lanceolata
Hedw., Spec. Muse. 63, 1801; Pottia lanceolata (Hedw.)
C. Mtill.; non Tortula lanceolata (Hedw.) P. Beauv.).
Tortula lanceola var. albidens (Corb.) Zand., comb. nov.
(Pottia lanceolata var. albidens Corb., Rev. Bryol. 22: 35,
1895), not seen.
Tortula lanceola var. angustata (B.&S. in BSG) Zand., comb.
nov. (Anacalypta lanceolata var. angustata B.&S. in
BSG, Bryol. Eur. 2: 48, 1843; Pottia lanceolata var.
angustata (B.&S. in BSG) C. Mtill.), not seen.
Tortula lanceola var. lejolisii (Corb.) Zand., comb. nov.
(Pottia lanceolata var. lejolisii Corb., Mem. Soc. Sci. Nat.
Cherbourg 26: 238, 1889), not seen.
Tortula lanceola var. /eucodonta (Schimp.) Zand., comb. nov.
224
GENERA OF THE POTTIACEAE
Plate 86. Tortula. 1-10 T. entosthodontllcea. 1. Habit. 2-4. Two leaves. 5. Leaf apex. 6. Upper marginal cells showing papillae. 7. Basal cells.
8. Transverse section at midleaf. 9. Portion of rudimentary peristome. 10. Operculum. 11-16. T. euryphylla. 11-13. Three leaves. 14. Leaf apex.
15. Transverse section at midleaf. 16. Peristome. 17-22. T. lanceola. 17-18. Two leaves. 19. Leaf apex, lateral view. 20. Transverse section at
midleaf. 21. Autoicous bud. 22. Peristome. 23-25. T. transcaspica. 23-24. Two leaves. 25. Leaf apex.
POTIIOIDEAE
225
Plate 87. Tortulo. 1-4. T. /oureri. 1-2. Two leaves. 3. Leaf apex. 4. Transverse section at midleaf. 5-10. T. leucostoma. 5-6. Two leaves. 7.
Leaf apex. 8. Transverse section at midleaf. 9-10. Two examples of peristome teeth. 11-17. T. lingu/ota. 11. Habit. 12. Transverse section of
stem. 13-15. Two leaves. 16. Leaf apex. 17. Two transverse sections at midleaf. 18-22. T. marginata. 18-19. Two leaves. 20. Leaf apex. 21.
Transverse section at midleaf. 22. Peristome. 23-27. T. porteri. 23-24. Two leaves. 25. Leaf apex. 26. Peristome. 27. Theca, operculum and
calyptra.
226
GENERA OF THE POTTIACEAE
(Pottia lanceolata var. leucodonta Schimp., Syn. ed. 2:
158, 1876), not seen.
Tortula lanceola var. macrophylla (Warnst.) Zand., comb. nov.
(Pottia lanceolata var. macrophylla Warnst., Hedwigia 58:
133, 1916), not seen.
Tortula lanceola var. microphylla (Warnst.) Zand., comb. nov.
(Pottia lanceolata var. microphylla Warnst., Hedwigia 58:
134, 1916), not seen.
Tortula lanceola var. mucronata (Amann) Zand., comb. nov.
(Pottia lanceolata var. mucronata Amann, Bull. Soc.
Vaudoise Sci. Nat. 53: 85, 1920), not seen.
Tortula lanceola var. ovalifolia (Warnst.) Zand., comb. nov.
(Pottia lanceolata var. ovalifolia Warnst., Hedwigia 58:
131, 1916), not seen.
Tortula lanceola var. papillosa (Corb.) Zand., comb. nov.
(Pottia lanceolata var. papillosa Corb., Mem. Soc. Sci. Nat.
Cherbourg 26: 237, 1889), not seen.
Tortula lanceola var. rigidior (Schwaegr.) Zand., comb. nov.
(Encalypta lanceolata var. rigidior Schwaegr., Spec. Muse.
Suppl. 1(1): 61, 1811.)
Tortula laureri var. setschwanica (Broth.) Zand., comb. nov.
(Desmatodon setschwanicus Broth., Symb. Sin. 4: 43, 1929;
Desmatodon laureri var. setschwanicus (Broth.) Chen), not
seen.
Tortula maritima (R. Br. ter) Zand., comb. nov. (Dendia
maritima R. Br. ter, Trans. New Zealand Inst. 30: 411,
1898; Pottia maritima (R. Br. ter) Broth.).
Tortula minor (C. Miill.) Zand., comb. nov. (Beccaria minor C.
Miill., Nuov. Giom. Bot. Ital. 4: 11, 1872; Pottia minor (C.
Miill.) Wijk & Marg.), not seen.
Tortula minor var. elatior (C. Miill.) Zand., comb. nov.
(Beccaria elatior C. Miill., Nuov. Giom. Bot. Ital. 4: 11,
1872; Pottia minor var. elatior (C. Miill.) Wijk & Marg.).
Tortula modica Zand., nom. nov. (Gymnostomum intermedium
Tum., Muse. Hib. 7, 1804; Pottia intermedia (Tum.)
Fuernr.).
Tortula modica var. corsa (Fleisch. & Warnst.) Zand., comb.
nov. (Pottia intermedia var. corsa Fleisch. & Warnst., Bot.
Centralbl. 65: 299, 1896), not seen.
Tortula modica var. gymnandra (Schiffn.) Zand., comb. nov.
(Pottia intermedia var. gymnandra Schiffn., Oesterr. Bot.
Zeitschr. 47: 55, 1897), not seen.
Tortula modica var. gymnogyna (Schiffn.) Zand., comb. nov.
(Pottia intermedia var. gymnogyna Schiffn., Oesterr. Bot.
Zeitschr. 48: 389, 1898), not seen.
Tortula modica var. revoluta (Schiffn.) Zand., comb. nov.
(Pottia intermedia var. revoluta Schiffn., Oesterr. Bot.
Zeitschr. 47: 55, 1897), not seen.
Tortula modica var. stenocarpa (Velen.) Zand., comb. nov.
(Pottia intermedia var. stenocarpa Velen., Rozpravy Cesk.
Ak. Ved. Tr. 2, 6(6): 148, 1897), not seen.
Tortula modica var. tenuis (Vent.) Zand., comb. nov. (Pottia
intermedia var. tenuis Vent., Muscin. Trent. 31, 1899), not
seen.
Tortula nevadensis (Card. & Ther.) Zand., comb. nov. (Pottia
nevadensis Card. & Ther., Bot. Gaz. 37: 365, 1904).
Tortula pal/ida (Lindb.) Zand., comb. nov. (Pottia pal/ida
Lindb., Oefv. K. Vet. Ak. Foerh. 21 : 220, 1864).
Tortula pal/ida var. longicuspis (Warnst.) Zand., comb. nov.
(Pottia pal/ida var. longicuspis Warnst., Hedwigia 58:
113, 1916).
Tortula porteri (Jam. in Aust.) Zand., comb. nov.
(Desmatodon porteri Jam. in Aust., Musci Appal. 123,
1870).
Tortula protobryoides Zand., nom. nov. (Phascum bryoides
Dicks., Pl. Crypt. Brit. 4: 3, 1801; Pottia bryoides (Dicks.)
Mitt.).
Tortula protobryoides var. brevifolia (De Not.) Zand., comb.
nov. (Phascum bryoides var. brevifolium De Not., Atti
Univ. Genova 1: 734, 1869; Pottia bryoides var. brevifolia
(De Not.) Wijk & Marg.), not seen.
Tortula protobryoides var. thornhillii (Wils.) Zand., comb.
nov. (Phascum bryoides var. thornhillii Wils., Bryol. Brit.
33, 1855; Pottia bryoides var. thornhillii (Wils.)
Braithw.), not seen.
Tortula randii (Kenn.) Zand., comb. nov. (Pottia randii
Kenn., Rhodora 1: 78, 1899; Desmatodon randii (Kenn.)
Laz.).
Tortula raucopapillosa (X.-j. Li) Zand., comb. nov. (Desmatodon raucopapillosus X.-j. Li, Acta Bot. Yunnan. 3:
105, 1981 "raucopapillosum").
Tortula rhodonia Zand., nom. nov. (Desmatodon wilczekii
Meyl., Bull. Soc. Vaudoise Sc. Nat. 52: 383, 1919), not
seen.
Tortula sainsburyana Zand., nom. nov. (Pottia stevensii R. Br.
ter, Trans. N. Z. Inst. 26: 291, 1894).
Tortula solomensis (Broth.) Zand., comb. nov. (Desmatodon
solomensis Broth., Rev. Bryol. n. ser. 2: 2, 1929), not
seen.
Tortula splachnoides (Homsch.) Zand., comb. nov. (Phascum
splachnoides Homsch., Horae Phys. Berol. 57, 1820;
Pottia splachnoides (Homsch.) Broth.).
Tortula thompsonii (C. Miill.) Zand., comb. nov. (Trichostomum thompsonii C. Miill., Bot. Zeit. 22: 359, 1854; Desmatodon thompsonii (C. Miill.) Jaeg.).
Tortula tonkinensis (Besch.) Zand., comb. nov. (Desmatodon
tonkinensis Besch., Bull. Soc. Bot. France 41: 80, 1894),
not seen, cf Brotherus (1924: 297).
Tortula truncata var. brevirostris (Lisa) Zand., comb. nov.
(Gymnostomum truncatum var. brevirostre Lisa, Elenco
Muschi Torino 16, 1837; Pottia truncata var. brevirostris
(Lisa) De Not.), not seen.
Tortula truncata var. illyrica (Latz.) Zand., comb. nov. (Pottia
illyrica Latz., Beih. Bot. Centralbl. 48(2): 481, 1931;
Pottia truncata var. illyrica (Latz.) Podp.), not seen.
Tortula truncata var. littoralis (Mitt.) Zand., comb. nov.
(Pottia littoralis Mitt., J. Bot. 9: 4, 1871; Pottia truncata
var. littoralis (Mitt.) Warnst.), not seen.
Tortula truncata var. minutissima (Warnst.) Zand., comb. nov.
(Pottia truncata var. minutissima Warnst., Hedwigia 58:
117, 1916), not seen.
Tortula ucrainica (Laz.) Zand., comb. nov. (Desmatodon
ucrainicus Laz., Bull. Jard. Bot. Kieff 4: 34, 1926).
Tortula willisiana Zand., nom. nov. (Phascum drummondii
POTTIOIDEAE
Wils., London J. Bot. 7: 26, 1848; Pottia drummondii
(Wils.) Willis).
Tortula willisiana var. obscura (Willis) Zand., comb. nov.
(Pottia drummondii var. obscura Willis, Viet. Nat. 70: 171,
1954), not seen.
Tortula wilsonii (Hook.) Zand., comb. nov. (Gymnostomum
wilsonii Hook., Bot. Misc. 1: 143, 41, 1829; Portia wilsonii
(Hook.) BSG).
Tortula wilsonii var. asperula (Mitt.) Zand., comb. nov. (Portia
asperula Mitt., J. Bot. 9: 4, 1871; Portia wilsonii ssp.
asperula (Mitt.) Kindb.), not seen.
Tortula wilsonii var. crinita (Wils. ex B.&S.) Zand., comb. nov.
(Portia crinita Wils. ex B.&S. in BSG, Bryol. Eur. 2: 43,
1843; Portia wilsonii var. crinita (Wils. ex B.&S.) Wamst.),
not seen.
Tortula wilsonii var. mucronifolia (Bruch in F. A. Miill.) Zand.,
comb. nov. (Entosthymenium mucronifolium Bruch in F. A.
Mtill., Flora 12: 387, 1829; Portia wilsonii var. mucronifolia
(Bruch in F. A. Mtill.) Wamst.), not seen.
Tortula zoddae Zand., nom. nov. (Portia cuneifolia Solms ex
Schimp., Syn. ed 2, 154, 1876).
TORTULA Sect. TORTULA
Tortula sect. Tortula Hedw., Sp. Muse. 122, 1801, nom. cons.
non Roxburgh, 1800. Lectotype: Tortula subulata Hedw.
Desmatodon Brid., Mant. Muse. 86, 1819. Lectotype: Desmatodon latifolius (Hedw.) Brid. fide Venturi, Comm.
Fauna Fl. Gea Venezia 1: 123, 1868.
Zygotrichia Brid., Bryol. Univ. 1: 520, 1826. Type:
Zygotrichia leucostoma (R. Br.) Brid.
Dermatodon Htib., Muse. Germ. 14: 109, 1833,p.p.
Pachyneurum Amann, F. Mouss. Suisse 2: 112, 1912.
Tortula subg. Desmatodon (Brid.) Lindb., Musci Scand. 20,
1879.
Tortula subg. Zygotrichia (Brid.) Lindb., Musci Scand. 20,
1879.
Tortula subg. Tortula C. Jens., Medd. Groen1and 3: 379,
1887, nom. illeg.
Didymodon subg. Desmatodon (Brid.) Kindb., Eur. N. Amer.
Bryin. 2: 273, 1897.
Barbula subg. Zygotrichia Kindb., Eur. N. Amer. Bryin. 2:
245, 1897.
Barbula subg. Tortula Kindb., Eur. N. Amer. Bryin. 2: 246,
1897, hom. illeg.
Tortula subg. Pachyneurum (Amann) C. Jens., Skand.
Bladmfl. 200, 1939.
Portia subg. Pseudodesmatodon Medel. in C. Jens., Skand.
Bladmfl. 207, 1939, nom. inval. descr. suec.
Tortula subg. Eutortula C. Jens., Skand. Bladmfl. 199, 1939,
nom. illeg.
Barbula sect. Tortulae Rebent., Prodr. Fl. Neomarch. 257,
1804.
Tortula sect. Subulatae De Not., Mem. R. Ace. Sc. Torino 40:
287, 1838. Type: Tortula subulata Hedw.
Barbula sect. Subulatae B.&S. in BSG, Bryol. Eur. 2: 98,
1842 (fasc. 13-15 Mon. 36).
Trichostomum sect. Desmatodon (Brid.) C. Miill., Syn. 1:
227
588, 1849.
Barbula sect. Crassinerves (De Not.) Milde, Bryol. Soles.
112, 1869. Type: Barbula nervosa (BSG) Milde.
Tortula sect. Desmatodon (Brid.) Mitt., J. Linn. Soc. Bot.
12: 145, 164, 1869.
Tortula sect. Zygotrichia (Brid.) Mitt., J. Linn. Soc. Bot. 12:
145, 168, 1869.
Barbula sect. Crassicostatae Schimp., Syn. ed. 2: 194,
1876.
Desmatodon sect. Eudesmatodon Jur., Laubmfl. Ost.
Ungam 129, 1882, nom. illeg.
Desmatodon sect. Crassicostati Jur., Laubmfl. Ost.-Ungam
135, 1882. Type: Desmatodon atrovirens (Sm.) Jur.
Desmatodon sect. Subulati Jur., Laubmfl. Ost. Ungam 129,
1882. Lectoype nov. : Desmatodon subulatus (Hedw.) Jur.
Portia sect. Didyctium C. Miill. in Broth., Hedwigia 34:
124, 1895. Type: Portia asperula C. Mtill. in Broth.
Barbula sect. Crassinervia Lazaro e lbiza, Bot. Oeser.
Comp. Fl. Esp. 1: 506, 1896, p.p. Crossidium & p.p.
Aloina.
Barbula sect. Muraliformes Kindb., Eur. N. Arner. Bryin. 2:
246, 1897. Type: Barbula muralis (Hedw.) Crome.
Barbula sect. Canescentes Kindb., Eur. N. Amer. Bryin. 2:
245, 1897. Type: Barbula canescens (Mont.) B.&S.
Barbula sect. Limbatae Kindb., Eur. N. Amer. Bryin. 2:
246, 1897. Type: Barbula marginata B.&S.
Barbula sect. Subulataeformes Kindb., Eur. N. Amer.
Bryin. 2: 245, 1897. Type: Barbula subulata (Hedw.) P.
Beauv.
Barbula sect. Tortula Herib., Mem. Ac. Sc. ClermontFerrand ser. 2, 14: 364, 1899, nom. illeg.
Tortula sect. Tortula Broth., Nat. Pfl. 1(3): 429, 1902, nom.
illeg.
Tortula sect. Eutortula Broth., Act. Hort. Gothoburg 1: 191,
1924, nom. illeg.
Syntrichia sect. Zygotrichia (Brid.) Monk., Laubm. Eur.
306, 1927.
Tortula sect. Crassicostatae (Schimp.) Podp., Consp. Muse.
Eur. 242, 1954.
Tortula sect. Crassinerves (Milde) Wijk & Marg., Taxon 8:
75, 1959. Type: Barbula nervosa (De Not.) Milde, hom.
illeg.
Tortula subsect. Desmatodon (Brid.) Braithw., Brit. Moss.
Fl. 3: 212, 1885.
Upper !aminal cells usually densely papillose and relatively
small (ca. 10-13 J.Lm in width), ventral costal cells 3-6 in
transverse section, usually forming a convex pad and usually
densely hollow papillose with simple or bifid papillae, leaves
usually with rounded apices and commonly with long hyaline
awns; costal sections with guide cells; peristomes usually
present.
Lindberg (1864) treated T. subulata and T. mucronifolia
as a single taxon. Steere ( 1940) pointed out that, although the
variable (Wamstorf 1912) T. subulata and T. mucronifolia
intergrade extensively in Europe, the two species appear to
have have distinct ranges and morphology in North America.
228
GENERA OF THE POTTIACEAE
Plate 88. Tortula. 1-9. T. maritima. 1. Habit. 2-3. Two leaves. 4. Leaf apex. 5. Two transverse sections at midleaf. 6-8. Capsule walls
dehiscing along circumferenciallines. 9. Spores. 10-14. T. mucronifolia. 10-11. Two leaves. 12. Leaf apex. 13. Transverse section at mid1eaf.
14. Peristome. 15-18. T. platyphylla. 15-16. Two leaves. 17. Leaf apex. 18. Peristome. 19-25. t. protobryoides. 19. Habit. 20-21. Two leaves.
22. Leaf apex. 23. Transverse section at midleaf. 24. Calyptra. 25-27. T. randii. 25-26. Two leaves. 27. Leaf apex.
POTIIOIDEAE
229
•'
Plate 89. Tortula. 1-4. T. r~~olvens. 1-2, Two leaves. 3. Leaf apex. 4. Transverse section at midleaf. 5-9. T. solmsii. 5-6. Two leaves. 7. Leaf
i .. apex. 8. Transverse section. at midleaf. 9. Peristome. 10. T. splachnoides. 10. Transverse section at midleaf. 11-15. T. truncata. 11. Habit.
· 12-13. Two leaves. 14. Leaf apex. 15. Transverse section at midleaf. 16-20. T. willisiana. 16. Habit. 17. Leaf. 18. Leaf apex. 19. Transverse
section at midleaf. 20. Calyptra. 21-25. T. wilsonii. 21. Habit. 22-23. Two leaves. 24. Leaf apex. 25. Transverse section at midleaf.
230
GENERA OF THE POTTIACEAE
Chen (1941) commented, significantly as it turns out, that T.
mucronifolia (as Syntrichia mucronifolia) has a leaf morphology quite like that of many Pottia species. This is similarly
unfortunate for nomenclatural purposes, since the closely
related T. subulata is the type of the genus and therefore of the
sect. Tortula. In the present study, the variation between T.
subulata and T. mucronifolia was found to parallel major differences in lamina! structure between sect. Tortula and sect.
Pottia. Thus, in spite of an apparent close relationship, these
two species are placed in different sections of the genus. This is
not unacceptable if the two are considered taxa whose ancestors
are inserted near the base of the lineage of Tortula species; this
needs to be analysed at the species level.
Tortula sect. Tortula includes species with papillose leaves
and small cells (e.g. T. muralis) that are nearly identical with
species with larger, smooth !aminal cells (e.g. T. ca/ifornica and
T. transcaspica, Pl. 86, f. 23-25), but these last are distinctly
awned and placed in sect. Tortula on that account. Perhaps T.
subulata, with its short awn, ought to be placed with the species
of section Pottia (a name change at the sectional level would be
necessary), but T. subulata otherwise has the bulk of characters
given above for the section. However, many species of sect.
Tortula with the !aminal morphology ofT. muralis, the species
probably most representative of sect. Tortu/a, also have short
awns or lack them altogether. The evidence that sect. Pottia
demonstrates far more sporophyte reduction than sect. Tortula
may be interpreted either as an additional character supporting
sect. Pottia as a distinctive line of evolution (just as Syntrichia
characteristically lacks sporophyte reduction) or that smooth
cells and reduced sporophytes are correlated at the end of several separate reduction series from sect. Tortula. In any case,
this should be further analysed at the species level.
Steere (1939a) felt that Tortula leucostoma (Pl. 87, f. 5-10)
and T. guepinii would better be regarded as subspecies of T.
euryphylla (all as Desmatodon spp., the last as D. latifolius).
Tortula revolvens (Pl. 89, f. 1-4) has the innermost cells of
its revolute upper margins hollow-papillose, the group somewhat differentiated as a photosynthetic organ like that of Hilpertia and species of Pseudocrossidium; T. revolvens is, however, closely related to T. muralis by the costa round in section,
with a single strong stereid band, and upper lamina highly
papillose, cells superficially flat, and KOH reaction yellow.
Tortula atrovirens (Pl. 85, f. 5-8) is near T. revolvens in
short-ovate leaf shape, thickened costa, and upper marginal
cells epapillose. Tortula muralis has a similarly thickened costa,
likewise often with two apparent stereid bands. Tortula
atrovirens is similar to Crossidium by its habit of low-growing,
gemmiform plants with spiralling ovate to obovate leaves, the
development of specialized ventral costal photosynthetic tissue
(the cells of which have lumens vertically elongate in section,
as was nicely illustrated by Flowers 1973a), the rounded dorsal
stereid band, the medial placement of the simple, hollow papillae on the leaf, upper lamina yellow but with a reddish blush
medially on the leaf in KOH, and a general similarity to Crossidium aberrans and C. seriatum of the areolation (upper cells
ca. 15-18 f.!m in width), the thickened dorsal superficial upper
median cell walls, and other features. Delgadillo (1975a, p. 282)
proposed that Crossidium was derived, along with Aloina,
from a Tortula or Desmatodon-!ike ancestor through elaboration of ventral costal outgrowths and reduction of leaves and
sporophytes. The cladograms do not support this otherwise
cogent inference, either placing Crossidium in a different tribe
(the Hyophileae, Cladograms 14-16) or having the immediate
ancestors of Crossidium inserted deeper in the cladogram than
those of Tortula (see Cladograms 9, 10 and 13). The costal
sections of C. aberrans and T. atrovirens are extraordinarily
similar. Tortula atrovirens has sporophyte maturation dates
(Zander 1979d) more characteristic of Crossidium than species previously placed in Desmatodon. As in the extension of
the Pseudocrossidium evolutionary series (Zander 1979f) to
include P. revolutum, which has no specialized ventral
photosynthetic tissue, it may be possible to include T.
atrovirens in Crossidium to fill it out as a natural genus. On
the other hand, Crossidium sect. Pseudocrossidium (no
nomenclatural relation to the genus Pseudocrossidium), which
includes C. aberrans and other species with similar costal filaments, may be more appropriately included in Tortula, perhaps as a section or recognized as a separate genus in its own
right. This needs further evaluation.
Tortula bogosica, because of its rather deep ventral costal
groove and lack of dorsal costal epidermal cells, actually may
well be Barbula indica or a closely related species lacking the
ventral stereid band; this also requires further study. The
little-papillose Tortula californica is placed in sect. Tortula
largely because of its evident relationship with T.
transcaspica, especially through the rounded leaf apex and
long, hyaline awn, but has the rather large upper !aminal cells
of sect. Pottia; the same is true for T. grandiretis, which has
somewhat longer leaves than the first two.
Species of sect. Tortula examined: T. a/tipes (H), T.
atrovirens, T. bogosica (NY), T. brevipes (BUF), T.
brevissima (US), T. ca/ifornica (BUF), T. canescens (BUF),
T. euryphylla, T. grandiretis (NY), T. guepinii (CANM), T.
leucostoma (NY), T. Ungulata (NY), T. peruviana (NY), T.
marginata (BUF), T. muralis, T. obtusifolius, T. platyphylla
(NY), T. p/inthobia, T. raucopapi/losa (NY), T. revolvens
(NY), T. sublimbata (NY), T. subulata (BUF), T.
thianschanica (H), T. trachyphylla (H), T. transcaspica (H),
T. vahliana (NY), T. wilsonii (BUF, NY).
TORTULA Sect. POTTIA
Tortula sect. Pottia (Ehrh. ex Reichenb.) Kindb., Bih. K.
Svensk. Vet. Ak. Hand!. 7(9): 131, 1883. Lectotype:
Pottia truncata (Hedw.) BSG, see Wareham in Grout,
Moss Fl. N. Amer. 1: 197, 1939.
Phascum L. ex Hedw., Spec. Muse. 19, 1801. Lectotype:
Phascum cuspidatum Hedw., see Wareham in Grout,
Moss Fl. N. Amer. 1: 195, 1939.
Anacalypta Rohl. ex Leman, Diet. Sci. Nat. ed. 2, 2 Suppl.
38, 1816. Type: Anaca/ypta /anceolata (Hedw.) Nees &
Hornsch.
Physedium Brid., Bryol. Univ. 1: 51, 1826. Type: Physedium splachnoides (Hornsch.) Brid.
POITIOIDEAE
Portia (Ehrh. ex Reichenb.) Ehrh. ex Fi.irnr., -Flora 12(2 Erg.):
10, 1829.
Mildeella Limpr., Laubm. Deutschl. 1: 191, 1885, hom. illeg.
non Trevisan, 1876. Type: Mildeella bryoides (Dicks.)
Limpr.
Mildea Warnst., Krypt. Fl. Brandenburg 2: 82, 1904, hom.
il/eg. non Grisebach, 1866.
Phascum subg. Phascum Sull. in A. Gray, Man. Bot. N. U.S.
ed. 2: 615, 1856.
Portia subg. Anacalypta (Rohl. ex Leman) Boul., Fl. Crypt.
Est Muscin. 509, 1872.
Tortula subg. Portia (Ehrh. ex Reichenb.) Lindb., Musci
Scand. 21, 1879.
Pottia subg. Eupottia Boul., Muscin. France 471, 1884, nom.
illeg.
Phascum subg. Euphascum Limpr., Laubm. Deutschl. 7: 185,
1885, nom. illeg.
Phascum subg. Mildee/la Kindb., Eur. N. Amer. Bryin. 2:
403, 1897. Type: Phascum bryoides Dicks.
Pottia subg. Portia (Ehrh. ex Reichenb.) Broth., Nat. Pfl 1(3):
423, 1902, hom. illeg.
Pottia subg. Mildee/la (Kindb.) Broth., Nat. Pfl. 1(3): 423,
1902.
Gymnostomum sect. Pottia Ehrh. ex Reichenb., Consp. Regn.
Veg. 1: 33, 1828.
Barbula sect. Cuneifoliae BSG, Bryol. Eur. 2: 93, 1842 (fasc.
13-14 Mon. 31).
Phascum sect. Annua Angstr. in Fries, Summ. Veg. Scand. 1:
97, 1846.
Pottia sect. Eupottia C. Miill., Syn. 1: 550, 1849, nom. illeg.
Pottia sect. Anacalypta (Rohl. ex Leman) C. Miill., Syn. 1:
547, 1849, nom. illeg. prior. ut gen.
Tortula sect. Cuneifoliae (B.&S. in BSG) Spruce, Ann. Mag.
Nat. Hist. ser. 2, 3: 375, 1849. Type: Tortula cuneifolia
(With.) Tum.
Trichostomum sect. Anacalypta (Rohl. ex Leman) C. Miill.,
Linnaea 42: 312,316, 1879.
Desmatodon sect. Cuneifolii (BSG) Jur., Laubmfl. Oest.
Ungarn 129, 1882. Lectotyp. nov.: Desmatodon cuneifolius
(With.) Jur.
Barbula sect. Camptopodiae Kindb., Eur. N. Amer. Bryin. 2:
245, 1897. Type: Barbula laureri (Schultz) Kindb.
Pottia sect. Mildeella (Kindb.) Monk., Laubm. Eur. 324,
1927.
Phascum sect. Euphascum Podp., Consp. Muse. Eur. 222,
1954, nom. illeg. Basionym: Phascum subg. Euphascum
Limpr., nom. illeg.
Pottia sect. Pseudodesmatodon Podp., Consp. Muse. Eur.
232, 1954, nom. inval. Type: Pottia randii Kenn.
Pottia sect. Portia (Ehrh. ex Reichenb.) Nyholm, Ill. Fl. Nordic Mosses 2: 81, 1989, nom. superfl.
Upper lamina! cells usually smooth or weakly papillose and
large (ca. 15-20 J..Lm in width), ventral costal cells 2-3 in transverse section, c&nmonly bulging individually or arranged in
longitudinal rowsas low lamellae and usually smooth or simply
once- or twice-papillose, leaves usually with acute apices and
231
short yellow-brown awns (rarely hyaline awned); costal section with guide cells; peristomes often rudimentary or absent.
Magill (1981) pointed out that Acaulon and Phascum are
not easily distinguishable in South Africa by the traditional
characters of size of capsule beak, curvature of leaf margins
and the papillosity of the !aminal cells. He defined Phascum
by its narrower, plane-margined, erect to spreading leaves,
and emergent capsule and cucullate calyptra, distinguishing
Acaulon by the bulbiform habit, broad, concave leaves and
small, mitrate calyptra. Acaulon is recognized here as a distinctive genus. Chamberlain (1978) suggested that Phascum
and Pottia are linked through Pottia recta and Pottia bryoides
and that it would be taxonomically more satisfactory if the
two genera were united. The two genera are here
synonymized with Tortula s. str., but Portia recta belongs
with Microbryum.
Holzinger (1925) thought Portia randii to be a
depauperate form of Desmatodon cernuus; this is also
reflected in the combination D. randii (Kenn.) Laz. of
Lazarenko (1963b). These two (Pl. 85, f. 15-20; 88, f. 25-27)
are kept together here in sect. Pottia. These and other species,
like T. laureri (Pl. 87, f. 1-4) and T. thompsonii, with bordered, mostly plane and weakly serrulate margined leaves and
large, superficially flat upper !aminal cells are quite like species of Hennedie/la, a genus otherwise only distinguished
from Tortula by commonly plane and serrate !aminal borders
and red !aminal KOH reaction. These species of sect. Pottia
(often placed in Desmatodon but not including the type of that
genus) commonly have summer sporophyte maturation dates
(Zander 1979d) rather than the more usual winter and spring
dates of traditional Tortula species, and, on revision, may
prove worthy of a section of their own.
Tortula lanceola (Pl. 86, f. 17-22) is similar to various
Crossidium species in the weakly twisted peristome of 16
split, triangular teeth, the short, bulging cells of the ventral
surface of the upper costa, the smooth areolation, and the
!aminal KOH reaction yellow with a red blush in the upper
medial portion of the leaf. It differs mainly in the narrow
recurvature of the !aminal margins and very weak development of the ventral costal bulge, not quite sufficient for one to
consider it a specialized photosynthetic organ. Another species similar to Crossidium, T. atrovirens, is here placed with
sect. Tortula.
Certain specimens of small-statured species with basal
!aminal cells commonly reddish in KOH (e.g. Tortula
truncata, Pl. 89, f. 11-15) may react red throughout the lamina giving the impression of the genus Microbryum; these are
apparently from unusually harsh environments, and usually
may be assigned to Tortula by a lack of !aminal papillae.
Species of sect. Pottia examined: T. atherodes, T. cernua
(NY), T. chungtienia (H, NY), T. cuneifolia (ALTA, BUF,
NY), T. deciduidentata (NY), T. modica (BUF), T. lanceo/a
(BUF), T. laureri (BUF, NY), T. minor (NY), T. mucronifolia
(BUF), T. nevadensis (US), T. pal/ida (NY), T. paulsenii (H),
T. planifolia (NY), T. protobryoides (BUF), T. randii (H,
NY), T. sainsburyana (NY), T. solmsii (BM), T. systylia
(NY), T. thompsonii (NY), T. truncata, T. ucrainica (NY), T.
232
GENERA OF THE POTTIACEAE
websteri (US), T. zoddae (NY).
TORTULA Sect. SCHIZOPHASCUM
Tortula sect. Schizophascum (C. Mtill.) Zand., comb. nov.
Phascum sect. Schizophascum C. Mtill., Flora 71: 6, 1888.
Type: Phascum disrumpens C. Mtill.
Dendia R. Br. ter, Trans. New Zealand lnst. 30: 411, 1898.
Type: Dendia maritima R. Br. ter
Pottia subg. Schizophascum (C. Mtill.) Broth., Nat. Pfl. 1(3):
423, 1901.
Portia sect. Schizophascum (C. Mtill.) Wareham in Grout,
Moss Fl. N. Amer. 1(4): 197, 1939.
Costal section lacking guide cells, and stereid band distinctly
central in the section, otherwise gametophyte similar to that of
sect. Portia; sporophyte with elongate seta and elliptical capsule, this being cleistocarpous or dehiscing along up to 8 circumferential lines, usually along weakened cell walls at butt
ends of longitudinally elongate rectangular cells (ca. 3-5: I)
arranged in even rows (or "pallisades").
Found on soil, often near the sea; South Africa, Australia
and New Zealand.
Brotherus (1902-09) sunk the monotypic Dendia (type species Dendia maritima R. Br. ter = Tortula maritima, Pl. 88, f.
1-9) into Pottia subg. Schizophascum, a move approved by
Dixon ( 1923 ), who regarded the curious tattered capsule
dehiscence of the two known collections as a merely
"unhealthy, not to say thoroughly rotten condition" associated
with "more or less complete immersion of the capsules from
time to time in sea-water." Tortu/a willisiana, Pl. 89, f. 16--20,
from Australia, however, has similar wall weakenings in its
exothecial cells. Characters allowing recognition at least at the
section level are the elongate, palisade-like exothecial cells that
have weak walls along the butt ends allowing dehiscence along
up to 8 circumferenciallines in at least two of the three species,
and the lack of guide cells in the costa in all species. The
hydroid strand is sometimes absent in some leaves of species in
this section. Species of other sections of Tortula, if cleistocarpous, dehisce along irregular lines, these not distinctly
arranged circumferencially, and have guide cells in their costae.
The type of Tortula sect. Schizophascum, Pottia disrumpens
(C. Mtill.) Broth., was referred to T. willisiana (as Portia
drummondii) by Willis (1954), a species very similar or the
same as T. maritima. Tortula sect. Schizophascum may be
apprehended as an end member of a reduction series through
sect. Pottia. This may possibly involve ancestors of T.
cuneifolia, which has nearly identical gametophytes but does
have guide cells, but not ancestors of the cleistocarpous T.
atherodes (Pl. 84, f. 16--22) (which, although it has a minute
seta and spherical capsule, has more strongly awned, narrower,
little concave leaves with a less lax areolation). Involved would
be loss of the costal guide cells and elaboration of a unique
dehiscence feature in some species. The spores are very large
(ca. 35-40 Jlm diam.), b11t this is a common condition of other
cleistocarpous (e.g. T. atherodes) and eperistomate species (e.g.
T. nevadensis). Note, however, that the various cladograms
demonstrate that reduced austral taxa are commonly relicts of
lineages inserted deeply on the tribal or subfamilial subclade.
Thus, sect. Schizophascum may be better recognized as a separate genus. Again, further analysis at the species level is
called for.
Tortula maritima (syntype NY! New Zealand, "Godley
Heads"), of which only two collections are known, is not
operculate, but neither is it exactly cleistocarpous since capsules are cleanly ruptured along the central one or two of five
to eight weak annular lines encircling the capsule from near
the base to near the apex. In nearly mature but undehisced
capsules, these annular lines are easily seen as minute cleavages between the cells in lateral view around the outline of the
somewhat flattened capsule on a microscope slide. No exothecial cells are differentiated, however, as smaller, specialized annular cells as in the case with stegocarpous mosses.
The exothecial cells are rectangular and have much the same
"pallisade" arrangement as those of Aschisma and Tetrapterum, (see Cladograms 14-16, but compare with other dadograms in the study). In Aschisma rectangular exothecial cells
(ca. 3-4: 1) are considered unusual because the capsule is
spherical, and, in taxa with spherical capsules, the exothecial
cells are generally short-rhomboidal or short-rectangular (e.g.
as in T. atherodes).
A significant character of all species here referred to sect.
Schizophascum is the costal section showing a central region
of stereid cells, this often including a ventrally or centrally situated hydroid strand, the stereid band surrounded by a ring of
substereid cells, then surrounded again by an epidermis of
thin-walled and often bulliform cells; thus, the guide cells are
either absent or somehow hidden amoung similar cells of different developmental origin. The costal section is somewhat
like that of Stegonia in the bulliform ventral cells and flattened dorsal surface, and similar to that of Saitoel/a, which
lacks guide cells and has bulliforin ventral epidermal cells,
but also differs in lacking a hydroid strand.
Magill (1981, p. 209) suggested that Tortula maritima of
New Zealand, T. splachnoides (Pl. 89, f. 10) of South Africa
(as Pottia) and T. willisiana of Australia (as Pottia
drummondii) may constitute a single, albeit variable
circumantarctic taxon (see also Catcheside 1980). He did not,
however, synonymize them, but referred South African specimens previously identified as T. maritima to T. sp/achnoides
(as Portia). An authentic specimen ofT. willisiana (NY!) has
the characters of T. maritima: concave leaves, upper !aminal
cells either epapillose or papillose medially, the costal section
with centrally located stereid band, and the sporophytes,
though immature, show a pallisade arrangement of cells with
weak breaks along circumferencial lines. On the other hand,
an isotype (NY! one plant) of Tortula splachnoides has exothecial cells ca. 2:1 in dimension, not in a pallisade arrangement, and breaks in the capsule wall are across cells, not
along cell walls; the costal section, however, is like that of the
other two species in the stereid (substereid) band positioned
centrally and lacking guide cells. Tortula splachnoides is
included here with other species (pending further study)
because of the like costal section, the fact that immature capsules of T. maritima have rather short exothecial cells, and in
POTTIOIDEAE
view of the short, concave, nearly smooth leaves. The non-type
specimens identified as T. splachnoides at BM have plane
leaves with rather papillose laminae (except for a smooth border), and a distinct guide cell layer with the stereid band not
central, but rather situated dorsally, thus there may be more
than one taxon referred to T. splachnoides in herbaria; these
non-type and as yet unidentified specimens, although cleistocarpous, lack a pallisade arrangement of exothecial cells but
have similarly large spores. The number of accepted species in
this section is three.
Additional literature: Brown (1898b).
Species of sect. Schizophascum examined: Tortula maritima
(NY), T. splachnoides (NY), T. willisiana (NY).
TORTULA Sect. HYOPHILOPSIS
Tortula sect. Hyophilopsis (Card. & Dix.) Zand., comb. et stat.
nov.
Hyophilopsis Card. & Dix., J. Bot. 49: 137, I9Il. Type: Hyophilopsis entosthodontacea Card. & Dix.
Near Tortula sect. Pottia but differing by the annulus being
revoluble, or persistent but strongly developed and bulgingreflexed when well wetted. Additional characters of value in
identification are upper laminal cells 2-3: I, usually weakly
papillose, costa percurrent, margins narrowly bordered by elongate cells, autoicous or rhizautoicous sexual condition; perigonia terminal on small, loosely foliate stems at base of
perichaetiate plants; seta elongate and capsule cylindrical but
peristome teeth rudimentary, consisting of several nubbins
barely rising above a low basal membrane which itself scarcely
exceeds the annulus; operculum conic, cells very weakly
twisted counterclockwise.
A single rare taxon found on walls, mortar, and lateritic
rocks in India's Western Ghats.
Hyophilopsis was found to be weakly distinguishable from
Tortula sect. Pottia, mainly by the peristome reduction unaccompanied by seta and capsule reduction (Pl. 86, f. 1-lO). The
gametophyte of this monotypic genus (type: India, Western
Ghats, Sedgwick 119, isotypes, NY, PC) is much like that of
Tortula laueri.
Literature: Dixon (19lla).
Species examined: T. entosthodontacea (NY, PC, US) .
63. WILLIA
Plates 90-91.
Willia C. Mtill. in Neum., Deutsch. Exp. Int. Polarforsch. 2:
3Il, I890. Type: Willia grimmioides C. Mtill.
Willia subg. Euwillia (C. Mtill.) Broth., Nat. Pfl. I(3): 417,
1902, nom. illeg.
Willia sect. Euwillia C. Miill., Gen. Muse. Fr. 424, 1900, nom.
illeg.
Named for Elie Jean Fran9ois Guillou, 1806-? of the voyage of
the French ships Astrolabe and :lelee, 1837-1840), whose collections are at NSW and P.
Plants forming cushions or turfs, green above, occasionally
blackened, reddish brown below. Stems branching occasionally,
233
ca. 1-2 em in length, transverse section rounded-pentagonal,
central strand usually present but weak, sclerodermis absent,
hyalodermis absent; axillary hairs 5-lO cells in length, basal
2-3 cells with thickened walls; rhizoids sparse. Leaves
appressed when dry, spreading when moist, ligulate to
spathulate, sometimes constricted medially, ca. 2-5 mm in
length, upper lamina concave or broadly channeled across
leaf, narrowly grooved along costa, margins plane, entire or
occasionally weakly denticulate near apex, often bordered
with 2-4 rows of cells with thicker walls, cells often elongaterhomboidal and hyaline near apex; apex broadly rounded to
rounded-acute; base weakly or strongly differentiated, elliptical to rhomboidal, occasionally clearly sheathing; costa
excurrent in a short or long awn, in immature branch leaves
represented only by an apiculus, costa with lamina inserted
laterally or ventrally, superficial cells quadrate to shortrectangular and papillose ventrally, dorsally elongate and
smooth to somewhat roughened, 3-5 rows of cells across
costa ventrally at midleaf, costal transverse section elliptical
to round, stereid band present dorsally, large and reniform to
semicircular in shape, ventral epidermis present, dorsal
absent, ca. 6 guide cells in 2 layers (ventrally 2 large cells,
dorsally ca. 4 smaller), hydroid strand present, distinct; upper
lamina/ cells subquadrate, rather small, IO-I5 IJ.m in width,
1(-2): 1, walls thin. to evenly thickened, superficially convex;
papillae solid or hollow, bifid, 4-8 per lumen; basal cells
strongly differentiated medially or occasionally across leaf in
lower 1/4-I/2 of leaf, rectangular, 15-23 jlm in width, mostly
3-4:I, walls thin, often bordered by 3-4 rows of narrower,
more sturdy cells. Dioicous. Perichaetia terminal, inner
leaves highly differentiated, long-ligulate to broadly lanceolate, 4-8 mm (inclusive of awn) in length, often secund,
loosely sheathing the seta, lower cells rectangular to inflatedrhomoidal, differentiated in lower I/2 to throughout leaf. Perigonia terminal, gemmate. Seta mostly rather short, I-7 mm in
length, I per perichaetium, yellow-brown, straight or twisted
clockwise; theca often short, 0.8-3.5 mm in length, yellowbrown, very short-elliptical to cylindrical, exothecial cells rectangular, ca. 15-25 IJ.m in width, 2-6:I, walls thin or somewhat thickened, stomates phaneropore, at base of theca, annulus of 2-4 rows of highly vesiculose cells, often everted, persistent; peristome absent or teeth of 32 short anastomosing
rami, filamentous, low-spiculose, ca. 800 IJ.m, with several
articulations, twisted very weakly counterclockwise, basal
membrane comparatively high, 400 jlm in height, lowspiculose. Operculum short- to long-conic, 0.6-1.6 mm in
length, cells twisted weakly counterclockwise. Ca/yptra
cucullate to long-mitrate, often lobed below, plicate below or
smooth, 1.5-4.0 mm in length. Spores 8-IO jlm in diameter,
light brown, essentially smooth to papillose. Lamina/ KOH
color reaction red.
Found mainly on rock and tree trunks in austral regions,
often near the coast.
The austral genus Wiflia is near Syntrichia in gross characteristics but differs largely in a tendency towards reduced
sporophytes (Pl. 90, f. I, 12; 9I, f. 1) and highly differentiated
perichaetialleaves (Pl. 90, f. IO, I9, 20; 9I, f. 8). Associated
234
GENERA OF THE POTTIACEAE
Plate 90. Willia. 1-11. W. austroleucophaea. I. Habit. 2-3. Two transverse sections of stem. 4-6. Three leaves. 7. Leaf apex. 8. Basal cells. 9.
Transverse section at midleaf. 10. Perichaetialleaf. 11. Calyptra. 12-20. W. calobolax. 12-13. Two habits. 14-16. Three leaves. 17. Leaf apex.
18. Transverse section at midleaf. 19-20. Two perichaetialleaves.
POTTIOIDEAE
characters are commonly panduriform cauline leaves often with
elongate (occasionally hyaline) laminal cells near the apex; the
leaf margins plane, weakly bordered by somewhat thickened
cells; the peristome absent or short but with a high basal membrane (Pl. 91, f. 9); the annulus in some species everted on capsule dehiscence; and the calyptra is usually large and lobed
below (Pl. 90, f. 11-12, but cf 91, f. 11).
Lightowlers (1985c) placed Tortula calobolax (Pl. 90, f.
235
12-20) in Willia because of its immersed, eperistomate capsules, the rigid, oblong, somewhat pandurate leaves with
plane margins, and the perichaetial leaves with longacuminate apices and differentiated marginal cells. A specimen (Prince Edward 1., Zinderen Bakker 671, NY) indicated
by him (1985c) to be this species additionally has the characteristic short, fleshy seta and short macrostomous capsules
also characteristic of W. austroleucophaea, and clearly
Plate 91. Willill. 1-11. W. brachychaete. 1. Habit of sporophyte-bearing plant. 2. Perigoniate plant. 3-4. Two leaves. 5. Leaf apex. 6. Basal
cells. 7. Transverse section at rnidleaf. 8. Perichaetialleaf. 9. Peristorne. 10. Operculum. 11. Calyptra.
:~.-;
236
GENERA OF THE POTTIACEAE
belongs with Wil/ia. The genus is expanded here with the addition of W. brachychaete (Pl. 91, f. 1-11, discussed by Kramer
1988). Clearly, W. brachychaete approaches Syntrichia by its
cucullate calyptra, presence of a peristome (albeit consisting
mostly of a high basal membrane), less strongly reduced seta
and capsule, and is itself approached by S. cavallii (Pl. 108, f.
7-12), which has a strongly differentiated perichaetium and
weakly pandurate leaves. Syntrichia cava/Iii differs from Willia
in the recurved upper leaf margins, somewhat cucullate leaf apices, and well developed sporophyte and peristome. Willia may
ultimately be acceptable as only a section of Syntrichia, but
more intensive study is needed to demonstrate this. Of especial
value would be an evaluation of the relationship of Willia to
plane-margined species of Syntrichia (sect. Aesiotortula). See
Cladograms 13 and 15 for character state changes of phylogenetic hypotheses of relationships based on a global evaluation
of all characters.
Additional literature: Bell (1974).
Number of accepted species: 3.
Species examined: W. austroleucophaea (BM, H, NY), W.
brachychaete (BUF), W. calobolax (NY).
New combination: Willia brachychaete (Dus.) Zand., comb.
nov. (Tortula brachychaete Dus., Bot. Not. 1905: 300, 1905).
64. SAITOELLA
Plate 92.
Saitoella Menzel, J. Hattori Bot. Lab. 71: 239, 1992.
Saitoa Zand., Phytologia 65: 430, 1989, hom. illeg. non
Saitoa Rajendran & Muthappa, Proc. Indian Acad. Sci.
sect. B, 89: 185, 1980. Type: Saitoella peruviana (Williams) Menzel.
Named for Japanese bryologist Kamezo Saito, author of "A
Monograph of Japanese Pottiaceae (Musci)" (1975a) and
many other papers on the Pottiaceae.
Small terete plants growing in dense clumps, deep reddish
brown above, reddish tan below. Stems branching occasionally, pseudodichotomous, to 4 mm in length, transverse section rounded-pentagonal, central strand distinct, sclerodermis
absent, hyalodermis absent; axillary hairs 2-4 cells in length,
all cells hyaline; sparsely radiculose. Leaves closely appressed
when dry, weakly spreading when moist, broadly oblongobovate, elliptic or suborbicular, 0.5-0.6 mm in length, upper
lamina concave and apex therefore somewhat cucullate, margins usually recurved at rnidleaf and below, crenulate in upper
half by projecting cell walls, bordered by ca. 15-18 rows of
smooth, rhomboidal, thick-walled cells with somewhat
rounded lumens; apex broadly rounded to weakly
Plate 92. Saitoella. 1-S. S. peruviana. 1. Habit. 2-3. Two leaves. 4. Lamina! areolation. 5. Transverse section at midleaf.
POTIIOIDEAE
emarginate; base not differentiated in shape; leaf widest above
middle; costa ending 2-3 cells below apex, thickest distally,
superficial cells ventrally quadrate to short-rectangular, papillose, dorsally elongate, smooth or papillose, 4-5 rows of cells
across costa ventrally at midleaf, ventral surface of upper costa
forming a bulging pad of I layer of papillose cells, costal transverse section elliptical to reversed semicircular (i.e. flattened
dorsally), stereid band present dorsally, strong, elliptical,
ventral epidermis strongly differentiated, of thin-walled, papillose cells, dorsal epidermis absent or of thick-walled bulging
cells, guide cells absent, hydroid strand absent; upper /aminal
cells subquadrate to short-rectangular, 6-10 Jlm in width, l-2:1,
walls thin to thickened, superficially equally weakly bulging on
both sides of lamina; papillae restricted to a small area of the
lamina near the costa, small, mostly bifid, hollow or solid,
about 4--6 per lumen; basal cells differentiated across the leaf
base, quadrate to short-rectangular, ca. 16-18 Jlm in width,
1-2: 1, walls thin. Sexual structures and sporophyte unknown.
Lamina/ KOH color reaction deep red.
Found on soil over volcanic rock at high elevations; Peru,
Ecuador, Mexico.
This genus contains only S. peruviana, which has been recognized in Globulinella for some time, being similar in the concave, ovate to rounded leaf shape and rather thick costa ending
before the apex. Saitoella differs considerably, however, in the
upper lamina bordered by many rows of rhomboidal, thickwalled cells, upper !aminal cells papillose in a medial patch (Pl.
92, f. 4), costa lacking guide cells (Pl. 92, f. 5), and red color
KOH reaction. The branching is pseudodichotomous (Zander
1976), a pattern apparently rare in mosses and as yet inadequately surveyed. The naked archegonia found in leaf axils inS.
peruviana are uncommon in Pottiaceae; they are also present in
Globulinella globifera and have been found in several other
genera of Pottiaceae and in other families (Zander 1976).
Saitoella is considered here to lack differentiated guide
cells, the superficial ventral parenchyma being of papillose
quadrate cells, and therefore probably not interpretable as guide
cells. Phascopsis is similar in lacking guide cells in at least
some leaf sections-in that genus cells in the guide cell position
become narrower and more thick-walled from the base of the
leaf to the apex. Scope/ophila ligulata either lacks guide cells
or, more probably, a ventral epidermis, since the ventral cell
layer is of inflated, smooth cells, but Scopelophi/a may be distinguished because it also lacks laminal papillae and a stem central strand. In Saitoella, putative guide cells cannot be distinguished anywhere in the costa. Bryoerthrophyllum co/umbianum has a similar ventrally bulging costa with the lamina
inserted laterally, but guide cells are present and two stereid
bands may be distinguished in at least some leaves. Tortula
maritima has a similar medial concentration of upper papillose
cells in the leaf and nearly identical costal section (guide cells
absent and epidermal cells bulliform), and Stegonia /atifolia has
similar rhomboidal upper laminal cells and a costal section
(epidermal cells bulliform but guide cells present) approaching
that of Saitoella; these two taxa are easily distinguished by their
yellow KOH reaction. Aloinella species may likewise have
papillae located only medially on the leaf, with a broad border
237
of non-papillose rectangular cells, but ventral costal filaments
are differentiated. Tortula revolvens is somewhat similar to
Saitoel/a in having upper !aminal cells papillose only medially, but may be distinguished by the presence of guide cells
in the costa, strongly revolute upper !aminal margins, and yellow KOH reaction.
A dorsal costal epidermis is clearly differentiated in some
leaves of S. peruviana but not in others. The epidermal presence, even partial, is of some value in separating it from Syntrichia, which lacks a dorsal costal epidermis.
Additional literature: Steere & Chapman (1946).
Number of accepted species: 1.
Species examined: S. peruviana (BUF, FH, MICH,
TENN,NY).
65. MICROBRYUM
Plate 93.
Microbryum Schimp., Syn. 10, 1860. Type: Microbryum
floerkeanum (Web. & Mohr) Schirnp.
Bryella Berk., Handb. Brit. Mosses 16: 300, 1863, Type:
Bryella recta (With.) Berk.
Cycnea Berk., Handb. Brit. Moss. 60, 301, 1863, hom. il/eg.
non Cycnia Griff., 1854. Type: Cycnea curvicolla
(Hedw.) Berk.
Pottiel/a (Lirnpr.) Gams, Krypt. Fl. Mitteleur. ed. 2, 1: 101,
1948.
Lydiaea Laz., Not. Syst. Sect. Cryptog. Inst. Bot. Nom.
Komar. Acad. Sci. URSS 12: 280, 1959. Type: Lydiaea
vlassovii (Laz.) Laz.
Phascum subg. Microbryum (Schirnp.) Limpr., Laubm.
Deutsch!. 1(3): 182, 1885.
Phascum subg. Pottie/la Limpr., Laubm. Deutsch!. 1: 188,
1885. Type: Phascum curvicol/um Hedw., /ectotyp . nov.
Pottia subg. Pottiella (Limpr.) Broth., Nat. Pfl. 1(3): 423,
1902.
Phascum sect. Pottiella (Limpr.) Par., Actes Soc. Linn. Bordeaux 51[1]: 66, 1897; Ind. Bryol. [4]: 1030, 1898.
Acaulon sect. Microbryum (Schimp.) C. Mtill., Gen. Muse.
Fr. 20, 1900.
Phascum sect. Microbryum (Schimp.) Podp., Consp. Muse.
Eur. 221, 1954.
Pottia sect. Pottiella (Limpr.) Nyholm, Ill. Fl. Nordic Mo.
2: 81, 1989.
Pottia subsect. Muticae C. Jens., Skand. Bladmfl. 210,
1939, nom. inval. descr. suec.
From JllKp6<;, small + o + bryum from ~puov, a moss.
Plants forming a low turf, scattered or gregarious, occasionally bulbiform, reddish brown above, brown below. Stems
seldom branching, extremely short, 0.2-0.4 mm in length,
transverse section round to rounded-pentagonal, central strand
present or absent, sclerodermis absent or weakly
differentiated, hyalodermis absent; axillary hairs 3-6 cells in
length, the basal 1-2 usually with thicker walls; rhizoids
sparse. Leaves appressed when dry, weakly spreading and tips
occasionally reflexed when moist, lanceolate, elliptical or
ovate, occasionally spathulate, short, 0.6-1.8 mm in length,
238
GENERA OF THE POTTIACEAE
Plate 93. Microbryum. 1-11. M. rectum. 1-2. Habits. 3. Transverse section of stem. 4-5. Two leaves. 6. Leaf apex. 7. Transverse section at
midleaf. 8. Stomates. 9. Calyptra. 10. Detail of calyptra. 11. Spores. 12-15. M. brevicaule. 12-13. Two leaves. 14. Transverse section at
midleaf. 15. Sporophyte. 16-19. M. curvicolum. 16-17. Two leaves. 18. Leaf apex. 19. Sporophyte. 20-22. Calyptrae and detail. 23-28. M.
jloerckeanum. 23-24. Two leaves. 25. Sporophyte. 26. Spores. 27. Calyptra. 28. Areolation of calyptra.
r
,
POTIIOIDEAE
upper lamina weakly concave to broadly channeled, margins
recurved at midleaf, commonly narrowly recurved to near apex,
entire or rarely serrulate near apex, marginal cells often less
papillose and somewhat thicker walled than the medial; apex
broadly acute; base not differentiated; costa excurrent as an
apiculus or forming a mucro or short awn, occasionally only
percurrent, costa with lamina inserted laterally, superficial cells
smooth or papillose, ventrally quadrate or short-rectangular or
elongate, dorsally short-rectangular to elongate, 2(4-6) rows of
cells across costa ventrally at midleaf, costal transverse section
usually round, stereid band present dorsally, round to semicircular in shape, ventral and dorsal epidermises present, guide
cells 2(-4) in 1 layer, hydroid strand present, occasionally centrally located in the stereid band, occasionally the ventral epidermis differentiated as a pad of enlarged parenchymatic cells;
upper lamina/ cells quadrate to hexagonal or short-rectangular,
occasionally rhomboidal, rather large, 11-20 jlm in width,
1-2:1, .walls thin to moderately and evenly thickened, superficially convex on both sides; papillae usually simple, seldom
bifid, hollow, 1-6 per lumen, occasionally branching and tall;
basal cells differentiated across leaf or higher medially, rectangular, ca. 18-30 jlm in width, 2-4:1, walls usually thin.
Monoicous, usually paroicous, occasionally synoicous. Perichaetia terminal, inner leaves often somewhat enlarged, otherwise little different from the cauline. Seta nearly absent to 4
mm in length, 1(-2) per perichaetium, yellow-brown, twisted
straight or counterclockwise below and clockwise above; theca
cleistocarpous or stegocarpous, ca. 0.5-1.1 mm in length,
brown or reddish or yellowish brown, ovate to short-elliptical,
apiculate when cleistocarpous, exothecial cells shortrectangular, ca. 18-25 jlm in width, mostly 2-5:1, thin-walled,
stomates phaneropore, at base of theca, annulus absent or of
1-2 rows of weakly vesiculose cells, persistent; eperistomate or
peristome teeth 16, irregular, often rudimentary, often apically
truncate, ligulate to triangular, spiculose, short, with few articulations, straight, basal membrane absent. Operculum when differentiated low-conic, ca. 0.1-0.2 mm in length, cells straight.
Calyptra mitrate to conic-cucullate, smooth or finely papillose,
ca. 0.2-0.8 mm in length. Spores ca. 20--30 jlm in diameter,
light brown, essentially smooth to warty or spiculose or hollowtuberculate. Lamina/ KOH color reaction red. Reported chromosome numbers: n =26, 27+m, 28, 30.
Found in the temperate zones worldwide, especially in
somewhat arid situations, mainly on soil.
Microbryum is distinctive in the combination of the small
habit, red KOH color reaction of the upper lamina, single round
to semicircular costal stereid band (Pl. 93, f. 7, 14), capsules
apiculate when cleistocarpous (Pl. 93, f. 19), peristomes when
present commonly apically truncate and seemingly large in
comparison with the size of the capsule; and calyptrae often
papillose (Pl. 93, f. 10, 22). If one considers the possibility that
taxa with single stereid bands evolved independently from taxa
with two stereid bands, then Microbryum might have been
derived through reduction from ancestors of Bryoerythrophyllum, which has similar areolation. Evidence for this is that there
are no other highly reduced taxa related to Bryoerythrophyllum
(see also discussion of Saitoella and Acaulon). Otherwise,
239
Microbryum could have come from progenitors similar to
Tortula sect. Tortula. Cladograms 11 and 14, however, indicate a different derivation.
Chamberlain's (1969, 1978) treatment of Pottia species
with erostrate opercula recognized M. starckeanum as a single
species including both tuberculate and papillose-spiculose
spored plants. The present study, however, supports the traditional arrangement of Corley et al. (1982) that distinguishes
material differing by the two spore ornamentations at the species level. Chamberlain's infraspecies are all recognized at the
varietal level, with the addition of M. davallianum var.
commutatum and M. starckeanum var. fosbergii. All varieties
of these two species that were recognized by Chamberlain
(1978) for Gt. Britain are also present in U.S.A. in California.
One collection, California, Pasadena, s.h., "g. 9", US,
includes M. starckeanum var. starckeanum, var.fosbergii, and
an intermediate with a capsule having a differentiated operculum and short peristome that is indehiscent even when
boiled in KOH solution. The intermediate ·is also found in
other collections (e.g. California, Ikenberry 369, CANM,
comm. T. Mcintosh). The type of Pottia arizonica (= M.
starkeanum var. starckeanum) has spores that appear to be
both papillose and tuberculate, but the "papillae" are loose in
the spore sac as well as partially coating the spore. The type
of Pottia fosbergii is more probematic, with spores that are
slightly wrinkled (very weakly tuberculate) and also weakly
papillose. In this case, the sporophytes and spores (if
unreduced) may be of hybrid origin; other specimens clearly
of var. fosbergii (operculum not differentiated at all) have the
spores typical of M. starckeana. Microbryum davallianum
var. conicum may have spores that are epapillose, but these
are never wrinkled. The two species may be seen as two series
of infraspecific peristome reduction.
The calyptrae are roughened with low, simple papillae in
many of the species with comparatively large calyptrae (e.g.
M. commutatum, M. rectum); calyptral papillae are apparently
absent in those species with much reduced sporophytes and
tiny calyptrae. The presence of calyptral papillae helps distinguish this genus from Acaulon (which, like highly reduced
members of Microbryum, has strongly bulging vaginulae),
Syntrichia and Tortula sect. Tortula. The new combination
Microbryum rufochaete reflects the strongly apiculate capsules and recurved upper laminal margins of this species.
Microbryum tasmanicum is similar but has simple !aminal
papillae. In M. rufochaete the perigoniate plants are about a
quarter to a third the size of the perichaetiate plants, and are
situated near the base of the perichaetiate plants (possibly
rhizautoicous).
Mcintosh (1989) discussed M. vlassovii for North America (as Phascum). Both M. vlassovii and M. raddei have an
enlarged pad of parenchymatic cells ventrally on the costa.
Carri6n et al. (1990) described the spore morphology of several species of Microbryum (as Phascum), and indicated that
the spore surface of M. vlassovii was rather different from that
of M. curvicolle (Pl. 93, f. 16-19) and M. jloerkeanum (Pl. 93,
f. 23-28), being more like that of Tortula atherodes (discussed asP. cuspidatum).
240
GENERA OF THE POTTIACEAE
Additional literature: Guerra et al. (1991, 1992).
Number of accepted species: 13.
New heterotypic synonymy: Pottia arizonica Wareham in
Grout = Microbryum starckeanum (Hedw.) Zand. var.
starckeanum. Pottia arizonica var. mucronulata Wareham in
Grout = Microbryum starckeanum var. brachyodus (BSG)
Zand.
Species examined: M. brevicaule (NY), M. curvicolle (NY),
M. davallianum (BUF, CANM, FH, US), M.floerkeanum (NY),
M. longipes (BUF), M. raddei (H), M. rectum (BUF), M.
rufochaete (NY), M. starckeanum (BUF, CANM), M.
subplanomarginatum (BUF), M . tasmanicum (BM), M.
vlassovii (BUF), M. zeelandiae (NY).
New combinations and statuses:
Microbryum brevicaule (Tayl.) Zand., comb. nov.
(Gymnostomum brevicaule Tayl., London J. Bot. 5: 42,
1846; Pottia brevicaulis (Tayl.) C. Muell.).
Microbryum curvicolle (Hedw.) Zand., comb. nov. (Phascum
curvicol/e Hedw., Spec. Muse. 21, 1801; Pottia curvicollis
(Hedw.) Mitt.).
Microbryum davallianum (Sm.) Zand., comb. nov.
(Gymnostomum daval/ianum Sm., Ann. Bot. 1: 577, 1805;
Pottia davalliana (Sm.) C. Jens.).
Microbryum davallianum var. commutatum (Limpr.) Zand.,
comb. nov. (Pottia commutata Limpr., Laubm. Deutsch!. 1:
537, 1888; Pottia dava/liana subsp. commutata (Limpr.)
Podp.).
Microbryum davallianum var. conicum (Schleich. ex
Schwaegr.) Zand., comb. nov. (Gymnostomum conicum
Schleich. ex Schwaegr., Sp. Muse. Suppl. 1(1) 25, 1811;
Pottia starckeana ssp. conica (Schleich. ex Schwaegr.)
Chamber!.).
Microbryum floerkeanum var. arbense (Loitl.) Zand., comb.
nov. (Phascum arbense Loitl., Verh. Zool. Bot. Ges. Wien
59: 55, 1909; Phascum floerkeanum var. arbense (Loitl.)
Podp.), not seen.
Microbryum longipes (Guerra, Martinez & Ros) Zand., comb.
nov. (Phascum /ongipes Guerra, Martinez & Ros, J. Bryol.
16: 55, 1990 as "longipedis" typographical error cf J.
Bryol. 16: 335, 1991).
Microbryum raddei (Broth.) Zand., comb. nov. (Tortula raddei
Broth., Bot. Centralbl. 34: 26, 1888).
Microbryum rectum (With.) Zand., comb. nov. (Phascum rectum With., Syst. Arr. Britt. Pl. ed. 4: 771, 1801; Pottia recta
(With.) Mitt.).
Microbryum rufochaete (Magill) Zand., comb. nov. (Acaulon
rufochaete Magill, Fl. S. Afr. I. Bryophyta 1: 201, 1981
[1982]).
Microbryum starckeanum (Hedw.) Zand., comb. nov. (Weisia
starckeana Hedw., Spec. Muse. 65, 1801; Pottia starckeana
(Hedw.) C. Miill.).
Microbryum starckeanum var. brachyodus (BSG) Zand., comb.
nov. (Anacalypta starckeana var. brachyodus BSG, Bryol.
Eur. 2: 47, 1843; Pottia starckeana var. brachyodus (BSG)
C. Miill.), commonly "brachyoda."
Microbryum starckeanum var. brevidens (Latz.) Zand., comb.
nov. (Pottia starckeana var. brevidens Latz., Beih. Bot.
Centralbl. 48(2): 483, 1931), not seen.
Microbryum starckeanum var. fosbergii (Bartr.) Zand., comb.
nov. (Pottia fosbergii Bartr., Bryologist 33: 18, 1930;
Pottia starckeanum var.fosbergii (Bartr.) Zand.).
Microbryum starckeanum var. leiostoma (Corb. in Corb. &
Pitard) Zand., comb. nov. (Pottia starckeana var.
leiostoma Corb. in Corb. & Pitard, Bull. Soc. Bot. France
56: ccxxiv, 1909), not seen.
Microbryum starckeanum var. subgymnostoma (De Not.)
Zand., comb. nov. (Anacalypta starckeana var.
subgymnostoma De Not., Atti Univ. Genova 1: 583, 1869;
Pottia starckeana var. subgymnostoma (De Not.) Grav.),
not seen.
Microbryum starckeanum var. submutica (Latz.) Zand., comb.
nov. (Pottia starckeana var. submutica Latz., Beih. Bot.
Centralbl. 48(2): 483, 1931), not seen.
Microbryum starckeanum fo. brevifolium (Limpr.) Zand.,
comb. nov. (Pottia starckeana fo. brevifolia Limpr.,
Laubm. Deutsch. 1: 536, 1888), not seen.
Microbryum starckeanum fo. dextrorsum (Limpr.) Zand.,
comb. nov. (Pottia starckeana fo. dextrorsa Limpr.,
Laubm. Deutsch. 1: 536, 1888), not seen.
Microbryum starckeanum fo. microphyllum (Wamst.) Zand.,
comb. nov. (Pottia starckeana fo. microphylla Warnst.,
Hedwigia 58: 144, 1917), not seen.
Microbryum subplanomarginatum (Dix.) Zand., comb. nov.
(Pottia subplanomarginata Dix., Trans. R. Soc. S. Afr.
18: 253, 1929).
Microbryum tasmanicum (Dix. & Rodw.) Zand., comb. nov.
(Phascum tasmanicum Dix. & Rodw., Pap. Proc. R. Soc.
Tasmania 1923: 25, 1923).
Microbryum vlassovii (Laz.) Zand., comb. nov. (Phascum
vlassovii Laz., J. Inst. Bot. Ac. Sc. R.S.S. Ukraine 26-27:
196, 1938).
Microbryum zeelandiae (R. Br. ter) Zand., comb. nov.
(Anacalypta zealandiae R. Br. ter, Trans. Proc. New Zealand Inst. 30: 413, 1898; Pottia zealandiae (R. Br. ter)
Par.).
66.CRUMIA
Plate 94.
Crumia Schof., Canad. J. Bot. 44: 609, 1966. Type: Crumia
latifolia (Kindb.) Schof.
Named for Howard Alvin Crum, 1922-, author of numerous
valuable bryological publications, including "Mosses of Eastem North America" (1981) with Lewis E. Anderson.
Plants forming cushions, green to reddish above, blackish
green to iridescent tan below. Stems branching often, to 3.0
em in length, transverse section rounded-pentagonal, central
strand absent or very weak, sclerodermis absent, hyalodermis
absent; axillary hairs ca. 6 cells in length, basal 1-2 cells yellowish; sparsely radiculose. Leaves appressed, weakly contorted when dry, spreading when moist, spathulate, to 4.0 mm
in length, upper lamina flat, shallowly channeled along costa,
margins recurved along 1 or both margins below midleaf,
POTIIOIDEAE
241
Plate 94. Crumia. 1-10. C. latifolia. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves, one enlarged. 6. Leaf apex. 7. Upper marginal
cells. 8. Basal cells. 9. Transverse section at midleaf. 10. Portion ofperistome.
242
GENERA OF THE POTTIACEAE
entire, bordered by ca. 6 rows of enlarged, thicker walled,
rounded rhomboidal to rectangular cells, except the extreme
1-2 marginal rows; apex broadly acute to rounded-acute, usually broadly apiculate; base scarcely differentiated in shape,
bordered by a few rows of long-rectangular cells; costa
percurrent or ending 1-2 cells below the apex, superficial cells
elongate on both sides, 5-{) rows of cells across costa ventrally
at midleaf, costal transverse section semicircular, one stereid
band present, strong, epidermis present ventrally, weakly developed dorsally, guide cells 3-4 in 1 layer, hydroid strand absent;
upper !aminal cells in medial portion of leaf hexagonal to shortrectangular, 13-20 )lm in width, 1-2:1, walls thin, superficially
weakly convex on both sides; papillae small but distinct, simple
to bifid, 8-12 per lumen, scattered; basal cells differentiated
medially, inflated, rectangular, wider than the upper cells,
3-5:1, walls thin, brown. Dioicous. Perichaetia terminal, inner
leaves little different from cauline leaves, not sheathing. Perigonia reported as termin\11, bracts not sheathing. Seta ca. ~15
mm in length, 1(-2) per perichaetium, reddish brown, twisted
clockwise below, counterclockwise above; theca 2.5-3.0 mm in
length, reddish brown, cylindrical, exothecial cells rectangular,
thin-walled, stomates phaneropore, at base of theca, annulus of
ca. 2 rows of vesiculose cells; peristome teeth 16, cleft to near
base into two branches, linear, rami much perforated and anastomosing (but articulations held together by a hyaline membrane), densely spiculose, ca. 200 )lm in length, with ca. 7 articulations, twisted very weakly counterclockwise, basal membrane
low, crazed. Operculum conic, 0.8-1.1 mm in length, cells
weakly twisted counterclockwise. Calyptra cucullate, smooth,
ca. 3.0 mm in length. Spores ca. 15-18 )lm in diameter, light
brown, weakly papillose. Lamina/ KOH color reaction reddish
orange.
Found on soil or rock, usually in wet areas, especially along
streams, often on calcareous substrates; found in southwestern
Canada and western United States.
Crumia is much like Scopelophila in the often blackish coloration of the plants, the lack of both a sclerodermis and a
hyalodermis in the stem (Pl. 94, f. 2), leaves spathulate, often
with a broad apiculus, costa percurrent, stereid band single,
hydroid strand absent (Pl. 94, f. 9), and perichaetialleaves little
differentiated from the cauline. It differs significantly from
Scopelophila in the presence of a stem central strand (though
generally only weakly developed or occasionally absent), a
more strongly differentiated laminal border of thick-walled cells
with one or two rows of much smaller cells on the extreme margin (Pl. 94, f. 7), upper !aminal cells papillose (rarely mostly
smooth), and peristome (Pl. 94, f. 10) present, though rarely
fruiting. Schofield (1966) added a few other distinctions for
Crumia, such as always entire leaf margins, fruiting in spring
(rather than late autumn for Scopelophila), stomata well developed (versus rudimentary or obscure), exothecial cells elongate,
and annular cells weakly developed, but these are either minor
characters or are based too strongly on only one species of
Scopelophila. His observation, however, that Crumia is found
on alkaline substratt:s poor in metallic ions, while Scopelophila
is largely restricted to acid habitats often rich in sulfur and iron,
copper or other metallic ions, has considerable weight, given
the other features. The leaf shape of Crumia is similar to that
of Tortula but the upper marginal cells of the former are
short-elliptical and swollen in section, with lumens larger than
those of the medial cells, while the marginal cells of Tortula
species are either undifferentiated or nearly so or rectangular
and lumens smaller (as seen in section) than those of the
medial cells.
The peristome of Crumia is quite interesting, being
weakly twisted counterclockwise and composed of fragmented articulations of the two rami, these held together by a
very thin, hyaline membrane reminiscent of that of Cinclidotus (but here the 16 teeth are quite separate). The peristome is
quite unlike that of Tortula deciduidentata, previously placed
in Crumia, which is strongly twisted, composed of 32 integral
rami above a distinct basal membrane, and comes off with the
operculum (the columella elongates and pushes the operculum, with the peristome inside it, up and away from the
capsule mouth). Although the cauline leaves of T.
deciduidentata are similar to those of Crumia or Scopelophila, the perichaetialleaves bear an awn, which is curiously
blunt or flattened apically. Other characters that would be
anomalous in Crumia for Tortula deciduidentata are the
latter's strong central strand, presence of a hydroid strand in
the costa, and the paroicous or autoicous sexual condition.
Additional literature: Sharp & Iwatsuki (1969), Vitt and
Zander (1978).
Number of accepted species: 1.
Species examimed: C. latifolia (BUF, NY).
67. HENNEDIELLA
Plates 95-98.
Hennediella Par., Ind. Bryol. 557, 1896, nom. nov. for
Hennedia R. Br. ter. Type: Hennediella macrophylla (R.
Br. ter) Par.
Hennedia R. Br. ter, Trans. New Zealand Inst. 25: 285,
1893, hom. illeg. non Harv., 1860. Type: Hennedia
macrophylla R. Br. ter, lectotyp. nov.
Beckettia C. Mi.ill., Hedwigia 37: 77, 1898. Type: Beckettia
bruchioides C. Mi.ill.
Neobarbula Dus., Bot. Not. 1905: 299, 1905. Type:
Neobarbula magellanica Dus.
Bauriella Wamst., Hedwigia 57: 88, 1915, nom. inval. prov.
Type: Tortula polyseta (C. Mi.ill.) Wamst.
Henediella Paris ex Fleisch., Musci Fl. Buitenzorg 4: 1697,
1923, orthogr. var.
Willia subg. Schistidiella (C. Mtill.) Broth., Nat. Pfl. 1(3):
417, 1902. Type: Willia marginata (Hook. f. & Wils.) C.
Mi.ill.
Hennediella subg. Beckettia (C. Mi.ill.) Roth, Aussereur.
Laubm. 219, 1911.
Hennediella subg. Hennedia Roth, Aussereur. Laubm. 220,
1911, nom. illeg. incl. typ. gen.
Willia sect. Schistidiella C. Miill., Gen. Muse. Fr. 424,
1900. Type: Willia marginata (Hook. f. & Wils.) C. Mtill.
Tortula sect. Neobarbula (Dus.) Card., Wiss. Ergebn.
Schwed. Stidpol. Exp. NordenskjOld 4(8): 95, 1908. Type:
Tortula magellanica Dus.
POTIIOIDEAE
243
Plate 95. Hennediello. 1-8. H. macrophylla. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex, dorsal view. 6. Basal cells.
7. Transverse section of costa at midleaf. 8. Calyptra. 9-11. H. acutidentata. 9-10. Two leaves. 11. Transverse section of costa at midleaf.
12-19. H. acletoi. 12. Transverse section of stem. 13-14. Two leaves. 15. Leaf apex. 16. Transverse section of costa at midleaf. 17. Perichaetial
leaf. 18. Peristome. 19. Operculum.
244
GENERA OF THE POTTIACEAE
Plate 96. Hennediella. 1-7. H. austrogeorgica. 1. Habit. 2. Plant with immature sporophyte and cladautoicous branch. 3-5. Three leaves. 6.
Operculum. 7. Calyptra. 8-10. H. bellei. 8-9. Two leaves. 10. Leaf apex. 11. H. densifolia. 11. Habit. 12-17. H. denticuloto. 12. Transverse
section of stem. 13-14. Two leaves. 15. Leaf apex. 16. Peristome. 17. Operculum. 18-21. H. heimei. 18-19. Two leaves. 20. Transverse section
of costa at midleaf. 21. Systylious capsule. 22-26. H. heteroloma. 22. Transverse section of stem. 23-24. Two leaves. 25. Leaf apex. 26.
Transverse section of costa at midleaf.
POTIIOIDEAE
Named for Roger Hennedy, 1809-1877, Glascow phycologist,
who was Brown's professsor +-ella, diminutive.
Plants forming loose turf or cushions, green above, reddish
brown below. Stems branching irregularly, ca. 0.5-1.0 em in
length, transverse section rounded-pentagonal or elliptical, central strand present, sclerodermis absent or represented as an
outer cortex of substereid cells, hyalodermis absent or weakly
developed, not collapsed; axillary hairs of ca. 5 cells, the basal
1-2 cells occasionally somewhat thick-walled; rhizoids usually
common. Leaves appressed or incurved when dry, spreading
and sometimes reflexed at midleaf when moist, ovate to longlanceolate, occasionally ligulate or spathulate, comparatively
large, (1.5-)3-4(-7.0) mm in length, upper lamina plane or
broadly channeled across leaf, margins plane or only rarely
narrowly recurved, often dentate above, less commonly entire,
with a border (occasionally inframarginal) of short-rectangular
to elongate cells, these usually less papillose or smooth and
occasionally thick-walled but rarely bi- or tristratose; apex
acute or rarely obtuse, rarely fragile; base not differentiated in
shape or ovate and somewhat sheathing, commonly bordered by
cells similar to those of the upper border; costa percurrent or
short-excurrent as a flattened, denticulate mucro, rarely ending
3-4 cells below apex, costa with lamina inserted laterally,
superficial cells quadrate to short-rectangular and papillose
ventrally, dorsally elongate and smooth or papillose, 4-5 rows
of cells across costa ventrally at midleaf, costal transverse section round to elliptical, stereid band one, of substereid cells and
elliptical to round in shape, ventral epidermis present, dorsal
epidermis usually present (occasionally present only laterally
on the costa or sometimes absent, guide cells 2-4 per layer in
(1-)2 layers, hydroid strand present, usually large; upper
/aminal cells relatively large, quadrate to hexagonal or shortrectangular, ca. 18-24 Jlm in width, 1-2:1, walls thin to evenly
thickened, superficially flat to weakly convex; papillae hollow,
simple to bifid, usually 6-8 or more per lumen, occasionally
absent; basal cells differentiated across leaf, rectangular, 18-30
Jlm in width, (2-)4-6: 1, walls thin. Autoicous, paroicous,
cladautoicous, or dioicous. Perichaetial leaves somewhat larger
than the cauline leaves. Perigonia in dioicous species borne terminally on plants somewhat smaller than the perichaetiate. Seta
extremely various in size among species, 0.05-25 em in length,
1 per perichaetium, yellow-brown to dull brown, twisted counterclockwise; capsule occasionally systylious, theca 0.8-3.5 mm
in length, yellow-brown to dark brown, ovate to cylindrical,
occasionally microstomous or macrostomous, exothecial cells
rectangular, 25-35 Jlm in width, 3-5:1, walls thin to irregularly
thickened, stomates phaneropore, at base of capsule, annulus of
2-4 rows of vesiculose cells; peristome teeth absent, rudimentary, or of 32 long, filamentous teeth, densely spiculose, when
well developed to 1.5 mm in length, with many articulations,
twisted counterclockwise, basal membrane occasionally to 500
Jlm in height, spiculose. Operculum conic to rostrate, often narrowly so, (0.6-)1.2-1.8 mm in length, cells twisted counterclockwise. Calyptra cucullate, occasionally flaring below and
not split, smooth, 2.0-4.5 mm in length. Spores sometimes quite
large, 8-30 Jlm in diameter, yellowish, essentially smooth,
papillose or tuberculate. Lamina/ KOH color reaction red.
245
Reported chromosome numbers: n = 25+m, 26, 26+m,
26+2m, 50.
Found on soil and rock, commonly in wet areas, nearly
worldwide but most diverse in temperate areas of the austral
region.
This previously monotypic genus has been treated as a
synonym of Pottia (Dixon 1923; Sainsbury 1955; van der
Wijk et al. 1959-69); however, a clear sporophyte reduction
series involving other species (here transferred to Hennediella) and the distinctive morphology of gametophyte warrant
recognition at the generic level. Support for recognition of
distinctive characters of at least the gametophyte includes
Sainsbury's (1955) comment that Hennediella be treated as "a
section of Pottia distinguished by the toothed and bordered
leaves and by the very large, usually mitriform, calyptra ... ";
A. Fife's comment (pers. comrn.) that Pottia macrophylla (Pl.
95, f. 1-8) deserves recognition (as Hennediella); Corley et al.
(1981) suggested Tortula stanfordensis (Pl. 98, f. 7-11)
should be in a separate section or genus; and Matteri (1977a,
1977b, 1988a) pointed out that Pottia austrogeorgica (Pl. 96,
f. 1-7), which she regarded as a good species, was
vegetatively similar both to the P. heimii (Pl. 96, f. 18-21)
complex and to Hennediella macrophylla. Recently, Mishler
(1990), in a phylogenetic analysis of 23 Tortula s. lat. species
from Mexico, found that T. stanfordensis, T. polyseta (Pl. 98,
f. 1-6) and T. leiostoma, all here recognized in Hennediella,
are "likely to be monophyletic." Blockeel (1991) appreciated
the relationships of H. macrophylla, T. stanfordensis and P.
austrogeorgica, and provided a detailed evaluation recognizing Hennediella as a good genus for at least these three species, making the appropriate combinations (and mentioned my
sketchy generic key that included Hennediella as conceived
here, Zander 1989, as support).
Characters here considered of most importance are the
often large and commonly broadly lanceolate leaves (PI 95, f.
3-4) bordered by a band of usually elongate thick-walled
cells, with usually serrulate to dentate and almost ~;~lways
plane (narrowly recurved in H. limbata, PI. 97, f. 1-6) upper
lamina! margins, upper medial lamina! cells large, superficially rather flat (Pl. 95, f. 7, 11, 16), thin-walled and strongly
hollow-papillose (rarely epapillose, as in H. denticulata, Pl.
96, f. 12-14), costa usually with a dorsal epidermis, stereid
band elliptical, or semicircular to rounded (Pl. 95, f. 7, 11,
16-like Tortula and unlike Syntrichia, which has lunate dorsal stereid band sections), sporophytes in a reduction series
(also like Tortula and unlike Syntrichia), operculum often
long and narrowly rostrate (Pl. 98, f. 6), calyptra enlarged,
sometimes inflated (Pl. 95, f. 8; 97, f. 7; 98, f. 17), and KOH
color reaction red. The perigonia are borne terminally on lateral branchlets on the archegoniate stems of H. macrophylla
and H. longirostris. Hennediella densifolia is also monoicous
but has gemmate perigonia sessile on the main axis. Dixon
(1923) noted that the calyptra of H. macrophylla is campanulate but tears basally as it matures, while Sainsbury
(1955) observed that in this species the inflated calyptra may
appear campanulate in some collections and cucullate in others with much variability and intergradation in shape.
246
GENERA OF THE POTTIACEAE
Plate 97. Hennediella. 1-6. H. limbata. 1-2. Two leaves. 3. Leaf apex. 4. Transverse section of costa at midleaf. 5. Theca. 6. Operculum. 7-9.
H. magellanica. 7. Leaf. 8. Leaf apex. 9. Portion of peristome. 10-12. H. oedipodioides. 1~11. Two leaves. 12. Leaf apex. 13-17. H.
marginata. 13. Habit. 14. Transverse section of stem. 15. Leaf apex. 16. Transverse section at midleaf. 17. Sporophyte, with operculum and
calyptra.
POTTIOIDEAE
The costal guide cells are in what may be interpreted as two
layers, the ventral layer being of smaller and somewhat thicker
walled cells than those of the dorsal guide cell layer. Because
the stereid band is made up of substereid cells, the ventral layer
of guide cells, although intermediate in appearance between the
larger layer of guide cells and the dorsal substereids, may be
interpreted as indication of a second stereid band. This, plus the
common lanceolate leaf shape may imply a relationship to the
Barbuleae. Its closest relationship, however, is probably with
Tortula species having leaves with similar lax areolation and
differentiated border (e.g. Tortula solmsii), but which differ by
the generally broader leaves, more convex superficial surface of
the !aminal cells, lack of marginal teeth, and yellow KOH reaction (see, however, Cladograms 11 and 14 for alternative
hypotheses). The upper !aminal cells of Hennediella are rather
large, but the genus may be distinguished from Chenia, Dolotortula and Sagenotortula, similar genera with red KOH reactions and very large upper !aminal cells, by the papillose upper
!aminal cells, except in the case of the epapillose H. denticulata,
247
which may be distinguished by its unistratose border of elongate cells that are denticulate in the upper portion of the leaf.
Like Pseudocrossidium, Hennediella is an essentially austral genus with extensions of some species into the north temperate zone probably through migration routes along northsouth cordilleras.
Number of accepted species: 20.
Species examined: H. acletoi (US), H . acutidentata (NY),
H. angustifolia (NY), H. arenae (NY, PC), H . austrogeorgica
(NY), H. be/lei Bartr. (FH), H . densifolia (NY), H .
denticulata (NY, BM), H. heimii, H. heteroloma (BUF,
TENN), H. kunzeana (NY), H . limbata (NY), H .
longipedunculata (NY), H. longirostris (BM), H. macrophylla
(BM), H . marginata (BM), H. oedipodioides ~NY), H.
polyseta (BUF, FH, TENN), H. serrulata (NY), H .
stanfordensis (BUF, NY), H . steereana (BUF).
New combinations:
Hennediella acletoi (Robins.) Zand., comb. nov. (Tortula
Plate 98. Hennediellll. 1-6. H. polyseta. 1. Transverse section of stem. 2- 3. Two leaves. 4. Transverse section of costa at midleaf. 5. Theca. 6.
Operculum. 7-11. H. stanfordensis. 1. Transverse section of stem. 8-9. Two leaves. 10. Leaf apex. 11. Transverse section of costa at midleaf.
248
GENERA OF THE POTTIACEAE
acletoi Robins., Phytologia 12: 390, 1971).
Hennediella acutidentata (Card. & Ther.) Zand., comb. nov.
(Pottia acutidentata Card. & Ther., Bull. Mus. Hist. Nat.
Paris 22: 338, 1916).
Hennediella angustifolia (Herz.) Zand., comb. nov. (Calyptopogon angustifolius Herz., Beih. Bot. Centralbl. 26(2): 63,
1910; Tortula angustifolia (Herz.) Herz., nom. illeg.; Tortuta herzogii Zand.).
Hennediella arenae (Besch.) Zand., comb. nov. (Barbula
arenae Besch., Bull. Soc. Bot. France 32: 59, 1885; Tortula
arenae (Besch.) Broth.).
Hennediella arenae var. petriei (Broth. ex Beck.) Zand., comb.
et stat. nov. (Tortula petriei Broth. ex Beck., Trans. New
Zealand Inst. 29: 441, 1897; Tortula arenae subsp. petriei
(Broth.) Lightowlers, J. Bryol. 13: 371, 1985), not seen.
Hennediella be/lei (Bartr.) Zand., comb. nov. (Desmatodon
be/Iii Bartr., Bull. Brit. Mus. Nat. Hist. Bot. 2: 56, 1955).
Hennediella densifolia\ (Hook. f. & Wils.) Zand., comb. nov.
(Barbula densifolia London J. Bot. 3: 543, 1844; Tortula
densifolia (Hook. f. & Wils.) Hook. f. & Wils.).
Hennediella denticulata (Wils. in Mitt.) Zand., comb. nov.
(Barbula denticulata Wils. in Mitt., Kew J. Bot. 3: 50,
1851; Tortula denticulata (Wils. in Mitt.) Mitt.). This
peristomate species is probably the same as Tortula
densifolia (not seen).
Hennediella heimii (Hedw.) Zand., comb. nov. (Gymnostomum
heimii Hedw., Spec. Muse. 32, 1801; Pottia heimii (Hedw.)
Hampe).
Hennediella heimii var. alpina (Amann) Zand., comb. nov.
(Pottia heimii var. alpina Amann Bull. Soc. Vaudoise Sc.
Nat. 54: 42, 1921), not seen.
Hennediella heimii var. arctica (Lindh.) Zand., comb. nov.
(Pottia heimii var. arctica Oefv. K. Vet. Ak. Foerh. 23: 551,
1867), not seen.
Hennediella heimii var. brachyphylla (Warnst.) Zand., comb.
nov. (Pottia heimii var. brachyphylla Warnst., Hedwigia 58:
100, 1916), not seen.
Hennediella heimii var. brevicuspis (Warnst.) Zand., comb. nov.
(Pottia heimii var. brevicuspis Warnst., Hedwigia 58: 98,
1916), not seen.
Hennediella heimii var. breviseta (Warnst.) Zand., comb. nov.
(Pottia heimii var. breviseta Warnst., Hedwigia 58: 99,
1916), not seen.
Hennediella heimii var. drummondii (Warnst.) Zand., comb.
nov. (Pottia heimii var. drummondii Warnst., Hedwigia 58:
96, 1916), not seen.
Hennediella heimii var. eurystoma (Card. & Broth.) Zand.,
comb. nov. (Pottia heimii var. eurystoma Card. & Broth., K.
Svensk. Vet. Ak. Handl. 63(10): 20, 1923), not seen.
Hennediella heimii var. guessfeldtii (Schlieph.) Zand., comb.
nov. (Pottia guessfeldtii Schlieph., Ber. Deutsch. Bot. Ges.
2: 461, 1884; Pottia heimii var. guessfeldtii (Schlieph.)
Warnst.), not seen.
Hennediella heimii var. ianceolata (Warnst.) Zand., comb. nov.
(Pottia heimii var. lanceo/ata Warnst., Hedwigia 58: 98,
1916), not seen.
Hennediella heimii var. magellanica (Warnst.) Zand., comb.
nov. (Pottia heimii var. magellanica Warnst., Hedwigia
58: 95, 1916).
Hennediella heimii var. maxima (Card.) Zand., comb. nov.
(Pottia heimii var. maxima Car., Bull. Herb. Boiss. ser. 2,
5: 1002, 1905), not seen.
Hennediella heimii var. spegazzinii (C. Mtill.) Zand., comb.
nov. (Pottia spegazzinii C. Mtill., Flora 68: 414, 1885;
Pottia heimii var. spegazzinii (C. Mi.ill.) Warnst.), not
seen.
Hennediella heimii var. thaxteri (Card. & Ther.) Zand., comb.
nov. (Pottia heimii var. thaxteri Card. & Ther., Rev.
Bryol. n. ser. 2: 165, 1930), not seen.
Hennediella heteroloma (Card.) Zand., comb. nov. (Tortula
heteroloma Card., Rev. Bryol. 37: 127, 1910).
Hennediella heteroloma var. eckeliae (Zand.) Zand., comb.
nov. (Tortula eckeliae Zand., Bryologist 88: 354, 1985
[1986]).
Hennediella kunzeana (C. Mtill.) Zand., comb. nov. (Barbula
kunzeana C. Mi.ill., Linnaea 17: 586, 1843; Tortula
kunzeana (C. Mi.ill.) Mont. in Gay).
Hennediella limbata (Mitt.) Zand., comb. nov. (Barbula
limbata Mitt., Kew J. Bot. 3: 354, 1851; Tortula limbata
(Mitt.) Mitt., hom. illeg.).
Hennediella longipedunculata (C. Mi.ill.) Zand., comb. nov.
(Barbula longipedunculata C. Mi.ill., Syn. Muse. 1: 630,
1849).
Hennediella longirostris (Hampe ex C. Mi.ill.) Zand., comb.
nov. (Pottia longirostris Hampe ex C. Mi.ill., Syn. 1: 552,
1849; Tortula longirostris (Hampe ex C. Mi.i1l.) Broth. in
Par., hom. illeg.).
Hennediella marginata (Hook. f. & Wils.) Zand., comb. nov.
(Schistidium marginatum Hook. f. & Wils., London J.
Bot. 3: 539, 1844; Streptopogon marginatus (Hook. f. &
Wils.) Mitt.; Willia marginata (Hook. f. & Wils.) C.
Mi.ill.)."
Hennediella oedipodioides (C. Mi.ill.) Zand., comb. nov.
(Pottia oedipodioides C. Mtill., Bot. Jahrb. 5: 79, 1883).
Hennediella polyseta (C. Mtill.) Zand., comb. nov. (Barbula
polyseta C. Mi.ill., Nuov. Giom. Bot. ltal. n. ser. 4: 414,
1897; Tortula polyseta (C.Mi.ill.) Warnst.).
Hennediella serrulata (Hook. & Grev.) Zand., comb. nov.
(Tortula serrulata Hook. & Grev., Edinburgh J. Sc. 1:
299, 1824).
Hennediella steereana (Zand. & Crum) Zand., comb. nov.
(Desmatodon steereanus Zand. & Crum, Bryologist 80:
638, 1977).
Plate 99.
68. DOLOTORTULA
Dolotortula Zand., Phytologia 65: 425, 1989. Type: Dolotortula mniifolia (Sull.) Zand.
From dolus, from &5A.o<;, artifice, deceit, guile+ o + Tortula,
a genus; mimicking the genus Tortula.
Plants in a loose turf, green above, brownish green below.
Stems seldom branching, to 1.5 em in length, transverse section round, central strand present, distinct, sclerodermis and
POTIIOIDEAE
hyalodermis absent; axillary hairs of clear cells, several cells in
length; weakly rhizoidiferous. Leaves contorted, appresSed
when dry, widely spreading and somewhat rosulate when moist,
spathulate, 2.5-4.0 mm in length, upper lamina flat, very
weakly channeled along the costa, margins weakly recurved
below midleaf, entire, with a narrow cartilaginous border of
stereid cells to 4 cells in thickness and ending at or just before
apex; apex rounded or emarginate, often bluntly apiculate; base
little differentiated in shape; costa thin, percurrent or ending up
to 4 cells below apex, costa with lamina inserted laterally,
superficial cells ventrally long-rectangular, dorsally narrowly
elongate, 2 rows of cells across costa ventrally at midleaf, costal
transverse section round, stereid band weak ar,d rounded in
shape, ventral and dorsal epidermis present, the latter only laterally, guide cells 2 in 1 layer, hydroid strand present, small;
upper /aminal cells hexagonal to short-rectangular, large, ca.
22-28 Jlm in width, 1-2:1, walls thin, often weakly trigonous,
superficially weakly convex on both sides; papillae absent;
basal cells scarcely differentiated, short-rectangular, ca. 30 Jlm
in width, 2-3:1, walls thin. Dioicous. Perichaetia terminal, inner
leaves slightly larger than the cauline. Perigonia not seen. Seta
1.0-1.3 em in length, 1 per perichaetium, red-brown, twisted
clockwise; theca ca. 1.7-2.0 mm in length, red- or yellowbrown, cylindrical, exothecial cells thin-walled, hexagonal to
short-rectangular, 25-35 J..Lm in width, stomates at base of theca,
phaneropore, annulus of 3-4 rows of vesiculose cells, persistent; peristome teeth ca. 32, filamentous, anastomosing at base,
densely spiculose, ca. 1000 J..Lm in length, with many articulations, weakly twisted counterclockwise, basal membrane ca. 70
J..Lm in height, densely spiculose-papillose. Operculum not seen
(blunt-conic ex descr.). Calyptra not seen. Spores 10-13 J..Lm in
diameter, yellowish, weakly papillose. Lamina/ KOH color
reaction red.
A rare taxon of scattered distribution, being found on soil,
often calcareous, at moderate elevations; Mexico, West Indies,
Central America and the Andes of South America.
Among the three genera with epapillose, very large upper
!aminal cells (Chenia, Dolotortula and Sagenotortula), Dolotortula is unusual in its strong, multistratose border of stereid
cells (Pl. 99, f. 6, 8). Species of Tortula sect. Pottia and Tortula
sect. Hyophilopsis with bordered leaves and large upper !aminal
cells have somewhat the same appearance, but these have
(broadly) acute leaf apices, uni- to bistratose borders of
substereid cells, at least weakly papillose upper lamina! cells,
and have a yellow !aminal KOH reaction. Quite possibly,
through elaboration of the marginal border, loss of upper
lamina! papillae, and modification of cell wall chemistry to
enable the red color reaction (all apparently advanced character
states), Dolotortula was derived from shared ancestors of such
taxa (see Cladogram 14). It appears unrelated to Sagenotortula
because of the Syntrichia-1ike costal section of the latter genus,
while Chenia may be derived from borderless, serrate-margined
species of Tortula sect. Pottia (but if so then distantly, see
Cladograms 11 and 14). Tortula domingensis, which Crum and
Steere (1957) indicated was "doubtfully distinct" from D.
mniifolia (discussed as Tortula), is actually a Brachymenium
(new combination made in section on Excluded Taxa).
249
Additional literature: Bartram (1949), Zander (1989).
Number of accepted species: 1.
Species examined: D. mniifolia (BM, BUF, FH, TENN).
Plate 100.
69. PHASCOPSIS
Phascopsis Stone, J. Bryol. 11: 17, 1980. Type: Phascopsis
rubicunda Stone.
From Phascum, a genus + cl'ljfcrt~. -E~, appearance; resembling the genus Phascum.
Plants in dense clumps, green above, brown below. Stems
branching often, 1- 5 mm in length, cells of stem apex filled
with oil globules, transverse section rounded~pentagonal , central strand present but generally very weak, sclerodermis
absent, hyalodermis absent or possibly present but weak;
axillary hairs of 3-4 cells, basal 1-2 cells somewhat thickerwalled; rhizoids sparse. Leaves appressed, somewhat incurved
when dry, weakly spreading when moist, ca. 1.3 mm in
length, oblong to spathulate, upper lamina flat to broadly
channeled across leaf, margins plane, entire, weakly bordered
by 1(-4) rows of slightly thicker-walled cells; apex rounded to
broadly acute; base scarcely differentiated in shape; costa
short-excurrent as a mucro, superficial cells quadrate to
short-rectangular ventrally, elongate dorsally, ca. 4 rows of
cells across costa ventrally at midleaf, costal transverse section semicircular to round, stereid band strong and mostly circular in section, epidermis present ventrally, absent dorsally,
guide cells 2-4 (occasionally absent) in 1(-2) layers, hydroid
strand very strong or absent, often encircled by stereid cells,
costa often greatly bulging ventrally; upper /aminal cells
quadrate to irregularly rhomboidal or occasionally shortrectangular, ca. 13-15Jlm in width,1(-2):1, walls thin, superficially weakly convex on both sides; papillae hollow, ohshaped, simple or occasionally bifid, rather large, 4-6 per
lumen; basal cells differentiated across leaf, rising higher
medially, short-rectangular, somewhat wider than upper cells,
2-3:1, walls thin, hyaline. Cauline leaves with unusually
thickened costae apparently act as propagula. Dioicous. Perichaetia weakly differentiated, inner leaves oblong, apex
emarginate, somewhat shorter in length, not sheathing, cells
long-rhomboidal in lower 4/5. Perigonia terminal (but overtopped by subperigonial innovations on smaller plants). Seta
very short, ca. 0.2 mm in length, 1 per perichaetium, yellowish brown, not twisted; capsule cleistocarpous, ca. 1 mm in
length, yellowish brown, short-elliptical, apiculate, exothecial
cells thin-walled, hexagonal to short-rhomboidal to rectangular, stomates phaneropore, at base of theca, annulus absent.
Calyptra mitrate, lobed, smooth, ca. 0.7 mm in length. Spores
round to elliptical, 20-23 Jlm in longest diameter, hyaline,
weakly papillose. Lamina/ KOH color reaction red.
Known to date from only a few collections in arid southem and western Australia and in New Zealand, on clay,
calcareous or sandy soil.
Phascopsis is characterized by the cleistocarpous, nearly
globose, apiculate capsule (Pl. 100, f. 1); lobed, mitrate calyptra; broad, entire leaves with stout costa (Pl. 100, f. 3-5) and
250
GENERA OF THE POTTIACEAE
large, hollow-papillose upper laminal cells (Pl. 100, f. 11);
transverse section of costa showing only one stereid band, a
very strong hydroid strand, and occasional absence of distinct
guide cells (Pl. 100, f. 8-10); and dioicous sexual condition.
Stone (1980a) described, illustrated and discussed this taxon
exhaustively. There is a certain resemblance to Aschisma, but
that genus clearly has two stereid bands. Phascopsis may be
closely related, as the end of a reduction series, to Syntrichia
Plate 99. Dolotortula. 1-9. D. mniifolw. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Leaf apex. 7. Basal cells. 8. Transverse
section at midleaf. 9. Portion of peristome.
POITIOIDEAE
sect. Aesiotortula, but Cladograms 13 and 14 indicate its ancestors are more primitive (inserted deeper in the Pottieae
subclade) than those of Syntrichia.
The morphology exhibited in the transverse section of the
costa of Phascopsis is curious in its variability. A build up of
stereid and parenchymatous cells ventral to the hydroid strand,
possibly associated ultimately with occasional production of
propaguloid cauline leaves with much swollen costae (Pl. 100,
f. 12), results in encirclement of the hydroid strand (Pl. 100, f.
9) so that it appears in the center of the stereid band. This also
may result in apparent absence of guide cells. Sections made
below midleaf in larger leaves (Pl. 100, f. 8) and at midleaf in
smaller leaves, however, demonstrate morphology consistent
with other Pottieae. Crossidium aberrans, C. convolutum and
Tortula muralis may have a similar encirclement of the hydroid
strand by stereid cells, though there is no immediate phylo-
251
genetic relationship between Phascopsis and these taxa
(Cladograms 13 and 14). Stone (1980) observed a weak line
of dehiscence of an operculum in some capsules of Phascopsis.
New Zealand collections of P. rubicunda include one
labeled as Pottia stevensii [R. Br. ter], New Zealand, North 1.,
Hawke Bay, near Wairoa, Whakama.hia, sandy soil adjoining
sea beach, Sainsbury, 3 Oct. 1931, US ex F and another determined as Pottia longifolia R. Br. ter by H. Dixon, North 1.,
Wellington, Berggren, 1874, NY. These names predate that of
P. rubicunda and their types should be examined. The upper
costa is only weakly swollen in collections other than the type
and the combination of plane, unbordered laminal margins
and red KOH reaction is here considered diagnostic.
Number of accepted species: 1.
Species examined: P. rubicunda (MELU, NY, US).
Plate 100. Phascopsis. 1-13. P. rubicunda. 1. Habit (and calyptra). 2. Transverse section of lower stem. 3-5. Three leaves. 6. Leaf apex. 7.
Basal cells. 8. Transverse section of leaf base. 9-10. Transverse sections near midleaf. 11. Upper lamina! papillae. 12. Propaguloid cauline leaf.
13. Apiculus of capsule.
252
GENERA OF THE POTTIACEAE
Plate 101.
70.STONEA
Stonea Zand., Phytologia 65: 431, 1989. Type: Stonea
oleaginosa (Stone) Zand.
Named for Ilma G. Stone, an Australian bryologist well known
for her detailed morphological and anatomical studies of
Australasian arid habitat mosses.
Plants gregarious, mostly buried in soil, green above, reddish brown below. Stems seldom branching, very short, to 0.3
mm in length, transverse section rounded, central strand absent,
sclerodermis absent, hyalodermis absent; axillary hairs 2-4
cells in length, basal cells firm-walled; lower stem clothed with
fine rhizoids. Leaves incurved when dry, weakly spreading
when moist, obovate or short-lingulate, occasionally wider than
long, short, ca. 0.4-0.5 mm in length, upper lamina broadly
and deeply concave, margins plane, entire or dentate at apex;
apex usually broadly and often also sharply apiculate, usually
cucullate; base not differentiated in shape; costa percurrent or
occasionally ending 1-3 cells below apex, costa with lamina
inserted nearly laterally, superficial cells papillose, quadrate
and usually bulging ventrally, dorsally elongate, sharply papillose near apex dorsally, ca. 3 rows of cells across costa ventrally at midleaf, costal transverse section semicircular to circular, stereid band weak and rounded in shape, ventral epidermis
present, often strongly bulging, dorsal epidermis present or
absent, guide cells 2 in one layer or absent, hydroid strand
absent, costa often expanded as a large, ventrally bulging,
rounded, oil-rich excrescence nearly as wide as the leaf but this
affecting only the deciduous, uppermost leaves; upper lamina!
cells quadrate, ca. 13 Jlm in width, 1: 1, walls thin, superficially weakly convex on both sides of lamina; /aminal papillae present only dorsally near costa at leaf apex, 1-3 per
lumen, simple, hollow to solid; basal cells only weakly
differentiated, quadrate to short-rectangular, ca. 15 Jlm in
width, 1-2:1, walls thin. Apparently dioicous (naked axillary
archegonia reported in original description). Sporophytes and
androgametophytes unknown. Lamina/ KOH color reaction
red. (Because of paucity of available material, details of the
costal section are from the original description and illustrations.)
Found on soil or thin soil over limestone in dry areas of
southern Australia.
Stonea oleaginosa has a swollen, oil-rich lenticular knob
on the ventral surface of the upper costa of the very uppermost leaves on the stem. Leaves farther down on the stem
lack this excrescence, but instead have protruding, bottleshaped, papillose cells on the ventral costal surface much like
those on the costa of Crossidium aberrans. Stonea is entirely
red in KOH and has plane margins, features absent in Crossidium. As in the case of Gymnostomiella ((q.v.), the leaves of
Stonea may have attained their general lack of distinctive
characters through paedomorphosis. This genus is probably
closely related to Syntrichia; S. caninervis has a similar
spinose-papillose dorsal surface of the costa, a very small
immature habit, and similar oil globules in its upper laminal
cells, but Stonea differs by its obovate leaves with plane
Plate 101. Stonea. 1-6. S. oleaginosa. 1. Habit. 2. Ventral surface of a propaguloid leaf from near plant apex. 3. Apex of lower leaf, dorsal
view. 4. Propaguloid leaf. 5-6. Lower leaves, dorsal and ventral views.
POTTIOIDEAE
margins and upper laminal cells smooth except for the extreme
upper marginal cells and the tip of the costa. There is some similarity with Chenia leptophylla, which also has a strongly papillose and sharply apiculate leaf apex, but which has larger
laminal cells and a very narrow costa.
Literature: Stone (1978).
Number of accepted species: 1.
Species examined: Stonea oleaginosa (MELU).
Plate 102.
71. ACAULON
Acaulon C. Mi.ill., Bot. Zeit. 5: 99, 1847. Lectotype: Acaulon
muticum (Hedw.) C. Mi.ill. see Limpr., Laubm. Deutsch!. 1:
178. 1885.
Sphaerangium Schimp., Syn. 12, 1860, nom. illeg. incl. gen .
prior.
Phascum subg. Acaulon (C. Mi.ill.) Wils. in Hook. f., Fl. Nov.
Zel. 2: 58, 1854.
Acaulon subg. Alaticosta Stone, J. Bryol. 9: 213, 1976, nom.
inval. holotyp. non cit.
Acaulon sect. Sphaerangium C. Mi.ill., Gen. Muse . Fr. 20,
1900.
Subg. Alaticosta Stone, J. Bryol. 9: 573, 1977. Type: Acaulon
chrysacanthum Stone.
Subg. Acaulonopsis Stone, J. Bryol. 15: 746, 1989. Type:
Acaulon robustum Broth. ex Roth.
From the alpha privitive + KauA.o<;, stalk, stem; stemless.
Plants very small, gemmate , gregarious or scattered, reddish
or occasionally yellowish brown above, brown below. Stems
not branching, very short, to 0.5 mm in length, transverse section rounded, central strand absent, sclerodermis absent, hyalodermis absent; axillary hairs ca. 5 cells in length, proximal cell
walls sometimes thickened; sparsely radiculose. Leaves strongly
oppressed and apices usually reflexed when dry, appressed to
weakly spreading when moist, ovate, small, 0.5-1.75 mm in
length, lamina broadly channeled and usually deeply concave,
margins plane, entire to serrulate or dentate; apex broadly
acute; base not differentiated in shape; costa excurrent in a
sharp apiculus or stout mucro, occasionally only percurrent or
as a short, sometimes dentate awn, costa with lamina inserted
laterally, superficial cells elongate and smooth ventrally, dorsally elongate and usually smooth, 3-4 rows of cells across
costa ventrally at midleaf, costal transverse section round,
stereid bands ventrally generally absent, dorsally present (but
usually weak) and rounded in shape, ventral and dorsal epidermises present, guide cells 2-4 in l layer or rarely absent,
hydroid strand usually present, occasionally apparently centered
in the stereid band, costal outgrowths sometimes present, of ca.
2 longitudinal lamellae formed on ventral surface of the costa;
upper /aminal cells rounded-quadrate to rhomboidal, ca. 13-15
Jlm in width, 1-4:1, walls evenly thickened, ocGasionally highly
thickened on dorsal walls, superficially convex on both sides of
lamina; papillae absent or occasionally large and simple, 1(-2)
253
per lumen ; leaf base not differentiated in shape, basal cells
rectangular, little wider than upper cells, 3-4:1, walls thin.
Dioicous and perigoniate plants smaller than the perichaetiate
(often much reduced), or else monoicous and usually
paroicous. Perichaetia terminal, inner leaves somewhat
enlarged. Seta very short, to 0.2 mm in length, 1 per
perichaetium, light brown; theca cleistocarpous, spherical,
apiculus lacking, ca. 0.4-D.7 mm in diameter, light brown,
exothecial cells quadrate to rhomboidal, 25-50 Jlm in width,
1: 1, thin-walled, stomates phaneropore, occurring at base of
capsule. Operculum absent. Calyptra mitrate, often lobed,
smooth, ca. 0.1--0.4 mm in length. Spores rather large, ca.
25-35 Jlm in diameter, spherical to weakly elliptical, light
brown, lightly papillose to irregularly warty or spiculose.
Lamina/ KOH color reaction red. Reported chromosome
number n :::: 26.
Found on soil, a widely distributed genus mainly found in
temperate regions of low rainfall.
Acaulon is clearly similar to Microbryum, differing in the
even smaller habit size, capsules spherical and lacking an
apiculus (Pl. 102, f. 28), upper laminal margins plane, and
papillae lacking in most species. Acaulon schimperianum (Pl.
102, f. 29) has upper laminal papillae somewhat like those of
Microbryum vlassovii and M. floerckeanum (one to two over
each lumen, simple or occasionally branching apically, hollow, often rather tall). Those species of Microbryum with
short-ovate leaves and no papillae also, like Acaulon, have
rectangular ventral quadrate cells. Acaulon may be an end
member of a series including Tortula sect. Tortula and Microbryum through reduction of gametophyte and sporophyte size
and complexity, and differentiation of red coloration. This is
supported somewhat by Cladograms 12 and 14 to the extent
that Tortula is lower on the tree than the other genera.
In A. eremico/a (Pl. 102, f. 14-15), A. muticum and A.
schimperianum, the perigoniate plants are about a quarter to a
third the size of the perichaetiate plants, and are situated near
the base of the perichaetiate plants (possibly rhizautoicous).
The costa may appear, in section, to have two stereid
bands in species (e.g. A. chrysacanthum, Pl. 102, f. 11-13)
that develop strong awns. Stone (1976b, 1977b, 1979, 1988,
1989) has discussed the morphology of Australasian Acaulon
species in a particularly incisive manner. Casas et al. (1990)
gave a key to the five species of the genus on the Iberian Peninsula, while Sergio (1992) noted that the European A.
piligerum belongs in the lamellate subgenus Alaticosta, previously known only from Australia.
Additional literature: Bryan (1956), Cardenas (1988),
Crum and Anderson (1965), Grout (1945), Hill (1982), Sergio
(1972a, 1992).
Number of accepted species: 15.
Species examined: A . chrysacanthum (NY), A . eremicola
(MELU), A. integrifolium (BUF, NY), A . leucochaete (BUF,
NY), A. muticum (BUF), A. robustum (NY), A . schimperianum, A . triquetrum (BUF), A. uleanum (BM).
254
GENERA OF THE POTTIACEAE
Plate 102. Acaulon. 1-10. A. uleanum. 1. Habit. 2. Perigoniate plant. 3-5. Three leaves. 6. Leaf apex. 7. Leaf section. 8. Sporophyte with
calyptra. 9. Exothecial cells. 10. Spores. 11-13. A. chrysacanthum. 11. Habit. 12. Leaf apex. 13. Calyptra. 14-15. A. eremicola. 14. Perigoniate
plant. 15. Leaf apex. 16. Leaf section. 17-19. A. inaequalifolium. 17-18. Two leaves. 19. Spores. 20-21. A. integrifolium. 20-21. Two leaves.
22-25. A. leucochaete. 22-23. Two leaves. 24. Leaf apex. 25 . Leaf section. 26-28. A. robustum. 26-27. Two leaves. 28. Sporophyte with
vaginula. 29. A. schimperianum. 29. Leaf apex. 30-33. A. triquetrum. 30-31. Two leaves. 32. Leaf section. 33. Spores.
POTIIOIDEAE
Plate 103.
72. SARCONEURUM
Sarconeurum Bryhn, Nyt Mag. Naturvid. 40: 204, 1902. Type:
Sarconeurum antarctium Bryhn.
From crdp~, crcxpK6cr, flesh + o + VE~ov, nerve, sinew, tendon; the costa is thickened, appearing meaty to some.
Plants forming dense cushions, blackish green above, reddish brown below. Stems often branching, to 3 em in length,
rounded-pentagonal in transverse section, central strand absent
to strong, sclerodermis absent, hyalodermis indistinct; axillary
hairs of ca. 10 cells, the basal 2-3 firm-walled, yellow. Leaves
appressed-incurved when dry, spreading-recurved when moist,
ligulate-lanceolate, to 1.5 mm in length, broadly channeled
across the ventral surface; margins plane, entire; lamina constricted below apex, which ends in a deciduous, cylindrical,
sharply apiculate propagulum; base scarcely differentiated in
shape to ovate; costa running into the apical propagulum,
epidermal cells quadrate to short-rectangular on both leaf surfaces, ca. 4 rows of cells across costa ventrally at midleaf, in
transverse section with one stereid or substereid band dorsally
(this generally lacking in small/eaves), 2-3 guide cells in 1(-2)
layers, hydroid strand usually present, a ventral stereid band
occasionally present near the leaf base in large leaves, epidermis present ventrally, absent dorsally; upper !aminal cells quadrate to rectangular, often transversely elongated, especially
along the margins, 15-20 IJ.m wide, 1-2:1, walls evenly thickened or collenchymatous, superficially flattened, papillae low,
small, punctiform or bifid, apparently solid, several per lumen;
basal cells differentiated medially, thin-walled, bulgingrectangular, to 23 j..lm in width, 3-5:1. Sexual organs and
sporophyte unknown. Lamina[ KOH color reaction red.
Found on sandy and volcanic soil or on lava at low elevations in Antarctica, and, rarely, southernmost South America.
A report (Matteri 1982) of sporophytes of Sarconeurum
glaciale is based on specimens (Argentina: trunks of
Nothofagus, TBPA B109, TBPA 3486, BA!) that can be
referred to Tortula pygmaea Dix., which is a good species (BM!
and see Lightowlers 1985b) found in southern South America
and New Zealand. Greene (1975) reported Sarconeurum from
South America apparently from specimens of T. pygmaea, a
species curiously similar to S. glaciate in general habit and the
constricted leaf apex bearing a deciduous, sharply apiculate
propagulum (Pl. 103, f. 3-6), but differs in the smaller (ca. 10
IJ.m in width) leaf cells with large, solid bifid papillae and the
deep medial groove along the ventral surface of the costa. On
the other hand, Lightowlers (1985b) demonstrated that the type
("Fuegia septentrionalis") of the South American T. lithophila
at Sis actually S. glaciate.
Sarconeurum tortelloides has been transferred to Tortella,
leaving Sarconeurum monotypic.
Hilpert (1933) referred Sarconeurum to the Pottieae near
Tortula (without further discussion) but probably because of the
generally single stereid band in the costa (Pl. 103, f. 8-10) and
the large upper lamina! cells. There is considerable similarity to
Bryoerythrophyllum, however, in the essentially oblonglanceolate leaf shape, the presence of apical !aminal propagulum in the related genus Mironia (but cf. Tortella
255
tortelloides, which has caducous leaf apices), the bifid papillae, the occasional second stereid band, the medially
differentiated basal cells (Pl. 103, f. 7) and the red color in
KOH. The cladistic analysis places Sarconeurum in the Pottieae (see Cladograms 13 and 14), surprisingly, near Acaulon.
Additional literature: Greene et al. (1970), SaviczLjubitzkaja & Smirnova (1961), Zander (1978h), Zander &
Hoe (1979).
Number of accepted species: 1.
Species examined: S. glaciate (BM, NY, US).
Plate 104.
73.CHENIA
Chenia Zand., Phytologia 65: 424, 1989. Type: Chenia
subobliqua (Williams) Zand.
Phascum sect. Leptophascum C. Mtill., Flora 71: 7, 1888.
Type: Phascum leptophyllum C. Mtill.
Phascum subg. Leptophascum (C. Mtill.) Roth, Aussereur.
Laubm. 2:214, 1911.
Named for Chen Pan-chieh, 1907-1970, author of "Studien
tiber die ostasiatischen Arten der Pottiaceae" (1941) and also
author or editor of "Genera Muscorum Sinicorum" (1963,
1978).
Plants forming turfs, occasionally rosulate, green above,
brownish below. Stems seldom branching, 0.3-1.0 em in
length, transverse section rounded, central strand weak to
strong, sclerodermis present, hyalodermis absent; axillary
hairs small, of 3-4 cells, basal cell firm-walled; rhizoids usually few. Leaves appressed and somewhat contorted when dry,
spreading when moist, ligulate to spathulate, 1.5-2.5 mm in
length, upper lamina occasionally grooved along costa, plane
or broadly channeled across leaf, margins plane above,
weakly recurved below, sharply crenulate to irregularly
dentate above with sharp mid-marginal wall projections usually ending in a weak simple papilla, marginal cells often
smaller than the medial; apex rounded to broadly acute, often
sharply apiculate by a distinctive thick-walled cell or cells;
base rectangular or not differentiated in shape; costa weak,
ending several (6-9) cells before the apex or percurrent, costa
with lamina inserted ventrally to laterally, ventral and dorsal
superficial cells short-rectangular, narrower than !aminal
cells, 2 rows of cells across costa ventrally at midleaf, costal
transverse section rounded to elliptical, stereid band very
weak or occasionally absent, rounded in shape, ventral and
dorsal epidermis present, guide cells 2 in 1 layer, hydroid
strand present, small to large; upper /aminal cells large,
bulging-hexagonal, 15-18 IJ.m in width, 1:1, walls thin,
weakly trigonous, superficially convex on both sides; papillae
absent (upper marginal teeth may be interpreted as ending in
sharp, simple papillae); basal cells differentiated across the
leaf base (except for one row of marginal cells similar to the
upper cells), rectangular, ca. 20-25 j..lm in width, 2-4:1, walls
thin, weakly trigonous. Propagula when present borne on
rhizoids in soil, irregularly rounded to clavate, ca. 100-130
j..lm in longest dimension. Dioicous. Perichaetia terminal,
inner leaves little different from the cauline, slightly
256
GENERA OF THE POTIIACEAE
larger.Seta ca. 0.1-1.2 em in length, 1 per perichaetium, reddish
brown, twisted clockwise; when present theca 0.7-2.0 mm in
length, brown, nearly spherical (then with a narrow beak to 0.25
mm in length) or short-ovate or cylindrical, exothecial cells rectangular, 20-45 Jlm in width, 1-4:1, thin-walled, stomates at
base of theca, phaneropore, cleistocarpous or stegocarpous,
annulus when present of 2-3 layers of strongly vesiculose
cells, persistent; when present peristome teeth 32, filamentous, somewhat anastomosing, densely branchingspiculose, ca. 400 Jlm in length, with ca. 5 articulations,
nearly straight or weakly twisted counterclockwise, basal
membrane 25-35 Jlm in height, low spiculose. Operculum
Plate 103. Sarconeurum. 1-10. S. glaciale. 1. Habit. 2. Transverse section of stem. 3-5. Three leaves. 6. Propaguloid leaf apex. 7. Basal cells.
8-10. Transverse sections near midleaf.
POITIOIDEAE
257
Plate 104. Chenia. 1-8. C. subobliqua. I. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Peristome. 9-13. C. /eptophylla. 9-10. Two leaves. 11. Leaf apex. 12. Transverse section at midleaf. 13. Propagula.
14-16. C. lorentzii. 14-15. Two leaves. 16. Leaf apex.
258
GENERA OF THE POTTIACEAE
when differentiated short- to long-conic, 500-650 J..Lm in length,
cells twisted weakly counterclockwise. Calyptra cucullate or
mitriform and then three-lobed, smooth, 1.5-2.0 mm in length.
Spores 10-20 J..Lm in diameter, light brown, nearly smooth to
finely papillose. Lamina/ KOH color reaction red. [Information
on the cleistocarpous sporophyte of C. leptophylla from Arts
and Sollman 1991]
Found on soil from sea level to nearly 3000 meters elevation; North and South America, Europe, eastern Asia and Australia.
Corley et al. (1981) suggested that Chenia leptophylla (as
Tortula rhizophylla) "is completely anomalous in Tortula." The
Andean C. obliqua (Pl. 104, f. 1-8) is closely related and differs
from C. rhizophylla in the peristomate sporophyte, broad leaf
apex, margins strongly dentate in the upper 1/3 of the leaf and
lacking a leaf apiculus of thick-walled cells. Although these two
species are rather different, a third species, C. lorentzii (Pl. 104,
f. 14-18), is sterile but otherwise intermediate in character, with
narrower leaves, not or moderately dentate, and the apiculus is
occasionally differentiated and thick-walled. Small, sterile
plants mixed in with the type of C. lorentzii are quite like C.
leptophylla. The autoicous condition ascribed to C. lorentzii in
the original description cannot be confirmed as no antheridia
were found in the NY isotype.
Like Stegonia, Chenia is a small genus that includes both
peristomate and cleistocarpous species. Chenia is easily distinguished from other Tortula-like species by the combination of
the dentate upper leaf margins, large, epapillose upper laminal
cells, a thin costa (Pl. 104, f. 12), and red coloration in KOH.
The last may have to be determined by examination of the
upper laminal cell walls under high magnification because the
dense yellow-green of the chlorophyll overwhelms the color of
the very thin cell walls. The leaf teeth are each usually tipped
with a single, simple papilla. Phascum leptophyllum of central
and southern Africa, an earlier name than Physcomitrium
rhizophyllum Sak., was reported by Arts and Sollman (1991) to
be the same as C. rhizophylla, along with Pottia denticulata
Dix. & Yarde of India and P. splachnobryoides C. Mtill. of
Asia; they describe and illustrate the sporophyte with its cleistocarpous capsule, and review the literature on this widely distributed species. Chenia leptophylla is, as is the case with Didymodon australasiae var. umbrosus (cf Crundwell & Whitehouse
1978, Eckel 1986a, Preston & Whitehouse 1985, Synnott &
Robinson 1990, and others), apparently spread by human
agency through rhizoid-borne propagula (Pl. 104, f. 13) in the
soil. Chenia is similar to Hennediella, especially H. serrulata
(which has a quite similar gametophytic appearance); also,
Tortula paulsenii has somewhat similar morphology. Phylogenetic analysis indicates a close relationship, however, to Syntrichia (see Cladograms 13 and 14).
Additional literature: Cortini and Aleffi (1989), lwatsuki
and Saito (1972), Martinez et al. (1989), Neumann ( 1972),
Pedrotti and Aleffi (1989), Reese (1967, 1968), Smith and
Whitehouse (1974), Sollman (1979), Stone (1980a).
Number of accepted species: 3.
Species examined: C. lorentzii (NY), C. leptophylla (BUF,
PAC), C. subobliqua (NY).
New combinations: Chenia leptophylla (C. Mtill.) Zand.,
comb. nov. (Phascum leptophyllum C. Mtill., Flora 71: 6,
1888). Chenia lorentzii (C. Mtill.) Zand., comb. nov. (Barbula
lorentzii C. Mtill., Linnaea 42: 346, 1879 ["-i"]; Tortula
lorentzii (C. Mtill.) Broth.
74. SYNTRICHIA
Plates 105-111.
Syntrichia Brid., J. Bot. (Schrader) 1(2): 299, 19 April 1801
(vide Sayre 1959). Type: Syntrichia ruralis (Hedw.) Web.
& Mohr, lectotype fide Zander, Phytologia 65: 432, 1989.
Tortula subg. Syntrichia (Brid.) Chev., Fl. Gen. Env. Paris
2: 52, 1827.
Barbula subg. Syntrichia (Brid.) BSG, Bryol. Eur. 2:10,
1851 (fasc. 46-47 Consp. 2: Ill).
Tortula sect. Syntrichia (Brid.) Lam. & Cand., Syn. 100,
1806.
Tortula sect. Rurales De Not., Mem. R. Ace. Sc. Torino 40:
286, 1838. Type: Tortula ruralis (Hedw.) Gaetrn., Meyer
& Scherb.
Barbula sect. Rurales BSG, Bryol. Eur. 2: 101, 1842 (fasc.
13-15 Mon. 39).
Barbula sect. Syntrichia (Brid.) C. Mi.ill., Syn. I: 632, 1849.
Barbula sect. Vallidens C. Mtill., Linnaea 42: 347, 1879,
nom. inval. Type: Barbula percarnosa C. Mi.ill.
Barbu/a sect. Syntrichiae Lesq. & James, Man. N. Amer.
Moss. 130, 1884, nom. illeg.
Barbula sect. Ruraliformes Kindb., Eur. N. Amer. Bryin. 2:
245, 1897, nom. illeg. incl. sect. prior. Type: Barbula
ruralis Hedw.
Syntrichia sect. Eusyntrichia Moenk., Laubm. Eur. 306,
1927, nom. illeg.
From crUV, together, with + 9pix, 'tplxoc;, hair; the peristome teeth are fused at the base into a cone.
Plants usually coarse, forming often deep turf, green,
occasionally reddish or blackish above, reddish brown below.
Stems branching occasionally, 1-4(-12) em in length, transverse section rounded-pentagonal, central strand present or
absent, sclerodermis absent or occasionally of 1-2 layers of
substereid cells, hyalodermis absent or present, sometimes
collapsed; axillary hairs ca. 3-10 cells in length, basal 1-3
cells thicker walled or all hyaline; rhizoids common, often
dense. Leaves appressed to weakly spreading when dry,
spreading to squarrose when moist, narrowly ligulate to
broadly spathulate, occasionally broadly lanceolate,
(1.5-)4-7 mm (inclusive of awn) in length, upper lamina
broadly channeled to keeled, often narrowly grooved along
costa, margins plane to recurved, rarely broadly incurved,
usually entire, occasionally serrulate to dentate above, occasionally bordered with thick-walled, less papillose cells, these
occasionally intrairlarginal or rarely elongate; apex rounded
to broadly acute, occasionally narrowly acute or cucullate;
base not differentiated in shape to elliptical; costa often stout,
commonly prominent dorsally, ending several cells before
apex to excurrent as a long, often hyaline, often serrate, occasionally flattened awn, costa with lamina inserted at 45',
POITIOIDEAE
259
Plate 105. Syntrichia. 1-11. S. ruralis. 1. Habit. 2. Transverse section of stem. ~. Two leaves. 5. Leaf apex. 6. Basal cells. 7. Transverse
section at midleaf. 8. Paraphyses. 9. Capsule with peristome. 10. Operculum. 11. Calyptra. 12-16. S. ammondsiana. 12-13. Two leaves. 14.
Leaf apex. 15. Transverse section at midleaf. 16. Cluster of propagula.
260
GENERA OF THE POTTIACEAE
Plate 106. Syntrichia. 1-8. S. amphidiacea. l-4. Four leaves. 5. Leaf with propagula. 6. Leaf apex. 7. Transverse section at midleaf. 8.
Peristome. 9-13. S. anderssonii. 9. Transverse section of stem. 10-11. Two leaves. 12. Leaf apex. 13. Transverse section at midleaf. 14-21. S.
andicola. 14. Transverse section of stem. 15. Leaf. 16. Leaf apex. 17. Upper lamina! papillae. 18. Transverse section at midle"af (internally pored
lamina! cells). 18. Perichaetialleaf. 20. Peristome. 21. Calyptra.
POTTIOIDEAE
costal superficial cells rounded-quadrate, papillose, occasionally elongate and smooth near apex ventrally, dorsally elongate,
papillose to denticulate or spinose, occasionally nearly smooth,
2-4 rows of cells across costa ventrally at midleaf, costal transverse section round, occasionally elliptical or semicircular,
stereid bands ventrally absent or very rarely of a few substereid
or stereid cells, dorsally present and crescent-shaped, occasionally semicircular, ventral epidermis present, the dorsal usually
absent, guide cells 2-4 per layer in 1-4 layers, hydroid strand
absent or present, occasionally multiple; upper lamina/ cells
rounded-quadrate, ca. 11-15(-20) Jlm in width, 1(-2):1, occasionally bistratose entirely or in patches, walls thin or occasionally collenchymatous and interiorly porose, superficially
strongly convex on both sides; papillae ca. (1-)4-8 per lumen,
usually bifid, occasionally simple, solid or hollow, rarely simple; basal cells differentiated across base, rising higher medially, often sharply differentiated from the upper cells, occasionally inflated medially, 18-30 J..lm in width, 2-6:1, walls thin,
sometimes irregularly porose, 3-6 rows of basal marginal cells
usually narrowly rectangular and thicker walled. Propagula
occasionally present, variously clusters of small, deciduous
leaves; rhizoid-borne tubers; very fragile cauline leaves or leaf
tips (rarely deciduous); obovate or clavate or raspberry-shaped
propagula found on ventral surface of costa or medially or marginally on upper lamina. Dioicous or monoicous (then usually
autoicous or synoicous). Perichaetia terminal, inner leaves usually little differentiated in size and shape, rarely enlarged and
sheathing, lower cells rhomboidal, rarely throughout leaf. Perigonia gemmate, terminal or lateral as stalked buds, paraphyses
often clavate. Seta elongate, (0.8-)1.0-3.0 em in length, 1(-2)
per perichaetium, reddish brown, twisted counterclockwise;
theca 2-6 mm in length, reddish brown, cylindrical, commonly
slightly curved, exothecial cells rectangular, 20-30 J..lm in
width, ca. 4-6: 1, walls thin to evenly thickened, capsule neck
occasionally distinct, stomates phaneropore, at base of theca,
annulus of 2-4 rows of vesiculose cells, persistent; peristome
teeth 32, filamentous, densely spiculose, ca. 1000-2000 J..lm in
length, with many articulations, twisted counterclockwise ca.
once, basal membrane usually present, ca. 100-800 J..lm in
height, papillose to spiculose. Operculum conic, rather large,
1.2-2.2 mm in length, cells twisted counterclockwise. Calyptra
cucu/late, smooth, rather large, 3.0-4.5 mm in length. Spores
8-15 J..lm in diameter, light brown, lightly papillose. Lamina/
KOH color reaction brick red. Reported chromosome number n
=6+m, 12, 12+m, 13, 13+m, 24, 24+2m, 26, 28, 32+2m, 48.
Found on rock, soil and bark on all continents, most commonly in temperate areas but also characteristic of dry climates.
Syntrichia is segregated from Tortula s. lat. by the combination of red KOH reaction of the upper !aminal cells, lack of narrowly elongate upper !aminal marginal cells, the crescentshaped transverse section of the stereid band, and the general
lack of differentiated dorsal costal epidermal cells. This concept
is essentially that of Kramer (1980, 1988), who emphasized the
exposed dorsal stereid band (Pl. 105, f. 7, 15), not covered dorsally by parenchymatous or otherwise differentiated epidermal
cells. Recently Ochyra (1992) supported this concept (Zander
1989) as "a natural group that deserves recognition as a genus
261
of its own" with a series of new combinations appropriate for
the Polish flora. Syntrichia is distinguished from Hennediella
(likewise KOH red) by the leaves ligulate to spathulate or
very seldom lanceolate, margins usually recurved and seldom
dentate or bordered, upper !aminal cells smaller and superficially strongly convex, costa commonly excurrent as an awn,
almost always lacking a dorsal costal epidermis, stereid band
semicircular to crescent-shaped, and sporophytes not in a
reduction series within the genus.
Note that in some few species (e.g. S. brandisii) a dorsal
epidermis is very weakly differentiated (seen as somewhat
wider lumens in costal transverse section), but these species
are placed here rather than with Hennediella because of their
recurved, unbordered margins and small upper !aminal cells.
Syntrichia costesii (Pl. 107, f. 17-19), like certain species of
Hennediella, has an intramarginal border of differentiated
cells, but those cells are isodiametric and the costa lacks a
differentiated dorsal epidermis. The existence of a weakly
differentiated dorsal epidermis in some species may indicate a
derivation of Syntrichia from Bryoerythrophyllum, which differs, in traditional characters, in the presence of a second
stereid band and usually narrower (broadly lanceolate to
ligulate) leaves. Most cladograms in the phylogenetic analysis
show Syntrichia and Bryoerythrophyllum to be quite distant.
A few species, like S. laevipila (Pl. 109, f. 4-5), may occasionally and perhaps abnormally have one or two stereid or
substereid cells present ventrally between the guide cells and
the ventral epidermis as a tiny second stereid band, while in
the type (isotype, NY) of S. rubra (Pl. Ill, f. 5-8) and a second specimen seen (as Tortula rubra var. subantarctica,
Campbell 1., Sorensen 1946, isotype, NY), the ventral stereid
band is of several stereid cells, the ventral epidermis is lacking, and superficially, the ventral surface of the costa is similar to the dorsal surface, being of superficially elongate, simply papillose cells. Syntrichia percarnosa (Pl. 109, f. 20-22)
has much the same leaf morphology as Trichostomum
crispulum, and the former species may well belong with that
genus; the substereid cells of the costa are, however, arranged
more like those of Syntrichia.
Certain species of Didymodon sect. Vineales (KOH red)
may lack a ventral stereid band, but have lanceolate leaves
and narrowly rectangular basal cells. Bryoerythrophyllum and
Mironia species are also KOH red and may have long-ligulate
leaves, but the transverse section of the costa is reniform in
section and the stem has a sclerodermis. Syntrichia
geheebiaeopsis (Pl. 108, f. 13-17, cf. discussion by
Lightowlers 1985a as Tortula) has many of the characters of
Bryoerythrophyllum and Mironia, including broadly lanceolate, dentate leaves with a sheathing leaf base, small upper
!aminal cells with low, crowded papillae. It further differs
from Syntrichia by the very narrowly rectangular basal cells.
Bryoerythrophyllum and Mironia have semicircular to
reniform costal sections, however, with usually two distinct
stereid bands and a differentiated dorsal epidermis, but S.
geheebiaeopsis has the typical costal morphology of Syntrichia: a nearly circular costal section, with a single stereid
band and no differentiated dorsal epidermal cells. It may
262
GENERA OF THE POTTIACEAE
Plate 107. Syntrichill. 1-4. S. bipedicellata. 1-2. Two leaves. 3. Leaf apex. 4. Peristome. S-6. S. cainii. 5. Leaf. 6. Leaf apex. 7-12. S. cavalii.
7-8. Two leaves. 9. Leaf apex. 10. Perichaetium. 11. Perichaetialleaf. 12. Perigonia. 13-16. S. chisosa. 13. Leaf. 14. Leaf apex, dorsal view.
15. Transverse section at midleaf. 16. Propagulum. 17-19. S. costesii. 17-18. Two leaves. 19. Leaf apex. 20-23. S. didymodontoides. 20-21.
Two leaves. 22. Leaf apex. 23. Transverse section at midleaf.
POTTIOIDEAE
263
~21
18
Plate 108. Syntrichia. 1-9. S. epilosa. 1-2. Two leaves. 3. Leaf apex. 4. Transverse section at midleaf. 5. Perichaetialleaf. 6-7. Peristomes. 8.
Operculum. 9. Calyptra. 10-12. S.jragilis. 10-11. Two leaves. 12. Transverse section at midleaf. 13-17. S. geheebiaeopsis. 13-14. Two leaves.
15. Leaf apex. 16. Basal cells. 17. Transverse section at midleaf. 18-21. S. gemmascens. 18-19. Two leaves. 20. Transverse section at midleaf.
21. Propagula. 22-24. S. obtusissima. 22. Leaf. 23. Leaf apex. 24. Transverse section at midleaf.
264
GENERA OF THE POTTIACEAE
prove to be a good genus in the Merceyoideae apparently having lost, like Streptopogon, the ventral stereid band.
Syntrichia cava/Iii (Pl. 107, f. 7-12) is unusual in its
strongly differentiated perichaetial leaves that are similar to
those of S. papillosa (sect. Collotortula); it is, however, apparently related to S. percarnosa, instead. Syntrichia cavallii differs from Calyptopogon, in which perichaetial leaves are
differentiated, in that propagula are lacking and the upper
laminal papillae are multiplex. Willia also has strongly
differentiated perichaetial leaves, but these are usually somewhat secund, and the sporophyte is distinctively reduced in
length and complexity. Syntrichia cavallii is here placed in sect.
Syntrichia, not sect. Collotortula, because of the lack of
collenchymatous thickenings in the upper laminal cell walls.
Eventually, perhaps, further study may require that a separate
section or genus be devised for S. cavallii (see also discussion
under Willia).
Syntrichia sect. Collotortula Zand., sect. nov. Type: Syntrichia
andicola (Mont.) Zand.
A· sectione typica cellulis laminalibus in regione mediana
superna plerumque distincte in angulis incrassatis vel
tumescentitrigonis, marginibus folii p/erumque recurvis,
papillis interdum simplicibus, propagulis si effectis clavatis vel
e/lipticis differt.
Differs from the typical section by the following combination of characters: medial upper laminal cells usually somewhat
thickened at the corners or even trigonous (i.e. with bulging
knots), leaf margins usually recurved, papillae sometimes simple, and propagula when present clavate or elliptical. Examination of transverse leaf sections indicates that the interior upper
laminal cell walls may have large, round central pores (staining
helps define this) in at least some species (Syntrichia aculeata,
S. amphidiacea, S. andicola (Pl. 106, f. 19), S. bogotensis, S.
gemmascens, S. gromschii and S. papillosa), while in other species, including those of other sections of Syntrichia, the pores
are absent or what appear to be interior pores are occasional,
irregular and possibly artifacts. Features correlated with !aminal
cell collenchymatous thickenings are the upper leaf margin usually serrate and the stem central strand often lacking, but these
are not unique characters. The section may not be sharply distinct from sect. Syntrichia because certain species of that section may have weakly thickened cell corners (appearing as
bright triangular points of light under high magnification). The
counterclockwise-twisted seta of Syntrichia papillosa commented upon by Dixon (1923) as unusual is not 'taxonomically
significant, since long setae in the Pottiaceae are generally
twisted clockwise below and counterclockwise above, and short
setae are untwisted or merely twisted clockwise. The seta of S.
papillosa (Pl. 109, f. 19-from a specimen misidentified as
Tortula panduraefolia, Tasmania, Weymouth 2821, NY) is only
about 5 mm in length. Species (some surely synonyms of S.
andicola) belonging to sect. Collotortula include: S. acu/eata,
S. alpestris, S. amphidiacea (Pl. 106, f. 1-8), S. andicola (Pl.
106, f. 14-20), S. antarctica (possibly belongs here, but lamina!
cell walls are only weakly collenchymatous), S. bogotensis, S.
cainii (Pl. 107, f. 5-6), S. ciliata, S. conferta, S. fontana
(laminal cells flat superficially, as in Hennediella), S.
gemmascens (Pl. 106, f. 18-21), S. goudotii, S. gromschii, S.
mollis, S. papillosa (Pl. 109, f. 14-19), S. rivularis (Pl. 109, f.
20-23), S. robusta (Pl. Ill, f. 1-4), S. rubra (Pl. Ill, f. 5-8),
and S. subaristata. Curiously, S. rigescens has propagula (Pl.
110, f. 18-19) borne ventrally on the costa, as in sect.
Co/lotortula, but is clearly in sect. Syntrichia near S.
caninervis. Note also Syntrichia leucostega, not seen, for
which ventral costal propagula are illustrated by Kramer
(1988).
Syntrichia sect. Aesiotortula Zand., sect. nov. Type: S.
pagorum (Milde) Amann.
A sectione typica plantis parvioribus quam congeneribus,
foliis ligulatis vel spathulatis, marginibus omnino planis,
papillis bifidis, propagulis si effectis foliaceis, in termino
caulis vel in apicibus foliorum deciduis portatis, costa supra
regione folii mediana perincrassata et in sectione
transversalis semicircu/aribus differt.
Differs from the typical section by the following combination of characters: plants rather small for the genus, leaves
ligulate to spathulate, margins plane throughout, papillae
bifid, and propagula when present leaf-like, borne terminally
on the stem or as deciduous leaf apices. The costa is often
very strongly thickened above midleaf and semicircular in
transverse section. This group of species includes S.
ammonsiana (Pl. 105, f. 12-16), S. baileyi, S. bartramii, S.
chisosa (Pl. 107, f. 13-16), S. epilosa (Pl. 108, f. 1-9), S.
pagorum (Pl. 109, f. 10-11), S. phaea (Pl. 109, f. 23-27) and
S. pygmaea (Pl. 110, f. 10-16). Syntrichia papillosa, although
here placed with sect. Collotortula because of the clearly
trigonous upper !aminal cells and clavate to spherical propagula, also has plane margins and a much thickened costa.
Section Aesiotortula (as the "Tortula laevipila-Tortula
pagorum-Gruppe") was not dealt with by Kramer (1980), who
viewed it as a complex requiring special study. Certain of the
infraspecific taxa of S. laevipila may also belong here, but not
var. laevipila (Pl. 109, f. 1-9) itself. Syntrichia pygmaea is
only tentatively placed in this section; it is propaguliferous,
however, by a deciduous leaf apex and has plane margins. On
the other hand, species of similarly small stature in many genera have a tendency to lose or exhibit less distinctly some
important characters, including recurvature of the lower margins. Syntrichia pygmaea is also unusual for its ventral leaf
surface very narrowly grooved along the costa and ventral
costal cells elongate, which may be an indication of a relationship with the Bryoerythrophylleae for this species. The
upper !aminal cells of the most highly derived of the species
of sect. Aesiotortula are relatively large for the genus, generally 13-15 Jlm in diameter; species with upper !aminal cells
small (ca. 8-10 Jlm in diameter), such as S. bartramii and S.
chisosa, are less distinctive. The cleistocarpous genus Phascopsis could be seen as having been derived from this section
of Syntrichia, but is apparently a relict of a more primitive lineage (see Cladograms 13 and 14).
The New World austral species Syntrichia epilosa is much
the same as S. bartramii of western North America and may
POTIIOIDEAE
265
Plate 109. Syntrichia. 1-9. S. lluvipila. l. Section of stem. 2-3. Two leaves. 4. Leaf apex. 4. Leaf section. 5. Leaf section, with small ventral
stereid band. 7. Perichaetium. 8. Stalked perigonial bud. 9. Capsule with peristome. 10-11. S. pagorum. 10-11. Two leaves. 12. Leaf section.
13. Propagula. 14-19. S. papillosa. 14-15. Two leaves. 16. Leaf apex. 17. Leaf section. 18. Perichaetialleaves. 19. Capsule, calyptra. 20-22. S.
percarnosa. 20-21. Two leaves. 22. Leaf section. 23-27. S. phaea. 23. Section of stem. 24-25. Two leaves. 26. Leaf apex. 27. Leaf section.
266
GENERA OF THE POTTIACEAE
Plate 110. Syntrichill. 1-5. S. pseudodesertorum. l. Transverse section of stem. 2-3. Two leaves. 4. Leaf apex. 5. Transverse section at
midleaf. 6-9. S. pseudorobusta. 6-1. Two leaves. 8. Leaf apex. 9. Transverse section at midleaf. 10-16. S. pygmaea. 10. Stem apex with young
propaguliferous leaves. 11-13. Three leaves. 14-15. Propagula, as deciduous leaf apices. 16. Transverse section at midleaf. 17-19. S. rigescens.
17-18. Two leaves, with propagula. 19. Transverse section at midleaf, with propagula. 20-23. S. rivularis. 2~21. Two leaves. 22. Leaf apex.
23. Transverse section at midleaf.
POITIOIDEAE
be synonymous; a parallel southern South America and southwestern North America distribution is that of the austral
Pseudocrossidium crinitum and its Arizona population previously known as Tortula aurea (see treatment of Pseudocrossidium).
Additional, selected literature: Allorge (1938), Anderson
(1943), Barkman (1963), Bartram (1924b, 1926b), Bewley
(1972, 1973a,b), Bewley et al. (1974), Bizot (1954, 1956),
Blomquist (1930), Boudier (1992), Casas de Puig (1975a),
Casas de Puig and Molinas (1975), Catcheside (1980, 1992),
Dedkov et al. (1989), Doei et al. (1985), El-Oqlah et al. (1988),
Hedenas (1989b), Kalenov (1977), Kramer (1978), Lazarenko
(1959), Lightowlers (1985a, 1986a,b,c), Magill et al. (1983),
Maya (1986), Mishler (1984a, 1985a,b, 1986a, 1987a, 1990),
Mishler and Newton (1987, 1988), Mishler and Oliver (1991),
Mishler and Scheirer (1983), Oliver and Mishler (1988, 1990),
Ovezova (1989), Saito (1973a), Side and Whitehouse (1974),
Steere (1939a, 1940), Stone (1971), Studlar et al. (1984), Toth
(1987), Tuba (1984, 1985), Willis (1964 ), Zander (1989).
Number of accepted species: 82.
New heterotypic synonymy: Tortula abruptinervis Dix. =
Syntrichia pygmaea (Dus.) Zand. Tortula kingii Robins. = Syntrichia percarnosa (C. Mull.) Zand. Tortula lemniscata Zand. =
Syntrichia aculeata (Wils.) Zand. Tortula nigra Zand. =Syntrichia percarnosa (C. Miill.) Zarid. Tortula tanganyikae Dix. =
Syntrichia amphidiaceus (C. Mull.) Zand.
Species examined: S. aculeata (BUF, NY}, S. alpestris
(BM), S. ammonsiana (BUF), S. amphidiaceus (BM, BUF,
TENN), S. amplexa (CU), S. anderssonii (NY), S. andico/a, S.
antarctica (NY), S. baileyi (NY), S. bartramii (BUF), S.
bipedicellata (NY), S. bogotensis (NY), S. bolanderi (BUF), S.
brandisii (NY), S. cainii (BUF, TRTC), S. caninervis (BUF, H),
S. cavallii (NY, US), S. chisosa (BUF), S. ciliata (H), S.
conferta (NY), S. costesii (NY), S. didymodontoides (H), S.
epilosa (COLO), S. filaris (H, US), S. flagel/aris (BUF), S.
fontana (S) as Tortula rivularis, S. fragitis, S. fuscoviridis (NY),
S. geheebiaeopsis (NY), S. gemmascens (NY), S. gromschii
(US), S. inermis, S. intermedia (BUF), S. jaffuelii (NY), S.
tacerifolia (NY), S. /aevipila (BUF), S. latifolia, S. limensis
(NY), S. linguifolia (L), S. longimucronata (NY), S. mongolica
(NY), S. mol/is (US), S. norvegica, S. obtusissima (BM, BUF,
TENN), S. papillosa (BUF, NY), S. percarnosa (BUF, H, US),
S. phaea (BUF), S. pichinchensis (US), S. princeps, S. prostrata
(US), S. pseudodesertorum (S), S. pseudorobusta (NY), S.
pygmaea (BM, BUF, MICH, NY), S. ramosissima (NY), S.
reflexa (NY), S. rigescens (FU), S. robusta (NY), S. rubella
(NY), S. rubra (NY), S. ruralis, S. saxico/a (NY), S. scabrella
(NY), S. scabrinervis (US), S. schnyderi (NY), S. serrata (NY),
S. serripungens (NY), S. sinensis (NY), S. socialis (S), S.
subaristata (NY), S. virescens (BUF), S. viriduta (NY).
New combinations and statuses:
Syntrichia acuteata (Wils .) Zand., comb. nov. (Barbula
aculeata Wils., Kew J. Bot. 3: 51, 1851; Tortula aculeata
(Wils.) Mitt.).
Syntrichia a/pestris (Dix. in Herz.) Zand., comb. nov. (Tortuta
alpestris Dix. in Herz., Rep. Spec. Nov. Reg. Veg. 38: 103,
267
1935).
Syntrichia amphidiacea (C. Mtill.) Zand., comb. nov. (Barbula amphidiacea C. MUll., Linnaea 42: 332, 1879; Tortuta
amphidiacea (D. Miill.) Broth.).
Syntrichia amptexa (Lesq.) Zand., comb. nov. (Barbula
amplexa Lesq., Trans. Am. Phil. Soc. n. ser. 13: 5, 1865;
Tortula amplexa (Lesq.) Steere in Grout).
Syntrichia anderssonii (Angstr.) Zand., comb. nov. (Tortula
anderssonii Angstr., Oefv. K. Vet. Ak. Foerh. 29(4): 6,
1872).
Syntrichia anderssonii var. fagicola (C. Miill.) Zand., comb.
nov. (Barbula conotricha var. fagicota C. Mtill., Flora 68:
416, 1885), not seen.
Syntrichia antarctica (Hampe in C. Miill.) Zand., comb. nov.
(Barbula antarctica Hampe in C. Miill., Syn. 1: 642,
1849; Tortula antarctica (Hampe in C. Miill.) Wils. in
Hook. f.).
Syntrichia austroafricana (Kramer) Zand., comb. nov. (Tortuta austroafricana Kramer, J. Hattori Bot. Lab. 65: 92,
1988), not seen but cf. Kramer, J. Hattori Bot. Lab. 65: 84,
1988.
Syntrichia baiteyi (Broth.) Zand., comb. nov. (Tortula baileyi
Broth., Oefv. Finsk. Vet. Soc. Foerh. 33: 97, 1891).
Syntrichia bartramii (Steere in Grout) Zand., comb. nov.
(Tortuta bartramii Steere in Grout, Moss Fl. N. Amer. 1:
241, 1939).
Syntrichia bipedicellata (Britt.) Zand., comb. nov. (Tortula
bipedicellata Britt., Bull. Torr. Bot. Cl. 23:431, 1896).
Syntrichia bogotensis (Hampe) Mitt. (Barbula bogotensis
Hampe, Ann. Sc. Nat. Bot. ser. 5, 3: 349, 1865; Tortuta
bogotensis (Hampe) Mitt.).
Syntrichia bolanderi (Lesq. & James) Zand., comb. nov.
(Barbula botanderi Lesq. & James, Trans. Am. Phil. Soc.
n. ser. 13: 5, 1865; Tortuta botanderi (Lesq. & James)
Howe).
Syntrichia brachyclada (Card.) Zand., comb. nov. (Tortula
brachyclada Card., Bull. Herb. Boiss. ser. 2, 5: 1002,
1905), not seen but cf Kramer, J. Hattori Bot. Lab. 65: 84,
1988.
Syntrichia brandisii (C. Miill.) Zand., comb. nov. (Barbuta
brandisii C. Miill., Flora 61: 82, 1878; Tortula brandisii
(C. Miill.) Broth.).
Syntrichia brevisetacea (F. Miill.) Zand., comb. nov. (Barbuta
brevisetacea F. Miill., Analyt. Draw. Austral. Moss. Tab.
4, 1864; Tortuta brevisetacea (F. Miill.) Ther.), not seen
but cf Kramer, J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia cainii (Crum & Anders.) Zand., comb. nov. (Tortuta cainii Crum & Anders., J. Elisha Mitchell Sci. Soc.
74: 35, 1958).
Syntrichia campestris (Dus.) Zand., comb. nov. (Tortuta
campestris Dus., Ark. Bot. 6(8): 25, 1906), not seen but
cf Kramer, J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia caninervis var. spuria (Amann) Zand., comb. nov.
(Tortuta spuria Amann, Bull. Murithienne 39: 351, 1916;
Tortuta caninervis ssp. spuria (Amann) Kramer), not
seen.
Syntrichia caninervis var. abranchesii (Luis.) Zand., comb.
268
GENERA OF THE POTTIACEAE
Plate 111. SyntrichitJ. 1-4. S. robusta. 1-2. Two leaves. 3. Leaf apex. 4. Transverse section at midleaf. S-8. S. rubra. 5-6. Two leaves. 7. Leaf
apex. 8. Transverse section at midleaf. 9-12. S. saxicola. 9-10. Two leaves. 11. Leaf apex. 12. Transverse section at midleaf. 13-16. S. se"ata.
13-14. Two leaves. 15. Leaf apex, lateral view. 16. Transverse section at midleaf. 17-19. S. virescens. 17-18. Two leaves. 19. Transverse
section at midleaf.
POTIIOIDEAE
nov. (Tortula abranchesii Luis., Broteria ser. Bot. 14: 115,
1916; Tortula caninervis var. abranchesii (Luis.) Kramer),
not seen.
Syntrichia caninervis var. spuria (Amann) Zand., comb. nov.
(Syntrichia spuria Amann, Fl. Mouss. Suisse 2: 119, 384,
1918).
Syntrichia chisosa (Magill, Delg. & Stark) Zand., comb. nov.
(Tortula chisosa Magill, Delg. & Stark, Ann. Missouri Bot.
Gard. 70: 200, 1983).
Syntrichia ciliata (Broth.) Zand., comb. nov. (Tortula ciliata
Broth., Bot. Jahrb. 49: 175, 1912).
Syntrichia conferta (Bartr.) Zand., comb. nov. (Tortula conferta
Bartr., Bryologist 60: 140, 1957; Tortula princeps var.
conferta (Bartr.) Lightow1ers).
Syntrichia costesii (Ther.) Zand., comb. nov. (Tortula costesii
Ther., Rev. Chil. Hist. Nat. 25: 298, 1921).
Syntrichia didymodontoides (Broth. in Dryg.) Zand., comb. nov.
(Tortula didymodontoides Broth. in Dryg., Deutsch.
Siidpolar Exp. 8: 86, 1906).
Syntrichia epilosa (Broth. ex Dus.) Zand., comb. nov. (Tortula
epilosa Dus., Ark. Bot. 6(8): 25, 1906.
Syntrichia epilosa var. pilifera (Ther.) Zand., comb. nov. (Tortula epilosa var. pilifera Ther., Rev. Chi!. Hist. Nat. 30: 343,
1926), not seen.
Syntrichia filaris (C. Miill. in Neum.) Zand., comb. nov. (Barbula filaris C. Miill. in Neum ., Deutsch. Exp . Int.
Polarforsch. 2: 309, 1890; Tortula filaris (C. Miill. in
Neum.) Broth.).
Syntrichia flagel/aris (Schimp.) Zand., comb. nov. (Barbula
flagellaris Schimp., Ann. Sci. Nat. Bot. ser. 2, 6: 146, 1836;
Tortulaflagellaris Mont. in Gay).
Syntrichia flagellaris var. densiretis (Ther.) Zand., comb. nov.
(Tortula flagellaris var. densiretis Ther., Recueil Publ. Soc.
Havraise Etud. Div. 1917: 7, 1917), not seen.
Syntrichia fontana (C. Miill. in Neum.) Zand., comb. nov.
(Barbula fontana C. Miill. in Neum., Deutsch. Exp. Int.
Polarforsch. 2: 308, 1890; Tortula fontana (C. Miill.)
Broth.).
Syntrichia fuscoviridis (Card.) Zand., comb. nov. (Tortula
fuscoviridis Card., Bull. Herb. Boiss. ser. 2, 6: 6, 1906).
Syntrichia geheebiaeopsis (C. Mii11.) Zand., comb. nov. (Barbula geheebiaeopsis C. Mii1l., Bot. Jahrb. 5; 80, 1883; Tortula geheebiaeopsis C. Miill.) Broth.).
Syntrichia gemmascens (Chen) Zand., comb. nov. (Desmatodon
gemmascens Chen, Hedwigia 80: 297, 1941; Didymodon
gemmascens Broth., Symb. Sin. 4: 38, 1929, hom. illeg.).
Syntrichia gemmascens var. hopeiensis (Chen) Zand., comb.
nov. (Desmatodon gemmascens var. hopeiensis Chen,
Hedwigia 80: 229, 1941).
Syntrichia g/acialis (Kunze ex C. Miill.) Zand., comb. nov.
(Barbula g/acialis Kunze ex C. Miill., Syn. 1: 634, 1849;
Tortula glacialis (Kunze ex C. Miill.) Mont. in Gay).
Syntrichia gromschii (Ther.) Zand., comb. nov. (Tortula
gromschii Ther., Rev. Bryol. Lichenol. 7: 173, 1935).
Syntrichia jaffuelii (Ther.) Zand., comb. nov. (Tortula jaffuelii
Ther. , Rev. Chil. Hist. Nat. 27: 9, 1923).
Syntrichia lacerifolia (Williams) Zand., comb. nov. (Tortula
269
lacerifolia Williams, Bull. Torr. Bot. Cl. 43: 326, 1916).
Syntrichia leucostega (C. Miill.) Zand., comb. nov. (Barbula
leucostega C. Miill., Syn. 1: 641, 1894; Tortula
leucostega (C. Miill.) Broth.), not seen but cf Kramer, J.
Hattori Bot. Lab. 65: 106, 1988.
Syntrichia leucostega var. trachyneura (Dix.) Zand., comb.
nov. (Tortula trachyneura Dix., Trans. R. Soc. S. Africa
8: 195, 1920; Tortula leucostega var. trachyneura
Kramer), not seen but cf Kramer, J. Hattori Bot. Lab. 65:
107, 1988.
Syntrichia limensis (Williams) Zand., comb. nov. (Tortula
limensis Williams, Bull. Torr. Bot. Cl. 42: 398, 1915).
Syntrichia linguifolia (Herz.) Zand., comb. nov. (Tortula
linguifolia Herz., Biblioth. Bot. 87:49, 1916).
Syntrichia longimucronata (X.-j. Li) Zand., comb. nov. (Tortula longimucronata X.-j. Li, Act. Bot. Yunnan. 3:
107- 109, 1981).
Syntrichia magel/anica (Mont. in Gay) Zand., comb. nov.
(Tortula magel/anica Mont. in Gay, Hist. Fis. Polit. Chile
Bot. 7: 145, 1850; Tortula princeps var. magel/anica
(Mont. in Gay) Lightowlers), not seen but cf Kramer, J.
Hattori Bot. Lab. 65: 110, 1988.
Syntrichia mol/is (B.&S. ex C. Miill.) Zand., comb. nov.
(Barbula mol/is B.&S. ex C. Miill., Syn. 1: 637, 1849;
Tortula mol/is (B.&S. ex C. Miill.) Broth.).
Syntrichia obtusissima (C. Miill.) Zand., comb. nov. (Barbula
obtusissima C. Miill. , Syn. 1: 640, 1849; Tortula
obtusissima (C. Miill.) Mitt.).
Syntrichia papillosa var. chilensis (Ther.) Zand., comb. nov.
(Tortula papi/losa var. chilensis Ther., Rev. Chil. Hist. ,
Nat. 25: 297, 1921), not seen.
Syntrichia percarnosa (C. Miill.) Zand., comb. nov. (Barbula
percarnosa C. Miill., Linnaea 42: 347, 1879; Tortula
percarnosa (C. Miill.) Broth.).
Syntrichia phaea (Hook. f. & Wils.) Zand., comb. nov.
(Trichostomum phaeum Hook. f. & Wils., Fl. Nov. Zel. 2:
72, 1854; Tortula phaea (Hook. f. & Wils.) Dix.).
Syntrichia pichinchensis (Tayl.) Zand., comb. nov. (Tortula
pichinchensis Tayl., London J. Bot. 6: 333, 1847).
Syntrichia princeps var. brachycarpa (De Not.) Zand., comb.
nov. (Tortula princeps var. brachycarpa De Not., Atti
Univ. Genova 1: 538, 1869).
Syntrichia princeps var. echinata (Shiffn.) Zand., comb. nov.
(Tortula echinata Schiffn., Oesterr. Bot. Zeitschr. 65: 4-5,
1915; Tortula princeps ssp. echinata (Schiffn.) Kramer).
Syntrichia prostrata (Mont.) Zand. , comb. nov. (Tortula
prostrata Mont., Ann. Sc. Nat. Bot. ser. 3, 4: 104, 1845).
Syntrichia pseudorobusta (Dus.) Zand., comb. nov. (Tortula
pseudorobusta Dus., Ark. Bot. 6(8): 19, 1906).
Syntrichia pygmaea (Dus.) Zand., comb. nov. (Tortula
pygmaea Dus., Ark. Bot. 6(10): 8, 1907).
Syntrichia ramosissima (Ther.) Zand., comb. nov. (Tortula
ramosissima Ther., Rev. Chil. Nat. 33: 136, 1926).
Syntrichia reflexa Zand., nom. nov. (Tortula reflexa X.-j . Li,
Acta Bot. Yunnan. 3: 109, 1981, hom. illeg. non Brid.).
Syntrichia robusta (Hook. & Grev.) Zand., comb. nov. (Tortula robusta Hook. & Grev., Edinburgh J . Sc. 1: 299,
270
GENERA OF THE POTTIACEAE
1824).
Syntrichia robusta var. recurva (Lightowlers) Zand., comb. nov.
(Tortula robusta var. recurva Lightowlers, Brit. Ant. Surv.
Bull. 64: 64, 1984).
Syntrichia rubella (Hook. f. & Wils.) Zand., comb. nov. (Tortula rubella Hook. f. & Wils., Fl. Tasman. 2: 176, 1859).
Syntrichia rubra (Mitt. in Hook. f.) Zand., comb. nov. (Tortula
rubra Mitt. in Hook. f., Handb. New Zealand Fl. 419,
1867).
Syntrichia rubra var. subantarctica (Sainsb.) Zand., comb. nov.
(Tortula subantarctica Sainsb., Svensk. Bot. Tidskr. 44: 72,
1950; Syntrichia rubra var. subantarctica (Sainsb.)
Lightowlers).
Syntrichia ruralis var. gigantea (Lesq.) Zand., comb. nov.
(Barbula ruralis var. gigantea Lesq., Mem. California
Acad. Sci. 1: 13, 1868; Tortula ruralis var. gigantea (Lesq.)
L. Koch.), not seen.
Syntrichia ruralis var. gracilis (C. Jens.) Zand., comb. nov.
(Tortula ruralis var. gracilis C. Jens., Medd. Groenland 15:
409, 1898), not seen.
Syntrichia ruralis var. spiralis (Herz.) Zand., comb. nov. (Tortula ruralis var. spiralis Herz., Biblioth. Bot. 87: 49, 1916),
not seen.
Syntrichis ruralis var. submamillosa (Kramer) Zand., comb.
nov. (Tortula ruralis var. submamillosa Kramer, Bryophyt.
Biblioth. 21 : 127, 1980), not seen.
Syntrichia ruralis var. subpapillosissima (Biz. & Pierr.) Zand.,
comb. nov. (Tortula ruralis var. subpapillosissima Biz. &
Pierr., Acta Bot. Acad. Sci. Hung. 18: 10, 1973; Tortula
ruraliformis var. subpapillosissima (Biz. & Pierr.) Kramer),
not seen.
Syntrichia saxicola (Card.) Zand., comb. nov. (Tortula saxicola
Card., Bull. Herb. Boiss. ser. 2, 5: 1002, 1905).
Syntrichia scabrella (Dus.) Zand., comb. nov. (Tortula
scabrella Dus., Ark. Bot. 6(10): 4, 1907).
Syntrichia scabrinervis (C. MUll.) Zand., comb. nov. (Barbula
scabrinervis C. Miill., Syn. 1: 634, 1849; Tortula
scabrinervis (C. Miill.) Mitt.).
Syntrichia schnyderi (C. MUll.) Zand., comb. nov. (Barbula
schnyderi C. Miill., Linnaea 43: 434, 1882; Tortula
schnyderi (C. MUll.) Broth.).
Syntrichia serrata Zand., comb. nov. (Tortula serrata Dix.,
New Zealand Inst. Bull. 3(3): 146, 1923, nom. legit. contrary to Index Muscorum, also cf Lightowlers, J. Bryol. 13:
373, 1985).
Syntrichia serripungens (Lor. & C. Miill.) Zand., comb. nov.
(Barbula serripungens Lor. & C. Miill., Linnaea 42: 360,
1879; Tortula serripungens (Lor. & C. Miill.) Broth.).
Syntrichia serripungens var. excesa (C. MUll.) Zand., comb.
nov. (Barbula serripungens var. excesa C. Miill., Linnaea
42: 351, 1879.
Syntrichia socialis (Dus.) Zand., comb. nov. (Tortula socialis
Dus., Ark. Bot. 6(10): 6, 1907).
Syntrichia subaristata (B.&S. ex C. Miill.) Zand., comb. nov.
(Barbula subaristata B.&S. ex C. Miill., Syn. 1: 644, 1849;
Tortula subaristata (B.&S. ex C. MUll.) Broth.).
Syntrichia virescens var. bizotiana (Kramer) Zand., comb. nov.
(Tortula virescens ssp. bizotiana Kramer, J. Hattori Bot.
Lab. 65: 123, 1988, nom. nov. for Tortula virescens ssp.
bizotii (Laz.) Kramer, Bryophyt. Biblioth. 21: 102, 1980,
comb. inval. basion. inval.; Tortula bizotii Laz., Vopr.
Evol. Biogeogr. Genet. Sel. 145, 1960, nom. inval.), not
seen but cf Kramer, J. Hattori Bot. Lab. 65: 84, 1988.
Syntrichia virescens var. iranica (Kramer) Zand., comb. nov.
(Tortula virescens var. iranica Kramer, Bryophyt.
Biblioth. 21 : 101 , 1980), not seen.
Syntrichia viridula (C. Miill.) Zand., comb. nov. (Barbula
viridula C. Miill., Nuov. Giorn. Bot. Ital. n. ser. 4: 114,
1897; (Tortula viridula (C. Miill.) Broth.; the isotype at
NY is probably S.fragilis).
75. HILPERTIA
Plate 112.
Hilpertia Zand., Phytologia 65: 427, 1989. Type: Hilpertia
velenovskyi (Schiffn.) Zand.
Named for Friedrich Hilpert, 1907-, originally of
Hildburghausen, Germany, student ofT. Herzog and author of
"Studien zur Systematik der Trichostomaceen" (1933) as a
doctoral dissertation.
Plants growing in loose cushions, greenish brown above,
light brown below. Stems branching irregularly, to 1.0 em in
length, transverse section rounded-pentagonal, central strand
distinct, sclerodermis not or weakly differentiated, hyalodermis absent; axillary hairs to 8 cells in length, all hyaline or
the basal one yellow-brown; rhizoids rare. Leaves crowded,
larger above, appressed and tightly spiralled when dry,
weakly spreading when moist, ovate to circular, 1.3-2.0 mm
in length (including awn), upper lamina flat to more usually
quite concave, margins strongly revolute (to 2 times), entire
or broadly toothed at or near the base of the awn; apex
broadly acute, hyaline in an apical patch or triangle; base not
differentiated in shape; costa narrow but broader above,
excurrent as a hyaline awn, costa with lamina inserted laterally, superficial cells long-rectangular and smooth on both
sides, 2-4 rows of cells across costa ventrally at midleaf,
costal transverse section rounded, stereid band rounded in
shape, ventral epidermis present but dorsally absent, guide
cells 2 in 1 layer, hydroid strand present; upper lamina/ cells
hexagonal to short-rectangular or rhomboidal, 14-25 Jlm in
width, 2-4:1, internal walls thin to thickened and porose, dorsal superficial walls much thickened, weakly convex superficially on both sides, cells of leaf apex rhomboidal to fusiform,
smooth, cells of revolute margin enlarged, strongly chlorophyllose; papillae absent medially, usually hollow-papillose
on revolute margins; basal cells weakly differentiated, rectangular, ca. 16--18 Jlm in width, 2-3:1, walls thin. Propagula
when present (1-)3-4 celled, brown, spherical to elliptical,
mostly 30-50 Jlm in length, borne on basal rhizoids.
Synoicous, paroicous, autoicous or apparently dioicous but
probably rhizautoicous. Perichaetia terminal, inner leaves
usually differentiated, long-oval, margins usually little
differentiated, to 1. 7 mm in length, basal portion of leaf
sheathing, lower cells rectangular, very thin-walled. Perigonia
r
t
POITIOIDEAE
lateral or occasionally terminal on a separate plant. Seta 3.5-4.0
mm in length, 1 per perichaetium, yellow-brown, twisted counterclockwise above, clockwise below; theca 1.2-1.5 mm in
length, yellow-brown, elliptical, occasionally weakly
ventricose, exothecial cells ca. 16-23 Jlm in width, 2-3: 1, thinwalled, stomates phaneropore, on capsule neck, annulus of 3
rows of smaller, quadrate, highly vesiculose cells; peristome
teeth 32, linear, densely branching-spiculose, 300-700 Jlm in
length, with many articulations, twisted counterclockwise about
1/2 tum, basal membrane short, to 45 Jlm in height, papillosespiculose. Operculum broadly short-conic to long-conic,
0.4-1.0 mm in length, cells twisted 1/2 tum counterclockwise.
Calyptra cucullate, smooth, ca. 2.8 mm in length. Spores 13-16
271
Jlm in diameter, light brown, indistinctly papillose. Lamina/
KOH color reaction red.
Found on soil in Canada (Northwest Territories) and
countries of eastern central Europe.
Corley et al. (1981) suggested that Tortula ve/enovskyi is
probably best placed in a separate section of Tortula. It is here
treated, together with T. scotteri, as a separate genus, Hi/per-
ria.
Schiffner (1893) reviewed the extensive variation in sexuality in the type species. The modification of the upper
!aminal margins of Hilpertia into tubes of photosynthetic tissue (Pl. 112) is paralleled in species of Pseudocrossidium and
to a lesser extent in Tortula revolvens (Pl. 89, f. 4). Hilpertia
Plate 112. Hilpertia. 1-7. H. velenovskyi. 1. Habit. 2. Leaf. 3. Leaf apex. 4. Transverse section of costa at midleaf. 5. Transverse section of
costa near apex. 6. Dissected plant apex. 7. Perichaetialleaf. 8-10. H. scotteri. 8-9. Two leaves. 10. Leaf apex.
272
GENERA OF THE POTTIACEAE
Plate 113. Sagenotortulil. 1-10. S. quitoensis. 1. Habit. 2. Transverse section of stem. 3-4. Two leaves. 5-6. Two leaf apices, one enlarged. 7.
Basal cells. 8. Transverse section at midleaf. 9. Propagulum on rhizoid. 10. Peristome.
POTTIOIDEAE
has many of the features of species of Acaulon including the
ovate (rather concave) leaf shape, thin costa, upper !aminal cells
often hyaline apically and dorsally superficially thick-walled (as
easily seen in section, Pl. 112, f. 4-5), a hyaline awn, and red
KOH color reaction. The concave leaves may indicate an ancestry of taxa with a somewhat bulbiform habit, thus Hilpertia may
well be derived from Acaulon-Iike ancestors through
desuppression of the sporophyte and elaboration of the leaf
margins (but cf Cladograms 13 and 14). Hilpertia is easily distinguished from Acaulon by its revolute leaf margins, elongate
stems, elongate setae, and peristomate, cylindrical capsules. On
the other hand, the leaves of Stegonia latifolia and S.
hyalinotricha (Pl. 69) are quite similar morphologically to those
of Hilpertia, thus Hilpertia may have been derived from ancestors shared with Stegonia by development of photosynthetically
specialized !aminal margins, differentiated perichaetial leaves,
and red !aminal KOH reaction. Cladistic analysis (see Cladogram 14), contrarily, indicates that Stegonia is best hypothesized as rather distantly related, and that Acaulon does not share
immediate ancestors with Hilpertia.
Additional literature: Boros (1941), Karczmarz (1961), Kuc
(1960), Peciar (1960b), PospiSil (1977), Waclawska (1958),
Zander (1989), Zander and Steere (1978).
Number of accepted species: 2.
Species examined: Hilpertia scotteri (BUF), H. velenovskyi
(FH, NY).
Plate 113.
76. SAGENOTORTULA
Sagenotortula Zand., Phytologia 65: 429, 1989. Type: Sagenotortula quitoensis (Tayl. in Hook.) Zand.
From sagena,-ae, fish-net+ o + Tortula, a genus; a TortulaIike moss with a net-like areolation.
Plants loosely caespitose or turf-forming, green above,
greenish brown below. Stems branching occasionally, especially from just below the perichaetia, to 2.0 em in length, transverse section round to elliptical, central strand present, often
strong or stem centrally hollow, sclerodermis absent, hyalodermis absent; axillary hairs several cells in length, all hyaline;
stem thickly matted with rhizoids. Leaves appressed, little
changed to incurved but little contorted when dry, erect to
weakly spreading when moist, broadly ligulate to spathulate,
3-5 mm in length, upper lamina flat or very broadly and shallowly channeled, margins plane, occasionally broadly incurved
above, entire (occasionally distantly bluntly toothed, especially
evident in young leaves); apex broadly acute, occasionally with
a broad, blunt apiculus; base not differentiated in shape; costa
ending ca. 4 cells below apex to percurrent, costa with lamina
inserted ventrally at about 45", superficial cells quadrate to hexagonal or short-rectangular ventrally, dorsally rectangular,
2(-3) rows of cells across costa ventrally at midleaf, costal
transverse section somewhat rounded, stereid band little developed, weakly substereid and reniform in shape, ventral epider-
273
mis present, dorsal absent, guide cells 2 in 1 layer, hydroid
strand present, often large; upper /aminal cells hexagonal,
occasionally quadrate or short-rectangular, very large ,
(25-)40-60(-75) J..Lm in width, 1(-2):1, walls thin, occasionally evenly thickened, often weakly trigonous, superficially
convex on both sides; papillae absent; basal cells
differentiated in lower 1/4 of leaf across leaf or just medially,
rectangular, ca. 35-60 J..Lm in width, 2-4:1, walls thin to somewhat thickened. Propagula consisting of red brood bodies
borne on tomentum of perigoniate plants, elliptical to spherical, 50-65 J..Lm in longest dimension, bearing superficially
scattered short peg-like projections. Dioicous. Perichaetia terminal, inner leaves slightly larger than the cauline, otherwise
little different, to 6 mm in length. Perigonia terminal, comparatively large, inner leaves ovate. Seta ca. 2.5 em in length,
1-3 per perichaetium, red-brown, twisted counterclockwise
above, occasionally clockwise below; theca 3-4 mm in
length, red-brown, cylindrical, exothecial cells longrectangular, thin-walled, 20-25 J..Lm in width, stomates at base
of theca, phaneropore, annulus of 2-3 rows of vesiculose
cells, apparently deciduous in pieces; peristome teeth 32, filamentous, low-spiculose, ca. 1000 J..Lm, with many articulations, twisted counterclockwise once, basal membrane to 200
J..Lm in height, low-spiculose to crazed. Operculum conic,
1.5-1.7 mm in length, cells twisted counterclockwise. Calyptra cucullate, smooth, ca. 4.5 mm in length. Spores 10-13 J..Lm
in diameter, tan, essentially smooth. Lamina/ KOH color
reaction red.
Found on rocks and soil at rather high elevations; Mexico
and the Andes of South America.
Sagenotortula is distinguished from other genera of Pottiaceae by the unique red brood bodies (Pl. 113, f. 9) that have
numerous short, blunt, papilla-like projections (unlike the
armed or caltrop-like propagula of Hyophila species that have
cell walls protruding). The brood bodies were found only on
perigoniate plants, being uncolored and transparent when
young, and are distributed in the tomentum along the entire
length of the stem. The peg-like projections develop one per
superficial cell and resemble simple papillae when small;
rhizoids are occasionally produced apically from the longer
protrusions. The genus differs from other genera that have
large, epapillose upper laminal cells by the mostly entire,
unbordered margins and the extremely large upper lamina!
cells (Pl. 113, f. 5).
The apparent lack of a dorsal costal epidermis (Pl. 113, f.
8) indicates that this genus could have immediate ancestors
shared with Syntrichia, with derivation through loss of papillae, inflation of the !aminal cells, and elaboration of the
unique brood body. Cladograms 13 and 14 support this.
Additional literature: Brotherus (1924-25).
Number of accepted species: 1.
Species examined: Sagenotortula quitoensis (NY, TENN).
EXCLUDED AND UNTREATED TAXA
GENERA
ARVILDIA
Arvildia Ignatov, Palaeontograph. B. 217: 150, 1990. Type:
Arvildia elenae Ignatov.
The two species of this Permian fossil genus were referred to
either the Pottiaceae or Trichostomaceae by Ignatov ( 1990),
although he indicated that other, closely related families were also
possibile dispositions. The published figures, however, are reminiscent of the Dicranaceae because of the very broad costae and
the bulging, thin-walled laminal cells, which lack papillae. Additional study is needed.
BRYOBARTRAMIA
Bryobartramia Sainsb., Bryologist 51: 10, 1948. Type: Bryobartramia robbinsii Sainsb.
Bryobartramia is a genus of arid areas of Australia and South
Africa. The single species, B. novae-valesiae (Broth.) Stone, has
been placed in its own family, the Bryobartramiaceae Sainsb., but
is here referred to the Encalyptaceae because of the large calyptra,
yellow KOH reaction of the upper lamina, and large, coarsely
papillose upper laminal cells. Stone (1977a) indicated that Bryobartramia had more characters in common with the Encalyptaceae than with any other family and described these in detail.
Bryobartramia differs from other genera of Encalyptaceae by,
among other features, the non-dehiscing calyptra (epigonium);
stomates scattered over the capsule, which is spherical and
cleistocarpous; exothecial cells transparent and nearly hyaline
(very weakly tinted yellow); seta very short; and perichaetial
leaves subulate, not papillose, elaminate above (rather like the
long-awned perichaetialleaves of Diphyscium). The inflated, nondehiscing calyptra may act as a flotation device as does, perhaps,
the thecal air sac ofTetrapterum.
CLADOPHASCUM
Cladophascum Dix. ex Sirn, Bryoph. S. Afr. 143, 1926. Type:
Cladophascum gymnomitrioides (Dix.) Dix. ex Sirn.
This genus was referred to the Pottiaceae in Crosby and Magill's
( 1977, 1978, 1981) Dictionary of the Mosses, but has been placed
in the Dicranaceae subf. Trematodontoideae by Magill (1981) in
his treatment of the bryoflora of South Africa.
ENTOSTHYMENIUM
Entosthymenium Brid., Bryol. Univ. 1: 761. 1827. Type: Entosthymenium tristichum Brid.
Paris (1906) and van der Wijk et al. (1959-Q9) recognized this
genus. Of the four combinations, the Index Muscorum (van der
Wijk et al. 1959-1969) reported two names assigned to the synonymy of Desmatodon convolutus (Brid.) Grout, and one to Pottia
wilsonii var. mucronifolia (Bruch) Warnst. Nothing seems known
of the generitype, E. tristichum. The type of E. tristichum, collected in southeastern France, is absent from the De Candolle
Herbarium at G (P. Geissler, pers. comm.). C. Miiller (1849-51)
and W. Schultze-Motel (pers. comm.) have indicated that the type
is not to be found in the Bridel Herbarium (B); the former
referred the genus to "Pottiacea suspectissima." L. Bertrand
(pers. comm.) wrote me that it is also not in the Grateloup
herbarium at Montpellier (MPU). Brotherus (1902-09,
1924-1925, Corley et al. (1981), Husnot (1884-1894), Jaeger
and Sauerbeck (1870-1880), Lindberg (1864), Miiller (1901),
Podpi!ra (1954) and Reed and Robinson (1972), perhaps understandably, neither acknowledged nor disposed of the genus in
their treatments, which include the Pottiaceae of France. The
original description is extremely vague. The genus is here set
aside pending further study that might lead to adequate lecto- or
neotypification.
ERPODIOPSIS
Erpodiopsis C. Miill., Flora 73: 470, 1890. Type: Erpodiopsis
kilimandscharica C. Miill. (= Phascum kilimandscharicum
(C. Miill.) Roth).
Brotherus (1902-{)9: 1239) placed the type of this genus very
near Bryum argenteum Hedw. (Bryaceae), while Roth (1911)
indicated that it is close to Phascum hyalinotrichum (here transferred to Stegonia and in fact with much the appearance of B.
argenteum). Inasmuch as the type (not seen) is, according to
both aforementioned authors, sterile and Roth's (1911) illustration is strikingly bryaceous, a new combination in Bryum would
be an effective disposition pending evaluation by a specialist in
that group.
New combination: Bryum kilimandscharicum (C. Miill.)
Zand., comb. nov. (Erpodiopsis kilimandscharica C. Miill.,
Flora 73: 470, 1890; Phascum kilimandscharicum (C. Miill.)
Roth).
KLEIOWEISIOPSIS
Kleioweisiopsis Dix., Smithsonian Misc. Coll. 72(3): 18, 1920.
Type: Kleioweisiopsis denticulata Dix., Kenya, Aberdare
Mts., Allan 395a, holotype, BM, isotype, US.
Because of the lack of characteristic papillae, the rectangular,
evenly thickened upper laminal cell walls, and the subulate perichaetial leaves, this genus is excluded from the Pottiaceae. For
the time being, it should be recognized in the Orthotrichaceae
near Amphidium. Kleioweisiopsis denticulata has dentate upper
laminal margins reminiscent of A. cyathicarpum but laminal cell
ornamentation consists of longitudinal lines of indistinct,
minute verrucae rather than small elliptical verrucae. The capsule of Kleioweisiopsis is spherical and the operculum is
differentiated but persistent. There is also some resemblance to
Dicranaceae subfam. Rhabdoweisioideae as noted by Dixon in
the original description, but the upper leaf cells of that group are
generally only as long as wide, and verrucae are apparently
absent. The holotype at BM is annotated by Saito as a possible
combination in Pleuridium; a second species K. involuta (PC!
an invalid name fide Crosby et al. 1992) proved to be Weissia
subg. Phasconica, see treatment of Weissia.
MELOPHYLLUM
Melophyllum Herz., Rep. Soc. Nov. Regn. Veg. 74: 98, 1939.
Type: Melophyllum radiculosum Herz.
EXCLUDED AND UNTREATED TAXA
275
This genus is a synonym of Astomiopsis C. Miill. (Ditrichaceae)
according to Buck and Zander (1980).
= Barbula amp/exifolia (Mitt.) Jaeg.
PSEUDOHYOPHILA
Pseudohyophila Hilp., Hedwigia 73: 68, 1933. Type: Pseudohyophila peruviana (Williams) Hilp., Peru, Juliaca, Williams
2874, holotype, NY.
Sebil/ea Biz., Rev. Bryol. Lichenol. 40: 120, 1974, nom. inval.
fide Crosby et al., 1992. Type: Sebil/ea brasiliensis Biz.,
nom. inval.
SEBILLEA
Originally published as Hyophila peruviana Williams (1903), this
monotypic genus has been recognized in the Grimmiaceae. Churchill (1981), in a treatment of the Grimmiaceae, indicated that
Pseudohyophi/a does not fit well in that family. The somewhat
tomentose stems; entire, lanceolate leaves with epapillose, quadrate upper laminal cells, often bistratose in patches on the upper
margins; weak, flattened, deeply grooved costa; sheathing perichaetial leaves; autoicous sexual condition; and capsule with distinct, furrowed neck are all characters shared by Dicranoweisia
Lindh. (cf. D. cirrata (Hedw.) Lindh. ex Milde), and, although
Pseudohyophila is eperistomate, it clearly should be placed near
that genus in the Dicranaceae.
Neither the genus and species names were validly published
(I.C.B.N., Art. 37 and 42 fide Crosby et al. 1992); the original
description merely cited "Type: Herbiers Sebille et Bizot" without indication of holotype. Although the herbarium of Sebille
has been incorporated into that of Bizot now at PC, according to
H. Bischler (in litt.), the type is not present in PC or at Dijon,
and may have been lost with certain other material during transfer of Bizot's herbarium to Paris. The illustration of this taxon is
reminiscent of Rhabdoweisia (Dicranaceae).
SERPOTORTELLA
Serpotortella Dix., J. Bot. 80: 44, 1942. Type: Serpotortel/a
madagassa Dix., Malagassy Republic, sin coli., "Herb.
Mus. Brit.," "Ref. No. 15," BM, lectotype nov.
PSEUDOTIMMIELLA
Pseudotimmiella Biz., Cryptogamie Bryol. Lichenol. 1: 425,
1980. Type: Pseudotimmiella pocsii Biz., Tanzania, Uluguru
Mts., above Morogoro, Pocs, Mwanjabe & Sharma 6549-H,
isotype, NY.= Diphyscium Mohr.
Characters indicating correct placement of this taxon (the type
collection lacks sporophytes) in Diphyscium (Diphysciaceae) are
the rigid, subulate perichaetical leaves, the long-ligulate cauline
leaves that develop an excurrent costa just below the perichaetium, and the lamina entirely bistratose, not unistratose marginally as in Timmiella.
New combination: Diphyscium pocsii (Biz.) Zand., comb.
nov. (Pseudotimmiel/a pocsii Biz., Cryptogamie Bryol. Lichenol.
1: 425, 1980).
RHAMPHIDIUM
Rhamphidium Mitt., J. Linn. Soc. Bot. 12: 45, 1869. Type: Rhamphidium macrostegium (Sull.) Mitt.
Trichostomum subg. Rhamphidium (Mitt.) Besch., Ann. Sc. Nat.
Bot., ser. 6(3): 198, 1876.
Norris and Koponen (1989) noted that the gametophyte of Rhamphidium was very similar to that of Dicranel/a (Dicranaceae), and
encouraged a restudy of familial placement. Eddy (1990) found
that Rhamphidium "appears to combine Pottioid sporophytic features with gametophytes of more Dicranoid character." According
to Hilpert (1933), Rhamphidium is better placed in the Ditrichaceae and he suggested a close relationship to Chei/othe/a
(Lindh.) Broth. of that family. Cheilothe/a chilensis (Mont.)
Broth., in face, has the characteristic sheathing leaf base of
Rhamphidium. In addition, the closely spirally ridged and often
spiculose peristome teeth rising from a low, weakly ornamented
and interiorly thickened plate of Rhamphidium are closely
approached if not matched by those of Sae/ania Lindh. (Ditrichaceae). Although the Ditrichaceae and the Pottiaceae are similar
in many gametophyte and peristome traits, the relationship of
Rhamphidium is clearly with the former family.
New heterotypic synonymy: Rhamphidium mussuriense Dix.
Serpotortel/a consists of two species of the island of Madagascar and is extraordinary for a number of features otherwise
characteristic of many species of Macromitrium (Orthotrichaceae). These features (some pointed out by Dixon 1942 in the
original description) include: the creeping main axis with distant leaves; erect branches with crowded leaves; leaves crisped
when dry, plicate, yellow in KOH, tendency towards a border of
elongate cells at the leaf base; upper larninal cells rounded in
section (with the appearance of a chain of beads), with columnar, simple to bifid papillae (although rather massive in Serpotortella). Serpotortella is similar to Tortel/a in the following
combination of characters: stem central strand present (absent in
the Orthotrichaceae fide Brotherus 1924-25 and cf. Zander &
Vitt 1979); costa with one layer of guide cells and a ventral
stereid band (Macromitrium has two layers of guide cells and
ventral stereid band absent or substereid at most); basal cells
inflated and sharply differentiated in a vee reaching up the leaf
margins. The strongly corrugate upper lamina of one species is
reminiscent of that of a variant of Anoectangium aestivum
(Zander 1976) and of Macromitrium incrustatifolium Robins.
(Robinson 1968; Gangulee 1969-80) and other Macromitrium
species, but the cells are not bistratose as they are in these latter
species, and the extra cell growth involved may be a malformation. Although Tortella species may sometimes send up new
shoots from an occasional prostrate shoot, this is always secondary, occurring after damage such as crushing of a turf. Serpotortella has a definite prostrate axis with distant leaves somewhat differently shaped than those of the crowded branches.
Although Mitthyridium Robins. of the Calymperaceae has such
a prostrate main axis, the differentiated, sheathing perichaetial
leaves and the stem central strand do not occur in the Calymperaceae.
Several characteristics that in combination led W. D. Reese
and me to place Serpotortella in its own monogeneric family,
Serpotortellaceae Reese & Zand. (1987, 1988b), include creeping main stems, with central strand; leaves with only a few
porose basal cells; sheathing perichaetialleaves; peristome teeth
smooth, with transverse bars, reflexed when dry; gemmae
276
GENERA OF THE POTTIACEAE
absent. The lectotype should be based on material seen by the
original author, and so is changed above from S. chenagonii (Ren.
& Card.) Reese & Zand.
New heterotypic synonymy: Serpotortella madagassa Dix. =
Serpotortella chenagonii (Ren. & Card.) Reese & Zand.,
Bryologist 90: 234, 1987 [1988]. Serpotortella marginata Dix. =
Serpotortel/a chenagonii (Ren. & Card.) Reese & Zand.,
Bryologist 90: 234, 1987 [1988].
SPLACHNOBRYUM
Splachnobryum C. Miill., Verh. Zool. Bot. Ges. Wien 19: 503,
1869. Lectotype: Splachnobryum obtusum (Brid.) C. Miill.
This genus has been removed (A. Koponen 1981) from the Pottiaceae to the Splachnobryaceae A. Kop., Ann. Bot. Fenn. 18: 128,
1981, type: Splachnobryum C. Miill. There has been considerable
support for this. Andrews (1949) first used the term
"Splachnobryaceae" but did not actually accept it as a family.
Corley et al. (1981) also felt that Splachnobryum "would be best
placed in a new family, Splachnobryaceae" but did not take the
necessary steps to do so.
TRIDONTIUM
Tridontium Hook. f. in Hook., Icon. Pl. Rar. 3: 148, 1840. Type:
Tridontium tasmanicum Hook. f., Australia, Tasmania, ex
herb. Hooker, isotype, NY.
Plants forming turfs or cushions, greenish brown above, blackish
brown to dark brown below. Stems branching irregularly, ca. 3.0
em in length, transverse section rounded-pentagonal, central
strand present, strong, sclerodermis present, hyalodermis absent;
axillary hairs to 5 cells in length, basal cell brown; occasionally
sparsely radiculose. Leaves incurved, contorted when dry, spreading when moist, ligulate-lanceolate, ca. 3.0 mrn in length, upper
lamina broadly channelled, margins plane to erect above,
recurved in lower 114, entire, ca. 4 rows of enlarged cells forming
inframarginal border in lower 1/3-l/2, border sometimes absent;
apex broadly rounded to occasionally rounded-acute, somewhat
cucullate; base scarcely differentiated to long-elliptical; costa
strong, flattened, ending ca. 6 cells below apex, superficial cells
elongate on both sides, ca. 6 rows of cells across costa ventrally at
midleaf, costal transverse section semicircular to elliptical, stereid
band absent to weak ventrally, present dorsally, epidermis present
dorsally and absent ventrally, guide cells 4(-6) in 1layer, hydroid
strand absent; upper /aminal cells irregularly hexagonal, heterogeneous in shape, ca. 9-13 jlm in width, approximately 1:1, walls
often irregularly thickened, superficially flat, lumens usually
round to elliptical; papillae absent; basal cells differentiated
across leaf from intramarginal border to costa, rhomboidal to rectangular, ca. 15 jlm in width, 3-4: I, walls evenly thickened.
Dioicous. Perichaetia terminal, archegonia long, to 1.5 mm, inner
perichaetialleaves long-oblong, to 4 mrn in length, sheathing seta,
cells long-rhomboidal and inflated in lower 1/2. Perigonia terminal, gemmate, paraphyses filiform, of uniseriate cells. Seta ca. l
em in length, I per perichaetium, dark brown, twisted clockwise
above; theca fleshy, ca. 1.2-1.5 mrn in length, dark brown,
urceolate to short-ovoid, often with a circumstomal ring, exothecial cells short-rectangular, ca. 20 jlm in width, 2: l, walls
evenly thickened, stomates phaneropore, at base of theca, usually
sunken in pits, annulus 4-6 rows of vesiculose, persistent cells;
peristome teeth 16, lanceolate, cleft 2-3 times halfway or to
base, often variously perforate, yellow, low-papillose to densely
spiculose, to 450 jlm in length, with several articulations,
straight, basal membrane absent. Operculum rostrate, oblique,
ca. 2 mm in length, cells straight. Calyptra cucullate, smooth,
ca. 3.8 mm in length. Spores ca. 25-30 jlm in diameter, yellow,
obscurely papillose. Lamina! KOH color reaction orange.
Found in wet, generally limy areas in New Zealand,
Tasmania, and Auckland and Macquarie Islands.
This critical monotypic genus is treated here in detail to
give some perspective on its familial relationship.
Clavate propagula in leaf axils were reported for Tridontium
by Norris and Koponen (1989). Hilpert (1933) emphasized a
relationship with the Pleuroweisieae (a tribe whose members
are here assigned elsewhere in the Pottiaceae in other
subfamilial groups), specifically with Eucladium (now in the
Trichostomoideae), but a careful comparison of the characters
italicized in the discriptions given here demonstrates that the
similarity of habitat has probably been given undue weight.
Although variation in appearance of the leaf apex is considerable in many taxa of Pottiaceae as correlated with variation in
ratio of length and width of the leaf, only a high level of character similarity in other respects or a clear phyletic series of
related species can replace utility of this feature in taxonomic
evaluations in the family. Tridontium has no apparent relationship with Eucladium. I agree with Dixon (1923) and Sainsbury
(1955) that the marginal border may be practically absent in
some specimens (e.g. type of Weissia lancifolia (BM, see
below), and I also fmd, as did Sainsbury, that the peristome is
not so regularly trifid as figured by Brotherus (1924-25).
Although Dixon appreciated a resemblance to Cinclidotus
P. Beauv. (Cinclidotaceae fide Saito 1975a), Tridontium is usually placed with the Pottiaceae, near Leptodontium (note the
absence of a costal ventral epidermis, this a characteristic of the
Leptodontium group), Erythrophyllopsis or Eucladium. Eddy
(1990) noted that Tridontium "stands well apart from the other
Pottiaceae.... " Although (1) the peristome of Tridontium is
quite unlike the net-like peristomial cone of Cinclidotus, (2) the
opercular cells are rectangular, not isodiametric as in Cinclidotus, and (3) stomates are present, it is because of the fleshy
theca, large spores, and above attributes of the gametophyte that
there is considerable resemblance to that similarly aquatic
genus.
Tridontium has a striking resemblance to Scouleria
(Grimmiaceae, or Scouleriaceae Churchill in Funk & Brooks,
Advances Cladistics 143, 1981, type: Scouleria Hook. in
Drumm. see Churchill 1985), including the hydric habitat,
thick-walled exothecial cells, lanceolate leaves with
intramarginal border, rounded hexagonal upper !aminal cells,
and cucullate calyptra. Character states listed by Murray (1984)
for certain other genera of Grimmiaceae (subfam. Coscinodontoideae), Jndusiella, Aligrimmia and Coscinodontella, such as
macrostomous capsules with cleft or rerforate peristome teeth,
strong stem central strand, ovate-lanceolate leaves with obtuse
apices, leaves with involute upper margins and leaf cells with
angular lumina, also occur in Tridontium. These taxa of the
Grimmiaceae are distinct, however, in the plicate calyptrae
(cucullate calyptrae are rare in Grimrniaceae) and monoicous
sexuality. It is quite possible that both Tridontium and Cinclidotus are taxa bridging the Pottiaceae and the Grimmiaceae.
EXCLUDED AND UNTREATED TAXA
Tridontium is here placed in the Grimmiaceae in the
Scoulerioideae pending some future evaluation of the relationship
of the two families.
Number of accepted species: 1.
Species examined: T. tasmanicum (ALTA, BM, BUF, DUKE,
MO,NY).
New heterotypic synonymy: Weissia lancifolia (R. Br. ter)
Wijk & Marg. (Dicranum lancifolium R. Br. ter) = Tridontium
tasmanicum Hook. f. in Hook.
TISSERANTIELLA
Tisserantiella P. Yarde, Bull. Soc. Bot. France 88: 469, 1941.
Type: Tisserantiella spathulata P. Yarde, Cameroon, LeMauf,
1937, holotype, PC.
Macroglossum Hilp., Beih. Bot. Centralbl. 50(2): 670, 1933,
nom. illeg. non Copeland, 1909. Type: Macroglossum
pulchellum Ther. & Hilp., Congo, Overlaet, 1923, isotype,
cu.
Potier de la Yarde (1941) in the original description of this genus
noted its similarity to papillose species of Uleastrum Buck (as
Ulea C. Mtill.). Tisserantiella lacks a peristome but nonetheless
clearly belongs to the Rhachitheciaceae by the arborescent substrate, presence of a stem central strand, sclerodermis absent;
leaves spathulate with costa ending 15-30 cells below apex;
!aminal papillae very small, simple, solid; basal cells much
differentiated, elongate, smooth; seta rather short, 0.6-l.O mm in
length; annulus strongly vesiculose; and !aminal KOH color reaction yellow. The type of Weissia minutissima (NY, see below)
from Brazil is also a member of this genus, differing in the longer
costa.
The family Rhachitheciaceae Robins. is distinguished from
the Pottiaceae (emending Robinson's 1964 description, also cf.
Iwatsuki & Sharp 1976 and Allen & Pursell 1991) by the following combination of characters: small corticolous plants usually of
low elevations in tropical areas; without stem sclerodermis, small
or absent central strand; leaves ligulate, upper lamina! cells often
with solid or hollow papillae, basal cells inflated or much elongate, smooth; costa usually ending several cells below the apex,
often reaching only to 2/3 or 3/4 the leaf length; perichaetial
leaves usually sheathing; seta stout, often kinked just below the
capsule; theca ovoid, smooth or 8-ribbed; peristome teeth seldom
absent, spreading when wet, strongly incurved when dry, 16 and
paired, smooth, articulations generally close; calyptra cucullate,
smooth; and KOH lamina! color reaction yellow. Multi-ribbed
thecae are also found in the Pottiaceae in Ganguleea angulosa
and in Weisiopsis plicata. The genera of the Rhachitheciaceae are
an apparent link between the Pottiaceae and the Orthotrichaceae.
Three genera are here recognized in the Rhachitheciaceae:
Hypnodontopsis Iwats. & Nog., Rhachithecium Broth. ex Le Jolis,
and Tisserantiella. Jonesiobryum Biz. & Pocs ex Allen & Pursell
apparently belongs in this family, too (cf. Allen & Purselll99l).
New heterotypic synonymy: Tisserantiella spathulata P.
Yarde= Tisserantiella pulchellum (Ther. & Hilp.) Zand.
New combinations in Tisserantiella: Tisserantiella pulchella
(Ther. & Hilp.) Zand., comb. nov. (Macroglossum pulchellum
Ther. & Hilp., Beih. Bot. Centralbl. 50(2): 671 , 1933; Hyophila
pulchella (Ther. & Hilp.) Wijk & Marg.). Tisserantiella
277
minutissima (Mitt.) Zand., comb. nov. (Weissia minutissima
Mitt., J. Linn. Soc. Bot. 12: 138, 1869; Hyophila minutissima
(Mitt.) Jaeg.; Macroglossum minutissimum (Mitt.) Hilp.).
New combination in Rhachithecium: Rhachithecium welwitchii (Duby) Zand., comb. nov. (Zygodon welwitchii Duby,
Mem. Soc. Phys. Hist. Nat. Geneve 21: 44, 1871; Ulea welwitschii (Duby) Broth. in Par.).
ULEASTRUM
Uleastrum Buck, Candollea 40: 203, 1985. Type: Ulea
palmicola C. Mtill.
Ulea C. Mtill., Hedwigia 36: 102, 1897, hom. illeg. non
Schroeter, 1892. Type: Ulea palmicola C. Mtill.
Spruceella C. Mtill., Gen. Muse. Fr. 396, 1900, hom. illeg.
non Pierre, 1890. Type: Spruceella octoblepharis (Jaeg.) C.
Mtill.
Except for the sharply differentiated basal cells, the combination of the the ligulate leaves with acute apices, smooth
bistratose upper !aminal cells and other gametophyte characters
are much like Ptychomitrium incurvum (Schwaegr.) Spruce of
the Ptychomitriaceae. Uleastrum differs from other genera of
the Ptychomitriaceae, however, in the cucullate calyptra and
smooth peristome teeth. Chen (1941: 39) suggested that this
genus is closely related to Rhachithecium and there is indeed
some evidence of relationship. The costa is relatively weak and
generally ends below the leaf apex. Uleastrum palmicola, the
generitype, has propagula borne on the dorsal side of the basal
cells, as has Rhachithecium perpusillum . Uleastrum
octoblepharis has a kinked seta similar to that of species of
Rhachithecium and Hypnodontopsis, but other species of Uleastrum lack this, and, in fact, probably should belong in another
genus. Pending further study, Uleastrum can be recognized in
the Orthotrichaceae, a disposition making the least necessary
emendation of recognized family limits.
Species examined: U. octoblephare (BUF, H, NY), U.
nitidum (PC), U. palmicola (H, NY), U. paraguense (H).
New heterotypic synonymy: Zygodon palmarum C. Miill.
(isotype H!) = Uleastrum palmicola (C. Mtill.) Zand.
New combinations: Uleastrum nitidum (Ther. in Felipp.)
Zand., comb. nov. (Ulea nitida Ther. in Felipp., Rev. Bryol. n.
ser. 2: 214, 1930). Uleastrum octoblephare (Spruce ex Jaeg.)
Zand., comb. nov. (Pottia octoblepharis Spruce ex Jaeg., Ber. S.
Gall. Naturw. Ges. 1871-72: 343, 1873 (Ad. 1: 191); Weissia
octoblepharis Spruce ex Mitt., J. Linn. Soc. Bot. 12: 40, 1869,
hom. illeg.; Ulea octoblepharis (Spruce ex Jaeg.) C. Mtill.,
Hedwigia 37: 234, 1898). Uleastrum palmicola (C. Mtill.)
Zand., comb. nov. (Ulea palmicola C. Mtill., Hedwigia 36: 102,
1897).
ULEOPSIS
Uleopsis Ther., Rev. Bryol. Lichenol. 9: 20, 1936. Type: Uleopsis mamillosa Ther..
The type of this species (Ecuador, rochers du Condorguachana,
Benoist 3154, PC) is Oreoweisia brasiliensis Hampe (sensu
Griffin 1986a). Uleopsis Ther. thus becomes ~ a synonym of
Oreoweisia De Not.
GENERA OF THE POTTIACEAE
278
SPECIES
Tortula montana Mitt., J. Linnean Soc. Bot. 12: 156, 1869 (Didymodon montanus (Mitt.) Broth.) = Rhamphidium montanus (Mitt.)
Zand., comb. nov.
Gymnostomum lessonii Besch. = Racopilum sp. (Racopilaceae).
Phascum carinatum Hampe, Vid. Medd. Naturh. For. Kjoebenh.
ser. 4, 1: 76, 1879 = Bruchia carinata (Hampe) Zand., comb. nov.
The type (BM) of this Brazilian species consists of (a very few)
plants with the characteristic enlarged capsule neck and rectangular !aminal cells of Bruchia (Bruchiaceae).
Pottia macrocarpa Schimp., Ann. Sc. Nat. Bot. ser. 2, 6: 145,
1836 = Funaria macrocarpa (Schimp.) Zand., comb. nov. A fragmentary type at NY is apparently a Funaria by the large,
eperistomate capsule, subpercurrent costa, bluntly dentate upper
leaf margins, lax upper lamina! cells, and yellow KOH reaction.
Pottia mirabilis Broth. & Par., Rev. Bryol. 31: 118, 1904 =
Physcomitrium mirabile (Broth. & Par.) Zand., comb. nov. The
gametophytes of the type at H are soft, glossy and otherwise
funariaceous or bryaceous in general appearance. The specimen
keys (Brotherus 1924-25) to Physcomitrium sect. Cryptopyxis
C. Miill. (Funariaceae) , and a new combination in
Physcomitrium is appropriate here.
Tortula domingensis Ther., Rev. Bryol. Lichenol. 14: 12, 1944
= Brachymenium domingense (Ther.) Zand., comb. nov.
lsotypes at NY and US have the characters of Brachymenium
Schwaegr. (Bryaceae). One may especially note the dense red
tomentum, lack of differentiated basal lamina! cells, and upright
capsule with an apparently bryaceous peristome (hyaline and
poorly developed in the rather young capsules). gametophytes
of this species are similar to those of Tortula sect. Pottia but are
more densely yellow in KOH solution. The operculum is large
for the genus, being stoutly long-conic.
AN UPDATED LIST OF GENERA, SPECIES AND INFRASPECIFIC TAXA
OF THE POTTIACEAE
This is a list of presently recognised genera and specific and infraspecific taxa and their distributions, as an extension to Index
Muscorum and its first supplement (Vander Wijk et al. 1959-1969), and later supplements (Crosby 1977, 1979; Crosby & Bauer
1981, 1983, 1986; Crosby, Magill & Bauer 1992). Other supraspecific taxa and thier synonyms are given in the family and generic
treatments. Only italicized (correct) names were extracted from the Index Muscorum for inclusion here-this list does not provide
all synonymy; included is new synonymy made since publication of that work, plus citations of formae, largely from Podpt\ra
(1954). Synonyms with which the author disagrees or which are superseded are placed in square brackets.
Series of new combinations at the species level that would put all epithets into some other genus were usually not made. Combinations at the species level were made only after seeing authentic material or illustrations of authentic material that show distinctive
characters. For example, all epithets of correct names in Astomum and Hymenostomum (now in synonymy with Weissia) were not
transferred to Weissia because, although the generitypes belong in Weissia, many species probably belong with Trichostomum.
Even the Index Muscorum has many "orphaned" specific and infraspecific epithets languishing in combinations with otherwise
synonymized generic and specific names. Revisionists are encouraged to examine types of these names and make the appropriate
combinations or synonymy. Complete series of new combinations for many infraspecific epithets were made, however, with the
assumption that their relationship with the typical variety is secure.
In the past, I have used varietal names for widely distributed taxa (e.g. segregates of Didymodon australasiae s. lat., D. rigidulus
s. lat., D. vinealis s. lat., Leptodontium viticulosoides s. lat.) that intergrade in morphology in major parts of their geographic range
(for further discussion see Zander 198lc). Since there is apparently a resistence of the part of other bryologists to using these admittedly ungainly three-part names for widely distributed taxa, and because in some parts of the range of these species they apparently
do not intergrade, the names of such taxa are given here as correct at the species level. Widespread infraspecific taxa of D. rigidulus
s. lat. are excepted since the abundant variation in these populations precludes morphological sorting of such names so as to assign
types to even vaguely circumscribed species limits. This argument is also true for weakly distinguishable local populations or
"facies" of New World Anoectangium aestivum (see Zander 1977c), which are not recognized as separate taxa.
In accordance with Art. 73.9 of the 1987 I.C.B.N. (Greuter 1988), hyphens are eliminated from all epithets unless the connected
words may stand independently. Also included here are the names of hybrid formulas, with authors given for each name and the
reporting author after "fide"; the usage of the Index Muscorum is apparently not in consonance with the Code. The abbreviation
used here for rank not cited (see Art. 35.1; also W. Margadant pers. comm.) is "nom. inval. dispon. non cit."
The floristic distribution codes of the Index Muscorum were updated at the correct name whenever new synonymy was added,
but, inasmuch as the taxonomy of the Pottiaceae was the priority for the time allotted to complete this study, no general survey of
the floristics literature since the publication of Index Muscorum was made to update all distributions. Afrl includes North Africa,
Madeira, Azores, Canary Islands; Afr2 includes Central Africa, St. Helena; Afr3 includes Madagascar, Mauritius, Reunion; Afr4
includes South Africa, Kerguelen; Ami includes North America, Greenland, Aleutians, Bermuda; Am2 is Central America; Am3 is
the West Indies; Am4 includes Venezuela, Colombia, Peru, Bolivia, Ecuador, Galapagos; Am5 includes Brazil, Paraguay, Guiana,
Trinidad, Tobago; Am6 includes Chili, Argentina, Uruguay, Falklands; Ant is Antarctica; Asl includes Northern Asia, Sakhalin;
As2 includes China, Mongolia, Japan, Korea, Formosa; As3 includes India, Pakistan, Ceylon, Burma, Thailand, Indochina; As4
includes Indonesia, Malaya, Philippines, New Guinea; As5 includes Asiatic portion of Middle East, Cyprus; Austrl includes Australia, Tasmania; Austr2 is New Zealand and nearby islands; Oc includes the Pacific islands.
This list recognizes as correct names in the Pottiaceae a total of 1457 species, 31 subspecies, 536 varieties, 339 formae and 7
subformae.
ACAULON C. Miill.
Acaulon capense C. MUll. [see Magill, Fl. S. Afr. I. Mosses 1: 201,
1981 (1982)] Afr4
Acau/on casasianum Brugu6s & Crum, Lindbergia 10: 1, 1984 Eur
Acau/on chrysacanthum Stone,]. Bryol. 9: 213, 1976 [1977] Austrl
Acau/on crassinervium C. Miill. Austr1 Austr2
Acaulon dertosense Casas, S6rgio, Cros & Brugu6s, Anales Jard. Bot.
Madrid 42: 299, 1986 Eur
Acaulon eremico/a Stone, J. Bryol. 10: 467-474. 1979 [1980] Austr1
Acau/on fontiquerianum Casas & S6rgio, Cryptogamie Bryol.
Lichenol. 11: 61, 1990 Eur
Acau/on granulosum Stone, J. Bryol. 15: 257. Austr1
Acaulon integrifolium C. Miill. Austr1
var. aristatum (Willis) Willis Austr1
Acaulon /eucochaete Stone, J. Bryol. 9: 217, 1976 [1977] Afr4 Austr1
[Acau/on mediterraneum Limpr. Eur
Acaulon muticum var.
=
mediterraneum (Limpr.) S6rgio fide S6rgio, Bol. Soc. Brot. 46:
460, 1972]
[Acaulon minus (Hook. & Tayl.) Jaeg. Eur Afr1 =Acaulon muticum
(Hedw.) C. Miill.fide Hill, J. Bryol. 12: 11, 1982]
Acaulon muticum (Hedw.) C. Miill. Eur Am1 As1 As5 Afrl Afr2
Afr4
var. mediterraneum (Limpr.) S6rgio fide S6rgio, Bol. Soc. Brot.
46:460,1972
[var. minus (Hook. & Tayl.) B.&S. in BSG Eur Afr1 = Acaulon
muticum (Hedw.) C. Miill.fide Hill, J. Bryol. 12: 11, 1982]
var. rufescens (Jaeg.) Crum, Bryo1ogist 72: 240, 1969 Am1
fo. subintegrifolium C. Jens., Skand. Bladmfl. 216, 1939 nom.
inval. descr. suec.
Acaulon nanum C. Miill. Am5
[Acaulon piligerum (De Not.) Limpr. Eur = Acaulon triquetrum
(Spruce) C. Miill. fide Corley et al., J. Bryol. 11: 621, 1981
GENERA OF THE POTfiACEAE
280
(1982)]
Acaulon recurvatum Magill, Fl. S. Afr. I. Mosses 1: 199, 1981 [1982]
Afr4
Acaulon robustum Broth. ex Roth Austr1
[Acaulon rufescens Jaeg. Am1
Acaulon muticum var. rufescens
(Jaeg.) Crumfide Crum, Steere & Anderson 1973]
[Acaulon rufochaete Magill, Fl. S. Afr. I. Mosses I: 20I, I98I [I982]
Afr4 = Microbryum rufochaete (Magill) Zand., see treatment of
Microbryum]
Acaulon schimperianum (Sull.) Sull. in Sull. & Lesq. Ami Am2
Acaulon sphaericum Shaw (Lsee Magill, Fl. S. Afr.l. Mosses 1: 20I,
I98I [I982]) Afr4
Acaulon triquetrum (Spruce) C. Miill. Eur Afri Ami As I Austri
var. desertorum (Besch.) Jelenc Afri
Acaulon uleanum C. Miill. Am5
Acaulon vesiculosum C. Miill. Am6
=
ALOINA Kindb.
Aloina a/aides (Schultz) Kindb. Eur As I As2 As5 Afri Ami
var. ambigua (BSG) Craig in Grout Eur Asi As2 As5 Afri Ami
Am2 Austri
[Aloina ambigua (BSG) Limpr. = Aloina rigida var. ambigua (BSG)
Craig]
fo. microphylla (Latz.) Podp., Consp. Muse. Eur. 238, I954 (Aloina
ericaefolia fo.) Eur
Aloina apiculata (Bartr) Delgad. Am3
Aloina bifrons (De Not.) Delgad. Eur Asi As2 As5 Afri Afr4 Ami
Am2Am3
Aloina brevirostris (Hook. & Grev.) Kindb. Eur As I As2 As5 Ami
[var. breidleri (Limpr.) Limpr. Eur = Aloina brevirostris (Hook. &
Grev.) Kindb.fide Delgadillo, I973]
[var. elongata (Angstr.) Par. Eur = Aloina brevirostris (Hook. &
Grev.) Kindb.fide Delgadillo, I973]
[var. rotundifolia Warns!. Eur = Aloina brevirostris (Hook. &
Grev.) Kindb.fide Delgadillo, I973]
[Aloina calceolifolia (Mitt.) Broth. Am2 Am4 = Aloina rigida
(Hedw.) Limpr. var. rigidafide Delgadillo, I975]
Aloina catillum (C. Miill.) Broth. Am2 Am4
Aloina cornifolia Delgad., Bryologist 78: 265, I975 As2
[Aloina ericaefolia (Lindh.) Kindb. Eur =Aloina rig ida var. ambigua
(BSG) Craig]
[fo. microphylla Latz., Bot. Centralbl. Beih. 48(2): 484, I93I =Alaina ambigua fo. microphylla (Latz.) Podp., Consp. Muse. Eur.
238, I954]
Aloina hamulus (C. Miill.) Broth. Ami Am2
[Aloina longirostris Card. hom. illeg. Am2 = Aloina hamulus (C.
Miill.) Broth. fide Delgadillo I973]
[Aloina obliquifolia (C. Miill.) Broth. As2
Aloina rigida var.
obliquifolia (C. Miill.) Delgad.]
[Aloina pilifera (De Not.) Crum & Steere nom. superfl. Eur As5 Ami
Am2 = Aloina bifrons (De Not.) Delgad. see Corley et al., J.
Bryol. 11: 620, I98I (1982)]
Aloina recurvipatula (C. Miill.) Broth. Am6
Aloina rigida (Hedw.) Limpr. Eur Afri Afr4 Ami Am2 As I As2 As3
As5 Austri
[var. ambigua (BSG) Craig Eur As1 As2 As5 Afrl Ami Aloina
a/aides var. ambigua (BSG) Craig in Grout fide Crum, Steere &
Anderson I973]
[var. longirostris (Torka) Podp. Eur = Aloina rigida (Hedw.)
Limpr.fide Delgadillo, Bryologist 78: 160, 1975]
var. mucronulata (BSG) Limpr. Eur (see Delgadillo, Bryologist 78:
262, I975
var. obliquifolia (C. Miill.) Delgad., Bryologist 78: 264, I975
=
=
(Barbula) As2
fo. mucronata Monk., Laubm. Eur. 3I7, I927 Eur
[fo. mucronulata (BSG) Podp., Consp. Muse. Eur. 237, 1954
(Tortula rig ida var.) Eur Aloina rig ida fo. mucronata Monk.]
fo. obtusa (Jur.) Monk., Laubm. Eur. 3I7, I927 (Tortula rigida
var.) Eur
[fo. pilifera (De Not.) Monk., 3I7, I927 (Tortula rigida var.) Eur
As5 Aloina pilifera (De Not.) Crum & Steere nom. superfl. =
Aloina bifrons (De Not.) Delgad. see Corley et al., J. Bryol. 1I:
620, I98I (I982)]
Aloina rosei (Williams) Delgad., Bryologist 76: 273, I973 Am1
Am4
Aloina sedifolia (C. Miill.) Broth. Am6
Aloina suJJivaniana (C. Miill.) Broth. Austri Austr2 [= Aloina
bifrons fide Delgadillo, I973, but see Scott & Stone, Mo.
Southern Australia 454, I976 and fide Catcheside, Mosses S.
Austr. I55, 1980]
=
=
ALOINELLA Card.
Aloinella andina Delgad. Am4
[Alaine/la apiculata Bartr. Am3
Aloina apiculata (Bartr.)
Delgad.]
Alaine/la boliviana Broth. in Herz. Am4
Aloinella catenula Card. Am2 Am4
Aloine/la cucu/latifolia (C. Mii1l.) Broth. Am6
Alaine/la cucul/ifera (Mitt.) Steere Am4
Aloinella galeata (C. Miill.) Broth. Am6
[Aloinella hamulus (C. Miill.) Bartr. Am2 = Aloina hamulus (C.
Miill.) Broth.]
Aloinel/a venezuelana Griffin, Bull. Torrey Bot. Club 102: 26, 1975
Am4
=
=
[ANICTANGIUM Hedw. nom. rejic. Anoectangium Schwaegr.
nom. cons.]
[Anictangium ciliatum Hedw.
Hedwigia ciliata (Hedw.) P.
Beauv.]
var. incanum Sw. Eur
Anictangium orthotrichoides Gill. ex Grev. Am6
=
ANOECTANGIUM Schwaegr.
Anoectangium abyssinicum Hampe ex Geh. Afr2
Anoectangium aestivum (Hedw.) Mitt. Eur As2 As3 As4 Afri Afr2
Ami Am2 Am4 Am5 Austr2 Oc
var. glaciale (Lor. & Mol.) Wijk & Marg. Eur
var. pel/ucidum (Wils.) Braithw. Eur.
Anoectangium afrocompactum C. Miill. ex Dus. Afrl
[Anoectangium angustifolium Mitt. Afri = Anoectangium aestivum
(Hedw.) Mitt. fide Dirkse et al., Cryptogamie Bryol. Lichenol.
I4: I5, I993]
[Anoectangium anomalum Bartr. As4 = Hymenostylium
recurvirostrum (Hedw.) Dix.fide Norris & Koponen, Acta Bot.
Fenn. 137: I02, I989j
[var. trifarium Bartr., Lloydia 5: 255, I942 As4 =Hymenostylium
recurvirostrum (Hedw.) Dix.fide Norris & Koponen, Acta Bot.
Fenn. 137: I02, I989]
[Anoectangium apiculatum Schimp. in Besch. Am2 = Anoectangium aestivum (Hedw.) Mitt. fide Zander, Bryologist 80: 243,
I977]
[Anoectangium arizonicum Bartr. ex Grout Ami Am2 = Gymnostomum aeruginosum Sm. fide Zander, Bryologist 80: 259, I977]
[Anoectangium balfourei Mitt. Afr2 = Semibarbula orienta/is
(Web.) Wijk & Marg. fide Frey & Kiirschner, Nova Hedw. 46:
94, 1988 =Barbula indica (Hook.) Spreng]
GENERA, SPECIES AND INFRASPECIFIC TAXA
Anoectangium be/Iii Broth. ex Dix. Austrl Austr2
Anoectangium bicolor Ren. & Card. As3
Anoectangium borbonense Besch. Afr3
Anoectangium brachyphyllum Broth. in Herz. As4
[Anoectangium breutelianum BSG ex Besch. Am2 =?Hymenostylium
recurvirostrum (Hedw.) Dix. fide Zander, Bryologist 80: 265,
1977]
Anoectangium brotherusii Kis, Mosses South-east Trop. Afr., lnst.
Ecol. Bot. Hungarian Acad. Sci. 46, 1985 (nom . nov. for Anoectangium torquatum Broth. hom. illeg.) Afr2
[Anoectangium ca/idum Mitt. Am4 = Anoectangium aestivum
(Hedw.) Mitt. fide Zander, Bryologist 80: 243, 1977]
Anoectangium c/arum Mitt. As2 As3
[Anoectangium compactum Schwaegr. = Anoectar gium aestivum
(Hedw.) Mitt.]
[var. a/askanum Card. & Ther. Ami = Anoectangium aestivum
(Hedw.) Mitt., see treatment of Anoectangium]
var. madeirense Geh. in Geh. & Herz. Afr1
Anoectangium contortum Broth. Asl
Anoectangium crassinervium Mitt. As2
[Anoectangium crustatum J. Frtihl., Ann. Naturh. Mus. Wien 66:
35-36, 1962 (1963) Eur = Gymnostomum viridulum Brid. fide
Corley & Crundwell, J. Bryol. 16: 346, 1991]
[Anoectangium handelii Schiffn. AsS = Molendoa sendtneriana
(BSG) Limpr.fide Zander, Bryologist 80: 248, 1977]
Anoectangium hanningtonii Mitt. Afr2
Anoectangium harttiae Bartr. Oc
Anoectangium herzogii Broth. in Herz. Am4
Anoectangium hobsonii Mitt. AsS
Anoectangium humblotii Ren. & Card. in Ren. Afr3
Anoectangium hymenodontoides (C. Mi.ill.) Jaeg. As3
Anoectangium impressum Hampe Afr3
[Anoectangium incrassatum Broth. in Boerg. Am3 = Anoectangium
aestivum (Hedw.) Mitt. fide Zander, Bryologist 80: 243, 1977]
[Anoectangium incurvans (Schimp. ex Besch.) Bartr. Am2 = Molendoa sendtneriana (BSG) Limpr. fide Zander, Bryologist 80: 248,
1977]
[Anoectangium jamaicense (C. Mi.ill.) Par. Am3 = Anoectangium
aestivum (Hedw.) Mitt. fide Zander, Bryologist 80: 243, 1977]
Anoectangium kashmiriense Aziz & Vohra, Bull. Bot. Surv. India 35:
239, 1983 As3
Anoectangium keniae (P. Yarde) Zand. (Gymnostomum), see treatment of Anoectangium Afr2
[Anoectangium kilimandscharicum Broth. hom. il/eg.]
var. minutum Broth. Afr2
[Anoectangium laetevirens Besch. & Card. As2 = Anoectangium
thomsonii Mitt. fide Saito, J. Hattori Bot. Lab. 39: 458, 1975]
[Anoectangium /aetum Ren. & Card. = Anoectangium stracheyanum
Mitt.]
fo. henryi Tix., Rev. Bryol. Lichenol. 34: 132, 1966 As3
[Anoectangium lechlerianum Mitt. Am4 = Molendoa sendtneriana
(BSG) Limpr.fide Zander, Bryologist 80: 247, 1977]
var. laetius Hampe Am4
var. limbatulum Bartr. in Bauer Am6
[Anoectangium liebmannii Schimpr. ex Besch. Am2 =Anoectangium
aestivum (Hedw.) Mitt. fide Zander, Bryologist 80: 243, 1977]
[var. viride Card. Am2 = Molendoa sendtneriana (BSG) Limpr.
fide Zander, Bryologist 80: 247, 1977]
Anoectangium lineare (C. Mi.ill.) Kindb. Am4 Am6
[Anoectangium /ombokense Broth. As4 = Anoectangium aestivum
(Hedw.) Mitt. fide Touw, J. Hattori Bot. Lab. 71 : 341, 1992]
Anoectangium madagassum Ren. & Par. Afr3
Anoectangium mafatense Ren. & Card. Afr2 Afr3
281
Anoectangium magnirete Ren. & Card. Afr2 Afr3
Anoectangium microphyllum Card. As2
Anoectangium nigerianum Broth. & Par. Afr2
Anoectangium papuanum Fleisch. As4
Anoectangium patagonicum Card. & Broth. Arn6
[Anoectangium peckii (Sull.) Sull. ex Aust. Arn1 = Anoectangium
aestivum (Hedw.) Mitt. fide Crum, Steere & Anders.,
Bryologist 76: 109, 1973, see also Zander, Bryologist 80: 243,
1977]
[Anoectangium peruvianum Sull. Am4 = Molendoa sendtneriana
(BSG) Limpr.fide Zander, Bryologist 80: 247, 1977]
Anoectangium pflanzii Broth. Arn4
[Anoectangium platyphyllum Williams Am4 = Molendoa
p/atyphyl/um (Williams) Zand., see treatment of Molendoa]
Anoectangium p/euroweisioides J. Frtilich, Ann. Naturhist. Mus.
Wien 67: 151, 1964 As3
Anoectangium rhaphidostegium C. Mi.ill. ex Broth. Afr3
Anoectangium rivale Card. As2
[Anoectangium rubrigemmium Hoe & Crum Oc
Zygodon
rubrigemmius (Hoe & Crum) Zand. & Vitt., Canad. J. Bot. 57:
296, 1979]
Anoectangium rufoviride Besch. As2
var. eucollum Besch. Afr3
Anoectangium schimperi Mitt. Afr2
Anoectangium sellae Negri Afr2
Anoectangium shepherdae (Card. & Dix.) Zand. (Hymenostylium),
see treatment of Anoectangium As3
Anoectangium sikkimense Aziz & Vohra, Bull. Bot. Surv. India 30:
185, 1988 [1990] As 3
Anoectangium spathulatum Mitt. Afr2
Anoectangium stracheyanum Mitt. As2 As3
[var. gymnostomoides (Broth. & Yas.) Wijk & Marg. As2
Anoectangium aestivum (Hedw.) Mitt. fide Saito, J. Hattori Bot.
Lab. 39: 457, 1975]
[Anoectangium sublaetevirens Card. As2 = Anoectangium
thomsonii Mitt. fide Saito, J. Hattori Bot. Lab. 39: 458, 1975]
[Anoectangium taeniatifolium (Herz.) Hill, J. Bryol. 11: 600, 1981
(1982) =Molendoa taeniatifo/ia Herz.]
Anoectangium tapes Besch. Oc
Anoectangium tasmanicum Broth. Austrl
[Anoectangium termale Card. As2 = Gymnostomum aeruginosum
Sm. fide Saito, J. Hattori Bot. Lab. 39: 450, 1975, as
"thermale"]
Anoectangium thomsonii Mitt. As 2 As3
[Anoectangium torquatum Broth. hom. il/eg. Afr2 Anoectangium
brotherusii Kis, Mosses South-east Trop. Afr., lnst. Ecol. Bot.
Hungarian Acad. Sci. 46, 1985]
Anoectangium walkeri Broth. As3
[Anoectangium warburgii Crundw. & Hill, J. Bryol. 9: 435, 1977
[1978] Eur Molendoa warburgii (Crundw. & Hill) Zand., see
treatment of Molendoa]
Anoectangium weisioides C. Mi.ill. Am4
Anoectangium wilmsianum (C. Mi.ill.) Par. Afr4
=
=
=
ASCHISMA Lindb.
[Aschisma aethiopicum (Welw. & dub.) Lindb. Afr2 = Byroceuthospora aethiopica (Welw. & dub.) Zand., see treatment of Byroceuthospora]
Aschisma carniolicum (Web. & Mohr) Lindb. Eur Afrl
var. speciosum Limpr. Eur
Aschisma kansanum Andrews Am 1
[Aschisma occultum Roth Am3
Uleobryum occultum (Roth)
Zand., see treatment of U/eobryum]
=
GENERA OF THE POTI1ACEAE
282
[ASTOMUM Hampe = Weissia Hedw. fide Saito, J. Hattori Bot.
Lab. 39:417, 1975]
[Astomum abbreviatum (Thwait. & Mitt.) Fleisch. As3 As4 Weissia
abbreviata (Thwait. & Mitt.) Zand., see treatment of Weissia]
[Astomum acuminatum Dix. & Ther. As2 = Weissia crispa (Hedw.)
Weissia
Mitt. fide Saito, J. Hattori Bot. Lab. 39: 418, 1975
longifolia Mitt. fide Crundwell & Nyholm, J. Bryol. 7: 13, 1972]
Astomum alternifolium Spruce hom. il/eg. Eur
[Astomum austrocrispum (Beckett) Broth. Austrl Austr2 Weissia
austrocrispa (Beckett) Stone, J. Bryol. 11: 236, 1980 Trichostomum austrocrispum (Beck.) Zand., see treatment of Trichostomum]
[var. longifolium (R. Br. ter) Dix. Austr2
Trichostomum
austrocrispum var. longifolium (R. Br. ter) Zand., see treatment
of Trichostomum]
[Astomum borbonicum Biz. & Onraedt in Biz., Rev. Bryol. Lichenol.
Astomum
40: 116, 1974 nom. inval. holoty. non cit. Afr3
borbonicum Biz. & Onraedt ex Onraedt, Bull. Jard. Bot. Nat!.
Belgique 46: 356, I976]
Astomum borbonicum Biz. & Onraedt ex Onraedt, Bull. Jard. Bot.
Nat!. Belgique 46: 356, I976 Afr3
[Astomum brisbanicum (C. Miill.) Broth. = Trachycarpidium
brisbanicum (C. Miill.) Stone]
[Astomum chilense Williams Am4 Trichostomum williamsii Zand.
nom. nov., see treatment ofTrichostomum]
[Astomum crispum (Hedw.) Hampe Eur Asi As2 As3 AsS Afri
Weissia crispa (Hedw.) Mitt. = Weissia longifolia Mitt. fide
Crundwell & Nyholm, J. Bryol. 7: I3, I972]
[var. aciculatum (Mitt.) Podp. Eur Weissia crispa var. aciculata
(Mitt.) Dix. = Weissia longifolia Mitt. fide Crundwell & Nyholm,
J. Bryol. 7: 13, I972]
[var. angustifolium Baumg. in Ginzb. Eur Weissia longifolia var.
angustifolia (Baumg. in Ginzb.) Crundw. & Nyh., J. Bryol. 7: I4,
I972]
[var. brevifolium Card. & Copp. Eur = Weissia longifolia var.
angustifolia (Baumg. in Ginzb.) Crundw. & Nyholm, J. Bryol. 7:
14, I972]
var. exsertum Nog. As3
[var. laubacense (Roth) Podp. Eur = Weissia longifolia Mitt. fide
Crundwell & Nyholm, J. Bryol. 7: I3, I972]
[var. philibertii (Husn.) Wijk & Marg. Eur Afrl = Weissia levieri
(Limpr.) Kindb.fide Crundwell & Nyholm, J. Bryol. 7: 16, 1972]
[var. sterile (Nich.) Moenk. Eur = Weissia crispa subsp. sterilis
(Nich.) Dix. = Weissia sterilis Nich.fide Crundwell & Nyholm, J.
Bryol. 7: 11, 1972]
[fo. major-planifolia Breidl. in Limpr., Laubm. Deutsch. 3: 638,
I901 Eur = Astomum crispum var. philibertii (Husn.) Wijk &
Marg. fide Podp!ra, Consp. Muse. Eur. 184, 1954 = Weissia
levieri (Limpr.) Kindb.fide Crundwell & Nyholm, J. Bryol. 7: 16,
1972]
Astomum crispum (Hedw.) Hampe x Astomum crispata (Nees. &
Hornsch.) C. Miill.fide Nich., Rev. Bryol. 32: 20, 1905 Eur
Astomum crispum (Hedw.) Hampe x Astomum microstoma Hornsch.
ex Nees & Homsch.fide Nich., Rev. Bryol. 33: 1, 1906 Eur
[Astomum cryptocarpum Broth. Am5 =Torre/la cryptocarpa (Broth.)
Zand., see treatment of Torre/la]
[Astomumfruchartii (C. Miill.) Broth. Am5 Am6 Tortel/afruchartii
(C. Miill.) Zand., see treatment of Torre/la]
Astomum japonicum Roth. As2
[Astomum latifolium Broth. in Roth Am5 = Torre/la fruchartii (C.
Miill.) Zand., see treatment of Torre/la]
[Astomum lindigii (Hampe) Jaeg. Am4 = Trichostomum lindigii
(Hampe) Zand., see treatment of Trichostomum]
=
=
=
=
=
=
=
=
=
=
=
[Astomum lonchophyllum Roth Am5 = Trachycarpidium
lonchophyllum (Roth) Zand., see treatment of Trachycarpidium]
Astomum lorentzii (C. Miill.) Broth. Am6
[Astomum ludovicianum (Sull.) Sull. Aml = Weissia ludoviciana
(Sull.) Reese & Lemmon, Bryologist 68: 282, 1965]
Astomum minutum Dix. & P. Yarde As3
[Astomum mittenii BSG Eur = Weissia mittenii (BSG) Mitt.]
Astomum mollifolium (C. Miill.) Broth. Am5
[Astomum muehlenbergianum (Sw.) Grout As2 Aml
Weissia
muehlenbergiana (Sw.) Reese & Lemmon, Bryologist 68: 282,
1965]
[Astomum multicapsulare (Sm.) BSG Eur Asl = Weissia
multicapsularis (Sm.) Mitt.]
[Astomum neocaledonicum (Ther.) Andrews in Broth. Oc Weissia
neocaledonica (Ther.) Zand., see treatment of Weissia]
Astomum nicholsonii Roth [= Weissia crispata (Nees & Hornsch.)
C. Miill. x Astomum crispum (Hedw.) Hampe] Eur
[Astomum nitidulum (C. Mi.ill.) Sull. & Lesq. = Weissia
muehlenbergianum (Sw.) Grout]
var. pygmaeum Lesq. & Jam. Aml
Astomum novae-valesiae Broth. ex Roth Eur Afrl
[Astomum occidentale Flow. ex Crum, Bryologist 76: 286, 1973
Aml
Weissia occidentalis (Flow. ex Crum) Stoneburner,
Bryologist 88: 310, 1985]
[Astomum phascoides (Hook.) Grout Aml Eur = Weissia rostellata
var. phascoides (Hook.) Reese & Lemmon, Bryologist 68: 283,
1965]
[Astomum platystegium Dix. As4
Weissia platystegia (Dix.)
Zand., see treatment of Weissia]
[Astomum unguiculatum (Mitt.) Broth. Afr2 Weissia unguiculata
(Mitt.) Crundw. & Nyholm, J. Bryol. 8: 69, 1974 = Trichostomum unguiculatum (Mitt.) Zand., see treatment of Trichostomum]
[Astomum viride C. Mi.ill. Austrl
Pleuridium viride (C. Mi.ill.)
Kindb.]
Astomum wattsii Broth. ex Roth Austrl
=
=
=
=
=
=
BARBULA Hedw.
[Barbula aaronis (Lor.) Hilp. Eur As5 Afrl
Trichostomopsis
aaronis (Lor.) Agnew & Towns., Israel J. Bot. 19: 258, 1970
Didymodon aaronis (Lor.) Guerra in Guerra & Ros,
Cryptogamic Bryol. Lichenol. 8: 55, 1987]
[Barbula abbonii Ther. Am2 = Didymodon tophaceus (Brid.) Lisa
fide Zander, Cryptogamic Bryol. Lichenol. 2: 406, 1981
(1982)]
[Barbula aciphylla BSG =Tortula ruralis var. alpina Wahlenb.]
var. mucronata Sendtn. in Gerb. Eur
Barbula acrophylla C. Miill. Austrl
[Barbula acuta (Brid.) Brid. Eur Asi As3 AsS Afrl Ami Arn2 =
Didymodon rigidulus var. gracilis (Hook. & Grev.) Zand. fide
Zander, Cryptog. Bryol. Lichenol. 2: 393, 1981 (1982)]
[subsp. abbreviatifolia (H. Muell.) Kindb. Eur Barbula acuta
var. abbreviatifolia (H. Miill.) Podp.]
[subsp. icmadophila (Schimp. ex C. Miill.) Amann Eur Asl As3
AsS Ami Am2 = Didymodon rigidulus var. icmadophilus
(Schimp. ex C. Mi.ill.) Zand., Cryptogamic Bryol. Lichenol. 2:
394, I98I]
[var. bescherellei (Sauerb. ex Jaeg. & Sauerb.) Crum, Bryologist
72: 24I, I969 Am2 = Didymodon rigidulus Hedw. s. lat. fide
Zander, Cryptogamic Bryol. Lichenol. 2: 389, I981 (1982)]
var. abbreviatifolia (H. Miill.) Podp. Eur
[var. icmadophila (Schimp. ex C. Mi.ill.) Crum, Bryologist 72:
=
=
=
283
GENERA, SPECIES AND INFRASPECIFIC TAXA
241, 1969 Am1 = Didymodon rigidulus var. icmadophilus
(Schimp. ex C. Mi.ill.) Zand. fide Zander, Cryptogamie Bryol.
Lichenol. 2: 394, 1981 ( 1982)]
fo. brevifolia (Roth) Podp., Consp. Muse. Eur. 209, 1954 (Barbula
gracilis var.) Eur
fo. ca/abrica (Roth) Podp., Consp. Muse. Eur. 209, 1954 (Barbula
gracilis var.) Eur
fo. irrigata (H. Mi.ill.) Podp., Consp. Muse. Eur. 209, 1954 (Barbula gracilis var.) Eur
fo. multiseta (Limpr.) Podp., Consp. Muse. Eur. 209, 1954 (Barbula gracilis var.) Eur
fo. patens (Glow.) Podp., Consp. Muse. Eur. 209, 1954 (Barbula
gracilis var.) Eur
fo. pu/veriplena (Loeske) Podp., Consp. Muse. Eur. 209, 1954
(Barbula icmadophila fo.)
fo. rufescens (Limpr.) Podp., Consp. Muse. Eur. 209, 1954 (Barbula gracilis var.) Eur
fo. viridis (BSG) Podp., Consp. Muse. Eur. 209, 1954 (Barbula
gracilis var.) Eur
[Barbula acutata C. Mi.ill. Afr4 = Pseudocrossidium replicatum
(Tayl.) Zand. fide Frey & Ki.irschner, Cryptogamie Bryol.
Lichenol. 9: 99, 1988 also Sollman, Lindbergia 16: 22, 1990]
[Barbula afrqduriuscu/a C. Mi.ill. Afr3 = Serpotortel/a chenagonii
(Ren. & Card.) Reese & Zand., Bryologist 90: 234, 1987 (1988)]
Barbula afrofontana (C. Mi.ill.) Broth. Afr4
var. acutiuscula P. Yarde Afr2
Barbula agraria Hedw. Am1 Am2 Am3 Am5
[fo. invo/uta Biz. & Ther. Am3 = Barbula agraria Hedw. fide
Zander, Phytologia 44: 202, 1979]
Barbula alpicola C. Mi.ill. Am6
Barbula altipapillosa Bartr. As3
[Barbula altiseta Card. Am2 = Didymodon rigidulus Hedw. s. lat.
fide Zander, Cryptogamie Bryol. Lichenol. 2: 390, 1981 (1982)]
Barbula amoena C. Mi.ill. hom. illeg. Austr1
Barbula amplexifolia (Mitt.) Jaeg. As3 As4 Am1
Barbula anastomosans C. Mi.ill. Am4 Am6
Barbula anceps Card. As2
Barbula angustifolia C. Mi.ill. & Kindb. in Macoun, Cat. Canad. Pl. 6:
264, 1892 hom. il/eg.
Barbula arcuata Griff. As3 As4 Oc
[Barbula anderssonii (Angstr.) Jaeg. Am6 Syntrichia anderssonii
(Aongstr.) Zand., see treatment of Syntrichia]
[Barbula andreaeoides Kindb. Am 1 = Didymodon subandreaeoides
fide Zander, Phytologia 41: 23, 1978]
Barbula aneitensis Broth. & Watts. Oc
[Barbula anserinocapitata X.-j. Li, Acta Bot. Yunnan. 3: 103, 1981
As2 = Didymodon anserinocapitatus (X.-j. Li) Zand., see treatment of Didymodon]
[Barbula apiculata Hedw. = Barbula unguiculata fo. apiculata
(Hedw.) Monk.]
var. minor Mart. Eur
Barbula appressifolia (Mitt.) Jaeg. Am4
Barbula arctoamericana C. Mi.ill. Am 1
Barbula arcuata Griff. As3 As4 Oc Am2 Am3 Am5
[Barbula arenicola Dus. Am6 =Pseudocrossidium crinitum (Schultz)
Zand., see treatment of Pseudocrossidium]
Didymodon
[Barbula asperifolia Mitt.e Eur As1 As2 As3 Am.l
asperifolius (Mitt.) Crum, Steere & Ander.]
[var. gorodkovii (A. Abr. & I. Abr.) L. Savicz, Novosti Sist. Niz.
Rast. 6: 148, 1969 [1970] Asl =Didymodon rufus var. gorodkovii
A. Abr. & I. Abr. in I. Abr., Trudy Arktic. Antarktic. Naucno-lssl.
Ins!. 224: 220, 1963]
var. gracilis (Amann & Meyl.) Wijk & Marg. Eur
=
=
var. grauhauptiana (De Not.) Wijk & Marg. Eur
var. kneuckeri (Loeske & Osterw.) Wijk & Marg. Eur
var. spitsbergensis (Jones) Wijk & Marg. Eur
[Barbula aurea (Bartr.) Zand. in Zand. & Steere, Bryologist 81:
466, 1978 Aml Am2 = Pseudocrossidium aureum (Bartr.)
Zand. fide Zander, Phytologia 44: 207, 1979 = Pseudocrossidium crinitum (Schultz) Zand.]
Barbula aureola C. Mi.ill. Austr2
[Barbula australasiae (Hook. & Grev.) Brid. Am4 Am6 Austr1
Austr2 Trichostomopsis australasiae (Hook. & Grev.) Robins., Phytologia 20: 187, 1970
Didymodon australasiae
(Hook. & Grev.) Zand., Phytologia 41: 21 , 1978]
Barbula austrogracilis Dus. Am6
[Barbula austrorevoluta Besch. in Britt. Am4 = Pseudocrossidium
austrorevolutum (Besch.) Zand., see treatment of Pseudocrossidium]
[Barbula barbuloides (Herz.) U. Miz., J. Jap. Bot. 46: 320. 1971 =
Didymodon erosodenticulatus (C. Mi.ill.) Saito fide Saito J.
Hattori Bot. Lab. 39: 504, 1975]
Barbula bagelensis Fleisch. As4
[Barbula barbuloides (Herz.) Mizushima, Jap. J. Bot. 46: 320, 1971
(Erythrophyllum) As2 = Didymodon erosodenticulatus (C.
Mi.ill.) Saito fide Saito, J. Hattori Bot. Lab. 39: 504, 1975]
[Barbula bescherellei Sauerb. in Jaeg. Am1 Am2 Am3 = Didymodon rigidulus Hedw. s. lat. fide Zander, Cryptogamie Bryol.
Lichenol. 2: 389, 1981 (1982)]
[var. crassinervia TMr. Am2 = Didymodon rigidulus Hedw. s.
lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 390, 1981
(1982)]
[var. stenocarpa Card. Am2 = Didymodon rigidulus Hedw. s. lat.
fide Zander, Cryptogamie Bryol. Lichenol. 2: 390, 1981
(1982)]
Barbula bicolor (BSG) Loeske Eur
Barbula bistrata Rungby, Bot. Not. 112: 81, 1959 Am4
Barbula boliviana (Broth.) Hilp. Am4
Barbula bol/eana (C. Mi.ill.) Broth. Afr2
[Barbula bourgaeana Besch. Am2 = Didymodon vinealis (Brid.)
Zand. fide Zander, Cryptogamie Bryol. Lichenol. 2: 407, 1981
(1982)]
Barbula brachymenia (Mitt.) Jaeg. Am4
[Barbula brachyphylla Sull. in Whipp!. Aml = Didymodon
brachyphyl/us (Sull. in Whipp!.) Zand., Phytologia 41: 24,
1978]
[Barbula brunneola C. Mi.ill. Am2 = Didymodon nigrescens (Mitt.)
Saito fide Zander, Phytologia 41: 22, 1978]
Barbula bulbiformis Brid. As5
[Barbula byrdii Bartr. Ant = Bryoerythrophyllum byrdii (Bartr.)
Zand., see treatment of Bryoerythrophyllum]
[Barbula calcarea Ther. Am2 = Bryoerythrophyllum calcareum
(Ther.) Zand., Bryologist 83: 232, 1980]
Barbula calcicola (Hampe) Broth. Austr1
Barbula ca/odictyon Broth. in Schum. & Lauterb. As4
Barbula ca/ycina Schwaegr. [see Catcheside, Mosses S. Austr. 185,
1980 and Magill, Fl. S. Afr. I. Mosses 1: 241, 1981 (1982)] As3
Afr4 Am6 Austrl Austr2
Barbula calyculosa (Mitt.) Jaeg. Am4 Am5
[Barbula campylocarpa (Tayl.) C. Mi.ill. Am4 = Didymodon
taylorii Zand. nom. nov., see treatment of Didymodon]
[Barbula canalicu/ata (Dix.) R. S. Chopra, Taxon. Indian Mosses
138, 1975 (Didymodon) As3 = Didymodon canaliculatus Dix.]
[Barbula cancel/ata C. Mi.ill. Aml = Barbula indica (Hook.)
Spreng. fide Zander, Phytologia 44: 1979 and, see treatment of
Barbula]
=
=
GENERA OF THE POTTIACEAE
284
Barbula capi/lipes Broth. Am5
[Barbula cardotii Dus. Am6 Didymodon cardotii (Dus.) Zand., see
treatment of Didymodon]
Barbula catenulata Dix., Anniv. Vol. Bot. Gard. Calcutta 181, 1942
As3
[Barbula chloronotos (Brid.) Brid. Crossidium chloronotos (Brid.)
Limpr. Eur As2 As3 As5 Afr1 Am1 Austr2 = Crossidium
squamiferumfide Delgadillo, Bryologist 78: 276, 1975]
[fo. nivea Besch., Cat. Moun. Alg. 12, 1882 Afr1 = Crossidium
chloronotos var. niveum (Besch.) Jelenc = Crossidium
chloronotos fo. nivea (Besch.) Par.]
Barbula chlorotricha (Broth. & Geh.) Par. Austr1
Barbula chocayensis Broth. & Herz. Am4
Barbula chrysochaete C. Miill. Austr1
Barbula chrysopus C. MUll. Austr1
Barbula c/avicostata (Ren. & Card.) Zand. (Hyophila), see treatment
of Barbula Afr3
[Barbula columbiana (Herm. & Lawt.) Herm. & Lawt., Bull. Torrey
Bryoerythrophyllum columbianum
Bot. Club 99: 309, 1972
(Herm. & Lawt.) Zand., Bryologist 81: 548, 1978]
[Barbula comosa Dozy & Molk. = Barbula arcuata Griff. fide Saito,
J. Hattori Bot. Lab. 39: 496, 1977]
[var. japonica Broth. As2 = Barbula arcuata Griff. fide Saito, J.
Hattori Bot. Lab. 39: 496, 1975]
[Barbula commutata Jur. = Streblotrichum convolutum subsp.
commutatum (Jur.) Giac.
Barbula convolutum subsp.
commutata (Jur.) Boul.]
var. erosa Corb. Afr1
[Barbula concava Dietr., Filic. Jenens. 49, 1827 nom. il/eg. incl.
Barbula cuneifolia (Dicks.) Web. & Mohr Eur
Tortula
cuneifolia (Dicks.) Tum.]
Barbula confertifolia Mitt. As3
Barbula congoana Ther. Afr2
[Barbula consanguinea (Thwait. & Mitt.) Jaeg. As3 As4 = Barbula
javanica Dozy & Molk. fide Saito, J. Hattori Bot. Lab. 39: 495,
1975]
[Barbula constricta Mitt. As2 As3 As4 = Didymodon constrictus
(Mitt.) Saito fide Saito, J. Hattori Bot. Lab. 39: 514, 1975 (=
Didymodon vinealis (Brid.) Zand. fide Sollman, Bryologist 86:
271, 1983)]
[var.flexicuspis Chen As2 Didymodon constrictus var.flexicuspis
(Chen) Saito fide Saito, J. Hattori Bot. Lab. 39: 516, 1975 (=
Didymodon vinealis (Brid.) Zand. fide Sollman, Bryologist 86:
272, 1983)]
[Barbula convolutifolia Dix. = Barbula javanica Dozy & Molk. fide
K. Saito, J. Hattori Bot. Lab. 39: 495, 1975]
Barbula convoluta Hedw. Eur As1 As2 Afr1 Am1 Am2 Austr2
subsp. commutata (Jur.) Boul. Eur As5 Afr1
[subsp. austriaca (Schiffn. & Baumg.) Podp., Consp. Muse. Eur.
206, 1954 Eur subsp. austriacus (Schiffn. & Baumg.) Wijk &
Marg. Eur]
var. gallinula Zand., Phytologia 44: 195, 1979 Am1
var. obtusata C. Miill. & Kindb. In Macoun Am1
[var. propagu/ifera Glow. = Gymnostomum aeruginosum Sm. fide
R. Zander, Phytologia 44: 211, 1979]
[var. robusta Schimp. in Husn., Musci Gall. 11: 513A, 1875 =
Barbula revolvens Schimp.fide Boul., Rev. Bryol. 2(2): 20, 1875
=Tortula revolvens (Schimp.) Roth]
var. sardoa BSG Eur
fo. brevifolia Podp., Cas. Morav. Mus. Zemsk. 13: 52, 1913 Eur
[fo. brunnescens Podp., Cas. Morav. Mus. Zemsk. 13: 52, 1913 Eur
=Barbula convoluta fo. rufescens Loeske & Quelle]
fo. insolata Latz., Bot. Centralbl. Beih. 48(2): 480, 1931 Eur
=
=
=
=
=
=
=
fo. rufescens Loeske & Quelle, Moosfl. Harz. 175, 1903 Eur
fo. rufipes Bauer, Muse. Eur. Exs. 1586a, 1923 Eur
fo. uliginosa Limpr. in Cohn, Krypt. Fl. Schles. 1: 172, 1876 Eur
[Barbula cordata (Jur.) Loeske =Didymodon cordatus Jur.]
fo. brevicaulis (Roll) Podp., Consp. Muse. Eur. 205, 1954 nom.
il/eg. (Didymodon cordatus fo.) Eur Didymodon cordatus var.
brevicaulis Roll]
fo. gracilis (Roll) Podp., Consp. Muse. Eur. 205, 1954 (Didymodon cordatus var.) Eur Didymodon cordatus var. gracilis]
[fo. longicaulis (Roll) Podp., Consp. Muse. Eur. 206, 1954 (Didymodon cordatus fo.) Eur = Didymodon cordatus fo. /ongicaulis
Roll]
[fo. ramosa (Roll) Podp., Consp. Muse. Eur. 206 (Didymodon
cordatus fo.) Didymodon cordatus fo. ramosus Roll]
[fo. robusta (Roll) Podp., Consp. Muse. Eur. 206, 1954 (Didymodon cordatus fo.) Didymodon cordatus fo. robustus Roll]
[fo. stricta (Roll) Podp., Consp. Muse. Eur. 206, 1954 (Didymodon cordatus fo.) =Didymodon cordatus fo. strictus Roll]
[fo. typica (Roll) Podp., Consp. Muse. Eur. 205, 1954 nom. illeg.
(Didymodon cordatus fo.) Eur Didymodon cordatus Jur. fo.
cordatus
Barbula coreensis (Card.) Saito, J. Hattori Bot. Lab. 39: 484, 1975
(Barbula paludosa var.) As2
Barbula costa-ricensis Ren. & Card. Am2
Barbula costata (Mitt.) Jaeg. Am4
Barbula costesii Ther. Am6
[Barbula crassicostata Bartr. Am2
Didymodon crassicostatus
(Bartr.) Zand., see treatment of Didymodon]
[Barbula crassicuspis Robins. Am2 = Morinia crassicuspis (Robins.) Zand. fide Zander, Bryologist 81 : 556, 1978 = Mironia
crassicuspis (Robins.) Zand., see treatment of Mironia]
[Barbula crinita Schultz (good species fide Catcheside, Mosses S.
Austr. 180, 1980 and Magill, Fl. S. Afr. I. Mosses 1: 237, 1981
[1982]) Afr2 Afr4 Am6 Austrl Austr2
Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
Barbula crocea (Brid.) Web. & Mohr Eur As2 Afr1
var. funckiana (Schultz) Margadant, Lindbergia 1: 124, 1972
(Barbula) Eur
Barbula crozalsii (Philib.) Broth. Eur
[Barbula cruegeri Sond. ex C. Miill. Aml Am2 Am3 Am4 Am5 =
Barbula indica (Hook.) Spreng. fide Zander, Phytologia 44:
185, 1979]
var. laevinervis Broth. Am3
Barbula cucullata J. Frolich, Ann. Naturhist. Mus. Wien 67: 153,
1964 As3
[Barbula cucullifera (Mitt.) Jaeg. Am4 = Aloinella cucullifera
(Mitt.) Steere]
[Barbula curvipes C. MUll. Am6 Trichostomopsis curvipes (C.
MUll.) Robins., Phytologia 20: 186, 1970]
[Barbula curvirostris Lindh.
Hymenostylium recurvirostre
(Hedw.) Dix.]
[fo. commutata (Mitt.) Lindh., Musci Scand. 22, 1879 Eur =
Hymenostylium recurvirostre var. commutatum (Mitt.) Podp.]
[fo. laeviuscu/a Lindh., Musci Scand. 22, 1879 Eur = Hymenostylium recurvirostre fo. breviusculum (Lindh.) Podp.]
[fo. scabra Lindh., Musci Scand. 22, 1879 Eur Asl Aml
Hymenostylium recurvirostrum var. scabrum (Lindh.) Podp =
Hymenostylium recuvirostrum var. latifolium (Zett.) Wijk &
Marg.]
Barbula cylindrangia C. MUll. Austrl
[Barbula cylindrica (Tayl.) Schimp. in Boul. Eur As1 As5 Afr1
Aml
Didymodon vinealis var. jlaccidus (BSG) Zand.,
Phytologia 41: 26, 1978 = Didymodon vinealis (Brid.) Zand.
=
=
=
=
=
=
=
=
=
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
var. vinealisfide Sollman, Bryologist 86: 272, 1983]
[fo. rivularis Warnst., Krypt. Fl. Brandenb. 2(2): 253, 1904 =Barbuta vinealis fo. rivularis (Loeske) Podp., Consp. Muse. Eur. 210,
1954 (Barbula) Eur]
[fo. robusta Loeske, Moosfl. Harz. 174, 1903 Eur = Barbula
vinealis fo. robusta (Loeske) Podp., Consp. Muse. Eur. 210,
1954]
[fo. rubella (Schiffn.) Herz., Krypt. Forsch. 4: 279, 1919 (Barbula
cylindrica var.) Eur = Barbula vinealis fo. rubella (Schiffn.)
Podp., Consp. Muse. Eur. 210, 1954]
[fo. rufescens C. Jens., Bot. Faeroes 155, 1901 Eur = Barbula
vinealis fo. rubella (Schiffn.) Podp. fide Podpl!ra, Consp. Muse.
Eur. 210, 1954]
fo. viridis Warnst., Krypt. Fl. Brandenb. 2(2): 253, 1904 Eur
Barbula declivium C. Miill. Am6
[Barbula decolorans Hampe Am4 = Trichostomopsis australasiae
(Hook. & Grev.) Robins., Phytologia 20: 187, 1970 =Didymodon
australasiae (Hook. & Grev.) Zand., Phytologia 41: 21, 1978]
Barbula decurrens Laz. As1
[Barbula denticulata Dix. & P. Yarde As3 hom. illeg. = Barbula
vardei R.S. Chopra nom. nov., Bryologist 80: 544, 1977]
Barbula dharvarensis Dix. As3
Barbula dioritica C. Miill. Afr2
Barbula dissita C. Mi.ill. Oc
[Barbula ditrichoides Broth. As3 Am1 = Didymodon acutus var.
ditrichoides (Broth.) Zand., Phytologia 41: 20, 1978 (Barbula
ditrichoides) = Didymodon rigidulus var. ditrichoides (Broth.)
Zand., see treatment of Didymodon]
[Barbula divergens Broth. in Hall. As4 '= Barbula inaequalifolia
Tayl. fide Sollman, Lindbergia 10: 54, 1984 = Bryoerythrophyllum inaequa/ifolium (Tayl.) Zand.]
[Barbula dixonii R. S. Chopra, Taxon. Indian Mosses 139, 1975
(Didymodon obtusifolius Card. ex Dix. & P. Yarde hom. illeg.
non Schkurh) As3 = Barbula inaequalifolia Tayl. fide Sollman,
Lindbergia 10: 54, 1984 = Bryoerythrophyllum inaequa/ifolium
(Tayl.) Zand.]
Barbula dorrii Ren. & Card. Afr3
[Barbula dregeana C. Miill. Afr4 =Alvina bifrons (C. Mi.ill.) Kindb.
fide Magill, Fl. S. Afr. I. Mosses 1: 193, 1981 (1982)]
Barbula dusenii C. Mi.ill. ex Broth. Afr2
[Barbula ecuadoriensis Broth. Am4 = Trichostomopsis australasiae
(Hook. & Grev.) Robins., Phytologia 20: 187, 1970 = Didymodon
australasiae (Hook. & Grev.) Zand., Phytologia 41 : 21, 1978]
Barbula ehrenbergii (Lor.) Fleisch. Eur As2 AsS Afrl Am1 Am2
Am3 Austr1
var. algeriae (C. Mi.ill.) Vent. & Bott. Eur Afrl
[var. mexicana Ther. Am2 = Barbula ehrenbergii (Lor.) Fleisch.
fide Zander, Phytologia 44: 198, 1979]
Barbula e/ata Our. & Mont. ex C. Mi.ill. Afr1
[Barbula elbertii Broth. in Hall. As4 = Didymodon vinealis (Brid.)
Zand.fide Sollman, Lindbergia 10: 54, 1984]
Barbula elliottii Broth. Afr2
Barbula enderesii Garov. Eur As1
[Barbula erosa Hampe in C. Miill. Am2 Am4 = Barbula indica
(Hook.) Spreng. fide Zander, Phytologia 44: 185, 1979]
[Barbula erythropoda Schimp. ex Besch. Am2 = Didymodon
rigidulus Hedw. s. /at. fide Zander, Cryptogamie Bryol. Lichenol.
2: 389, 1981 (1982)]
Barbula eubryum C. Mi.ill. (good species fide Magill, Fl. S. Afr. I.
Mosses 1: 245, 1981 [I982]) Afr2 Afr4
Barbula eustegia Card. & Ther. Ami
[Barbula falcifolia C. Mi.ill. = Didymodon rigidicaulis (C. Mi.ill.)
Saito fide K. Saito, J. Hattori Bot. Lab. 39: 502, I975 = Didymo-
285
don ferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11: 599,
I981 (1982)]
[Barbulafallax Hedw. Eur As I As2 As3 AsS Afri Ami= Didymodonfallax (Hedw.) Zand., Phytologia 4I: 28, I978]
subsp. brevifolia (With.) Kindb. Eur Afrl Ami
var. atroviridis Hi.ib. Eur
[var. brevifolia (Dicks. ex With.) Schultz Eur Afl Ami = Didymodonfallax var. brevifolius (Dicks. ex With.) Ochyra, Fragm.
Fl. Geobot. 28: 449, I982 (I984)]
var.filescens Roll. Eur
var. laevifolia Hess. Eur
var. longifolia Wamst. & Fleisch. Eur
var. obtusifolia Amann Eur
[var. recurvifolia (Wils.) Husn. = Didymodon rigidicaulis (C.
Mi.ill.) Saito fide K. Saito, J. Hattori Bot. Lab. 39: 502, I975 =
Didymodonferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11:
599, I98I (1982)]
var. stricta Schultz Eur AsS
var. tristicha Brid. Ami
var. vinealoides March. Eur
fo. a/pina Loeske & Paul, Krypt. Forschung. Bayer. Bot. Ges. 5:
354, I920 Eur
fo. atrata Roll, Jahrb. Ak. Wiss. Erfurt n.ser. 41: I23, I9I5 Eur
fo. biseta Gyorf., Magyar Bot. Lapok 5: 340, I906 Eur
fo. brevicaulis (Schwaegr.) Podp., Consp. Muse. Eur. 206, I954
(Barbula) Eur Ami
fo. crispula (Warns!.) Podp., Consp. Muse. Eur. 206, I954 (Barbulafallax var.) Eur
fo. e/ata Loeske in Bauer, Muse. Eur. Exs. I590, I923 Eur
fo. fastigiata (Warnst.) Podp., Consp. Muse. Eur. 206, I954
(Barbulafallax var.) Eur
fo. robusta Podp., Consp. Muse. Eur. 207, I954 (Barbula fallax
var. robusta Warnst. hom. il/eg.) Eur
Barbula farriae Crum & Bartr. Am3
Barbulafendleri C. Miill. Am4
[Barbula ferrinervis C. Mi.ill. =Barbula arcuata Griff. fide Zander,
Phytologia 44: 199, I975]
[var. eggersiana C. Miill. Am3 = Barbula arcuata Griff. fide
Zander, Phytologia 44: 199, I975]
Barbulafide/is Crum & Steere Am3
[Barbula flaccidiseta Lor. Am2 = Didymodon rigidulus Hedw. s.
lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 389, I98I
(1982)]
[Barbula flaviseta (Herz.) Wijk & Marg. Am6 = Trichostomopsis
umbrosa (C. Mi.ill.) Robins., Phytologia 20: 185, I970 (= Didymodon australasiae (C. Mi.ill.) Zand., Cryptogamie Bryol.
Lichenol. 2: 400, I981 [I982]) = Didymodon umbrosus (C.
Mi.ill.) Zand.]
[Barbula fontana (C. Mi.ill.) Broth. hom. illeg. Afr2 = Barbula
meidensis Cufodontis, nom. nov.]
Barbula francii Ther. Oc
Barbulafrigida C. Mi.ill. Am4
Barbulafunalis Dix. & Badhw. As3
Barbulafurvofusca C. Mi.ill. Oc
[Barbula fusca C. Mi.ill. Am4 Am6 = Didymodon vinealis (Brid.)
Zand. fide Sollman, Bryologist 86: 271, I983; Dusen collections of this sp. at NY are also Didymodon vinealis]
Barbulafuscescens Wallich As2 As3
[Barbula fuscinervia (Mitt.) Jaeg. Am6 = Bryoerythrophyllum
fu scinervium (Mitt.) Zand., see treatment of Byroerythrophyllum]
Barbula fuscovirens Bartr. As3
Barbulafuscoviridis Broth. ex Ther. Am6
GENERA OF THE POTTIACEAE
286
[Barbula gangetica C. MUll. As3 = Barbula arcuata Griff. fide
Gangulee, Mosses E. India 3: 725, 1972]
Barbula gattefossei P. Yarde Afr1
Barbula geminata C. MUll. Austr1
Barbula geniculata (Mont.) C. MUll. Am6
Barbula glaucescens Hampe Am4
var./atifolia Herz. Am4
Barbula g/aucula C. MUll. Oc
Barbula goniospora C. MUll. Oc
Barbula gracilenta Mitt. As3
[Barbula gracilescens Schimp. ex Besch. Am2 = Didymodon
rigidulus Hedw. s. lat. fide Zander, Cryptogamie Bryol. Lichenol.
2: 389, 1981 (1982)]
[Barbula graciliformis Schimp. ex Besch. Am2 = Didymodon
rigidulus Hedw. s.lat.fide Zander, Cryptogamie Bryol. Lichenol.
2: 389, 1981 (1982)]
[Barbula graminicolor C. MUll. Am2 Am6 = Didymodon
australasiae (Hook. & Grev.) Zand. s. lat. fide Zander,
Cryptogamie Bryol. Lichenol. 2: 397, 1981 (1982)]
subsp. subgraminicolor Ther. Am6
Barbula granulosa Ther. Am4
[Barbula gregaria (Mitt.) Jaeg. As2 As3 = Barbula indica var.
gregaria (Mitt.) Zand., Cryptogamie Bryol. Lichenol. 2: 6, 1981]
Barbula grimmiacea C. MUll. Am6
Barbula gymnostoma C. MUll. As3
Barbula hampeana Par. Austr1
[Barbula haringae Crum Am1 = Barbula amplexifolia (Mitt.) Jaeg.
fide Zander, Phytologia 44: 193, 1979]
[Barbula hastata Mitt. As3 Didymodon hastatus (Mitt.) Zand., see
treatment of Didymodon]
Barbula hiroshii Saito, J. Hattori Bot. Lab. 39: 499, 1975 As2
Barbula hispaniolensis Buck & Steere, Moscosoa 2(1): 30, 1983
(nom. nov. for Barbula /eptodontioides Crum & Steere) Am3
[Barbula hookeri Steud. Afr4 = Barbula calycina Schwaegr. fide
Magill, Fl. S. Afr.l. Mosses 1: 241, 1981 (1982)]
[Barbula hornschuchiana Schultz Eur As5 Afr1 Afr4 Am1 =Pseudocrossidium hornschuchianum (Schultz) Zand. fide Zander,
Phytologia 44: 205, 1979]
var. incrassata Podp. Eur
[var. obtusula (Lindh.) Podp. Eur Am1
Pseudocrossidium
revolutum var. obtusulum (Lindh.) Tan, Zand. & T. Tayl.,
Lindbergia 7: 41, 1981]
fo. brevifolia Reim., Hedwigia 79: 268, 1940 Eur
[Barbula horrinervis Saito, J. Hattori Bot. Lab. 39: 486, 1975 As5 =
Barbula indica (Hook.) Spreng. fide Zander, Phytologia 44: 186,
1979]
Barbula hosseusii Ther. Am4
Barbula horricomis C. MUll. ex Gangulee, Nov. Hedw. 12: 423, 1966
[1967] As2
Barbula humboldtii Herz. Am4 = Didymodon humboldtii (Herz.)
Hegew. & Hegew.]
[Barbula husnotii Schimp. ex Besch. Am3 = Barbula agraria Hedw.
fide Zander, Phytologia 44: 202, 1979]
Barbula hyalinobasis Broth. Am4
Barbula hymenostylioides Broth. in Urban Am3
[Barbula icmadophila Schimp. ex C. MUll. Eur As2 As3 As5 Am1
Am2 = Didymodon rigidulus var. icmadophilus (Schimp. ex C.
MUll.) Zand. fide Zander, Cryptogamie Bryol. Lichenol. 2: 394,
1981 (1982)]
[fo. pu/veriplena Loeske, Hedwigia 49: 29, 1910 Eur Barbula
acuta fo. pulveriplena (Loeske) Podp., Consp. Muse. Eur. 209,
1954]
[Barbula imbricata Hermstadt & Heyn, Bryologist 94: 174, 1991
=
=
=
(nom . nov. for Didymodon luridus Hornsch. ex Spreng. Eur As5
Afr1 Afr2 Didymodon luridus Hornsch. ex Spreng.]
[Barbula imperfecta (C. MUll.) Broth. Am6 = Didymodon
imperfectum (C. MUll.) Zand., see treatment of Didymodon]
[Barbula imshaugii Yitt, Bryologist 74: 464, 1971 (1972) Austr2 =
Trichostomum imshaugii (Yitt) Zand., see treatment of Trichostomum]
[Barbula inaequalifolia Tayl. Am1 Am2 Am4 As2 As4 = Bryoerythrophyllum inaequalifolium (Tayl.) Zand., Bryologist 83: 232,
1980]
Barbula incerta Dix. hom. illeg. Austr1
Barbula inclinans Schimp. ex Besch. Afr3
[Barbula inclinata (Hedw. f.) Schwaegr. - Tortel/a inclinata
(Hedw. f.) Limpr.]
var. magel/anica Card. Am6
[fo. acuminata Farn., Muschi Prov. Pavia, Atti lstituto Bot. Univ.
Pavia 3: 17, 1891 = Tortel/a inclinata fo. acuminata (Farn.)
Par.]
Barbula indica (Hook.) Spreng. As2 As3 As4 Afr2 Afr3 Afr4 Am1
Am2 Am3 Am4 Austr 1
var. gregaria (Mitt.) Zand., Cryptogamie Bryol. Lichenol. 2: 6,
1981 (Barbula) As3 Am1 Am2
var. scaberrima Dix. Afr2
Barbula inflexa (Duby) C. MUll. As2 As3 As4 Oc
Barbula integrifolia (Williams) Zand., Bryologist 75: 277, 1972
(Leptodontium) Am4
Barbula isoindica Zand. (nom. nov. for Portia papillinervis Lor.),
see treatment of Barbula Afr2
[Barbula jacksharpii Crum, Bryologist 87: 204, 1984 Am3 =Gymnostomumjacksharpii (Crum) Allen, Bryologist 93 : 207, 1991]
Barbula javanica Dozy & Molk. As2 As3 As4
var. epapillosa Fleisch. As4
var. robusta Dix. As3
[Barbula johansenii Williams Am1 = Didymodon johansenii (Williams) Crum]
Barbulajuniperoidea C. MUll. Am6
Barbula kiaerii Broth. Afr3
Barbula kivuensis Leroy & Potier de Ia Yarde. in Demar. & Leroy,
Expl. Pare Nation. Albert 4 Mission Lebrun 6: 15, 1944 Afr2
[Barbula laevifolia Broth. & Yas. =Barbula subcomosa Broth. fide
K. Saito, J. Hattori Bot. Lab. 39: 493. 1975]
[Barbula /aevipila (Brid.) Garov. = Tortula laevipila (Brid.)
Schwaegr.]
var. populina Brockm. Eur.
fo. pagorum (Milde) Mol., Ber. Naturhis. Yer. Passau 10: 100,
1875 Eur
[Barbula lagunicola C. MUll. Am2 = Didymodon rigidulus Hedw.
s. lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 389, 1981
(1982)]
Barbula lambarenensis P. Yarde Afr2 [:: Semibarbula
lamharenensis (P. Yarde) Biz., Sv. Bot. Tidsk. 63: 453, 1969]
Barbula lamprocalyx C. MUll. Am6
Barbula lanceo/ata Schum hom. il/eg. Eur
Barbula /aureriana Lor. Afr4
Barbula laxiretis Broth. in Hall. As4
Barbula leiophylla Tix., Rev. Bryol. Lichenol. 34: 136, 1966 As3
[Barbula /eptocarpa Besch. Am2 = Didymodon rigidulus Hedw. s.
lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 389, 1981
(1982)]
[Barbula /eptodontoides Crum & Steere hom. illeg. Am3 Barbula
hispaniolensis Buck & Steere nom. nov.]
Barbula leucobasis Dix. in Dix. & Greenwood, Proc. Linn. Soc. N.
South Wales 55: 277, 1930 Oc
=
=
287
GENERA, SPECIES AND INFRASPECIFIC TAXA
Barbula /eucodontoides C. Miill. in Par. ex Gangu1ee, Nov. Hedw.
12: 425, 1966 As3
[Barbula ligularis (Mitt.) Jaeg. Am4 = Bryoerythrophyllum ligulare
(Mitt.) Zand. see treatment of Bryoerythrophyllum]
[Barbula lindigii Hampe = Didymodon lindigii (Hampe) Zand., see
treatment of Didymodon Am4
[Barbula linguaefolia Bartr. Am2 = Barbula calcarea lner. fide
Zander, Phyto1ogia 44: 183, 1979 = Bryoerythrophyllum
calcareum (Ther.) Zand., Bryo1ogist 83: 232, 1980]
[Barbula linearis Web. & Mohr Am3 Am5 = Torre/la linearis (Web.
& Mohr.) Zand., see treatment of Torre/la]
Barbula linguaecuspis Broth. Am4
Barbula lobayetensis Williams As4
[Barbula lombokensis Broth. in Hall. As4 = Bryoerythrophyllum
ferruginascens (Stirt.) Giac.fide Sollman in Touw, J. Hattori Bot.
Lab. 71: 342, 1992]
Barbula /onchodonta C. Miill. Am6
[Barbula lonchostega C. Miill. Am2 = Barbula indica (Hook.)
Spreng. fide Zander, Cryptogamie Bryol. Lichenol. 2: 418, 1981
(1982)]
Barbula longicostata X.-j. Li, Acta Bot. Yunnan. 3: 105, 1981 As2
[Barbula longirostris Hampe Am2 Am4 = Tortula leiostoma Herz. =
Hennediella limbata (Mitt.) Zand., see treatment of Hennediella
and cf. Mishler in Sharp et al., Moss Fl. Mex.]
[Barbula /ouisiadum Broth. Oc = Barbula consanguinea (Thw. &
Mitt.) Jaeg. fide Eddy, Handb. Males. Mosses 2: 178, 1991 =
Barbula javanica Dozy & Mo1k. fide Saito, J. Hattori Bot. Lab.
39: 495, 1975]
[Barbula lozanoi Card. Am2 = Didymodon vinealis (Brid.) Zand.fide
Zander, Cryptogamie Bryol. Lichenol. 2: 408, 1981 (1982)]
[Barbula luehmannii Broth. & Geh. Austr1 =Didymodon /uehmannii
(Broth. & Geh.) Catcheside, Mosses S. Austr. 175, 1980]
Barbu/a lurida Homsch. Am2 Am4 Am5
[Barbula lurida (Homsch.) Lindh. hom. illeg. Eur As5 Afr1 Afr2
Am1 =Didymodon luridus Homsch. ex Spreng.]
[fo. cuspidata (Schimp.) Monk., Laubm. Eur. 294, 1927 (Didymodon luridus var.) = Didymodon trifarius var. cuspidata (Schimp.)
Wijk & Marg.]
Barbula luteo/a (Mitt.) Par. (good species fide Catcheside, Mosses S.
Austr. 182, 1980) Austr1
[Barbula mamillosa Crundw., J. Bryol. 9: 163, 1976 [1977] Didymodon mamillosus (Crundw.) Hill]
[Barbula macrogonia Besch. Am3 = Barbula arcuata Griff. fide
Zander, Phyto1ogia 44: 199, 1979]
Barbula macassarensis Fleisch. As3 As4
Barbula majuscu/a C. Miill. As2
Barbula malagana Crum, Bryo1ogist 70: 235, 1967 Am4
Barbu/a marginans C. Mii11. Eur
Barbula marginatula C. Miill. ex Gangu1ee, Nov. Hedw. 12: 424,
1966 [ 1967] As3
[Barbula maschalogena Ren. & Card. As3
Didymodon
maschalogena (Ren. & Card.) Broth.]
[Barbula maxima Syed & Crundw., J. Bryol. 7: 527. 1973 [1974]
nov. nov. for Barbula reflexa var. robusta Braithw. Didymodon
maximus (Syed & Crundw.) M. Hill]
Barbula meidensis Cufodontis, Oesterr. Bot. Z. 98: 225, 1951 (nom.
nov. for Barbulafontana (C. Mii11.) Broth.) Afr2
[Barbula mendozensis (Mitt.) Jaeg. Am6
Pseudocrossidium
mendozense (Mitt.) Zand., see treatment of Pseudocrossidium]
[Barbula michiganensis Steere in Grout Ami. Am2, As2 = Didymodon michiganensis (Steere in Grout) Saito, J. Hattori Bot. Lab.
39: 517. 1975]
Barbula microcalycina Magill, Fl. S. Afr. I. Mosses 1: 241, 1981
=
=
=
=
[1982] Afr4
[Barbula microglottis C. Miill. Am3 = Barbula indica (Hook.)
Spreng. fide Zander, Phyto1ogia 44: 186, 1979]
Barbula microstoma (Dix. & Badhw.) R. S. Chopra, Bryo1ogist 80:
544, 1977 (Didymodon) As3
[Barbula mobilis C. MUll. Am4 = Didymodon rigidulus Hedw. s.
lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 389, 1981
(1982)]
[Barbula mosis (Lor.) Hilp. As5 =Gymnostomum mosis (Lor.) Jur.]
Barbula mucronulata Ren. & Card. hom. illeg. Afr3
Barbula mucronulata Lilj. hom. illeg. Eur
[Barbula muenchii Card. Am2 = Barbula indica (Hook.) Spreng.
fide Zander, Phyto1ogia 44: 186, 1979]
Barbula munyensis Williams Am4
[Barbula muralis (Hedw.) Crom. Tortula muralis Hedw.]
var. breviseta Opiz Eur
var. microcarpa HUb. Eur
var. tenuis Opiz Eur
[fo. stenocarpa Baker, Pacif. Slope Bryoph. 360, 1902 = Tortu/a
brevipes (Lesq.) Broth. fide Steere in Grout, Moss Fl. N. Am.
1(4): 232, 1939]
[Barbula mutica (Schultz) Kindb., Eur. N. Am. Bryin. 2: 247, 1897
hom. illeg. (Syntrichia laevipila var.) Am1 = Tortula latifolia
Hartm.]
Barbula nana C. MUll. Afr2
[Barbula nigrescens Mitt. As2 As3 Am1 Am2
Didymodon
nigrescens (Mitt.) Saito fide Saito, J. Hattori Bot. Lab. 39: 510,
1975]
[Barbula nipponica Nog. As2 = Didymodon constrictus (Mitt.)
Saito fide Saito, J. Hattori Bot. Lab. 39: 514, 1975 (= Didymodon vinealis (Brid.) Zand. fide Sollman, Bryologist 86: 27i,
1983)]
[var. gracilis Nog. As2 = Didymodon constrictus (Mitt.) Saito
fide Saito, J. Hattori Bot. Lab. 39: 514, 1975 (= Didymodon
vinealis (Brid.) Zand.fide Sollman, Bryo1ogist 86: 271, 1983)]
Barbula novae-caledoniae C. MUll. Oc
Barbula novoguinensis Broth. As4
Barbula novogranatensis Hampe Am4
var. gracilior Hampe Am4
[Barbula obscuriretis Dix. As3 As4 Oc = Barbula consanguinea
(Thw. & Mitt.) Jaeg.fide Eddy, Handb. Males. Mosses 2: 178,
1991 = Barbula javanica Dozy & Molk. fide Saito, J. Hattori
Bot. Lab. 39: 495, 1975]
Barbula obtusissima Broth. & Par. hom. illeg. Oc
Barbula occidenralis (Mitt.) Broth. Am4
[Barbula ochrocarpa Toyama As2 = Barbula indica (Hook.)
Spreng. fide Saito, J. Hattori Bot. Lab. 39: 488, 1975]
[Barbula olivacea (mitt.) Besch. Am2 =Didymodon vinealis (Brid.)
Zand.fide Zander, Cryptogamie Bryol. Lichenol. 2: 407, 1981
(1982)]
Barbula omissa Ther. Afr3
[Barbula orienta/is (Web.) Broth. = Barbula indica (Hook.)
Spreng. fide Saito, J. Hattori Bot. Lab. 39: 488, 1975]
var. scaberrima Dix.
Barbula orizabensis C. Miill. Am1 Am2 Am3
[Barbula pachydictyon Broth. in Hall. As4 = Didymodon vinealis
(Brid.) Zand.fide Sollman, Lindbergia 10: 54, 1984]
Barbula pachyloma Broth. l= Cinclidotus involutus Hi1p.] As4
Barbula pallidobasis Dix. As1
[Barbula papillinervis (Lor.) Broth. hom. illeg. Afr2 = Barbula
isoindica Zand. nom. nov., see treatment of Barbula]
[Barbula perlinearis C. Mii11. = Pseudocrossidium replicatum
(Tayl.) Zand.fide Zander, Phyto1ogia 44: 206, 1975]
=
=
288
GENERA OF THE POTTIACEAE
Barbula pernana C. Miill. Am6
[Barbula perobtusa (Broth.) Chen As2 = Didymodon perobtusus
Broth. fide Zander, Phytologia 41: 23, 1978]
[Barbula perrevoluta C. Miill. Am6 = Pseudocrossidium perrevolutum (C. Miill.) Zand., see treatment of Pseudocrossidium]
[var. acutifolia (C. Miill.) Par. Am6 = Pseudocrossidium perrevolutum var. acutifolium (C. Miill.) Zand., see treatment of Pseudocrossidium]
[var. linearifolia (C. Miill.) Par. Am6 = Pseudocrossidium perrevolutum var. linearifolium (C. Miill.) Zand., see treatment of
Pseudocrossidium]
Barbula pertorquescens Broth. Am6
Barbula peruviana (Mitt.) Jaeg. Am4
Barbula pflanzii (Broth.) Herz. Am4
var.falcatula Broth. in Herz. Am4
[Barbula pilifera (Hook.) Sim. As4 Afr2 Afr4 Am6 =Barbula crinita
Schultz fide Catcheside, Mosses S. Austr. 180, 1980 and Magill,
Fl. S. Afr. I. Mosses 1: 237, 1981 [1982] = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
var. gracilis Homsch. Afr4 As4
fo. elata Ther., Rev. Bryol. Lichenol. 7: 173, 1934 [1935] nom.
inval. descr. gall. Am4
[Barbula planifolia Broth. & Yas. As2 = Trichostomum
brachydontium Bruch fide Saito, J. Hattori Bot. Lab. 39: 431,
1975]
[Barbula platydictyon Broth. in Hall., Meded. Rijks Herb. Leiden 14,
Elbert's Sunda Exp. 19, 1912 As4 = Didymodon vinealis (Brid.)
Zand.fide Sollman, Lindbergia 10: 54, 1990]
[Barbula platyneura C. Miill. & Kindb. ex Macoun & Kindb. Am1 (=
Pseudocrossidium revolutum (Brid. in Schrad.) Zand. sensu
Zander, Phytologia 44: 204, 1975) = Pseudocrossidium
revolutum var. obtusulum fide Tan, Zand. & T. Tayl., Lindbergia
7: 41, 1981]
Barbula plebeja C. Miill. Am6
[Barbula porphyreoneura C. Miill. Afr2 Afr4 = Tortula
porphyreoneura (C. Miill.) Townsend, J. Bryol. 10: 576, 1979]
Barbula potaninii Broth. ex C. Miill. As2
[Barbula prionophylla Saito, J. Jap. Bot. 46: 142, 1971 = Barbula
barbuloides fide Iwatski & Nog. 1973 = Didymodon
erosodenticulatus (C. Miill.) Saito fide Saito, J. Hattori Bot. Lab.
39: 504, 1975]
Barbula prschewalzkii Broth. in C. Miill. As2
[Barbula pruinosa (Mitt.) Jaeg. Am4 = Didymodon pruinosus (Mitt.)
Zand., see treatment of Didymodon]
Barbula pseudoehrenbergii Fleisch. As3 As4 Afr2
var. aspera Baumg. & Frohl. As4
[Barbula pseudogracilis C. Miill. Afr2 = Barbula acuta (Brid.) Brid.
fide Townsend, Lindbergia 10: 177, 1985 = Didymodon rigidulus
var. gracilis (Schleich. ex Hook. & Grev.) Zand., Cryptogamie
Bryol. Lichenol. 2: 393, 1981]
Barbula pseudonigrescens Tix., Ann. Hist.-Nat. Mus. Nat!. Hungarici
66: 88, 1974 As3
[Barbula pseudopilifera C. Miill. & Hampe Austr1 Austr2 = Barbula
pilifera (Hook.) Brid. (hom. illeg.) fide _Weber, Lindbergia 1:
214-216, 1972 =Barbula crinita Schultz fide Catcheside, Mosses
S. Austr. 180, 1980 = Pseudocrossidium crinitum (Schultz)
Zand., see treatment of Pseudocrossidium]
[var. obscura (Dix.) Sainsb. Austr2 = Barbula pilifera (Hook.)
Brid. (hom. il/eg.) fide Weber, Lindbergia 1: 214-216, 1972 =
Barbula crinita Schultz fide Catcheside, Mosses . S. Austr. 180,
1980 = Pseudocrossidium crinitum (Schultz) Zand., see treatment
of Pseudocrossidium]
Barbula punae Herz. Am4
Barbula punctulata (Ren. & Par.) Broth. hom. il/eg. Afr3
[Barbu/a pungens (Hook. f. & Wils.) Jaeg. Austr1 = Barbula
luteola (Mitt.) Par. fide Catcheside, Mosses S. Austr. 182,
1980]
Barbula purpurascens Dus. Am6
[Barbu/a purpuripes C. Miill. Am3 = Barbula indica (Hook.)
Spreng. fide Zander, Phytologia 44: 186, 1979]
Barbula pycnophylla Card. Am6
Barbula pygmaea C. Miill. Afr2
Barbula rechingeri Broth. Oc
[Barbula rectifolia Tayl. Am2 Am4 = Didymodon vinealis (Brid.)
Zand. fide Zander, Cryptogamie Bryol. Lichenol. 2: 407, 1981
(1982)]
[Barbula recurva (Griff.) R. S. Chopra, Bryologist 80: 544, 1977
(Gymnostomum) inval. basion. non cit. Bryoerythrophyllum
recurvum (Griff.) Saito, Bull. Univ. Mus. Univ. Tokyo 8: 254,
1975 = Bellibarbula recurva (Griff.) Zand., see treatment of
Bellibarbula]
Barbula recurvopungens C. Miill. Am4
[Barbula recurvirostris (Hedw.) Dix. = Bryoerythrophyllum
recurvirostrum (Hedw.) Chen]
[var. latinervia Holmen Am1 = Bryoerythrophyl/um recurvirostrum var. latinervium (Holmen) Murray, Bryobrothera 1: 14,
1992.]
[Barbula recurvis (Griff.) R. S. Chopra, Taxon. Indian Mosses 139,
1975 (Gymnostomum) comb. inval. basion. non cit. As3 =
Bryoerythrophyllum recurvum (Griff.) Saito = Bellibarbula
recurva (Griff.) Zand., see treatment of Bellibarbula]
[Barbula recurvis (Griff.) R. S. Chopra ex R. S. Chopra, Bryologist
80: 544, 1977 (Gymnostomum) As3 = Bryoerythrophyllum
recurvum (Griff.) Saito = Bellibarbula recurva (Griff.) Zand.,
see treatment of Bellibarbula]
[Barbula reflexa (Brid.) Brid. Eur As1 As2 Am1 Am2 = Didymodon rigidicaulis (C. Miill.) Saito fide Saito, J. Hattori Bot. Lab.
39: 502, 1975 = Didymodon ferrugineus (Schimp. ex Besch.)
Hill, J. Bryol. 11: 599, 1981 (1982)]
var. obtusata Monk. Eur
[var. robusta Braithw. Eur = Barbula maxima Syed & Crundw., J.
Bryol. 7: 527. 1973 nov. nov. = Didymodon maximus (Syed &
Crundw.) M. Hill]
fo. robusta Roll, Hedwigia 56: 152, 1915 Eur
[Barbula reflexifolia Fleisch. As4 = Barbula inaequalifolia Tayl.
fide Zander, Bryologist 71: 421, 1968 = Bryoerythrophyllum
inaequalifolium (Tayl.) Zand., Bryologist 83: 232, 1980]
Barbula rehmannii C. Miill. [good species fide Magill, Fl. S. Afr. I.
Mosses 1: 245, 1981 (1982)] Afr2 Afr4
[Barbula replicata Tayl. Am4 Am6 = Pseudocrossidium replicatum
(Tayl.) Zand.fide Zander, Phytologia 44: 206, 1979.]
[Barbula revoluta Brid. in Schrad. Eur As5 Afr1 Afr4 = Pseudocrossidium revolutum (Brid.) Zand.fide Zander, Phytologia 44:
204, 1979]
var. propagulifera Amann Eur
fo. elata Herz., Ber. Ziirich. Bot. Ges. 15: 46, 1905 Eur
fo. macrophylla Wamst., Krypt. Fl. Brandenb. 2(2): 244, 1904
Eur
fo. mucronata Loeske, Moosfl. Harz. 175, 1903 Eur
fo. umbrosa Loeske in Bauer, Muse. Eur. Exs. 978, 1913 Eur
[fo. mucronata Loeske, Moosfl. Harz. 175, 1903 Eur = Barbula
hornschuchiana var. obtusula (Lindh.) Podp.fide Podp., Consp.
Muse. Eur. 212, 1954 = Pseudocrossidium revolutum var.
obtusulum (Lindh.) Tan, Zand. & T. Tayl.]
Barbula riebeckii C. Miill. Afr2
[Barbula rigida Hedw. =Aloina rigida (Hedw.) Limpr.]
=
289
GENERA, SPECIES AND INFRASPECIFIC TAXA
var. desertorum Frohl. As5
var. rostellifolia Brid. Eur
[Barbula rigidula (Hedw.) Mild. = Didymodon rigidulus Hedw.]
[var. desertorum Frohl., Ann. Naturhist. Mus. Wien 63: 31, 1959
As5 = Barbula trifaria var. desertorum (Frohl.) S. Agnew,
Feddes Rep. 86: 366, 1975]
var. perobtusa Broth. As2
[fo. biseta (Pet.) Podpera, Consp. Muse. Eur. 202, 1954 (DidymoDidymodon rigidulus fo. biseta Pet.,
don rigidulus fo.) Eur
Magyar Bot. Lapok 2: 292, 1903]
[fo. brevicaulis Roll, Deutsche Bot. Monatsschr. 3: 163, 1885
Didymodon rigidulus fo. brevicaulis (Roll) Roll]
[fo. brevifolia (Roll) Podp., Consp. Muse. Eur. 202, 1954 Didymodon rigidulus fo. brevifolius Roll, Hedwigia 42: 299, 1903]
fo. densa (BSG) Podp., Consp. Muse. Eur. 203, 1954 (Trichostomum rigidulum var.) Eur
[fo. gemmipara Herz., Wiener Bot. Z. 93: 39, 1944 Eur = Barbula
rigidula (Hedw.) Milde fide Crundwell & Nyholm, Svensk Bot.
Tidskr. 59(2): 211, 1965 = Didymodon rigidulus Hedw.]
[fo. laxa (Mol.) Monk., Laubm. Eur. 1927 (Didymodon rigidulus
var.) Eur =Didymodon rigidulus fo. laxus Mol.]
[fo. longicau/is Roll, Deutsche Bot. Monatsschr. 3: 163, 1885
Didymodon rigidulus fo. longicaulis (Roll) Roll]
[fo. longifolia Podp., Consp. Muse. Eur. (Didymodon rigidulus var.)
Eur. 203, 1954 Eur]
[fo. major (Podp.) Podp., Consp. Muse. Eur. 203, 1954 (Didymodon rigidulus var.) Eur =Didymodon rigidulus var. major Podp.]
[fo. propagulifera (Schiffn.) Podp., Consp. Muse. Eur. 203, 1954
Didymodon rigidulus fo.
(Didymodon rigidulus fo.) Eur
propaguliferus (Schiffn.) Limpr.]
[fo. tenuis (Hammerschm.) Podp., Consp. Muse. Eur. 203, 1954
(Didymodon rigidulus var.) Eur = Didymodon rigidulus var.
tenuis Hammerschm.]
[fo. viridis Roll, Deutsche Bot. Monatsschr. 3: 164, 1885 Didymodon rigidulus fo. brevicaulis (Roll) Roll fide Podpera, Consp.
Muse. Eur. 203, 1954]
Barbula riograndensis Bartr. Am5
Barbula riparia C. Mtill. Am6
[Barbula rivicola Broth. As2 = Didymodon rivicola (Broth.) Zand. in
Kop., Gao, Lou & Jarvinen]
Barbula robbinsii Bartr. As4
Barbula rothii Herz. Am4
Barbula rottensis Weyl. Eur, fossil
[Barbula rubella (Hiib.) Mitt. in Lindh. hom. illeg. Eur = Bryoerythrophyllum recurvirostrum (Hedw.) Chen]
fo. minor Amell in Lindh. & Amell, K. Svensk. Yet.-Ak. Hand!.
23(10): 72, 1890 As1
Didymodon occidentalis Zand.,
[Barbula rubiginosa Mitt. Am1
Phyto1ogia 41: 26, 1978 nom. nov. (= Didymodon vinealis var.
rubiginosus (Mitt.) Zand., Cryptogamie Bryol. Lichenol. 2: 417,
1981)]
[Barbula rubricaulis Ther. Am2 = Barbula arcuata Griff. fide
Zander, Phytologia 44: 200, 1979]
Barbula rubriseta Bartr. in Bauer Am6
Didymodon asperifolius (Mitt.) Crum,
[Barbula rufa (Lor.) Jur.
Steere & Anders.]
fo. laevis C. Jens., Bot. Faerties 155, 1901 [Skand. Bladmfl. 238,
1939] Eur
fo. sublaevigata (Herz.) Podp., Consp. Muse. Eur. 211, 1954 (Didymodon rufus fo.) Eur
[Barbula rufofusca Lawt. & Herm., Bull. Torrey Bot. Club 99: 307,
1972 Am1 = Didymodon nigrescens (Mitt.) Saito fide Zander,
Phytologia 41: 22, 1978]
=
=
=
=
=
=
=
=
Barbula scaberrima Broth. & Par. As3
Barbula semirosulata Zand. (nom . nov. for Gymnostomiella
rosulata P. Yarde), see treatment of Barbula Afr1
[Barbula sobolifera Aeisch. As4 = Barbula arcuata Griff. fide
Saito, J. Hattori Bot. Lab. 39: 496, 1975]
Barbula solfatariensis Fleisch. As4
Barbula somaliae C. Miill. Afr2
Barbula sordida Besch. As3
Didymodon
[Barbula spadicea (Mitt.) Braithw. Eur As1 As5
spadiceus (Mitt.) Limpr.]
var. squarrosa Latz. Eur
[var. vaginans (Lindh.) G. Roth in Bauer, Lotos 54(5): 15, 1906
(Barbula) Eur Barbula spadicea fo. vaginans (Lindh.) Podp.]
fo. bernensis (Culm.) Podp., Consp. Muse. Eur. 208, 1954 (Barbula spadicea var.) Eur
fo. brevifolia Latz., Magyar Bot. Lapok 29: 117, 1930 Eur
fo. debilis Latz., Magyar Bot. Lapok 29: 117, 1930 Eur
fo. leptoderma Podp., Consp. Muse. Eur. 208, 1954 (Barbula
spadicea var. leptoderma Jens. nom. inval.) Eur
fo. mol/is (Burchard) Podp., Consp. Muse. Eur. 208, 1954 (Didymodon spadiceus var.) Eur
fo. obtusifolia (Roll) Podp., Consp. Muse. Eur. 208, 1954 (Didymodon spadiceus var.) Eur
fo. vaginans (Lindh.) Podp., Consp. Muse. Eur. 208, 1954 (Didymodon) Eur
[Barbula sparsifolia Ren. & Card. Afr3 = Trichostomum
sparsifolium (Ren. & Card.) Card. in Grand.]
Barbula spathulifolia (Dix. & P. Yarde) Zand., see treatment of
Barbula (Merceyopsis) As3
Barbula speirostega C. Miill. Austr1
[Barbula spiralis Schimp. ex C. Miill. Am1 Am2 =Pseudocrossidium replicatum (Tayl.) Zand. fide Zander, Phytologia 44: 206,
1979]
[var. emarginata Card. Am2
Pseudocrossidium replicatum
(Tayl.) Zand.fide Zander, Phytologia 44: 206, 1979]
Barbula squarrosa Schultz hom. i/leg. Eur
Barbula stenocarpa Hampe Am4
[Barbula stenotheca Ther. Am2 (= Barbula orizabensis C. Miill.
fide Zander, Phytologia 44: 183, 1979) = Morinia stenotheca
(Ther.) Zand., Bryologist 86: 156, 1983 = Mironia stenotheca
(Ther.) Zand., see treatment of Mironia]
[Barbula stewartii Bartr. As3 =Didymodon stewartii (Bartr.) Zand.,
see treatment of Didymodon]
[Barbula stillicidiorum Card. Am2 = Barbula arcuata Griff. fide
Zander, Phytologia 44: 200, 1979]
[Barbula strictidens C. Miill. Am2 = Didymodon rigidulus Hedw. s.
lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 390, 1981
(1982)]
[Barbula strictifolia (Dix. & P. Yarde) R. S. Chopra, Taxon. Indian
Didymodon strictifo/ius Dix. & P.
Mosses 140, 1975 As3
Yarde As3 = Didymodon recurvus (Griff.) Broth. fide Robinson, Bryologist 71: 85, 1968 Bryoerythrophyllum recurvum
(Griff.) Saito = Bellibarbula recurva (Griff.) Zand., see treatment of Bellibarbula]
Barbula stulhmannii (Broth.) Broth. Afr2 Afr4
[Barbula subanomala C. Miill. As3 = Timmie/la anomala (BSG)
Limpr.]
Barbula subcespitosa (Hampe) Broth. Am4
Barbula subcalycina C. Miill. Austr1
Barbula subcernua Schimp. in Besch. Afr4
Barbula subcomosa Broth. As2
[Barbula subcontorta As2 = Didymodon vinea/is (Brid.) Zand.fide
Sollman, Bryologist 86: 272, 1983]
=
=
=
=
=
290
GENERA OF THE POTTIACEAE
Barbula subdenticulata Dix. As3
[Barbula suberythropoda C. Miill. Am2 =Barhula arcuata Griff. fide
Zander, Phytologia 44: 199, 1979]
Barbula subglaucescens C. Miill. Am4
[Barbula subglaucula Dix. ex Sainsb. nom. inval. descr. angl. Austr2
= Barbula unguiculata Hedw. fide Sollman, Lindbergia 10: 54,
1984]
Barbula subgracilis C. Miill. & Kindb. in Macoun Ami
Barbula subgrimmiacea Th6r. in Felipp. Am6
[Barbula sublaevifolia Toyama As2 = Barbula javanica Dozy &
Molk.fide Saito, J. Hattori Bot. Lab. 39: 495, 1975]
Barbula subobtusa Th6r. Afr3
Barbula subpel/ucida Mitt. As2 As3
Barbula subreflexifolia C. Miill. Am4
Barbula subreplicata Broth. Am4
[Barbula subrivicola Chen As2 = Didymodon nigrescens (Mitt.) Saito
fide Saito, J. Hattori Bot. Lab. 39: 510, 1975]
[var. densifolia Chen As2 = Didymodon nigrescens (Mitt.) Saito
fide Saito, J. Hattori Bot. Lab. 39: 510, 1975]
Barbula subrufa Broth. & C. Miill. Asl
Barbula subrunciata C. Miill. Am6
[Barbula subscabrinervis Dix. & Nav. Afr2 = Bryoerythrophyllum
recurvum (Griff.) Saito fide Sollman, Lindbergia 16: 22, 1990 =
Bellibarbula recurva (Griff.) Zand., see treatment of Bellibarbula]
[Barbula subteretiuscula Card. Am2 Am3 = Didymodon rigidulus
Hedw. s. lat. fide Zander, Cryptogamic Bryol. Lichenol. 2: 390,
1981 (1982)]
[Barbula subtorquata C. Miill. & Hampe Austrl
Didymodon
subtorquatus (C. Miill. & Hampe) Catcheside, Mosses S. Austr.
174, 1980]
[Barbula subulifolia Sull. Am2 Am3 Am4 Am5 = Barbula arcuata
Griff. fide Zander, Phytologia 44: 199, 1979]
Barbula su/cata Geh. in Geh. & Herz. Afr2
Barbula sumatrana Baumg. & Dix. As4
[Barbula svihlae Bartr. As3 = Hymenostylium recurvirostrum
(Hedw.) Dix. var. recurvirostrum, see treatment of Hymenostylium]
Barbula swartziana C. Miill. Am3
[Barbula tamakii Broth. As2 = Dichodontium pellucidum (Hedw.)
Schimp.fide Saito, J. Hattori Bot. Lab. 39: 528, 1975]
Barbula taylorii Bartr. & Steere hom. illeg. Am4 [= Hydrogonium
taylorii Weber nom. nov., Lindbergia 3: 81, 1975 (1976)]
[Barbula tectorum C. Miill. As2 Didymodon tectorum (C. Miill.)
Saito, J. Hattori Bot. Lab. 39: 517, 1975]
[Barbula tenii Herz. As2 = Barbula inaequa/ifolia Tayl. fide Zander,
Bryologist 71 : 41, 1968 = Bryoerythrophyllum inaequa/ifolium
(Tayl.) Zand., Bryologist 83: 232, 1980]
Barbula tenuicoma C. Miill. ex Broth. Am5
Barbula tenuirostris Brid. As3 As4 Oc
[Barbula teretiuscu/a Schimp. ex C. Miill. Am2 = Didymodon
rigidulus Hedw. s. lat. fide Zander, Cryptogamic Bryol. Lichenol.
2: 389, 1981 (1982)]
Barbula thelimitria C. Miill. As4
Barbula tisserantii (P. Yarde) P. Yarde Afr2
[Barbula tokyensis Besch. As2 = Barbula unguiculata Hedw. fide
Saito, J. Hattori Bot. Lab. 39: 491, 1975 as "tokyoensis"]
[Barbula tonkinensis (Besch.) Broth. As3 = Barbula indica (Hook.)
Spreng., see treatment of Barbula]
[Barbula tomaculosa Blockeel, J. Bryology 11: 583, 1981 (1982) Eur
= Didymodon tomaculosus (Blockeel) Corley in Corley et al., J.
Bryol. 11 : 649, 1981 (1982)]
[Barbula tophacea (Brid.) Mitt. Didymodon tophaceus (Brid.) Lisa]
=
=
=
[fo. acutifolia (Schimp.) Monk., Laubm. Eur. 295, 1927 (Trichostomum tophaceum var.) Eur
Didymodon tophaceus fo.
acutifolius Monk., Laubm. Eur. 295, 1927 Eur]
[fo. anatina (Hammerschm.) Podp., Consp. Muse. Eur. 201 , 1954
(Didymodon tophaceus var.) Eur = Didymodon tophaceus var.
anatinus Hammerschm.]
[fo. bosniaca (Glow.) Monk. Laubm. Eur. 296, 1927 (Didymodon) Eur = Didymodon tophaceus var. decurrens Card. & Ther.
fide Podp., Consp. Muse. Eur. 201, 1954]
fo. cylindrica (Boul.) Podp., Consp. Muse. Eur. 201, 1954
(Trichostomum tophaceum fo.) Eur
[fo. humilis (Schimp.) Monk., Laubm. Eur. 295, 1927 (Didymodon tophaceus var.) Eur = Didymodon tophaceus var. humilis
Schimp.]
[fo. taxa (Kindb.) Podp., Consp. Muse. Eur. 201, 1954 (Didymodon tophaceus fo.) Eur = Didymodon tophaceus fo. laxus
Kindb., Boll. Soc. Bot. ltal. 7: 15, 1896]
[fo. linearis (De Not.) Podp., Consp. Muse. Eur. 201, 1954
(Trichostomum tophaceum var.) Eur Didymodon tophaceus
var. linearis (De Not.) Limpr.]
fo. Ungulata (Boul.) Monk., Laubm. Eur. 295, 1927 (Trichostomum tophaceum fo.) Eur
[fo. propagulifera (Amann) Monk., Laubm. Eur. 296, 1927
(Didymodon tophaceus fo.) Eur
Didymodon tophaceus fo.
propagu/iferus Amann]
[fo. recurvifolia (Boul.) Demar., Bull. Jard. Bot. Nat!. Belgique
17: 335, 1945 (Trichostomum tophaceum fo .) Eur =Didymodon
tophaceus fo. recurvifolius De Willd., Prodr. 434, 1899]
[fo. scabrinervis (Podp.) Podp., Consp. Muse. Eur. 202, 1954
(Didymodon tophaceus fo.) Eur = Didymodon tophaceus fo.
scabrinervis Podp.]
fo . thermarum Boros in Bauer, Muse. Eur. Exs. 2011, 1928 Eur
fo. torrentium (Loeske) Podp., Consp. Muse. Eur. 202, 1954
(Barbula tophacea var.) Eur
fo. truncata (Boul.) Podp., Consp. Muse. Eur. 202, 1954 (Trichostomum tophaceum fo.) Eur
[Barbula torquata Tayl. Austrl Austr2 (= Trichostomopsis
australasiae (Hook. & Grev.) Robins. fide Robinson,
Phytologia 20: 184, 1970)
Didymodon torquata (Tayl.)
Catcheside, Mosses S. Austr. 174, 1980]
[Barbula torquatifolia Geh. Afr2 Afr4 = Tortula porphyreoneura
(C. Miill.) Townsend fide Magill, Fl. S. Afr. I. Mosses 1: 213,
1981 [1982] Pseudocrossidium porphyreoneurum (C. Miill.)
Zand., see treatment of Pseudocrossidium]
Barbula tortelloides C. Miill. Am6
[Barbula tortuosa (Hedw.) Web. & Mohr
Tarte/la tortuosa
(Hedw.) Limpr.]
[fo. a/pina Boul., Muscin. France 421, 1886 Eur = Tarte/la
tortuosa fo. alpina (Boul.) Podp.]
[fo. dentata Fam., Muschi Prov. Pavia, Atti lstituto Bot. Univ.
Pavia 3: 18, 1891 Eur = Tarte/la tortuosa fo. dentata (Farn.)
Podp.]
[fo. gracilescens Zett., K. Svensk Vet. Ak. Hand!. 5(10): 25, 1865
Eur =Torte /Ia tortuosa fo. gracil/escens (Zett.) Par.]
[Barbula tosaensis Broth., Oefv. Finsk. Vet. Sco. Foerh. 62A(9):
10, 1921 As2 = Barbula subcomosa Broth. fide Saito, J. Jap.
Bot.47: 11,1972]
Barbula translucens Salzm. ex Bruch Afr2
Barbula trichomanoides Broth. ex Ihs. As2
[Barbula trichostomacea C. Miill. Afr4 = Trichostomopsis
australasiae (Hook. & Grev.) Robins., Phytologia 20: 187,
Didymodon australasiae (Hook. & Grev.) Zand.,
1970
Phytologia 41: 21, 1978]
=
=
=
=
=
=
=
291
GENERA, SPECIES AND INFRASPECIFIC TAXA
[Barbula trifaria (Hedw.) Mitt. Eur As1 As2 As3 As5 Afr1 Am1
Am2 Oc = Saelania g/aucescens (Hedw.) Broth. in Bomanss &
Broth. fide Zander, Cryptog. Bryol. Lichenol. 2: 412, 1981
(1982)]
var. desertorum (Frohl.) S. Agnew, Feddes Rep. 86: 366, 1975
(Barbula rigidula var.) As2
[fo. brevifolia (Latz.) Podp., Consp. Muse. Eur. 204, 1954 Eur
(Didymodon luridus fo.) Eur = Didymodon luridus Co. brevifolius
Latz.]
[fo. rubella (Loeske in Bauer) Podp., Consp. Muse. Eur. 204, 1954
(Didymodon luridus fo.) Eur = Didymodon luridus Co. rubella
Loeske in Bauer]
[fo. subscabra (Linder) Podp., Consp. Muse. Eur. 204, 1954 (Didymodon luridus fo.) Eur
Didymodon luridus Co. subscabrus
Linder]
[fo. tophacea (Amann) Podp. , Consp. Muse. Eur. 204, 1954 (Didymodon luridus) = Didymodon luridus Co. tophaceus Amann, Fl.
Mousses Suisse Ad. 3: 26, 1935]
[fo. cuspidata (Schimp.) Monk., Laubm. Eur. 294, 1927 (DidymoDidymodon trifarius var. cuspidatus
don luridus var.) Eur
(Schimp.) Wijk & Marg.]
[Barbula trivia/is C. Mill!. Afr4 Trichostomopsis trivia/is (C. Mill!.)
Robins., Phytologia 20: 187, 1970 = Didymodon trivia/is (C.
Milll.) Guerra in Guerra & Ros, Cryptogamic Bryol. Lichenol. 8:
64, 1987]
Barbula tuberculosa (Ren. & Par.) Card. in Grand. Afr3
[Barbula umbrosa C. MUll. Afr6
Trichostomopsis umbrosa (C.
Milll.) Robins., Phytologia 20: 185, 1970
Didymodon
australasiae var. umbrosus (C. Mill!.) Zand. fide Zander,
Cryptogamic Bryol. Lichenol. 2: 400, 1981 (1982) Didymodon
umbrosus (C. Milll.) Zand.]
Barbula umtaliensis Magill in Magill & Schelpe, Mem. Bot. Surv. S.
Afr. 43: 5, 1979 (nom. nov. for Torre/la obtusifolia Dix.) Afr2
Barbula unguicu/ata Hedw. Eur As1 As2 As3 As5 Afr1 Am1 Am2
Am6 Austr1
var. e/ongata (Schultz) Brid. Eur
var. patagonica C. Mill!. Am6
[var. proligera Broth. As2 = Barbula indica (Hook.) Spreng. fide
Saito, J. Hattori Bot. Lab. 39: 488, 1975]
var. rigidula Roll Eur
var. rupestris Hiib. Eur
var. simplex Brid. Eur
[var. trichostomifolia (C. Milll.) Chen As2 = Barbula unguiculata
Hedw.fide Saito, J. Hattori Bot. Lab. 39: 490, 1975]
fo. apicu/ata (Hedw.) Monk., Laubm. Eur. 286, 1927 (Barbula) Eur
Am1
fo.? brachypus (Brid.) Podp., Consp. Cusc. Eur. 213, 1954 nom.
inval. dispon . non cit. (Barbula) Eur
fo.? breviseta (Farn.) Podp., Consp. Muse. Eur. 213, 1954 nom.
inval. dispon . non cit. (Barbula unguiculata var.) Eur
fo.? bulbifera (Schiff.) Podp., Consp. Muse. Eur. 213, 1954 nom.
inval. dispon. non cit. (Barbula unguiculata var.) Eur
fo. cuspidata (Schultz) Monk., Laubm. Eur. 286, 1927 (Barbula
cuspidata) Eur
fo. fastigiata (Schultz) Monk., Laubm. Eur. 286, 1927 (Barbula)
Eur
fo.jlaccida Roll, Jahrb. Ak. Wiss. Erfurt 41: 123, 1915 Eur
fo.? /anceo/ata (Hedw.) Podp., Consp. Muse. Eur. 213, 1954 nom.
inval. dispon. non cit. (Barbula) Eur Am1
fo.? microcarpa (Schultz) Podp., Consp. Muse. Eur. 213, 1954
nom. inval. dispon. non cit. (Barbula) Eur
fo. minus Tosco, Webbia 28: 284, 1973 "minor" Eur
fo.? nitidocostata (Farn.) Podp., Consp. Muse. Eur. 213, 1954 nom.
=
=
=
=
=
=
inval. dispon. non cit. (Barbula unguiculata var.) Eur
fo. obtusifolia Monk., Laubm. Eur. 286, 1927 (Barbula
obtusifolia Schultz hom. i/leg.) Eur!
fo.? pa/udosa (Roll) Podp., Consp. Muse. Eur. 214, 1954 nom.
inval. dispon. non cit. (Barbula unguicu/ata var.) Eur
fo. polyseta Peterfi, Mag. Bot. Lapok 1: 49, 1902 Eur
[fo. propagulosa Crum, Bryologist 72: 241, 1969 Ami= Barbula
indica (Hook.) Spreng. fide Zander, Photologia 44: 186, 1979]
fo. robusta (Lindb. ex Hag.) Podp. in Sap., Bot. Jahrb. Syst. 46
(Beibl. 105): 12, 1911 (Barbula unguiculata var.) Eur
fo. subsquarrosa Latz., Bot. Centralbl. Beih. 48: 2, 480, 1931 Eur
Barbula unguiculatula C. Mill!. Am4 Am6
[Barbula uruguayensis Broth. (hom. illeg. of Barbula uruguensis
Par. fide Margadant in !itt.) Am6 = Didymodon uruguayensis
(Broth.) Zand., see treatment of Didymodon]
[Barbula uruguensis Par., Act. Soc. Linn. Bordeaux 46: 111, 1893
[1894] Am6 Didymodon deciduus Zand. nom. nov., see treatment of Didymodon]
Barbula vaginata Warnst. Am6
[Barbula valida (Limpr.) Moll. Eur = Barbula acuta (Brid.) Brid.
fide Crundwell & Nyholm, Svensk Bot. Tidskr. 59(2): 214,
1965 = Didymodon rigidulus var. gracilis (Hook. & Grev.)
Zand. fide Zander, Cryptog. Bryol. Lichenol. 2: 393, 1981
(1982)]
Barbula validinervia C. Miill. Afr4
Barbula vardei R.S. Chopra, Bryologist 80: 544, 1977 (nom . nov.
for Barbula denticulata Dix. & P. Yarde) As3
Barbula ventanica C. Milll. Am6
[Barbula vinealis Brid. Eur Asl As2 As3 As5 Afrl Am1 Am2 Oc =
Didymodon vinealis (Brid.) Zand., Phytologia 41: 25, 1978]
[subsp. cylindrica (Tayl.) Podp. Eur As1 As5 Afr1 Am1 Am6 Oc
= Didymodon vinealis var. flaccidus (BSG) Zand., Phytologia
41: 25, 1978 Didymodon vinealis (Brid.) Zand. var. vinealis
fide Sollman, Bryologist 86: 272, 1983]
var. decipiens Meyl. in Amann Eur
[var. flaccida BSG Eur As1 As5 Afrl Am1 Am6 Oc Didymodon vinealis var. flaccidus (BSG) Zand., Phytologia 41: 25,
1978 = Didymodon vinealis (Brid.) Zand. var. vinealis fide
Sollman, Bryologist 86: 272, 1983]
var. propagulifera Amann Eur
fo. compacta Herz., Ber. Zilrich. Bot. Ges. 15: 45, 1905 Eur
fo. propagulifera (Amann) Podp., Consp. Muse. Eur. 210, 1954
(Didymodon vinealis var.) Eur
fo. rivularis (Loeske) Podp., Consp. Muse. Eur. 210, 1954 (Barbula) Eur
fo. robusta (Loeske) Podp., Consp. Muse. Eur. 210, 1954 (Barbula cylindrica fo.) Eur
fo. rubella (Schiffn.) Podp., Consp. Muse. Eur. 210, 1954 (Barbuta cylindrica var.) Eur
fo. viridis Podp., Consp. Muse. Eur. 210, 1954 Eur
Barbula vulcanica Lor. Am4
[Barbula whitehouseae Crum Am1 = Barbula eustegia Card. &
Ther.fide Zander, Phytologia 44: 197, 1979]
Barbu/a williamsii (Chen) lwats. & Tan, Kilikasan, Philippine J.
Bioi. 8: 186, 1979 (Hydrogonium) As4
[Barbula wisselii Dix. As4 Didymodon wisselii (Dix.) Norris & T.
Kop., Acta Bot. Fenn. 137: 127, 1989]
[Barbula wrightii Saurb. in Jag. Am3 = Barbula indica (Hook.)
Spreng. fide Zander, Phytologia 44: 186, 1979]
[Barbula xanthocarpa C. Mill!. Afr2 Afr4
Didymodon
xanthocarpus (C. Miill.) Magill, Fl. So. Afr. Bryoph. 1: 235,
1981]
Barbula yunnanensis Copp. As2
=
=
=
=
=
GENERA OF THE POTTIACEAE
292
Barbu/a zambesiaca Magill in Magill & Schelpe, Mem. Bot. Surv. S.
Afr. 43: 5, 1979 (nom. nov. for Semibarbula e/ongata Hilp.) Afr2
[Barbu/a zol/ingeri (Fleisch.) Broth. As4 = Barbula indica (Hook.)
Spreng. var. indica fide Norris & Koponen, Acta Bot. Fenn. 137:
114, 1989]
[BARNESIA Card. = Streptocalyptra C. Miill. fide Zander,
Lindbergia 8: 162, 1982 (1983)]
[Barnesia tortel/oides Card. Am2
Streptoca/ypta tortelloides
(Card.) Zand., Lindbergia 8: 163, 1982 (1983)]
=
BELLIBARBULA Chen
Bellibarbu/a kurziana Chen As3
var. purpurascens Gangulee, Nov. Hedw. 8: 148, 1964 As3
[Bellibarbula obtusicuspis (Besch.) Chen As2 =Bellibarbula recurva
(Griff.) Zand., see treatment of Bellibarbula]
Bellibarbula recurva (Griff.) Zand., see treatment of Bellibarbula
(Gymnostomum) As2 As3 Am1 Am2
BYROCEUTHOSPORA Crum & Anders.
Byroceuthospora aethiopica (Welw. & dub.) Zand. (Ephemerum), see
treatment of Byroceuthospora Afr2
Byroceuthospora mexicana (Bartr.) Crum & Anders. Am2
BRYOERYTHROPHYLLUM Chen
Bryoerythrophyllum a/pigenum (Vent.) Chen Eur As1 As2 As3 Am1
Austr2
Bryoerythrophyllum andersonianum Zand. & Sharp, Bryologist 84:
545, 1981 Am2
[Bryoerythrophyl/um angustulum (Herz.) Robins., Bryologist 70: 22,
1970 (Didymodon) Am4 = Trichostomopsis austra/asiae (Hook.
& Grev.) Robins. fide Robinson, Smiths. Contr. Bot. 27: 30, 1975
= Didymodon australasiae (Hook. & Grev.) Zand. fide Zander,
Phytologia41: 21, 1978]
[Bryoerythrophyllum arcuatum (Mitt.) Crum Am4 (= Bryoerythrophyllumjamesonii (Tayl.) Crumfide Zander, Bryologist 81: 549,
1978 [1979]) = Bryoerythrophyllum campylocarpum (C. Miill.)
Crumfide Zander, Bryologist 89: 15, 1986]
[Bryoerythrophyllum afrorubellum (Broth. & Wag.) De Sloover, Bull.
Jard. Bot. Nat. Belg. 49: 398, 1979 Afr4 (= Bryoerythrophyllum
jamesonii (Tayl.) Crum fide Magill, Fl. S. Afr. I. Mosses 1: 248,
1981 [1982]) = Bryoerythrophyllum campy/ocarpum (C. Miill.)
Crum from illustration by Magill, Fl. S. Afr. I. Mosses 1: 248,
1981 (1982); see Zander, Bryologist 89: 13, 1986]
[Bryoerythrophyllum atrorubens (Besch.) Chen As2 As3 = Bryoerythrophyllum wallichii (Mitt.) Chen fide Saito, J. Hattori Bot. Lab.
39: 479, 1975]
Bryoerythrophyllum binnsii (R. Brown ter) Wijk & Marg. Austr1
Austr2 (good species fide Zander, Bryo1ogist 89: 15, 1986)
Bryoerythrophyllum holivianum (C. Miill.) Zand., Bryo1ogist 81: 545,
1978 [1979] (Giobulina) Am2 Am4
Bryoerythrophyllum brachystegium (Besch.) Saito, J. Jap. Bot. 47: 14,
1972 As2
Bryoerythrophyllum byrdii (Bartr.) Zand. (Barbula), see treatment of
Bryoerythrophyllum Ant
Bryoerythrophyllum ca/careum (Thtr.) Zand., Bryologist 83: 232,
1980 (Barbula) Am2
Bryoerythrophyl/um campy/ocarpum (C. Miill.) Crum Eur As2 As3
Afr2 Afr4 Am2 Am3 Am4 Am6 [= Bryoerythrophyllum
jamesonii (Tayl.) Crum fide Zander, Bryologist 81: 549, 1978
(1979)] = Bryoerythrophyllum campylocarpum (C. Miill.) Crum
fide Zander, Bryologist 89: 15, 1986]
[Bryoerythrophyllum cavernarum (Mol.) Podp. =Bryoerythrophyllum
rubrum (Jur.) Chen]
fo. brevifolium (Herz.) Podp., Consp. Muse. Eur. 219, 1954
(Erythrophyllum ruhrum f.) Eur
Bryoerythrophyllum chimhorazense (Mitt.) Zand. (Tortula), see
treatment of Bryoerythrophyllum Am4
Bryoerythrophyllum co/umbianum (Herm. & Lawt.) Zand.,
Bryologist 81: 548, 1978 (Didymodon) Am1
[var. atacamense Zand. & Lewis in Lewis, Bryologist 84: 536,
1981 [1982] "atacamensis" Am4 = Bryoerythrophyllumfuscinervium (Mitt.) Zand., see treatment of Byroerythrophyllum]
[Bryoerythrophyllum dentatum (Mitt.) Chen As3 As4 = Leptodontium flexifolium (Dicks.) Hampe in Lindb. fide Zander,
Bryologist 75: 231, 1972 cf. Zander, Bryologist 84: 546, 1972]
Bryoerythrophyllumferruginascens (Stirt.) Giac. Eur As1 As4 Am1
Am2
Bryoerythrophyllumferrugineum Gangulee, Nov. Hedwigia 8: 147,
1964 As3
Bryoerythrophyllumfuscinervium (Mitt.) Zand. (Tortula), see treatment of Byroerythrophyllum Am4 Am6
Bryoerythrophyllum gymnostomum (Broth.) Chen As2
Bryoerythrophyllum hostile (Herz.) Chen As2
Bryoerythrophyllum inaequalifolium (Tayl.) Zand., Bryologist 83:
232, 1980 (Barhula) As1 As2 As3 As4 Afr1 Am1 Am2 Am4
[Bryoerythrophyllum linearifolium Saito, J. Hattori Bot. Lab. 39:
481, 1975 As2 = Bryoerythrophyllum recurvum (Griff.) Saito
fide Sollman, Lindbergia 16: 22, 1990 Bellibarhula recurva
(Griff.) Zand., see treatment of Bel/iharhula]
[Bryoerythrophyllum /usitanicum (Card. & Dix.) Hill, J. Bryol. 11:
600, 1981 [1982] (Hyophila) Eur = Bryoerythrophyllum
campylocarpum (C. Miill.) Crum fide Zander, Bryologist 89:
15, 1986]
Bryoerythrophyllumjamesonii (Tayl.) Crum Am2 Am4
Bryoerythrophyllum ligu/are (Mitt.) Zand. (Tortula), see treatment
of Bryoerythrophyllum Am4
Bryoerythrophyllum machadoanum (Sergio) Hill, J. Bryol. 11: 601,
1981 [1982] (Hyophila) Eur
Bryoerythrophyllum noguchianum (Gangulee) Saito, Bull. Univ.
Mus. Univ. Tokyo 8: 254, 1975 (Bryoerythrophyllum
yunnanense var.) As2
[Bryoerythrophyllum obtussissimum (Broth.) Chen As2 = Didymodon hrachystegius (Besch.) Broth. fide lwats. & Nog., J. Hattori
Bot. Lab. 37: 316, 1973 = Bryoerythrophyllum brachystegium
(Besch.) Saito fide Saito, J. Jap. Bot. 47: 14, 1972 and J. Hattori
Bot. Lab. 39: 477, 1975]
[Bryoerythrophyllum pergemmascens (Broth.) Chen As2 = Leptodontium flexifolium (Dicks. ex. With.) Hampe in Lindb. fide
Zander, Bryologist 75: 231, 1972]
[Bryoerythrophyllum recurvifolium (Tayl.) Zand., Bryologist 75:
Oxystegus recurvifolius (Tayl.) Zand.,
277, 1972 Eur Am1
Lindbergia 8: 187, 1982 = Trichostomum recurvifolium (Tayl.)
Zand., see treatment of Trichostomum]
Bryoerythrophyllum recurvirostrum (Hedw.) Chen Eur As1 As2
As3 As4 AsS Afr1 Afr2 Afr4 Am1 Am2 Austr1 Oc
var. aeneum (Ren. & Card.) Zand., Bryologist 79: 229, 1977
(Trichostomum) Am2
var. antarcticum Sav.-Ljub. & Z. Smirn. Ant
[var. angustifolium (P. Yarde) Wijk & Marg. Afr2 = Bryoerythrophyllum afrorubellum (Broth. & Wag.) De S1oover fide De
Sloover, Bull. Jard. Bot. Nat!. Belg. 49: 398, 1979]
var. hrevifolium (Lindb. & Am.) Podp. As1 Ami
[var. dentatum (Schimp.) Crum, Steere & Anders., Bryologist 67:
163, 1964 Am1 = Bryoerythrophyllum recurvirostrum (Hedw.)
Chenfide Crum, Steere & Anderson, Bryologist 68: 418, 1973]
=
=
293
GENERA, SPECIES AND INFRASPECIFIC TAXA
var. crassinerve (Herz.) Podp. Eur
var. latinervium (Holmen) Murray, Bryobrothera 1: 14, 1992
(Bryoerythrophyllum recurvirostrum var.) Am1
var. robustum Saito, J. Hattori Bot. Lab. 39: 474, 197S As2
var. serratum (Roll) Podp. Eur
fo. alpinum (Herz.) Podp., Consp. Muse. Eur. 2I8, 19S4
(Erythrophyllum rubellum f.) Eur
fo. apiculatum (Roll) Podp., Consp. Muse. Eur. 2I8, I9S4 (Didymodon rubellus var.) Eur
fo. brevirostre (Warnst.) Podp., Consp. Muse. Eur. 218, 19S4
(Didymodon rubellus f.) Eur
fo. confertum (Velen.) Podp., Consp. Muse. Eur. 2I8, I9S4 (Didymodon rubellus var.) Eur
fo. flaccidum (Roll) Podp., Consp. Muse. Eur. 218, 19S4 (Didymodon rubellus var.) Eur
fo. gracile (Limpr.) Podp., Consp. Muse. Eur. 218, 19S4 (Didymodon rubellus f.) Eur
fo. latifolium (Herz.) Podp., Consp. Muse. Eur. 218, 19S4
(Erythrophyllum rubellum f.) Eur
fo. longirostre (Warnst.) Podp., Consp. Muse. Eur. 2I8, I9S4
(Didymodon rube/Ius f.) Eur
fo. maius (Breidl. ex Matous.) Podp., Consp. Muse. Eur. 218, 19S4
(Didymodon ruhel/us f.] Eur
fo. obtusifolium (Roll) Podp., Consp. Muse. Eur. 218, 19S4 (Didymodon ruhel/us var.) Eur
fo. pallens (Ryan & Hag.) Podp., Consp. Muse. Eur. 2I8, I9S4
(Didymodon ruhellus var.) Eur
fo. pygmaeum (Meyl.) Podp., Consp. Muse. Eur. 218, 19S4 (Didymodon rubellus f.) Eur
fo. pulvinatum (Herz.) Podp., Consp. Muse. Eur. 218, 19S4
(Erythrophyllum rubellum f.) Eur
fo. viride (Sehlieph. ex Limpr.) Podp., Consp. Muse. Eur. 218,
I9S4 (Didymodon rubellus f.) Eur
[Bryoerythrophyllum recurvum (Griff.) Saito, Bull. Univ. Mus. Univ.
Tokyo 8: 2S4, 197S (Didymodon) As3 Aml Am2 Belliharbula
recurva (Griff.) Zand .., see treatment of Belliharhula]
Bryoerythrophyllum rotundatum (Lindh. & Am.) Chen As I
Bryoerythrophyllu'm rubrum (Geh.) Chen Eur As! As2
[var. minus Saito, J. Hattori Bot. Lab. 39: 476, I97S As2 =Bryoerythrophyllumferruginascens (Stirt.) Giac.fide Sollman, Bryologist
86: 271, I983J
Bryoerythrophyllum sharpii Zand., Bryologist 89: 13, 1986 Am2
[Bryoerythrophyllum tenerrimum (Broth.) Chen As2 = Bryoerythrophyllum recurvum (Griff.) Saito fide Sollman, Bryologist 86: 271,
1983 =Beflibarbula recurva (Griff.) Zand., see treatment of Bellibarbula]
Bryoerythrophyllum wallichii (Mitt.) Chen As2 As3
Bryoerythrophyllum yichunense Ch. Gao, Fl. Muse. Chinae Bor.-Or.
379, 1977 As2
Bryoerythrophyllum yunnanense (Herz.) Chen As2
[var. noguchianum Gangulee, Mosses of India and Adj. Reg., Fase.
3: 7SO, I972 As2 = Bryoerythrophyllum noguchianum Saito,
Bull. Univ. Mus. Univ. Tokyo 8: 2S4, 197SJ
var. pulvinans (Herz.) Chen As2
=
CALYMPERASTRUM Stone, J. Bryol. I4: 3IS, I986 [I987]
Calymperastrum latifolium Stone, J. Bryol. I4: 3IS, I986 [I987J
Austri
CALYPTOPOGON (Mitt.) Broth.
Calyptopogon mnioides (Sehwaegr.) Broth. Am4 Am6 Austri Austr2
var. anguste-limhatus Salm. Austr2
CHENIA Zand.
Chenia lorentzii (C. Miill.) Zand. (Barbula), see treatment of
Chenia Am6
Chenia leptophylla (C. Miill.) Zand. (Phascum) Eur As2 As4 Oc
Afri Afr2 Afr4 Ami Am2 Am4 AmS Am6 Austri
[Chenia rhizophylla (Sak.) Zand., Phytologia 6S: 42S, I989
(Physcomitrium) Eur As2 As4 Oe Afr2 Afr4 Ami Am2 Am4
AmS Am6 Austri = Chenia leptophylla (C. Miill.) Zand., see
treatment of Chenia]
Chenia subohliqua (Williams) Zand., Phytologia 6S: 42S, 1989
(Tortula) Am4
CHIONOLOMA Dix. [= Pseudosymblepharis Broth. fide Eddy,
Handb. Males. Mosses 2: IS3, I99I]
[Chionoloma angustata (Mitt.) Menzel, Willdenowia 22: I97, I992
Pseudosymblepharis angustata (Mitt.) Hilp.]
[Chionoloma bartramii (Ther. ex Bartr.) Menzel, Willdenowia 22:
I98, I992 = Pseudosymblepharis hartramii Ther. ex Bartr.] =
Pseudosymblepharis schimperiana (Par.) Crum, see treatment
of Pseudosymblepharis]
[Chionoloma duriuscula (Mitt.) Menzel, Willdenowia 22: 198,
I992 Pseudosymblepharis duriuscula (Mitt.) Chen]
Chionoloma induratum Dix. As3 [= Pseudosymblepharis
suhduriuscula (C. Miill.) Chen fide Eddy, Handb. Males.
Mosses 2: IS3, 1991]
Chionoloma latifolium Dix. As3 [= Pseudosymblepharis
subduriuscula (C. Miill.) Chen fide Eddy, Handb. Males.
Mosses 2: IS3, 1991]
Chionoloma longifolium Dix. As4 [= Pseudosymblepharis
subduriuscu/a (C. Miill.) Chen fide Eddy, Handb. Males.
Mosses 2: 1S3, I99I]
[Chionoloma schimperiana (Par.) Menzel, Menzel, Willdenowia
22: I98, I992
Pseudosymblepharis schimperiana (Par.)
Crum]
[Chionoloma socotrana (Mitt.) Menzel, Willdenowia 22: I98, I992
= Pseudosymblepharis socotrana (Mitt.) Ther. Afr2 = Weissia
artocosana (Mitt.) Zand. nom. nov., see treatment of Weissia]
[Chionoloma suhduriuscula (C. Miill.) Menzel, Willdenowia 22:
I98, I992
Pseudosymblepharis suhduriuscula (C. Miill.)
Chen As2 As3 As4 = Pseudosymblepharis angustata (Mitt.)
Hilp.fide Norris & Koponen, Acta Bot. Penn. 137: 94, 1987]
=
=
=
=
[CINCLIDOTUS P. Beauv. referred to Cinclidotaeeae]
CROSSIDIUM Jur.
Crossidium aberrans Holz. & Bartr. Afri Ami Am2
fo. epilosum Flow., Bryologist 76: 289, I973 Ami
Crossidium apiculatum Magill, Fl. S. Afr. I. Mosses 1: I9S, 198I
[I982] Afr4
Crossidium asirense Frey & Kiirsehner, J. Bryol. 13: 2S, 1984 AsS
[Crossidium chloronotos (Brid.) Limpr. Eur As2 As3 AsS Afr1
Am1 Austr2 = Crossidium squamiferum (Viv.) Jur. fide
Delgadillo, Bryologist 78: 276, 197S]
Crossidium crassinerve (De Not.) Jur. Eur As3 Afr1 Am1 Am2
[var. laevipilum (Ther. & Trab.) Delg., Bryologist 78: 27S, 197S
(Crossidium) AsS Afrl Crossidium laevipilum Ther. & Trab.
fide Frey & Kiirehner, Cryptogamic Bryol. Liehenol. 12:
441-4SO, 1991]
Crossidium davidai Cateheside, Mosses S. Australia IS2, I980
Austri
Crossidium deserti Frey & Kiirsehner, Nova Hedw. 46: 87, 1988
(nom. nov. for Crossidium desertorum Frey & Kiirsehner) AsS
=
294
GENERA OF THE POITIACEAE
[Crossidium desertorum Holz. & Bartr. Am1 Am2 = Crossidium
crassinerve (De Not.) Jur. fide Delgadillo, Sharp et al., Moss Fl.
Mex.]
[Crossidium desertorum Frey & Kiirschner, Nova Hedw. 45: 132,
1987 hom. il/eg. As5 = Crossidium deserti Frey & Kiirschner,
Nova Hedw. 46: 87, 1988]
[Crossidium elatum Williams Am4 = Pseudocrossidium e/atum (Williams) Delg., Bryologist 78: 278, 1975]
[Crossidium erosum Ho1z. & Bartr. Am1 = Crossidium crassinerve
(De Not.) Jur.fide Delgadillo, Sharp eta!., Moss Fl. Mex.]
Crossidium geheebii (Broth.) Broth. Austr1 Austr2
Crossidium laevipilum Ther. & Trab. Afr1 As5 [= Crossidium
crassinerve var. /aevipilum (Ther. & Trab.) Delg. fide Delgadillo,
Bryologist 78: 275, 1975 but good species fide Frey & Ktirchner,
Cryptogamie Bryol. Lichenol. 12: 441-450, 1991]
Crossidium /axefi/amentosum Frey & Kiirschner, Nova Hedw. 45:
130, 1987 As5
[Crossidium peruvianum Broth. Am4 = Aloina rosei (Williams)
Delg.]
[Crossidium rosei Williams Am4 Am6 = Aloina rosei (Williams)
Delg.
Crossidium seriatum Crum & Steere Eur Am1 Am2
[Crossidium spatulaefolium Holz. & Bartr. Am4 Am6 = Crossidium
aberrans Holz. & Bartr. fide Delgadillo, Bryologist 78: 270,
1975]
Crossidium spiralifolium Magill, Fl. S. Afr. I. Mosses 1: 197, 1981
Afr4
Crossidium squamiferum (Viv.) Jur. Eur Asl As2 As3 As5 Afr1 Am1
Austr2
var. brevisetum (Besch.) Par. Afr1
var. /ongipilum Amann Eur
var. pottioideum (De Not.) Monic. Eur As3 Afrl Am1 Am2
var. pumilum Amann Eur
Crossidium woodii (Delgad.) Zand. (Pseuda/oina), see treatment of
Crossidium As5
CRUMIA Schof.
[Crumia deciduidentata Sharp & Iwats., J. Hattori Bot. Lab. 32: 95,
1969 Am1 = Tortula deciduidentata (Sharp & lwats.) Zand., see
treatment ofTortula]
Crumia latifolia (Kindb. ex Macoun) Schof., Canad. J. Bot. 44: 610,
1966 (Merceya) Am1
[DESMATODON Brid. = Tortula Hedw., see treatment ofTortula]
[Desmatodon africanus P. Yarde Afr2 = Scope/ophila cataractae
(Mitt.) Broth., see treatment of Scope/ophila]
[Desmatodon altipes Broth. As1 = Tortula altipes (Broth.) Zand., see
·
treatment of Tortula]
[Desmatodon argentinicus Broth. Am6 = Tortula argentinica (Broth.)
Zand., see treatment of Tortula]
[Desmatodon austrogeorgicus (Card.) Ochyra in Ochyra, Vitt &
Hort., Cryptog. Bryol. Lichenol. 7: 57, 1986 = Hennediella
austrogeorgica (Card.) Blockeel, J. Bryol. 16: 191, 1991]
[Desmatodon be/Iii Bartr. Am4 = Hennediella be/lei (Bartr.) Zand.,
see treatment of Hennediella]
[Desmatodon bogosicus C. Miill. Afr1 Afr2 Afr4 = Tortula bogosica
(C. MUll.) Zand., see treatment of Tortula]
[Desmatodon capillaris Chen As2 = Tortula capil/aris (Chen) Zand.,
see treatment of Tortulu]
[Desmatodon cernuus (Hiib.) BSG Eur As1 As2 Ami = Tortula
cernua (Hueb.) Lindb., see treatment of Tortula]
[var. xanthopus Kindh. Am1 = Tortula cernua var. xanthopus
(Kindh.) Zand., see treatment of Tortula]
[Desmatodon coloradensis Grout Am1 = Desmatodon obtusifolius
(Schwaegr.) Schimp. fide Crum & Anderson, Mo. E. N. Amer.
371, 1981 = Tortula obtusifolia (Schwaegr.) Math. fide Corley
eta!., J. Bryol. 11: 620, 1981 (1982)]
[Desmatodon convolutus (Brid.) Grout Eur Am1 Am2 Am3 Am4
Am6 Austr1 Austr2 Oc Afr1 Afr2 Afr3 Afr4 = Tortula
atrovirens (Sm.) Lindb. fide Corley et a!., J. Bryol. 11: 620,
1981 (1982)]
[var. edentulus (BSG) Grout Eur Afr4 = Tortula atrovirens var.
edentula (BSG) Schimp.]
[var. gasi/ienii (Vent.) Wijk & Marg. Eur = Tortula atrovirens
var. gasi/ienii (Vent.) Limpr.]
[var. /eucodontus (Corh.) Wijk & Marg. Eur = Tortula atrovirens
var. /eucodonta (Corh.) Zand., see treatment of Tortula]
[Desmatodon el/esmerensis Brass., Bryologist 74: 208, 1971 Am1
(= Pseudocrossidium revolutum (Brid. in Schrad.) Zand. fide
Zander, Phytologia 44: 204, 1979) = Pseudocrossidium
revolutum var. obtusulum (Lindh.) Tan, Zand. & T. Tayl.,
Lindbergia 7: 41, 1981]
[Desmatodon fisherae Crum, Bryologist 75: 360, 1972 Am 1 =Desmatodon porteri Jam. in Aust. fide Crum & Anderson, Mo. E.
N. Amer. 371, 1981 = Hennediella porteri (Jam. in Aust.)
Zand., see treatment of Hennediella]
[Desmatodon gemmascens Chen As2 = Syntrichia gemmascens
(Chen) Zand., see treatment of Syntrichia]
[var. hopeiensis Chen As2 = Syntrichia gemmascens var.
hopeiensis (Chen) Zand., see treatment of Syntrichia As2]
[Desmatodon guepinii BSG Eur Am1 Am2 = Tortula guepinii
(B.&S. in BSG) Broth., see treatment of Tortula]
[Desmatodon haussknechtii (Jur. & Milde) Frohl., Ann. Nat. Mus.
Wien 67: 155, 1964 As5 = Didymodon haussknechtii (Jur. &
Mild.) Broth.]
[Desmatodon heimii (Hedw.) Mitt. = Pottia heimii (Hedw.) Fiirnr.
in Hampe= Hennediel/a heimii (Hedw.) Zand., see treatment of
Hennediella]
[var. arcticus (Lindb.) Crum, Bryologist 72: 242, 1969 "arctica"
Am1 = Hennediella heimii var. arctica (Lindh.) Zand., see
treatment of Hennediella]
[Desmatodon hendersonii (Ren. & Card.) Williams in Millsp. &
Nutt. Am1 = Didymodon tophaceus (Brid.) Lisa fide Anderson
et al., Bryologist 93: 478, 1990]
[Desmatodon involutus Bartr. hom. illeg. As3 = Plaubelia
perinvoluta Zand., see treatment of Plaube/ia]
[Desmatodon kabir-khanii Broth. As3 = Tortula kabir-khanii
(Broth.) Zand., see treatment of Tortula]
[Desmatodon /atifo/ius (Hedw.) Brid. Eur Asl As2 As3 As5 Afr1
Am1 = Tortula euryphyl/a Zand. nom. nov., see treatment of
Tortula]
[suhsp. brevifo/ius (Kindb.) Kindh. Eur = Tortula euryphylla
suhsp. brevifolia (Kindh.) Zand., see treatment of Tortula]
[var. eucalyptratus (Lindb.) Kaur. Eur = Tortula euryphyl/a var.
euca/yptrata (Lindb.) Zand., see treatment of T ortula]
[var. flavescens (Brid.) Steud. Eur = Tortula euryphylla var.
flavescens (Brid.) Zand., see treatment of Tortula]
[var. muticus (Brid.) Brid. Eur As1 Am1 Am3 = Desmatodon
latifo/ius fo. fide Anderson et a!. 93: 478, 1990 = Tortula
euryphylla Zand., see treatment of Tortula]
[var. pilifer (Dicks.) Rabenh. Eur = Desmatodon /atifo/ius
(Hedw.) Brid. var. latifolius fide Lawton, Moss Fl. Pacif.
Northw. 95, 1971 Tortula euryphylla Zand., see treatment of
Tortula]
[var. spelaeus (Amann) Podp. Eur
Tortula euryphylla var.
spelaeus (Amann) Zand., see treatment of Tortula]
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
[var. subobliquus Lindb. Eur = Tortula euryphylla var. subobliqua
(Lindb.) Zand., see treatment of Tortula]
fo. eucalyptratus (Lindb.) Hag., Skand. Bladmfl. 204, 1939 (Tortula) Eur
[Desmatodon /aureri (Schultz) BSG Eur As1 As2 Afr4 Am1 =Tortula laureri (Schultz) Lindb., see treatment of Tortula]
[var. setschwanicus (Broth.) Chen As2 = Tortula laureri var.
setschwanica (Broth.) Zand., see treatment of Tortula]
[Desmatodon leucostoma (R. Br.) Berggr. Eur Am1 As1 As2 =Tortula /eucostoma (R. Br.) Hook. & Grev., see treatment of Tortula]
var. muticus (Lindb.) Sav.-Ljubits., Novosti Sist. Niz. Rast. 6: 248,
1969 [1970] (Desmatodon obliquus var.) Eur
[Desmatodon lingulatus (Hook. f. & Wils.) Sainsb. Austr2 = Didymodon lingulatus (Hook. f. & Wils.) Broth., see treatment of Didymodon]
[Desmatodon longipedunculatus (C. Mtill.) Magill, Fl. S. Afr. I.
Mosses 1: 210, 1981 (1982) (Barbula) Afr4 = Hennediella
longipedunculata (C. Mtill.) Zand., see treatment of Hennediella]
[Desmatodon meridiana/is Luis. Eur = Tortula meridiana/is (Luis.)
Mach.]
[Desmatodon obtusifolius (Schwaegr.) Schimp. Eur As1 As2 As5
Afrl Am1 = Tortula obtusifolia (Schwaegr.) Math. fide Corley et
al., J. Bryol. 11: 620, 1981 (1982)]
var. brevifolius Schimp. Eur
var.limbatus (Herz.) Wijk & Marg. Eur
[Desmatodon plinthobius Sull. & Lesq. Am1 = Tortula plinthobia
(Sull. & Lesq.) Broth. fide Zander, Bryologist 82: 551, 1979]
[Desmatodon porteri Jam. in Aust. Am1 = Tortula porteri (Jam. in
Aust.) Zand., see treatment of Tortula]
[Desmatodon randii (Kenn.) Laz. Eur Am1 = Tortula randii (Kenn.)
Zand., see treatment of Tortula]
[Desmatodon raucopapil/osus X.-j. Li, Acta Bot. Yunnan. 3: 105,
1981 As2 = Tortula raucopapil/osa (X.-j. Li) Zand., see treatment
of Tortula]
[Desmatodon recurvatus (Hook.) Mitt. (good species fide Catcheside,
Mosses S. Austr. 150, 1980) Afr4 Austr1 = Tortula recurvata
Hook.]
[Desmatodon reflexidens (Hampe) Jaeg. Austr1 = Desmatodon
recurvatus (Hook.) Mitt. fide Catcheside, Mosses S. Austr.
148,1980]
[Desmatodon solomensis Broth. As2 = Tortula solomensis (Broth.)
Zand., see treatment of Tortula]
[Desmatodon spathulifolius Bartr. Am2 = Neohyophila sprengelii
(Schwaegr.) Crum s. lat. fide Zander, Bryologist 86: 135, 1983 =
Plaubelia sprengelii (Schwaegr.) Zand., see treatment of
Plaubelia]
[Desmatodon sprenge/ii (Schwaegr.) Williams Am1 Am2 Am3 =
Neohyophila sprengelii (Schwaegr.) Crum, Bryologist 68: 470,
1965 = Plaubelia sprengelii (Schwaegr.) Zand., see treatment of
Plaubelia]
[Desmatodon steereanus Zand. & Crum, Bryologist 80: 638, 1977
Am4 = Hennediel/a steereana (Zand. & Crum) Zand., see treatment of Hennediella]
[Desmatodon stomatodontus (Card.) Williams Am2 Am3 Am5 =
Neohyophila sprengelii var. stomatodonta (Card.) Zand.,
Bryologist 86: 138, 1983 = Plaubelia sprengelii var.
stomatodonta (Card.) Zand., see treatment of Plaubelia]
[Desmatodon suberectus (Hook.) Limpr. Eur As1 As2 Am1 = Tortula
leucostoma (R. Brown) Berggr.]
[var. limbatus Amann Eur =Desmatodon wilczekii Meyl.J
fo. apiculatus (Lindb.) Podp., Consp. Muse. Eur. 241, 1954
(Desmatodon obliquus var.) Eur
fo. muticus (Lindb.) Podp., Consp. Muse. Eur. 241, 1954
295
(Desmatodon obliquus var.) Eur
fo. pilifer (Lindb.) Podp., Consp. Muse. Eur. 241, 1954
(Desmatodon obliquus var.) Eur
[Desmatodon systylius Schimp. Eur As1 As3 Am1 = Tortula
systylia (Schimp.) Lindb., see treatment of Tortula]
[Desmatodon thomsonii (C. Miill.) Jaeg. As2 As3 = Tortula
thompsonii (C. Miill.) Zand., see treatment of Tortula]
[Desmatodon tonkinensis Besch. As3 = Tortula tonkinensis
(Besch.) Zand., see treatment of Tortula]
[Desmatodon ucrainicus Laz. Eur = Tortula ucrainica (Laz.) Zand.,
see treatment of Tortula]
[Desmatodon viridipilus (Dix. & Sainsb.) Sainsb. Austr2 = Tortula
viridipila Dix. & Sainsb.]
[Desmatodon wilczekii Meyl. Eur = Tortula rhodonia Zand. nom.
nov., see treatment of Tortula]
[Desmatodon yuennanensis Broth. As2 = Tortula chungtienia Zand.
nom. nov., see treatment of Tortula]
DIALYTRICHIA (Schimp.) Limpr.
[Dialytrichia brebissonii (Brid.) Limpr. = Dialytrichia mucronata
(Brid.) Broth.]
var. pumila Amann Eur
[Dialytrichia fragillima Dix. & Sak. As2 = Barbula subcomosa
Broth. fide Saito, J. Hattori Bot. Lab. 39: 493, 1975]
Dialytrichia mucronata (Brid.) Broth. Eur As2 As5 Afr1
var. conferta (Corb.) Corb. in Pitard Eur Afr1
var.fragilifolia Biz. & Roux, Rev. Bryol. Lichenol. 36: 110, 1968
[1969] inval. holotyp. non cit. Eur
DIDYMODON Hedw.
Didymodon aaronis (Lor.) Guerra in Guerra & Ros, Cryptogamie
Bryol. Lichenol. 8: 55, 1987 (Trichostomum) Eur As5 Afrl
[Didymodon acutus (Brid.) Saito, J. Hattori Bot. Lab. 39: 519,
1975. Eur As1 As2 Afr1 Am1 = Didymodon rigidulus var.
gracilis (Schleich. ex Hook. & Grev.) Zand. fide Zander,
Cryptogamie Bryol. Lichenol. 2: 393, 1981 (1982)]
[var. icmadophilus (Schimp. ex C. Miill.) Zand., Phytologia 41:
20, 1978 (Barbula) = Didymodon rigidulus var. icmadophilus
(Schimp. ex C. Miill.) Zand. fide Zander, Cryptogamie Bryol.
Lichenol. 2: 394, 1981 (1982)]
[var. ditrichoides (Broth.) Zand., Phytologia 41 : 20, 1978 (Barbula ditrichoides) As3 Ami = Didymodon rigidulus var.
ditrichoides (Broth.) Zand., see treatment of Didymodon]
[Didymodon afrorubellus Broth. & Wag. ex Dix. Afr4 = Bryoerythrophyllum afrorubellum (Broth. & Wag.) DeSloover, Bull.
Jard. Bot. Natl. Belg. 49: 398, 1979]
Didymodon alticaulis Bartr. Am2
Didymodon amblyophyllus (Hook.) Broth., Nat. Pfl. 1(3): 406, 1902
Am5 Am6
Didymodon ampliretis Card. & Broth. Am6
Didymodon andreaeoides Card. & Broth. Am6
[Didymodon angustifolius Bartr. hom. il/eg. Am4 = Didymodon
bartramii Zand. nom. nov., see treatment of Didymodon]
[Didymodon angustulus Herz. Am4 = Bryoerythrophyllum
angustulum (Herz.) Robins. fide Robinson, Bryologist 70: 22,
1967 = Bryoerythrophyllum campylocarpum (C. Miill.) Crum
fide Robinson, Smithsonian Contr. Bot. 27: 31, 1975]
Didymodon anserinocapitatus (X.-j. Li) Zand. (Barbula), see treatment of Didymodon As2
[Didymodon arcticus (Kaal.) Broth. Eur As1 Ami= Trichostomum
arcticum Kaal., cf. Frisvoll, J. Bryol. 13: 435, 1985]
Didymodon argentinicus (Par.) Par. Am4 Am6
Didymodon argentiniensis Warnst. Am6
296
GENERA OF THE POTTIACEAE
Didymodon asperifolius (Mitt.) Crum, Steere & Anders., Bryologist
67: 163, 1964 (Barbula) Eur Asl As2 As3 Ami
Didymodon australasiae (Hook. & Grev.) Zand., Phytologia 41: 21,
1978 (Trichostomopsis) Eur Afrl Afr4 Aml Am2 Am3 Am4
Am6 Austrl Austr2
[var. umbrosus (C. Miill.) Zand., Cryptogamie Bryol. Lichenol. 2:
400, 1981 [1982] (Barbula) Eur Ami Am2 Am4 Am6 Didymodon umbrosus (C. Miill.) Zand.]
fo. propaguliferus Brugues ex Guerra in Guerra & Ros,
Cryptogamie Bryol. Lichenol. 8: 55, 1987 Eur
Didymodon austroalpigena (C. Miill.) Broth. Afr4
[Didymodon atrorubens (Besch.) Broth. = Bryoerythrophyllum
wallichii (Mitt.) Chen fide Saito, J. Hattori Bot. Lab. 39: 479,
1975]
Didymodon barbuloides Libert ex March. Eur
Didymodon bartramii Zand. nom. nov. (Didymodon angustifolius
Bartr. hom. illeg.), see treatment of Didymodon Am4
Didymodon berthoanus Ther. Am6
Didymodon brachyphyllus (Sull. in Whipp!.) Zand., Phytologia 41:
24, 1978 (Barbula) Am1 l= Didymodon vinealis var.
brachyphyllus (Sull. in Whipp!.) Zand., Cryptogamie Bryol.
Lichenol. 2: 411, 1981 (1982)]
[Didymodon brachystegius (Besch.) Broth. As2 = Bryoerythrophyllum brachystegium (Besch.) Saito, J. Jap. Bot. 47: 14, 1972]
Didymodon brotheri (Ren. & Par.) Zand. (Trichostomum), see treatment of Didymodon Afr3
Didymodon brunneus (C. Miill.) Wamst. in Broth. Am6
Didymodon calycinus Dix. Austr2
[Didymodon calymperidictyon Broth. in Skottsb. Am6 =Bryoerythrophyllum campylocarpum (Tayl.) Crum fide Robinson, Smithsonian Contr. Bot. 27: 30, 1975]
Didymodon canaliculatus Dix. As3 [= Barbula canaliculata (Dix.) R.
S. Chopra, Taxon. Indian Mosses 138, 1975]
[Didymodon capil/aceus (Hedw.) Web. & Mohr
Distichium
capillaceum (Hedw.) BSGJ
var. elongatus Brid. ignot.
Didymodon capitatus Dix. ex Sappa & Piov. Eur?
Didymodon cardotii (Dus.) Zand., see treatment of Didymodon Am6
Didymodon catenulatus Dix. As3
Didymodon ceratodonteus (C. Miill.) Dix. Afr2 Afr4
Didymodon challaense (Broth. in Herz.) Zand., see treatment of DidymodonAm4
[Didymodon chimborazensis (Mitt.) Broth. in Par. Am4 Bryoerythrophyllum chimborazense (Mitt.) Zand., see treatment of Bryoerythrophyllum]
Didymodon coffeanus Norris & T. Kop., Acta Bot. Fenn. 137: 125,
1989 As4
[Didymodon columbianus Herm. & Lawt. Aml =Bryoerythrophyllum
columbianum (Herm. & Lawt.) Zand.fide Zander, Bryologist 81 :
548, 1987 (1979)]
Didymodon constrictus (Mitt.) Saito, J. Hattori Bot. Lab. 39: 514,
1975 As2 [= Didymodon vinea/is (Brid.) Zand. fide Sollman,
Bryo1ogist 86: 271, 1983 J
var.flexicuspis (Chen) Saito, J. Hattori Bot. Lab. 39: 516, 1975 As2
[= Didymodon vinealis (Brid.) Zand. fide Sollman, Bryo1ogist 86:
272, 1983]
Didymodon contortus Herz. Am4
Didymodon cordatus Jur. Eur
subsp. austriacus (Schiffn. & Baumg.) Wijk & Marg. Eur
var. austriacus (Schiffn. & Baumg.) Latz. Eur
var. latifolius (Roll) Roll Eur
var. longifolius Roll Eur
fo. brevicaulis Roll, Deutsche Bot. Monatsschr. 3: 47, 1885 Eur
=
=
=
fo. gracilis Roll, Deutsche Bot. Monatsschr. 3: 47, 1885 Eur
[fo. /atifolius Roll, Deutsch. Bot. Monatsh. 3: 46, 1885 Eur
Didymodon cordatus var. latifolius (Roll) Roll]
[fo. latifolius Roll, Deutsche Bot. Monatsschr. 3: 46, 1885 Eur
Barbula cordata var. latifolia (Roll) Podp. fide Podp~ra, Consp.
Didymodon cordatus var. latifolius
Muse. Eur. 205, 1954
(Roll) Roll]
fo. longicaulis Roll, Hedwigia 56: 145, 1915 Eur
fo. ramosus Roll, Deutsche Bot. Monatsschr. 3: 57, 1885 Eur
fo. robustus Roll, Deutsche Bot. Monatsschr. 3: 58, 1885 Eur
fo. strictus Roll, Deutsche Bot. Monatsschr. 3: 57, 1885 Eur
fo . tenellus Podp. in Latz., Bot. Centralbl. Beih. 48(2): 477, 1931
Eur
[fo. typicus Roll, Deutsche Bot. Monatsschr. 3: 47, 1885 nom.
illeg. Eur =Didymodon cordatus Jur. var. cordatus]
[Didymodon craspedophyllus Card. Am2 = Didymodon australasiae (Hook. & Grev.) Zand. s. lat. fide Zander, Cryptogamie
Bryol. Lichenol. 2: 397, 1981 (1982)]
Didymodon crassicostatus (Bartr.) Zand. (Barbula), see treatment
of Didymodon Am2
Didymodon deciduus Zand. (nom . nov. for Barbula decidua C.
Miill.), see treatment of Didymodon Am6
[Didymodon decolorans (Hampe) Williams Am4 = Trichostomopsis australasiae (Hook. & Grev.) Robins ., Phytologia 20:
187, 1970 Didymodon australasiae (Hook. & Grev.) Zand.,
Phytologia41: 21, 1978]
var. obtusus Herz. Am4
[Didymodon diaphanobasis Card. Aml = Trichostomopsis
australasiae (Hook. & Grev.) Robins., Phytologia 20: 187,
Didymodon australasiae (Hook. & Grev.) Zand.,
1970
Phyto1ogia41: 21, 1978]
[var. angustifolius Ther. in Bartr. Am1 = Trichostomopsis
australasiae (Hook. & Grev.) Robins., Phytologia 20: 187,
1970
Didymodon australasiae (Hook. & Grev.) Zand.,
Phytologia 41: 21, 1978]
[Didymodon dimorphus (C. Miill.) Broth. (not Gymnostomum
dimorphum (C. Mii11.) Simfide Magill & Schelpe, 1979) Afr4 =
Didymodon ceratodonteus (C. Miill.) Dix. fide Magill, Fl. S.
Afr. I. Mosses 1: 233, 1981 (1982)]
[Didymodon distans (Hampe in C. Miill.) Jaeg. Afr2 Trichostomum distans Hampe in C. Miill., see treatment of Trichostomum]
[Didymodon dixonii Wadhwa & Yohra, Curr. Sci. 32: 483, 1963
As3 (nom. nov. for Didymodon obtusifolius Card. ex Dix. & P.
Yard.) As3 = Barbula dixonii R. S. Chopra, Taxon. Indian
Mosses 139, 1975 nom. nov. = Barbula inaequalifolia Tayl.
fide Sollman, Lindbergia 10: 54, 1984 Bryoerythrophyllum
inaequa/ifolium (Tayl.) Zand.]
Didymodon dubius (Schwaegr.) Par. Austr1
Didymodon eckendorfii P. Yarde Afr2
[Didymodon ehrenbergii (Lor.) Kindb. Barbula ehrenbergii (C.
Miill.) Wijk & Marg.J
[fo. densiretis Amann, Fl. Mousses Suisse 3: 27, 1933 Eur
Hydrogonium ehrenbergii fo. densirete (Amann) Podp., Consp.
Muse. Eur. 199, 1954]
[fo . laxiretis Amann, Rev. Bryol. 51 : 10, 1924 Eur =
Hydrogonium ehrenbergii fo. /axirete (Amann) Podp., Consp.
Muse. Eur. 199, 1954]
Didymodon erosodenticu/atus (C. Miill.) Saito, J. Hattori Bot. Lab.
39: 504, 1975 (Barbula) As2
Didymodonfallax (Hedw.) Zand., Phytologia 41: 28, 1978 (Barbula) Eur Asl As2 As3 As5 Afr1 Ami
var. brevifolius (Dicks. ex With.) Ochyra, Fragm. Fl. Geobot. 28:
=
=
=
=
=
=
=
=
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
449, 1982 [1984] (Bryum) Eur
[var. reflexus (Brid.) Zand., Bryologist 83: 230, 1980 (Barbula) Eur
As1 As2 Am1 Am2 = Didymodon rigidicaulis (C. Mi.ill.) Saito
Didymodon
fide Saito, J. Hattori Bot. Lab. 39: 502, 1975
ferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11: 599, 1981
(1982)]
Didymodonferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11: 599,
1981 (1982) (Barbula) As1 As2 Am1 Am2 Am3 Eur Afr [=
Didymodon fa/lax var. reflexus (Brid.) Zand. fide Zander,
Bryologist 83: 230, 1980]
[Didymodon filicau/is Card. Am2 = Bryoerythrophyllum
recurvirostrum (Hedw.) Chen var. recurvirostrum fide Zander,
Cryptogamic Bryol. Lichenol. 2: 418, 1981 (1982)]
Didymodonfragilicuspis Broth. As2
[Didymodon fuscoviridis Card. Am1 Am2 Am3 = Didymodon
rigidu/us Hedw. s. lat.fide Zander, Cryptogamic Bryol. Lichenol.
2: 390, 1981 (1982)]
Didymodon gelidus Card. Ant
[Didymodon gemmascens Mitt. in Hunt, Mem. Litt. Phil. Soc. Manchester ser. 3, 3: 235, 1868 [as "gemmescens"] Eur Leptodontium gemmascens (Mitt. in Hunt) Braithw. good species fide
Zander, Bryologist 75: 236, 1972]
Didymodon giganteus (Funck) Jur. Eur As2 As3 Am1
[var. japonicus Broth. ex lhs. As2 = Barbula gigantea Funck fide
Iwats. & Nog., J. Hattori Bot. Lab. 37: 308, 1973 Didymodon
giganteus (Funck) Jur.]
Didymodon glaucoviridis (C. Mi.ill.) Broth. Am5
Didymodon glaucus Lindb. ex Roth Eur hom. il/eg.
[Didymodon godmanianus (C. Mi.ill.) Bartr. Am2 = Didymodon
rigidulus Hedw. s. lat. fide Zander, Cryptogamic Bryol. Lichenol.
2: 390, 1981 (1982)]
Didymodon gracilescens Bartr. in Bauer hom. illeg. Am6
Didymodon guineensis Broth. & Par. Afr2
Didymodon gymnus (C. Mi.ill.) Broth. Am4 Am6
Didymodon hampei Zand. (nom. nov. for Gyroweisia obtusifolia
Hampe), see treatment of Didymodon Am2
Didymodon hastatus (Mitt.) Zand. (Barbu/a), see treatment of Didymodon As3
Didymodon haussknechtii (Jur. & Mild.) Broth. As5
[Didymodon heribaudii Card. Am2 = Didymodon rigidu/us Hedw. s.
lat. fide Zander, Cryptogamic Bryol. Lichenol. 2: 390, 1981
(1982)]
[Didymodon homomallus Hedw. = Ditrichum heteromallum (Hedw.)
Britt.]
var. e/ongatus Hi.ib. Eur
Didymodon humbo/dtii (Herz.) Hegew. & Hegew., Nov. Hedw. 28:
747, 1976 (Barbula) Am4
Didymodon humidus (Mitt.) Zand. (Tortula), see treatment of DidymodonAm4
[Didymodon icmadophilus Schimp. ex C. Mi.ill.) Saito, J. Hattori Bot.
Lab. 39: 519, 1975 (as "icmadophy/lus") (Barbula) As1 As2 As3
Eur Am1 Am2 = Didymodon rigidulus var. icmadophilus
(Schimp. ex C. Mi.ill.) Zand. fide Zander, Cryptogamic Bryol.
Lichenol. 2: 394, 1981 (1982)]
Didymodon imperfectus (C. Mi.ill.) Zand. (Trichostomum), see treatment of Didymodon Am4
Didymodon incrassatolimbatus Card. Am2
Didymodon incrassatus (Broth.) Broth. Eur
[Didymodon insulanus (De Not.) Hill, J. Bryol. 11: 599, 1981 (1982)
(Tortula) Eur = Didymodon vinealis var. flaccidus (BSG) Zand.
cf. Corley et al., J. Bryol. 11: 622, 1981 (1982) (= Didymodon
vinealis (Brid.) Zand.fide Sollman, Bryologist 86: 272, 1983)]
Didymodon integrifolius Broth. in Mildbr. Afr2
=
=
=
297
[var. paucidentatus Ther. Afr2 = Bryoerythrophyllum
campy/ocarpum (C. Mi.ill.) Crum, see treatment of Bryoerythrophyllum]
Didymodon inundatus (Mitt.) Broth. Am4 Am6
Didymodon japonicus (Broth.) Saito, J. Hattori Bot. Lab. 39: 508,
1975 (Molendoa) As2
Didymodonjohansenii (Williams) Crum Eur Ami
Didymodonjuniperinus (C. Mi.ill.) Broth. Am4
[Didymodon killipii Williams Am4 = Morinia ehrenbergiana (C.
Mi.ill .) Ther., see treatment of Morinia = Mironia
ehrenbergiana (C. Mi.ill.) Zand., see treatment of Mironia]
[Didymodon knysnae Rehm. ex Sim Afr4 = Didymodon
xanthocarpus (C. Mi.ill.) Magill in Magill & Schelpe, Mem.
Bot. Surv. S. Afr. 43: 5, 1979]
Didymodon laevigatus (Mitt.) Zand., Phytologia 41: 29, 1978 (Tortula) Am4 Am6
Didymodon lamyanus (Schimp.) Ther. Eur
Didymodon leskeoides Saito, J. Hattori Bot. Lab. 39: 508, 1975 As2
Am1
[Didymodon linearis Broth. in Skottsb. hom. illeg. Am5 = Bryoerythrophyllum campylocarpum (C. Mi.ill.) Crum fide Robinson,
Smiths. Contr. Bot. 27: 30, 1975]
Didymodon lindigii (Hampe) Zand. (Hyophi/a), see treatment of
Didymodon Am4
Didymodon lingulatus (Hook. f. & Wils.) Broth. Austr2
[Didymodon lingulatus (Rehm. ex Sim) Magill, Mem. Bot. Surv. S.
Afr. 43 : 5, 1979 (Gymnostomum) hom. il/eg. Afr4 = Gymnostomum lingulatum Rehm. ex Sim fide Magill, Fl. S. Afr. I.
Mosses 1: 184, 1981 (1982)]
Didymodon loeskei Aeisch. As4
[Didymodon /ongifolius (Brid.) Hook. - Leucoloma longifolia
(Brid.) Wijk. & Marg.]
var. curvifolius Hook. f. & Wils. Am6
Didymodon /orentzianus (C. Mi.i11.) Broth. Am6
Didymodon luehmannii (Broth. & Geh.) Catcheside, Mosses S.
Australia 175, 1980 (Barbula) Austrl
Didymodon luridus Homsch. in Spreng. Eur As1 As2 As5 Afr1
Afr2 Am2 Am3 [= Didymodon vinealis var. luridus (Homsch.
in Spreng.) Zand.fide Zander, Cryptogamic Bryol. Lichenol. 2:
412, 1981 (1982)]
var. latifolius Hill Eur
fo. brevifolius Latz., Bot. Centralb. Beih. 48(2): 477, 1931 Eur
fo. cuspidatus (Schimp.) Roll, Hedwigia 56: 144, 1915 Eur
fo. rube/Ius Loeske in Bauer, Muse. Eur. Exs. 1581, 1923 Eur
fo. subscabrus Linder, Mitt. Bad. Landesvereins Naturk.
234-235: 265, 1909 Eur
fo. tophaceus Amann, Fl. Mousses Suisse Ad. 3: 26, 1935 Eur
[Didymodon luzonensis Bartr. As4 = Bryoerythrophyllum ferruginascens (Stirt.) Giac., see treatment of Bryoerythrophyllum]
Didymodon macrophyllus Broth. in Herz. Am4
Didymodon jackvancei Zand. nom. nov. (Husnotiella plicata
Magill), see treatment of Didymodon] Afr4
Didymodon mamillosus (Crundw.) Hill, J. Bryol. 11: 599, 1981
[1982] (Barbula) Eur
Didymodon marginatum (Robins.) Zand. (Trichostomum), see treatment of Didymodon Am4
Didymodon maschalogena (Ren. & Card.) Broth. As3
Didymodon maximus (Syed & Crundw.) Hill, J. Bryol. 11: 599,
1981 [1982] (Barbula) Am1 Eur
[Didymodon merceyoides Broth. in Herz. Am4 = Bryoerythrophyllum campylocarpum (C. Mi.ill.) Crum, see treatment of Bryoerythrophyllum]
[Didymodon mexicanus Besch. Am1 Am2 = Didymodon rigidulus
GENERA OF THE POTTIACEAE
298
Hedw. s. lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 389,
1981 (1982)]
[var. subulatus Ther. & Bartr. Am1 Am2 Didymodon rigidulus
var. subulatus (Ther. & Bartr.) Zand. fide Zander, Cryptogamie
Bryol. Licbenol. 2: 395, 1981 (1982)]
Didymodon michiganensis (Steere in Grout) Saito, J. Hattori Bot.
Lab. 39: 517, 1975 (Barbula) As2 As3 Am1 Am2
[Didymodon microstomus Dix. & Badhwar As3 = Bryoerythrophyllum recurvirostrum (Hedw.) Chenfide Gangulee, Mosses E. India
3: 742, 1972 also fide Sollman, Lindbergia 16: 22, 1990]
Didymodon microthecius (C. Miill.) Broth. Am6
Didymodon minusculus (Williams) Zand. (Tortula), see treatment of
Didymodon Am4
Didymodon mittenii Gangulee, Nov. Hedw. 8: 149, 1964. As3
[Didymodon montanus (Mitt.) Broth. Am4 Rhamphidium montanus
(Mitt.) Zand. see Excluded Taxa]
Didymodon monrevidensis Broth. in Felipp. Am5
Didymodon moritzianus (C. Miill.) Broth. Am4
Didymodon nicholsonii Culm. Eur Am1
Didymodon nigrescens (Mitt.) Saito, J. Hattori Bot. Lab. 39: 510,
1975 (Barbula) As2 Am1 Am2
[Didymodon obscurus (Mitt.) Dix. As3 hom. illeg. = Didymodon
mittenii Ganguleefide Gangulee, Mosses E. India 758, 1972]
[Didymodon obtusifolius Card. ex Dix. & P. Yarde As3 hom. illeg.
non Schkuhr = Barbula dixonii R. S. Chopra, Taxon. Indian
Mosses 139, 1975 nom. nov. = Barbula inaequalifolia Tayl. fide
Sollman, Lindbergia 10: 54, 1984
Bryoerythrophyllum
inaequallfolium (Tayl.) Zand.]
Didymodon obtussissimus Broth. = Bryoerythrophyllum
brachystegium (Besch.) Saito fide Saito, J. Jap. Bot. 47: 14, 1972
and J. Hattori Bot. Lab. 39: 477, 1975]
var. japonicus Broth. As2
Didymodon occidentalis Zand., Phytologia 41: 26, 1978 (nom. nov.
for Barbula rubiginosa) Aml [:: Didymodon vinealis var.
rubiginosus (Mitt.) Zand., Cryptogamie Bryol. Lichenol. 2: 417,
1981]
Didymodon orbignyanus (C. Miill.) Broth. Am6
[Didymodon ovatus (Mitt.) Jaeg. As3 = Bryoerythrophyllum
recurvum (Griff.) Saito fide Sollman, Lindbergia 16: 22, 1990
Bellibarbula recurva (Griff.) Zand., see treatment of Bellibarbula]
[Didymodon papillinervis (Dix. & Herz.) Demar. Afr2 = Bryoerythrophyllum recurvirostrum (Hedw.) Chen fide Sollman, Lindbergia
16: 22, 1990]
[Didymodon parvu/us (Kindb.) Britt. in Williams Am1 = Distichium
inc/inatum (Hedw.) BSG fide Steere & Crum, Mem. New York
Bot. Gard. 28(2): 76, 1977]
Didymodon patagonicus (Mitt.) Broth. Am6
[Didymodon patentifolius Ther. Am2 = Didymodon australasiae
(Hook. & Grev.) Zand. s. lat. fide Zander, Cryptogamie Bryol.
Licbenol. 2: 397, 1981 (1982)]
[Didymodon paucidentatus (C. Miill.) Broth. in Par. Afr2 = Bryoerythrophyllum campylocarpum (C. Miill.) Crum fide Sollman,
Lindbergia 16: 23, 1990]
[Didymodon pelichucensis Williams Am4 = Bryoerythrophyllum
campylocarpum (C. Miill.) Crum, see treatment of Bryoerythrophyllum]
[Didymodon percarinatus Dix. & Sak. As2 = Grimmia sp. fide Saito,
J. Hattori Bot. Lab. 39: 528, 1975]
Didymodon perexilis (C. Miill.) Broth. Am4
Didymodon perobtusus Broth. As2 Am1
[Didymodon perrevolutus P. Yarde Afr2 =Didymodon ceratodonteus
(C. Miill.) Dix. fide Magill, Fl. S. Afr. I. Mosses 1: 235, 1981
=
=
=
=
(1982)]
[Didymodon planifolius P. Yarde & Ther. Am3 = Trichostomum
brachydontium Bruch in F. Miill. fide Zander, Cryptogamie
Bryol. Lichenol. 2: 419, 1981 (1982)]
Didymodon planotophaceus Frohl. AsS
Didymodon polycephalus Mont. Am6
Didymodon pruinosus (Mitt.) Zand., see treatment of Didymodon
(Tortula) Am4
[Didymodon pusillus Card. hom. illeg. Am2 =Didymodon rigidulus
Hedw. s. lat. fide Zander, Cryptogamie Bryol. Lichenol. 2: 390,
1981 (1982)]
[Didymodon ramulosus (Schimp. ex Besch.) Card. in Broth. Am2 =
Didymodon rigidulus Hedw. s. lat. fide Zander, Cryptogamie
Bryol. Lichenol. 2: 389, 1981 (1982)]
[Didymodon recurvus (Griff.) Broth. As3 = Bryoerythrophyllum
recurvum (Griff.) Saito Bellibarbula recurva (Griff.) Zand.,
see treatment of Bellibarbula]
Didymodon reedii Robins., Bryologist 70: 323, 1967 Am1 Eur
Didymodon reflexus Ther. Am6
Didymodon reticularus Gillies ex Grev. Am6
Didymodon revolutus (Card.) Williams fide Zander, Cryptogamie
Bryol. Lichenol. 2: 401, 1981 [1982] (Husnotiella) Aml Am2
Am4
[Didymodon rigidicaulis (C. Miill.) Saito, J. Hattori Bot. Lab. 39:
502, 1975 (Barbula) As1 As2 Am1 Am2 Am3 Eur Afr =Didymodonfallax var. rejlexus (Brid.) Zand.fide Hill, J. Bryol. 11:
599, 1981 (1982) = Didymodon ferrugineus (Schimp. ex
Besch.) Hill, J. Bryol. 11:599, 1981 (1982)]
[Didymodon rigidifolius Dix. Afr2 =Bryoerythrophyllum recurvum
(Griff.) Saito fide Sollman, Lindbergia 16: 22, 1990 Bellibarbula recurva (Griff.) Zand., see treatment of Bellibarbula]
Didymodon rigidulus Hedw. Eur As1 As2 As3 AsS Afr1 Am 1 Am6
subsp. andreaeoides (Limpr.) Wijk & Marg. Eur As2
[subsp. validus (Limpr.) Loesk. Eur = Didymodon rigidulus var.
validus (Limpr.) R. Diill, J. Hattori Bot. Lab. 55: 263, 1984 =
Didymodon rigidulus var. gracilis (Schleich. ex Hook. &
Grev.) Zand. fide Zander, Cryptogamie Bryol. Licbenol. 2: 393,
1981 (1982)]
.
[subsp. verbanus (Nichol. & Dix.) Loesk. Eur = Didymodon
glaucus Ryan fide Corley et al., J. Bryol. 11: 622, 1981 = Didymodon rigidulus var. glaucus (Ryan) Wijk & Marg.]
var. acutus Biz., Acta Bot. Acad. Sci. Hung. 18: 21, 1973 Afr2
var. brevifolius Zodd. illeg. hom. Afr1
var. campicola (Broth.) Wijk & Marg. As1
var. compactus Roll Eur
[var. excurrens Hag. Eur = Didymodon rigidulus fo. excurrens
(Hag.) Diill, J. Hattori Bot. Lab. 55: 259, 1984]
var. flaccidus (Roll) Roll Eur
var. glaucus (Ryan) Wijk & Marg. Eur As1
var. gracilis (Schleich. ex Hook. & Grev.) Zand., Cryptogamie
Bryol. Lichenol. 2: 393, 1981 (Tortula) Eur As1 As3 As5 Afr1
Am1Am2
var. icmadophilus (Schimp. ex C. Miill.) Zand., Cryptogamie
Bryol. Licbenol. 2: 394, 1981 (Barbula) Eur As1 As3 AsS Am1 '
Am2
var. insidiosus (Jur. & Mild.) Roll, Hedwigia 56(1-3): 149, 1915
(Barbula rigidulus var.) Eur = Didymodon spadiceus (Mitt.)
Limpr.fide Limpricht, Laubm. 556, 1890 (1888)]
var. major Podp., Sitzungsber. K. Boehm. Ges. Wiss. 1899(44):
14, 1899 Eur
var. paludosus (Lang.) Par. Eur Am 1
var. rigidus (Roll) Roll Eur
var. subulatus (Ther. & Bartr.) Zand., Cryptogamie Bryol.
=
=
299
GENERA, SPECIES AND INFRASPECIFIC TAXA
Lichenol. 2: 395, 1981 (Didymodon mexicanus var.) Am1 Am2
var. tenuis Hamm. Eur
var. trifarius Hartm. Eur
[var. validus (Limpr.) R. Diill, J. Hattori Bot. Lab. 55: 263, 1984
Eur = Didymodon rigidulus var. gracilis (Schleich. ex Hook. &
Grev.) Zand. fide Zander, Cryptogamie Bryol. Lichenol. 2: 393,
1981 (1982)]
fo. bisetus Pet. Magyar Bot. Lapok 2: 292, 1903 Eur
fo. brevicaulis (Roll) Roll, Hedwigia 56: 147, 1915 (Barbula
rigidula fo.) Eur
fo. brevifolius Roll, Hedwigia 42: 299, 1903 Eur
fo. densus (B.&S. in BSG) Limpr., Laubm. Deutsch. 1: 556, 1888
(Trichostomum rigidulum var.) Eur
fo. excurrens (Hag.) Diill, J. Hattori Bot. Lab. 55: 259, 1984 (Didymodon rigidulus var.) Eur
fo. laxus Mol., AuHiu-Studien 65, 1864 Eur
fo. /ongicau/is (Roll) Roll, Hedwigia 56: 147, 1915 (Barbula
rigidula fo.) Eur
fo. propagu/iferus (Schiffn.) Limpr., Laubm. Deutsch. 3: 691, 1901
Eur
subfo. viridis (Roll) Roll, Hedwigia 56: 147, 1915 (Barbula
rigidula fo.) Eur
Didymodon rivico/a (Broth.) Zand. in Kop., Gao, Lou & Jarvinen,
Ann. Bot. Fenn. 20: 222, 1983 (Barbula) As2
[Didymodon rubellus BSG = Bryoerythrophyllum recurvirostrum
(Hedw.) Chen]
var. angustifolius P. Yarde Afr2 = Bryoerythrophyl/um
afrorubellum (Broth. & Wag.) De Sloover, Bull. Jard. Bot. Natl.
Belg. 49: 398, 1979]
[var. subdentatus Ther., Rev. Bot. 9: 486, 1892 Eur Didymodon
rubellus fo. subdentatus (Ther.) Herib., Mem. Ac. Sc. ClermontFerrand ser. 2, 14: 382, 1899]
[fo. brevirostris Warnst., Krypt. Fl. Brandenburg 224, 1904 Eur
Bryoerythrophyllum recurvirostrum fo. brevirostre (Warnst.)
Podp.]
[fo.flaccidus Roll, Hedwigia 38: 261, 1899 Eur =Bryoerythrophyllum recurvirostrum fo. flaccidum (Roll) Podp.]
[fo. gracilis Limpr., Laubm. Deutsch. 1: 547, 1888 Eur = Bryoerythrophyllum recurvirostrum fo. gracile (Limpr.) Podp.]
[fo. longirostris Warns!., Krypt. Fl. Brandenburg 224, 1904 Eur
Bryoerythrophyllum recurvirostrum fo.]
[fo. maior Breidl. ex Matous., Hedwigia 44: 29, 1904 Eur Bryoerythrophyllum recurvirostrum fo. maius]
[fo. obtusifolius Roll, Hedwigia 38: 261, 1899 Eur Bryoerythrophyllum recurvirostrum fo. obtusifolium (Roll) Podp.]
[fo. pygmaeus Meyl., Bull. Soc. Yaud. 57: 123, 1929 = Bryoerythrophy/lum recurvirostrum fo. pygmaeum (Meyl.) Podp.]
fo. subdentatus Ther. in Herib., Mem. Ac. Sc. Clermont-Ferrand
ser. 2, 14: 382, 1899 Eur
[fo. viridis Schlieph. ex Limpr., Laubm. Deutsch. 1: 547, 1888 Eur
Bryoerythrophyllum recurvirostrum fo. viride (Schlieph. ex
Limpr.) Podp]
Didymodon rubiginosus (C. Miill.) Broth. Austr1
Didymodon rufidu/us (C. Miill.) Broth. As2
[Didymodon rufus Lor. in Rabenh. = Didymodon asperifolius (Mitt.)
Crum, Steere & Anders.)]
var. gorodkovii A. Abr. & I. Abr. in I. Abr., Trudy Arktic.
Antarktic. Naucno-Issl. Inst. 224: 220, 1963 As1
fo. sublaevigatus Herz., Weiner Bot. Z. 93 : 39, 1914 Eur
Didymodon schilleri Herz. & Ther. Am6
[Didymodon schimperi (Mont.) Broth. Am6 = Tortula atrovirens
(Sm.) Lindh., see treatment of Tortula]
[Didymodon saxico/a Broth. ex Gangulee, Mosses E. India 774, 1972
=
=
=
=
=
=
nom. nud. in syn. = Desmatodon gemmascens Chen fide
Gangulee, Mo. E. India 3: 774, 1972]
[Didymodon semivaginatus (Britt.) Broth. Am4 =Erythrophyllopsis
boliviano Broth. in Herz., see treatment of Erythrophyllopsis]
Didymodon sinuosus (Mitt.) Delogn. Eur
Didymodon soaressii Luis. Eur
Didymodon spadiceus (Mitt.) Limpr. Eur As1 As5
subsp. poeninus (Amann) Wijk & Marg. Eur
var. siluricus Yelen. Eur
Didymodon spathulatolinearis (C. Miill.) Broth. Am6
[Didymodon stenopyxis Card. Am2 = Bryoerythrophyl/um
recurvirostrum var. aeneum (C. Miill.) Zand. fide Zander in
Sharp, Moss Fl. Mexico]
Didymodon stewartii (Bartr.) Zand. (Barbula), see treatment of
Didymodon As3
[Didymodon strictifolius Dix. & P. Yarde As3 = Qarbula strictifolia
(Dix. & P. Yarde) R. S. Chopra, Taxon. Indian Mosses 140,
1975 = Didymodon recurvus (Griff.) Mitt. fide Robinson,
Bryologist 71: 86, 1968 Bellibarbu/a recurva (Griff.) Zand.,
see treatment of Bellibarbula]
Didymodon subandreaeoides (Kindb.) Zand., Phytologia 41: 23,
1978 (Barbula) Am1
Didymodon subfontanus Dix. in Sim Afr4
[Didymodon sub/ingulatus Dix. Afr2 = Bryoerythrophyllum
campylocarpum (C. Mii11.) Crumfide Sollman, Lindbergia 16:
22, 1990]
Didymodon subrevolutus (Hampe) Broth. Afr3
Didymodon subtriquetrus Robins., Bryo1ogist 70: 319, 1967 Am4
[Didymodon subtophaceus Williams Am4 Am6 = Trichostomopsis
australasiae (Hook. & Grev.) Robins., Phytologia 20: 187,
Didymodon australasiae (Hook. & Grev.) Zand.,
1970
Phyto1ogia 41: 21, 1978]
Didymodon subtorquatus (C. Miill. & Hampe) Catcheside, Mosses
S. Australia 174, 1980 (Barbula) Austr1
Didymodon subulatus Cambessedes hom . il/eg. Eur
Didymodon taylorii Zand. (nom . nov. for Tortula campylocarpa
Tayl.) Am4
Didymodon tenellus Hedw. f. ex Brid. ignot.
Didymodon tectorum (C. Miill.) Saito, J. Hattori Bot. Lab. 39: 516,
1975 (Barbula) As2
Didymodon tomaculosus (B1ockeel) Corley in Corley et al., J.
Bryol. 11: 649, 1981 [1982] (Barbula) Eur
Didymodon tophaceopsis Zand. (nom. nov. for Gyroweisia
amplexicaulis Sim), see treatment of Didymodon Afr4
Didymodon tophaceus (Brid.) Lisa Eur As1 As3 As3 As5 Afr1
Am1Am2Am4
var. anatinus Hammerschm. Eur
var. bosniacus (Glow.) Latz. Eur = Didymodon tophaceus var.
decurrens Card. & Ther. fide Podp., Consp. Muse. Eur. 201,
1954]
var. breidleri Bauer Eur
var. brevifolius (BSG) Warns!. Eur
var. decurrens Card. & Ther. Eur
var. excurrens (Broth.) Wijk & Marg. As1
var. humilis (Schimp.) Warnst. Eur
var. linearis (De Not.) Limpr. Eur
var. riparius Amann Eur
fo. acutifolius (Boul.) Zodda, Malphighia 22: 509, 1908 (Trichostomum tophaceum fo.) Eur
[fo. anatinus (Hammerschm.) Marg., Lindbergia 1: 127, 1972 =
Didymodon tophaceus var. anatinus Hammerschm.]
fo. cylindrica (Boul.) Limpr., Laubm. Deutsch. 1: 554, 1888
(Trichostomum tophaceum fo.) Eur
=
=
GENERA OF THE POTTIACEAE
300
fo. elatus (Boul.) Artaria in E. Bauer, Musci. Eur. Am. Exsicc. 37:
n. 1823, 1925 (Trichostomum tophaceum fo.) Eur
fo. laxus Kindb., Bull. Soc. Bot. !tal. 7: 15, 1896 Eur
[fo. humilis (Schimp.) Marg., Lindbergia 1: 128, 1972 Didymodon tophaceus var. humilis (Schimp.) Wamst.]
fo.lingulatus (Boul.) Monk., Siisswasserfl. 14: 67, 1914 (Trichostomum tophaceum fo.) Eur
fo. lingulifolius (Roth in Zodda) Marg., Lindbergia 1: 128, 1972
Eur
fo. propaguliferus Amann, Bull. Soc. Vaud. Sc. Nat. 53: 86, 1920
Eur
fo. recurvifolius (Boul.) De Willd., Prodr. 434, 1899 (Trichostomum tophaceum fo.) Eur
fo. scabrinervis Podp., Acta Bot. Bohem. 1: 11, 1922 Eur
fo. truncatus (Boul.) Limpr., Laubm. Deutsch. 1: 554, 1888
(Trichostomum tophaceum fo.) Eur
fo. vulgaris Monk., Siisswasserfl. 14: 67, 1914 Eur
Didymodon torquatus (Tayl.) Catcheside, Mosses of South Australia
174, 1980 (Barbula) Austr1 Austr2
[Didymodon tosaensis Card. As2 = Barbula subcomosa Broth. fide
Saito, J. Jap. Bot. 47: 11, 1972]
[Didymodon trifarius (Hedw.) Rohl., the type, not the concept of
modern authors which is correctly Didymodon luridus (Hornsch.
in Spreng.)= Saelania glaucescens (Hedw.) Broth. in Bomanss &
Broth.fide Zander, Bryologist 81:421, 1978]
[subsp. nicholsonii (Culm.) Wijk & Marg. Eur
Didymodon
vinealis var. nicholsonii (Culm.) Zand., Cryptogarnie Bryol.
Lichenol. 2: 416, 1981 (1982) Didymodon nicholsonii Culm.]
var. angustifolius (Warnst.) Wijk & Marg. Eur
var. tuspidatus (Schirnp.) Wijk & Marg. Eur Am1
var. intermedius (Ruth.) Wijk & Marg. Eur
var. krimensis (Warns!.) Wijk & Marg. Eur
Didymodon trivia/is (C. Miill.) Guerra in Guerra & Ros, Cryptogarnie
Bryol. Lichenol. 8: 64, 1987 (Barbula) Eur Afr4
Didymodon umbrosus (C. Miill.) Zand., Phytologia 41 : 22, 1978
(Barbula) Eur Am1 Am2 Am4 Am6 [= Didymodon australasiae
var. umbrosus (C. Miill.) Zand. fide Zander, Cryptogamie Bryol.
Lichenol. 2: 400, 1981 (1982)]
Didymodon uruguayensis Zand. (nom. nov. for Barbula uruguayensis
Broth. hom. illeg. of Barbula uruguensis Par. fide Margadant in
litt.), see. treatment of Didymodon Am6
[Didymodon viridissimus Card. Am2 = Didymodon rigidulus Hedw.
s. lat. fide Zander, Cryptogarnie Bryol. Lichenol. 2: 390, 1981
(1982)]
Didymodon vinea/is (Brid.) Zand., Phytologia 41: 25, 1978 (Barbula)
Eur As1 As2 As3 AsS Afr1 Am1 Am2 Am4 Am6 Oc
[var. brachyphyl/us (Sull. in Whipp!.) Zand., Cryptogarnie Bryol.
Lichenol. 2: 411, 1981 (Barbula) Am1 = Didymodon
brachyphyllus (Sull. in Whipp!.) Zand., Phytologia 41: 24, 1978]
[var.fiaccidus (BSG) Zand., Phytologia 41: 25, 1978 (Barbula) Eur
As1 AsS Afr1 Am1 Am6 Oc [= Didymodon vinealis (Brid.) Zand.
var. vinealis fide Sollman, Bryologist 86: 272, 1983]
[var. luridus (Hornsch. in Spreng.) Zand., Cryptogamie Bryol.
Lichenol. 2: 412, 1981 (Didymodon) Eur As1 As2 AsS Afr1 Afr2
Am2 Am3 Didymodon luridus Hornsch. ex Spreng.]
[var. nicholsonii (Culm.) Zand., Cryptogarnie Bryol. Lichenol. 2:
416, 1981 (Didymodon) Eur Am1
Didymodon nicholsonii
Culm.]
[var. rubiginosus (Mitt.) Zand., Cryptogamic Bryol. Lichenol. 2:
417, 1981 (Barbula) Am1 = Didymodon occidentalis Zand.,
Phytologia 41 : 26, 1978]
Didymodon waymouthii (R. Br. ter) Zand. (Weissia) , see treatment of
Didymodon Austr2
=
=
=
=
=
Didymodon wildii (Broth.) Broth. Austr1
Didymodon wisse/ii (Dix.) Norris & T. Kop., Acta Bot. Fenn. 137:
127, 1989 (Barbula) As4
[Didymodon wollei (Aust.) Aust. Am1 = Trichostomum tenuirostre
var. holtii (Braithw.) Dix., see treatment of Trichostomum]
Didymodon xanthocarpus (C. Miill.) Magill in Magill & Schelpe,
Mem. Bot. Surv. S. Afr. 43: 5, 1979 (Barbula) Afr4
DOLOTORTULA Zand.
Dolotortula mniifo/ia (Sull.) Zand., Phytologia 65: 426, 1989
(Barbula) Am2 Arn3 Am4
ENTOSTHYMENIUM Brid.
Entosthymenium tristichum Brid., Bryol. Univ. 1: 761, 1827 ignot.
nom. dub. Eur
ERYTHROPHYLLASTRUM Zand., see treatment of Erythrophyl/astrum
Erythrophyl/astrum andinum (Sull.) Zand. (Trichostomum), see
treatment of Erythrophyl/astrum Am4
ERYTHROPHYLLOPSIS Broth. in Herz.
[Erythrophyl/opsis andina (Sull.) Zand., Bryologist 80: 159, 1977
Erythrophyl/astrum andinum (Sull.)
Am4 (Trichostomum)
Zand., see treatment of Erythrophyl/astrum]
[Erythrophyllopsis boliviano Broth. in Herz. Am4 Am6 (= Erythrophyllopsis andina (Sull.) Zand.fide Zander, Bryologist 80: 159,
1977, but, see treatment of Erythrophyllastrum) = Erythrophyl/opsisfuscula (C. Miill.) Hilp.fide Hilpert, Beih. Bot. Centralbl.
50(2): 639, 1933]
[Erythrophyllopsis challaensis (Broth. in Herz.) Hilp. Am4 Didymodon challaense (Broth. in Herz.) Zand., see treatment of
Didymodon]
Erythrophyllopsisfuscula (C. Miill.) Hilp. Am4 Am6
=
=
[ERYTHROPHYLLUM (Lindh.) Loeske = Bryoerythrophyllum
Chen]
[Erythrophyllum rubellum Hilp. = Bryoerythrophyllum
recurvirostrum (Hedw.) Chen]
[fo. alpinum Herz. , Weiner Bot. z. 93: 38, 1944 Eur Bryoerythrophyllum recurvirostrum fo.]
[fo. latifolium Herz., Wiener Bot. Z. 93: 38, 1944 Eur =Bryoerythrophyl/um recurvirostrum fo. latifolium (Herz.) Podp.]
[fo. pulvinatum Herz., Wiener Bot. Z. 93: 38, 1944 Eur = Bryoerythrophyllum recurvirostrum fo.]
Bryo[Erythrophyllum recurvirostrum (Hedw.) Loeske
erthrophyllum recurvirostrum (Hedw.) Chen]
[fo. serratum (Schirnp. ex Roell) Lazar., Opr. Listv. Mchov Ukr.
168. 1955 (Didymodon rubel/us var.) Eur = Bryoerythrophyllum recurvirostrum var. serratum (Roll) Podp.]
[fo. viride (Schlieph. ex Lirnpr.) Lazar., Opr. Listv. Mchov Ukr.
168, 1955 Eur Bryoerythrophyllum recurvirostrum fo. viride
(Schlieph. ex Limpr.) Podp.]
=
=
=
EUCLADIUM BSG
[Euc/adium irroratum (Mitt. in Hook.) Jaeg. Austr2 = Tetracoscinodon irrorata (Mitt. in Hook. f.) Zand. see treatments of
Tetracoscinodon and Eucladium]
Euc/adium verticil/arum (Brid.) BSG Eur As1 As2 As3 AsS Afr1
Afr4 Am 1 Am2
subsp. crassinervium Podp. Eur
subsp: styriacum (Glow.) Amman Eur
var. acuminatum Glow. Eur
GENERA, SPECIES AND INFRASPECIFIC TAXA
var. angustifolium Lindh. Eur Afrl As2
var. brevifolium Wamst. Eur
var. clinotheca (Besch.) Par. Afr1
var. commutatum (Glow.) Podp. Eur
var. crassinervium (Podp.) Podp. Eur
var. dalmaticum Par. Eur As2
var. latebrico/a Lamar!. & Maheu Eur
var. michelettii Fleisch. in Wamst. Eur
var. obtusifolium Wamst. Eur
var. penici/liforme Fam. Eur
var. recurvatum Dunk & Dunk, Herzogia 2: 419, 1973 Eur
var. recurvifolium Lindh. AsS
var. styriacum (Glow.) Glow. Eur
fo. crispum (Roll) Roll, Hedwigia 56: 139, 1915 (Eucladium
verticil/arum var.) Eur
fo. gracile Roll, Hedwigia 56: 139, 1915 Eur
fo. inundatum (Fam.) Podp., Consp. Muse. Eur. 174, 1954 (Euc/adium verticil/arum var.) Eur
[fo. lacustre Amann, Fl. Mousses Suisse ll: 37, 1918 Eur = fo.
inundatum (Fam.) Podp. fide Podp., Consp. Muse. Eur. 174,
1954]
fo. laetevirens (Zett.) Podp., Consp. Muse. Eur. 174, 1954 (Euc/adium verticillatum var.) Eur
fo. thermale Boros in Bauer, Muse. Eur. Exs. 2057, Sched. 42: 2,
1930 Eur
fo. viridissimum Bauer, Muse. Eur. Exs. 2059, Sched. 42: 2, 1930
Eur
GANGULEEA Zand.
Gangu/eea angulosa (Broth. & Dix.) Zand., Phytologia 65: 427, 1989
(Merceyopsis) As3 Am5
[GEHEEBIA Schimp. = Didymodon Hedw.fide Saito, J. Hattori Bot.
Lab. 39: 501, 1975]
[Geheebia gigantea (Funck) Boul. Eur As2 As3 Am1 = Didymodon
giganteus (Funck) Jur. fide Zander, Phytologia 41: 29, 1978 =
Didymodon fa/lax var. giganteus (Funck) Boul., see treatment of
Didymodon]
=
[GERTRUDIA Herz. Gertrudie/la (Herz.) Broth.]
[Gertrudia validinervis var. serrato-pungens Herz. Am4 Gertrudie/la validinervis var. serrato-pungens (Herz.) Zand., see treatment of Gertrudiella]
=
GERTRUDIELLA Broth.
[Gertrudie/la ferruginea (Herz.) Hilp. Am4 :: Didymodon herzogii
Zand. nom. nov., see treatment of Didymodon]
Gertrudiella validinervis (Herz.) Broth. Am4
var. serratopungens (Herz.) Zand., see treatment of Gertrudie/la
Am4
[GLOBULINA (C. Mtill.) Broth. = Bryoerythrophyllum Chen fide
Zander, Bryo1ogist 81: 541, 1978]
[G/obulina boliviano C. MUll., Nuovo Giom. Bot. Ita!. 4(1): 39, 1897
Am4 = Bryoerythrophyllum bolivianum (C. Mtill.) Zand.,
Bryologist 81: S4S, 1978]
GLOBULINELLA Steere in Steere & Chapman
Globulinella benoistii (Ther.) Magill, Bryologist 80: 79, 1977 Am4
Globuline/la globifera (Hampe) Steere Am1 Am2 Am3
[Giobuline/la peruviana (Williams) Steere Am2 Am4 = Saitoa
peruviana (Williams) Zand., Phytologia 6S: 431, 1989 = Sairoe/la
peruviana (Williams) Menzel]
301
GYMNOSTOMIELLA Fleisch.
Gymnostomie/la burmensis Bartr. As3
Gymnostomie/la /onginervis Broth. As2 As4
Gymnostomie/la monodii P. Varde Afr2
Gymnostomiella orcuttii Bartr. in Orcutt Am1 Am2 Am3 AmS
[Gymnostomie/la rosu/ara P. Varde Afr1 = Barbula semirosulata
Zand. nom. nov., see treatment of Barbula]
Gymnostomiel/a tanganyikae Sloover, Bull. Jard. Bot. Nat!.
Belgique 47: 146, 1977 Afr2
Gymnosromie/la vernicosa (Hook.) Fleisch. As2 As3 As4 Austr1
GYMNOSTOMUM Nees & Homsch.
Gymnostomum aeruginosum Sm. Eur As1 As2 As3 AsS Afr1 Afr4
Am1 Am2 Am3 Am6 Austr1 Austr2 Oc
var. cochlearifolium Karczmarz, Lindbergia 7: 127, 1981 [1982]
As3
fo. compactum (BSG) Wijk & Marg., Lindbergia 1: 128, 1972
Eur
fo. ramosissimum (BSG) Wijk & Marg., Lindbergia 1: 128, 1972
(Gymnostomum rupestre var.) Eur
fo. rigidum (Schimp.) Wijk & Marg., Lindbergia 1: 128, 1972
Eur
fo. ste/ligerum (Schimp.) Lazar.(Saviz.-Lub. & Smimova 1970)
Eur
[Gymnostomum angustifolium Saito, J. Hattori Bot. Lab. 36: 163,
1972 nom. inval. Am1 Am2 As2 Tuerckheimia angustifolia
(Saito) Zand., Misc. Bryol. Lichenol. 8: 27, 1978 = Tuerckheimia svihlae (Bartr.) Zand., see treatment of Tuerckheimia]
[Gymnostomum aurantiacum (Mitt.) Jaeg. = Hymenostylium
recurvirostrum var. aurantiacum (Mitt.) Gangulee, Mosses E.
India 648, 1972 (Hymenostylium) hom. i/leg. incl. var. prior.=
Hymenostylium recurvirostrum var. luzonense (Broth.) Bartr.
fide Gangulee, Mosses E. India 3: 648, 1972 = Hymenostylium
recurvirostrum var. cylindricum (Bartr.) Zand., see treatment of
Hymenostylium]
Gymnostomum bewsii Dix. Afr4 [good species, see Magill, Fl. S.
Afr. 1: 18S, 1981 (1982)]
Gymnostomum bescherellei Broth. & Geh. ex Herz. Afr2
Gymnosromum borea/e Nyholm & Hedenas, Lindbergia 12: 41,
1986 Eur
[Gymnostomum brachystegium Besch., J. Bot. (Morot) 12: 281,
1898 As2 :: Bryoerythrophyl/um brachystegium (Besch.) Saito,
J. Jap. Bot. 47: 14, 1972]
Gymnostomum ca/careum Nees & Homsch. Eur Asl As2 As3 AsS
Afr1 Afr4 Am1 Am2 Am6 Austrl Austr2 Oc [= Gymnostomum
aeruginosum Sm. fide Zander, Bryo1ogist 80: 2S9, 1977;
Magill, 1981, Fl. S. Afr. 1: 183, 1981 (1982)]
var. australe Broth. & Geh. Austr1 Austr2
[var. brevifolium Schimp. Eur Afr1 = Gymnostomum luisieri
(Sergio) Sergio ex Crundw. fide Sergio, Anales Bioi. (Univ.
Murcia) 2, Sec. Esp. 2) 1984: 361, 1984 = Gymnostomum
viridulum Brid. fide Whitehouse & Crundwell, J. Bryol. 16:
S63, 1991]
var. /ongifolium Dix. Austr1 Austr2
var. muticum Boul. Eur Afr1
var. perpusi/lum Sull. Am1
[var. viridulum (Brid.) B.&S. in BSG Eur = Gymnostomum
viridulum Brid. fide Whitehouse & Crundwell, J. Bryol. 16:
S63, 1991]
fo. cavernarum Grom, Acta Bot. Croat. 26-27: 249, 1967-68
[1968] Eur
fo. gracile (Breidl.) Podp., Consp. Muse. Eur. 170, 19S4 (Gymnostomum calcareum var.) Eur
=
302
GENERA OF THE POTTIACEAE
fo. intermedium (Schimp.) Podp., Consp. Muse. Eur. 170, I954
(Gymnostomum calcareum var.) Eur
fo. longifolium Meylan, Bull. Soc. Yaud. Sci. Nat. 60(249): 270,
I939 Eur
fo. subrotundifolium Latz., Beih. Bot. Centralbl. 48(2): 473, 193I
Eur
fo. tenellum (BSG) Podp., Consp. Muse. Eur. I70, I954 (Gymnostomum ca/careum var.) Eur
Gymnostomum carthusianum (Brid.) P. Beauv. Eur
Gymnostomum chenii Saito, J. Jap. Bot. 48: I64, I973 As3
Gymnostomum cirrhatum De Not. Eur
[Gymnostomum curvirostre Hedw. ex Brid. = Hymenostylium
recurvirostrum (Hedw.) Dix.]
[var. anoectangioides Ther. Eur = Hymenostylium recurvirostrum
var. anoectangioides (Ther.) Wijk & Marg.]
[Gymnostomum denticulatum Griff. As3 =Hyophila involuta (Hook.)
Jaeg., see treatment of Hyophila]
[Gymnostomum dimorphum (C. MUll.) Sim Afr4 = Didymodon
dimorphus (C. MUll.) Broth. fide Magill & Schelpe, Mem. Bot.
Surv. S. Afr. 43: 23, I979]
Gymnostomumfoliosum Rohl. Eur
[Gymnostomum gracile Dix. hom. i/leg. Afr2 Afr4 = Didymodon
ceratodonteus (C. Miill.) Dix.fide Magill, Fl. S. Afr. I. Mosses I:
233, I98I (I982)]
Gymnostomum hymenostylioides (Broth. & Dix.) Zand. (Merceyopsis), see treatment of Gymnostomum As3
[Gymnostomum insigne (Dix.) A. Sm., J. Bryol. 9: 279, 1976 (1977)
Eur As2 As3 Ami = Hymenostylium recurvirostrum var. insigne
(Dix.) Bartr.fide Zander & Eckel, Canad. J. Bot. 60: 1596, 1982]
Gymnostomum jacksharpii (Crum) Allen, Bryologist 93: 207, 1991
(Barbula) Am3
[Gymnostomum keniae P. Yarde Afr2 = Anoectangium keniae (P.
Yarde) Zand., see treatment of Anoectangium]
[Gymnostomum knightii Schimp. in Knight, Trans. New Zealand lnst.
7: 354, 1875 Austr2 = Didymodon tophaceus (Brid.) Lisa, see
treatment of Didymodon]
Gymnostomum /axirete (Broth.) Chen As2
Gymnostomum leptostomum Brid. hom. illeg. Afr3
[Gymnostomum lessonii Besch. Afr2 = Racopilum sp. see Excluded
Taxa]
Gymnostomum lingulatum Rehm. ex Sim Afr4 [= Didymodon
lingulatus (Sim) Magill, Mem. Bot. Surv. S. Afr. 43: 5, 1979 =
good species fide Magill, Fl. S. Afr. I. Mosses 1: 184, 1981
(1982)]
Gymnostomum ludovicae Broth. & Par. Oc
[Gymnostomum luisieri (Sergio) Sergio ex Crundw., J. Bryol. 11:
603, I98I [I982] (Gyroweisia) Eur = Gymnostomum viridulum
Brid.fide Whitehouse & Crundwell, J. Bryol. I6: 563, I99I]
[Gymnostomum microcarpon Nees & Homsch. = Hymenostylium
recurvirostrum (Hedw.) Dix.]
var. elongatum Nees & Homsch. Eur
Gymnostomum minutulum Saporta hom. i/leg., fossil Eur
Gymnostomum mosis (Lor.) Jur. As5
Gymnostomum mucronulatum Hedw. f. in Mohr nom. inval. Eur
[Gymnostomum ovatum Hedw. = Pterygoneurum ovatum (Hedw.)
Dix.]
var. longicapsu/um Chev. Eur
[Gymnostomum pottsii (Dix.) Sim = Didymodon dimorphus (C.
Miill.) Broth. fide Magill & Schelpe, Mem. Bot. Surv. So. Afr.
43: 23, 1979 = Didymodon ceratodonteus (C. Miill.) Dix. fide
Magill, Fl. S. Afr. I. Mosses I: 233, I98I (I982)]
[Gymnostomum recurvirostrum Hedw. = Hymenostylium recurvirostrum (Hedw.) Dix.]
[var. latifolium (Zett.) Flow. ex Crum, Bryologist 72: 243, 1969.
Eur Asi Ami = Hymenostylium recurvirostrum (Hedw.) Dix.
fide Zander, Bryologist 80: 253 , I977]
[Gymnostomum rupestre Schleich. ex Schwaegr. = Gymnostomum
aeruginosum Sm.]
var. minus De Not. Eur
var. obtusifolium Kindb, Nuov. Giom. Bot. ltal. 25: 116, 1893
Eur
fo. arboreum Geh., Bot. Centralb. Beih. 22(2): 98, I907 Eur
fo. cavernicola Boros., Bot. Kozlem. 32: 108, I935 Eur
[fo. compactum (BSG) Monk., Laubm. Eur. 25I, I927 (Gymnostomum rupestre var.) Eur = Gymnostomum aeruginosum fo.
compactum (BSG) Wijk & Marg.]
fo. crispatulum Roll, Hedwigia 38: 260, 1899 Eur
fo. curvisetum (Amann) Podp., Consp. Muse. Eur. 170, 1954
(Gymnostomum rupestre var.) Eur
fo. elatum Smarda, Cas. Morav. Mus. Zemsk. 32: 14, 1949 Eur
fo. intermedium Limpr., Laubm. 233, 1886 Eur
fo. ramosissimum (BSG) Monk., Laubm. Eur. 250, 1927 (Gymnostomum rupestre var.) Eur
fo. rigidum (Schimp.) Podp., Consp. Muse. Eur. 17I, 1954 (Gymnostomum rupestre var.) Eur
[fo. stel/igerum (BSG) Monk., Laubm. Eur. 25I , I927 (Gymnostomum rupestre var.) Eur Ami= Gymnostomum aeruginosum
fo. stel/igerum (BSG) Lazar.]
[Gymnostomum setifolium Hook. & Am. Am4 = Bartramidu/a
setifolia (Hook. & Am.) Fransen, Lindbergia 14: 31, 1988 =
Flowersia setifolia (Hook. & Am.) Griffin & Buck, Bryologist
92: 372, 1989]
Gymnostomum simplicissimum (P. Beauv.) Brid. As3
Gymnostomum splachnobryoides Biz., Cryptogamie Bryol.
Lichenol. 1: 425, 1980 Afr2
[Gymnostomum subrigidulum (Broth.) Chen As2 =Hymenostylium
recurvirostrum var. insigne (Dix.) Bartr. fide Zander & Eckel,
Canad. J. Bot. 60: I596, I982]
Gymnostomum tenerrimum (C. Miill.) Wijk & Marg., Taxon I7:
467, I968 (Anoectangium) Am6
[Gymnostomum truncatum (With.) Hedw. ex Schum. = Portia
truncata (Hedw.) BSG]
var. intermedium Lilj. Eur
Gymnostomum unguiculatum Philib. in Husn., Musci Gall. Herb.
12: n. 55 I, I879 Eur
[Gymnostomum valerianum (Bartr.) Zand. , Bryologist 80: 262,
1977 (Leptodontium) Am2 = Tuerckheimia valeriana (Bartr.)
Zand.fide Zander, Misc. Bryol. Lichenol. 8: 27, 1978]
[Gymnostomum venezue/ense (C. MUll.) Kindb. Am4 = Hymenostylium recurvirostrum (Hedw.) Dix., see treatment of Hymenostylium]
Gymnostomum viridulum Brid. good species fide Whitehouse &
Crundwell, J. Bryol. 16: 563, I99I Eur Afri
[Gymnostomum wageri Schelpe in Magill & Schelpe, Mem. Bot.
Surv. S. Afr. 43: 5, I979 (Gymnostomum gracile Dix. non (R.
Br.) Hook.) Afr4 = Didymodon ceratodonteus (C. MUll.) Dix.
fide Magill, Fl. S. Afr. I. Bryoph. 1: 235, 1981 (1982)]
GYROWEISIA Schimp.
[Gyroweisia amplexicaulis Sim Afr4 = Husnotiella latifolia (Dix.)
Zand. & Magill in Magill & Schelpe, Mem. Bot. Surv. S. Afr.
43: 23, I979 = Didymodon tophaceopsis Zand. nom. nov., see
treatment of Didymodon]
Gyroweisia barbulacea (C. Miill.) Broth. Am2 (ignot., type cannot
be not located fide Zander, Bryologist 80: 265, I977)
[Gyroweisia benoistii Ther. Am4 = Globuilinella benoistii (Ther.)
GENERA, SPECIES AND INFRASPECIFIC TAXA
Magill, Bryologist 80: 79, 1977]
[Gyroweisia boliviano Williams .(= Barbula sp. fide Hilpert 1933) =
Didymodon tophaceus (Brid.) Lisa, see treatment of Gyroweisia]
[Gyroweisia brevicaulis (C. Miill.) Broth. As4 Oc = Luisierel/a barbuta (Schwaegr.) Steere fide Zander, Bryologist 80: 266, 1977]
[Gyroweisia hildebrandtii (C. Miill.) Kindb. Afr2 Hymenostylium
hildebrandtii (C. Miill.) Zand., see treatment of Hymenostylium]
[Gyroweisia /atifolia Dix. Afr2 Husnotiella latifolia (Dix.) Zand. &
Magill in Magill & Schelpe, Mem. Bot. Surv. S. Afr. 43: 7, 1979
Didymodon tophaceopsis Zand. nom. nov., see treatment of
Didymodon]]
var. tanneri Biz., Rev. Bryol. Lichenol. 40: 120, 1974 Afr2
[Gyroweisia lindigii (Hampe) Broth. Am4
Didymodon lindigii
(Hampe) Zand., see treatment of Didymodon]
[Gyroweisia luisieri Sergio, Bol. Soc. Port. Cienc. Nat. 14: 82, 1972
Eur (= Gymnostomum aeruginosum fide Zander, Bryologist 80:
259, 1977) = Gymnostomum /uisieri (Sergio) Sergio ex Crundw.,
J. Bryol. 11: 603, 1981 (1982) = Gymnostomum viridulum Brid.
fide Whitehouse & Crundwell, J. Bryol. 16: 563, 1991]
Gyroweisia monterreia Zand. & Herm., Bryologist 89: 227, 1986
[1987] Am2
[Gyroweisia nigeriana Egun. & Olar., Bryologist 81: 443, 1978 Afr2
= Weisiopsis nigeriana (Egun. & Olar.) Zand., see treatment of
Weisiopsis]
[Gyroweisia obtusifolia Broth. Am2
Husnotiella obtusifolia
(Hampe) Zand., Bryologist 80: 265, 1977
Gyroweisia
obtusifolia Broth. fide Zander, Cryptogamic Bryol. Lichenol. 2:
419, 1981 (1982) = Didymodon hampei Zand. nom. nov., see
treatment of Didymodon]
[Gyroweisia papillosa Ther. Am2 = Husnotiella obtusifolia (Broth.)
fide Zander, Bryologist 80: 265, 1977 = Gyroweisia obtusifolia
Broth. fide Zander, Cryptogamic Bryol. Lichenol. 2: 419, 1981
(1982)] Didymodon hampei Zand. nom. nov., see treatment of
Didymodon]
[Gyroweisia pocsii Biz., Cryptogamic Bryol. Lichenol. 1: 424, 1980
Afr2 = Weisiopsis nigeriana Egun. & Olar. Zand., see treatment
of Weisiopsis]
Gyroweisia pusil/a Broth. Am1
Gyroweisia refiexa (Brid.) Schimp. Eur Afr1 Am1
Gyroweisia rohlfsiana (C. Miill.) Par. Afr1
Gyroweisia shansiensis Sak. As2
Gyroweisia tenuis (Hedw.) Schimp. Eur As5 Afr1 Am1
var. badia Limpr. Eur Afrl
var. cuspidata Roll Eur
var. lacustris Amann Eur
var. schisticola Roth in Zodda Eur
fo. compacta (Hag.) Podp., Consp. Muse. Eur. 171, 1954 (Gyroweisia tenuis var.) Eur
[fo. propagulifera Lirnpr. Eur = Gyroweisia tenuis var. compacta
(Hag.) Podp., Consp. Muse. Eur. 171, 1954]
[Gyroweisia tophico/a (C. Miill.) Kindb. Afr2 = Hymenostylium
recurvirostrum (Hedw.) Dix., see treatment of Hymenostylium]
Gyroweisia yuennanensis Broth. As2
=
=
=
=
=
=
=
HENNEDIELLA Par.
Hennediel/a acletoi (Robins.) Zand. (Tortula), see treatment of Hennediel/a Am4
Hennediella acutidentata (Card. & Ther.) Zand. (Pottia), see treatment of Hennediel/a Afr4
Hennediella angustifolia (Herz.) Zand. (Calyptopogon), see treatment
of Hennediella Am4
Hennediella arenae (Besch.) Zand. (Barbula), see treatment of Hennediel/a Austr2 Ant
303
var. petriei (Broth.) Zand. (Tortula), see treatment of Hennediel/a
Austr2
Hennediel/a austrogeorgica (Card.) Blockeel, J. Bryol. 16: 191,
1991 (Pottia) Am6
Hennediella be/lei (Bartr.) Zand. (Desmatodon), see treatment of
Hennediella Am4
Hennediella densifolia (Hook. f. & Wils.) Zand. (Barbula), see
treatment of Hennediella Am6 Ant
Hennediella denticulata (Wils. in Mitt.) Zand. (Barbula), see treatment of Hennediella Am3 Am4
Hennediella heimii (Hedw.) Zand. (Gymnostomum), see treatment
of Hennediella Eur As1 As2 Am1 Am6 Austr1 Austr2 Ant
var. alpina (Amann) Zand. (Pottia heimii var.), see treatment of
Hennediella Eur
var. arctica (Lindh.) Zand. (Pottia heimii var.), see treatment of
Hennediella Eur As! Am1
var. brachyphylla (Warnst.) Zand. (Pottia heimii var.), see treatment of Hennediel/a Eur
var. brevicuspis (Warns!.) Zand. (Pottia heimii var.), see treatment of Hennediel/a Eur
var. breviseta (Warnst.) Zand. (Pottia heimii var.), see treatment
of Hennediella Am1
var. drummondii (Warnst.) Zand. (Pottia heimii var.), see treatment of Hennediel/a Am1
var. eurystoma (Card. & Broth.) Zand. (Pottia heimii var.), see
treatment of Hennediella Am6
var. guessfeldtii (Schlieph.) Zand. (Pottia), see treatment of Hennediella Eur
var. lanceolata (Warns!.) Zand. (Pottia heimii var.), see treatment
of Hennediella Eur Ami
var. magellanica (Warns!.) Zand. (Pottia heimii var.), see treatment of Hennediella Am6
var. maxima (Card.) Zand. (Pottia heimii var.), see treatment of
Hennediella Am6
var. spegazzinii (C. Miill.) Zand. (Portia), see treatment of Hennediel/a Am6
var. thaxteri (Card. & Ther.) Zand. (Pottia heimii var.), see treatment of Hennediella Am6
Hennediella heteroloma (Card.) Zand. (Tortula), see treatment of
Hennediel/a Am2
var. eckeliae (Zand.) Zand., see treatment of Hennediel/a Am2
Hennediella kunzeana (C. Miill.) Zand. (Barbula), see treatment of
Hennediella Afr4 Am6
Hennediella limbata (Mitt.) Zand. (Barbula), see treatment of Hennediella Am2 Am4
Hennediella /ongipedunculata (C. Miill.) Zand. (Barbula), see treatment of Hennediel/a Afr4
Hennediella /ongirostris (Hampe ex C. Miill.) Zand. (Pottia), see
treatment of Hennediella Afr2
Hennediella macrophyl/a (R. Br. ter) Par. Am6 Austr2
Hennediel/a marginata (Hook. f. & Wils.) Zand. (Schistidium), see
treatment of Hennediella Afr4
Hennediel/a oedipodioides (C. Miill.) Zand. (Pottia), see treatment
of Hennediella Afr4
Hennediella polyseta (C. Miill.) Zand. (Barbula), see treatment of
Hennediella Am2 Am4
Hennediella serrulata (Hook. & Grev.) Zand. (Tortula), see treatment of Hennediella Am6
Hennediella stanfordensis (Steere) Blockeel J. Bryol. 16: 191, 1991
(Tortula) Eur Am1 Austr1
Hennediella steereana (Zand. & Crum) Zand. (Desmatodon), see
treatment of Hennediella Am4
304
GENERA OF THE POTTIACEAE
HILPERTIA Zand.
Hilpertia scotteri (Zand. & Steere) Zand., Phytologia 65: 428, 1989
(Tortula) Am1
Hilpertia velenovskyi (Schiffn.) Zand., Phytologia 65: 428, 1989
(Tortula) Eur Am6
[HUSNOTIELLA Card. = Didymodon fide Zander, Cryptogamic
Bryol. Lichenol. 2: 384, 1981 (1982)]
[Husnotiel/a baueri Bartr. in Bauer Am6 = Didymodon tophaceus
(Brid.) Lisa see treatments of Didymodon and Bryoerythrophyllum]
Husnotiel/a glossophylla Herz. Am4 (type apparently lost, see treatment of Bryoerythrophyllum)
[Husnotiella latifolia (Dix.) Zand. & Magill in Magill & Schelpe,
Mem. Bot. Surv. S. Afr. 43 : 7, 1979 (Gyroweisia) Afr4 = Didymodon tophaceopsis Zand. nom. nov., see treatment of Didymodon]
[Husnotiella obtusifolia (Hampe) Zand., Bryologist 80: 265, 1977
(Trichostomum) Am2 = Gyroweisia obtusifolia Broth. fide
Zander, Cryptogamic Bryol. Lichenol. 2: 419, 1981 (1982) =
Didymodon hampei Zand. nom. nov., see treatment of Didymodon]
[Husnotiella plicata Magill, Fl. S. Afr. I. Mosses 1: 222, 1981 [1982]
Afr4 = Didymodon jackvancei Zand. nom. nov., see treatment of
Didymodon]
[Husnotiella revoluta Card. Ami Am2 Am4 = Didymodon revolutus
(Card.) Williams fide Zander, Cryptogamic Bryol. Lichenol. 2:
401, 1981 (1982)]
[var. palmeri (Card.) Ther. Am1 = Didymodon revolutus (Card.)
Williamsfide Zander, Cryptogamic Bryol. Lichenol. 2: 401, 1981
(1982)]
fo. e/ata Ther., Smiths. Misc. Coli. 85: 7, 1931 Am2
[Husnotiella torquescens (Card.) Bartr. Am1 Am2 Am4 = Didymodon australasiae (Hook. & Grev.) Zand. s. lat. fide Zander,
Cryptogamic Bryo1. Lichenol. 2: 397, 1981 (1982)]
[HYDROGONIUM (C. Miill.) Jaeg. =Barbulafide Saito, J. Hattori
Bot. Lab. 39: 492, 1975]
[Hydrogonium afrofontanum (C. Miill.) Hilp. Afr4 = Barbula
afrofontana (C. Miill.) Broth.]
[Hydrogonium amplexifolium (Mitt.) Chen As3 = Barbula
amplexifolia (Mitt.) Jaeg.]
[Hydrogonium anceps (Card.) Herz. & Nog. As2 = Barbula anceps
Card.]
[Hydrogonium aneitense (Broth. & Watts) Schultze-Motel,
Willdenowia 7: 55, 1973 = Barbula aneitensis Broth. & Watts.]
[Hydrogonium arcuatum (Griff.) Wijk & Marg. As3 As4 Oc =Barbula arcuata Griff.]
[Hydrogonium bolleanum (C. Miill.) Jaeg. Afr2 = Barbula bol/eana
(C. Miill.) Broth.]
[Hydrogonium brotheri (Ren. & Par.) Hilp. Afr3 = Didymodon
brotheri (Ren. & Par.) Zand. (Trichostomum), see treatment of
Didymodon]
[Hydrogonium clavicostatum (Ren. & Card.) Hilp. Afr3 = Barbula
c/avicostata (Ren. & Hilp.) Zand., see treatment of Barbula]
(Hydrogonium comosum (Dozy & Molk.) Hilp. As4 = Barbula
arcuara Griff. fide Saito, J. Hattori Bot. Lab. 39: 496, 1977]
[var. japonica Broth. As2 = Barbula arcuata Griff. fide Saito, J.
Hattori Bot. Lab. 39: 496, 1977]
[Hydrogonium consanguineum (Thwait. & Mitt.) Hilp. As3 As4 =
Barbula consanguinea (Thwait. & Mitt.) Jaeg. = Barbula
javanica Dozy & Molk. fide Saito, J. Hattori Bot. Lab. 39: 495,
1975]
[Hydrogonium crozalsii (Philib.) Podp. Eur = Barbula crozalsii
(Philib.) Broth.]
[Hydrogonium decolyi Broth. ex Gangulee, Nov. Hedw. 12: 426,
1966 As3 = Bryoerythrophyllum recurvum (Griff.) Saito fide
Sollman, Lindbergia 16: 23 , 1990]
Hydrogonium dicranel/oides Gangu1ee, Nov. Hedw. 12: 427, 1966
As3
Hydrogonium dixonianum Chen As2
[Hydrogonium dorrii (Ren. & Card.) Hilp. Afr3 = Barbula dorrii
Ren. & Card.]
[Hydrogonium ehrenbergii (Lor.) Jaeg. Eur As2 As5 Afr1 Am1
Am2 = Barbula ehrenbergii (Lor.) Fleisch.]
[var. algeriae (C. Miill.) Wijk & Marg. Eur Afr1 = Hydrogonium
ehrenbergii var. algeriae (C. Miill.) Vent. & Bott.]
fo. densirete (Amann) Podp., Consp. Muse. Eur. 199, 1954
(Didymodon ehrenbergii fo.) Eur
fo. laxirete (Amann) Podp., Consp. Muse. Eur. 199, 1954 (Didymodon ehrenbergii fo.) Eur
[Hydrogonium fontanum (C. Miill.) Jaeg. Afr2 = Barbula fontana
(C. Miill.) Broth.= Barbula meidensis Cufodontis nom. nov.]
[Hydrogonium gangeticum (C. Miill.) Chen As3 = Barbula
gangerica C. Miill. = Barbula arcuara Griff. fide Gangulee,
Mosses E. India 3: 725, 1972]
[Hydrogonium gracilentum (Mitt.) Chen As3 = Barbula gracilenta
Mitt.]
Hydrogonium heterophyllum Frohl., Ann. Naturhist. Mus. Wien 67:
154, 1964 As3
Hydrogonium hygrophilum Hilp. As4
[Hydrogonium javanicum (Dozy & Molk.) Hilp. As2 As3 As4 =
Barbulajavanica Dozy & Molk.J
[var. convolutifolium (Dix.) Chen As2 = Barbula javanica Dozy
& Molk .fide Saito, J. Hattori Bot. Lab. 39: 495, 1975]
[var. epapillosum (Fleisch.) Hilp. As4 = Barbula javanica var.
epapillosa Fleisch.]
var. kurzii (C. Miill.) Gangulee, Mosses E. India 3: 738, 1972 As3
[Hydrogonium /aevifolium (Broth. & Yas.) Chen As2 = Barbula
/aevifolia Broth. & Yas.J
[Hydrogonium leucobasis (Dix. in Dix. & Greenwood) SchultzeMotel, Willdenowia 7: 371, 1974 Oc (Barbula) = Barbula
/eucobasis Dix. in Dix. & Greenwood]
Hydrogonium leucodontoides Gangulee, Nov. Hedw. 12: 425, 1966
[1967] As3
[Hydrogonium /eucodonroides (C. Miill. in Par. ex Gangulee)
Chopra, Taxon. Indian Mosses 145. 1975 (Barbula) As3 =
Barbula leucodonrioides C. Miill. in Par. ex Gangulee, Nov.
Hedw. 12: 425, 1966]
[Hydrogonium louisiadum (Broth.) Schultze-Motel, Willdenowia 7:
55, 1973 (Barbu/a) Oc = Barbula consanguinea (Thw. & Mitt.)
Jaeg.fide Eddy, Handb. Males. Mosses 2: 178, 1991 =Barbula
javanica Dozy & Molk . fide Saito, J. Hattori Bot. Lab. 39: 495,
1975]
[Hydrogonium majusculum (C. Miill.) Chen As2 = Barbula
majuscu/a C. Miill.]
Hydrogonium mamatkulovii Laz., Dopov. Akad. Nauk Ukr. RSR,
Ser. B 8: 753, 1967
Hydrogonium mussoorianum Vohra, J. Bombay Nat. Hist. Soc. 63:
464, 1967. As3
[Hydrogonium novoguinense (Broth.) Chen As4 = Barbula
novoguinense Broth.]
Hydrogonium patulifolium Frohl., Ann. Naturhist. Mus. Wien 67:
154, 1964 As3
[Hydrogonium pseudoehrenbergii (Fleisch.) Chen As2 As4 Afr2 =
Barbula pseudoehrenbergii Fleisch.]
305
GENERA, SPECIES AND INFRASPECIFIC TAXA
[Hydrogonium rechingeri (Broth.) Schultze-Motel, Willdenowia 7:
55, 1973 Oc =Barbula rechingeri Broth.]
[Hydrogonium setschwanicum (Broth.) Chen As2
Barbula
setschwanica Broth. = Barbula indica (Hook.) Spreng. fide Saito,
J. Hattori Bot. Lab. 39: 488, 1975]
[Hydrogonium sobo/iferum (Fleisch.) Hilp. As4 = Barbula sobolifera
Fleisch. = Barbula arcuata Griff. fide Saito, J. Hattori Bot. Lab.
39: 496, 1975]
[Hydrogonium sordidum (Besch.) Chen As3
Barbula sordida
Besch.]
[Hydrogonium subcomosum (Broth.) Chen As2 Barbula subcomosa
Broth.]
[Hydrogonium subpel/ucidum (Mitt.) Hilp.
Barbula subpel/ucida
(Mitt.) Broth.]
var. hya/oloma Herz. As2
[Hydrogonium tay/orii W. Weber, Lindbergia 3: 81, 1975 [1976] =
Barbula taylorii Bartr. & Steere hom. illeg. non Lindh.]
[Hydrogonium tisserantii (P. Yarde) Schultze-Motel, Willdenowia 7:
491, 1975 (Didymodon) Afr2 = Barbula tisserantii (P. Yarde) P.
Yarde]
[Hydrogonium williamsii Chen As4
Barbula williamsii (Chen)
Iwats. & Tan, Kilikasan 8: 186, 1979]
=
=
=
=
=
[HYMENOSTOMUM R. Br. = Weissia Hedw.fide Saito, J. Hattori
Bot. Lab. 39: 417, 1975]]
[Hymenostomum anomalum Broth. in Herz. Am4
Trichostomum
ovatifolium Zand. nom. nov., see treatment of Trichostomum]
Hymenostomum aristatulum Par. & Broth. Oc
[Hymenostomum ayresii (Schimp.) Broth. Afr2 Afr3 = Weissia
ayresii Schimp. in Besch.]
[Hymenostomum balansaeanum Besch. Am5 Am6 = Weissia
ba/ansaeana (Besch.) C. Miill.]
var. densirete Ther. Am6
[Hymenostomum brachype/ma (C. Miill.) Kindb. Afr2 = Weissia
brachypelma C. Miill. Afr2
[Hymenostomum breutelii (C. Miill.) Kindb. Am2 Am3 Am5 Weissia breutelii C. Miill. Am2 Am3 Am4 Am5
[Hymenostomum castaneum Crum & Steere, Amer. Midland Nat. 60:
12, 1959 Am3 = Trichostomum castaneum (Crum & Steere)
Zand., see treatment of Trichostomum]
[Hymenostomum ch/oropus (Besch.) Broth. Afr2 Afr3 = Hymenostomum ayresii (Schimp. ex Besch.) Broth. fide Een, Lindbergia
3: 125, 1976 =Weissia ayresii Schimp. in Besch.]
[Hymenostomum cucu/latum (C. Miill.) Kindb. Afr4
Weissia
cucu/lata C. Miill. Afr4
Hymenostomum densirete Ther. Am4
[Hymenostomum eckendorffii P. Yarde Afr2 Tortel/a eckendorffii
(P. Yarde) Zand., see treatment of Tarte/la]
[Hymenostomum edentulum (Mitt.) Besch. As3 As4 Oc = Weissia
edentula Mitt. As3 As4 Oc
[Hymenostomum eurybasis Dix. Afr2 = Weissia /atiuscula C. Miill.
fide Magill, Fl. S. Afr. I. Mosses 1: 267, 1981 (1982)]
[Hymenostomum exsertum (Broth.) Broth. As2 = Weissia exserta
(Broth.) Chen]
Hymenostomumfasciculatum Hampe Am5
[Hymenostomumflavescens Britt. inN. Britt. & Millsp. Am1 Am3
Weissia flavescens (Britt. in N. Britt. & Millsp.) Reese,
Bryologist 94: 54, 1991 = Trichostomum brittonianum nom. nov.,
see treatment of Trichostomum]
Hymenostomumfrancii Ther. Oc
Hymenostomum goyazense (Broth.) Broth. Am5
Hymenostomum guineense Broth. & Par. Afr2
[Hymenostomum guyazensis (Broth.) Broth., Nat. Pfl. 1(3): 386, 1902
=
=
=
=
=
err. pro Hymenostomum goyazense (Broth.) Broth.]
[Hymenostomum gymnostomum (Besch.) Nog. As2 = Weissia
edentula Mitt. fide Saito, J. Hattori Bot. Lab. 39: 421, 1975]
[Hymenostomum humico/a (C. Miill.) Par. Afr2 Afr4
Weissia
humicola C. Miill.]
[Hymenostomum inopercu/atum Crum, Madroiio 14: 74, 1957 Am1
Weissia inopercu/ata (Crum) Crum, Steere & Anders.,
Bryologist 67: 164, 1973]
[Hymenostomum jamesonii (Arnott) Hampe Am2 Am5 = Weissia
jamesonii Tayl.]
[Hymenostomum krassavinii Laz. As1 = Weissia krassavinii (Laz.)
Laz. ex Ochyra]
[Hymenostomum kunzeanum (C. Miill.) Broth. Am6 = Weissia
kunzeana C. Miill.]
Hymenostomum latifolium Nog. As2
Hymenostomum laxirete (Broth.) Broth. Am6
Hymenostomum leratii Par. & Broth. Oc
var. acuminatum Ther. Oc
[Hymenostomum lineaefolium (C. Miill.) Par. Afr2 = Weissia lineaefolia C. Miill.]
[Hymenostomum malayense Fleisch. As2 As4 = Weissia malayensis
(Fleisch.) Manuel, Fed. Mus. Jour. 26: 161, 1981 = Barbula
indica (Hook.) Spreng. fide Iwatsuki & Noguchi, J. Hattori Bot.
Lab. 37: 355, 1973 as Barbula cruegeri Sond. ex C. Miill.]
[Hymeilostomum mexicanum Card. Am2 = Weissia controversa
Hedw.fide Zander in Sharp et al., Moss Fl. Mex.]
[Hymenostomum meylanii Amann = Hymenostomum squarrosum
Nees & Homsch. fide Crundwell & Nyholm, J. Bryology 7: 9,
1972 Weissia squarrosa (Nees & Homsch.) C. Miill.]
[Hymenostomum micaceum (Schlecht.) Hampe As2 As3 As4 Am2
Am3 Am5 Oc Weissia micacea (Schlecht.) C. Miill.]
[Hymenostomum microstomum (Hedw.) R. Br. in Nees & Homsch.
Weissia microstoma
Eur As1 As2 As3 As5 Afr1 Aml
(Hedw.) C. Miill. Eur As1 As2 As3 As5 Afr1 Aml hom. illeg.
= Weissia brachycarpa (Nees & Hornsch.) Jur. var.
brachycarpafide Corley et al., J. Bryol. 11 : 650, 1981 (1982)
and Koponen, Isoviita & Lammes, Flora Fennica 6: 61, 1977]
[var. brachycarpum (Nees & Homsch.) Hiib. Eur
Weissia
brachycarpa (Nees & Homsch.) Jur. fide Corley et al., J. Bryol.
11 : 650, 1981 (1982) and Koponen, lsoviita & Lammes, Flora
Fennica 6: 61, 1977]
[var. obliquum (Nees in Nees & Homsch.) Hiib. Eur = Weissia
brachycarpa var. obliqua (Nees in Nees & Homsch.) Hill nom.
illeg.]
fo. e/atum (B.&S. in BSG) Podp., Consp. Muse. Eur. 186, 1954
Eur
fo. planifolium Fleisch., Beitr. Laubmoosfl. Ligur. Atti Congr.
Botan. Intern. 1892: 268, 1893 Eur
[Hymenostomum newcomeri Bartr. As2 = Weissia newcomeri
(Bartr.) Saito]
Hymenostomum noumeanum Ther. Oc
Hymenostomum obscurissimum Dix. As3
[Hymenostomum obtusatum (C. Miill.) Broth. Am5 - Weissia
obtusata C. Miill.]
Hymenostomum olivaceum C. Miill. ex Geh. Austr1
[Hymenostomum opacum Wag. & Dix. Afr4 = Barbu/a indica
(Hook.) Spreng. fide Magill, Fl. S. Afr. I. Mosses 1: 243, 1981
(1982)]
[Hymenostomum ovale Williams Oc
Weissia ova/is (Williams)
Bartr.]
[Hymenostomum papi/losissimum Lazar., Dopov. Akad. Nauk Ukr.
R.S.R. ser. B 8: 752, 1967 nom. inval. in syon. Eur Weissia
papillosissima Lazar.]
=
=
=
=
=
=
=
=
GENERA OF THE POTTIACEAE
306
[Hymenostomum papillosissimum (Lazar.) Savicz-Ljubitsk., Novosti
Sist. Niz. Rast. 6: 248, 1969 [1970] Eur Weissia papillosissima
Lazar.]
Hymenostomum patulum (Knight) Dix. Austr2
[Hymenostomum perpusil/um (C. Mtill.) Par., Acta Soc. Linn. Bordeaux 49: 272, 1895 [1896], Ind. Bryol. [2]: 596, 1896 Austr1
Weissia perpusilla (C. Miill.) Stone, J. Bryol. 11: 231, 1980]
[Hymenostomum pulicare Besch. Afr3 = Trichostomum pulicare
(Besch.) Zand., see treatment of Trichostomum]
[Hymenostomum riograndense Broth. Am5 = Weissia riograndensis
(Broth.) Zand., see treatment of Weissia]
[Hymenostomum rostel/atum (Brid.) Schimp. Eur = Weissia rostel/ata
(Brid.) Lindh.]
[subsp. meylanii (Amann) Podp. Eur = Hymenostomum squarrosum
Nees & Hornsch. fide Crundwell & Nyholm, J. Bryology 7: 9,
1972 Weissia squarrosa (Nees & Hornsch.) C. Miill.]
[var. meylanii (Amann) Podp. Eur = Hymenostomum squarrosum
Nees & Hornsch. fide Crundwell & Nyholm, J. Bryology 7: 9,
1972 = Weissia squarrosa (Nees & Hornsch.) C. Mtill.]
[var. phascoides (Hook.) Card. Eur Am1 = Weissia rostel/ata var.
phascoides (Hook.) Reese & Lemmon, Bryologist 68: 283, 1965]
[Hymenostomum semidiaphanum Ther. Am2 = Weissia semidiaphana
(Ther.) Zand., Monogr. Syst. Bot. Missouri Bot. Gard. 11 : 197,
1985]
[Hymenostomum semiinvolutum (C. Miill.) Kindb. Am6 = Weissia
semiinvoluta C. Miill.]
Hymenostomum siamense Dix. As3
[Hymenostomum socotranum (Mitt.) Broth. Afr2 Weissia socotrana
Mitt.]
[Hymenostomum squarrosum Nees & Hornsch. Eur
Weissia
squarrosa (Nees & Hornsch.) C. Miill.]
[Hymenostomum striatum Geh. & Hampe Am6 = Weissia glaziouii
Zand. nom. nov., see treatment of Weissia]
[Hymenostomum strictifolium Dix. & Sak. As2 = Weissia edentula
Mitt. fide Saito, J. Hattori Bot. Lab. 39: 421, 1975]
[Hymenostomum subacaule (Mitt.) Par., Acta Soc. Linn. Bordeaux
49: 272, 1895 (1896), Ind. Bryol. (2): 596, 1896 Am4 Weissia
subacaulis (Mitt.) Par.]
[Hymenostomum submicaceum (C. Mtill.) Par., Acta Soc. Linn. Bordeaux 49: 273, 1895 [1896], Ind. Bryol. [2] : 597, 1896 Am5 =
Weissia submicacea C. Miill.]
Hymenostomum subrostellatum Schimp. ex Basch. Afrl
Hymenostomum sullivanii C. Miill. ex Geh. Austr1
[Hymenostomum termitarum (C. Miill.) Broth. Am5 Trichostomum
termitarum (C. Miill.), see treatment of Trichostomum]
[Hymenostomum termitidarum (C. Miill.) Par., Acta Soc. Linn. Bordeaux 49: 273, 1895 (1896), Ind. Bryol. (2): 597, 1896 Afr2
Weissia termitidarum C. Miill.]
[Hymenostomum tortile (Schwaegr.) B.&S. in BSG Eur As1 As3 As5
Afr1 Austr2 Am1 Am2 = Weissia condensa (Voit) Lindh. fide
Corley eta!., J. Bryol. 11 : 623, 1981 (1982)]
var. brevifolium Schiffn. Eur
[var. intermedium (Monk.) Podp. Eur = Weissia torti/is var.
intermedia Monk.]
var. pseudocrispatum Podp. Eur
var. subalpinum Kern Eur
[var. subcy/indricum BSG Eur Weissia torti/is var. subcy/indrica
(BSG) Dix.]
var. tunetana Besch. in Besch. & Pat. Afr1
[fo. brevifolium Amann, Fl. Mouss. Suisse 3: 7, 1923 hom. il/eg.
non Hymenostomum tortile var. brevifolium Schiffn. EurJ
fo. rufidulum Podp., Bul. Griid. Bot. Univ. Cluj 11 : 55 , 1931 Eur
[Hymenostomum urceolare (Hampe) Hampe Am5
Trichostomum
=
=
=
=
=
=
=
=
=
=
urceo/are (Hampe) Zand., see treatment of Trichostomum]
HYMENOSTYLIELLA Bartr.
Hymenostyliella alata (Herz.) Robins., Phytologia 21: 3, 1971
(Timmie/la) Am4
Hymenosty/iel/a calcarea (Dix.) lwats., J. Hattori Bot. Lab. 40:
146, 1976 (Diphyscium) As3
[Hymenostyliel/a involuta (Card. & Ther.) Bartr. As4 = Hymenostyliella 1/anosii (Broth.) Robins. fide Robinson, Phytologia 21 : 2,
1971]
[Hymenostyliella japonica (Broth.) Saito, J. Jap. Bot. 46: 145, 1971
As2 = Didymodon japonicus (Broth.) Saito, J. Hattori Bot. Lab.
39: 508, 1975]
Hymenostyliella 1/anosii (Broth.) Robins., Phytologia 21 : 2, 1971
(Barbula) As2 As3
HYMENOSTYLIUM Brid.
[Hymenostylium annotinum Mitt. ex Dix. As3 = Hymenostylium
recurvirostrum var. insigne (Dix.) Bartr. fide Zander & Eckel,
Canad. J. Bot. 60: 1596, 1982]
[Hymenostylium anoectangioides (C. Miill.) Broth. As2 = Hymenostylium recurvirostrum (Hedw.) Dix. fide Chen, Hedwigia 80:
62, 1941]
[Hymenostylium aurantiacum Mitt. As2 As3 As4 = Hymenostylium
recurvirostrum var. aurantiacum (Mitt.) Gangulee, Mosses E.
India 648, 1972 (Hymenostylium) hom. il/eg. incl. var. prior. =
Hymenostylium recurvirostrum var. luzonense (Broth.) Bartr.
fide Gangulee, Mosses E. India 3: 648, 1972 =Hymenostylium
recurvirostrum var. cylindricum (Bartr.) Zand., see treatment of
Hymenosty/ium]
[Hymenostylium col/enchymaticum Baumg. & Frohl. As4 =
Hymenostylium recurvirostrum (Hedw.) Dix. fide Sollman in
Touw, J. Hattori Bot. Lab. 71: 343, 1992]
Hymenostylium congoanum Dix. & Nav. Afr2
Hymenostylium contextum Herz. Am4
[Hymenostylium courtoisii Broth. & Par. As2 = Hymenostylium
recurvirostrum var. aurantiacum (Mitt.) Gangulee, Mosses E.
India 648, 1972 hom. illeg. incl. var. prior = Hymenostylium
recurvirostrum var. luzonense (Broth.) Bartr. fide Gangulee,
Mosses E. India 3: 648, 1972 = Hymenostylium recurvirostrum
var. cylindricum (Bartr.) Zand., see treatment of Hymenostylium]
Hymenostylium crassinervium Broth. & Dix. Afr2 Afr4
Hymenostylium crispulum Broth. & Par. Afr2
[Hymenostylium curvirostre Mitt. = Hymenostylium recurvirostrum
(Hedw.) Dix.]
Hymenostylium recurvirostrum var.
[var. bicolor Dix. As3
bicolor (Dix.) Crum ]
[fo. commutatum (Mitt.) Hag., Musci Norv. Bor. 2, 1899
(Hymenostomum) Eur
Hymenostylium recurvirostrum var.
commutatum (Mitt.) Podp.]
[fo. megalosporum Latz., Hedwigia 66: 138, 1926 Eur Hymenosty/ium recurvirostrum fo. mega/osporum (Latz.) Podp.]
[fo. scabrum (Lindh.) Hag., Musci Norv. Bor. 2, 1899 Eur =
Hymenostylium recurvirostrum var. latifo/ium (Zett.) Wijk &
Marg. = Hymenostylium recurvirostrum (Hedw.) Dix. fide
Zander in Sharp et al., Moss Fl. Mex.]
[fo. serrulatum Roll , Hedwigia 38: 260, 1899 Eur = Hymenostylium recurvirostrum fo. serrulatum (Roll) Podp.]
Hymenostylium dicranelloides Broth. ex Dix. As3
Hymenostylium diversifolium Frohl., Ann. Naturhist. Mus. Wien
67: 152, 1964 As3
Hymenostyliumfiliforme Dix. As3
=
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
[Hymenosryliumfirmum (C. MUll.) Broth. in Bartr. Oc = Hymenosrylium recurvirostrum (Hedw.) Dix. var. recurvirostrum, see treatment of Hymenostylium]
[Hymenosrylium glaucum (C. MUll.) Broth. = Hymenostylium
recurvirostrum (Hedw.) Dix.]
[var. cylindricum Bartr. Am3 (= Hymenosrylium recurvirostrum
(Hedw.) Dix. fide Zander, Bryologist 80: 253, 1977) = Hymenosry/ium recurvirostrum var. cylindricum (Bartr.) Zand. in Zand. &
Eckel, Canad. J. Bot. 60: 1982]
Hymenosry/ium grandirete Dix., Anniv. Vol. Bot. Gard. Calcutta 178,
1942 As3
Hymenosrylium hildebrandtii (C. MUll.) Zand. (Weissia) Afr2
[Hymenostylium insigne (Dix.) Podp. Eur
Hymenostylium
recurvirostrum var. insigne (Dix.) Bartr. fide Zmder & Eckel,
Canad. J. Bot. 60: 1596, 1982]
Hymenosrylium kunzeanum (C. MUll.) C. Miill. nom. il/eg. Am6
[Hymenostylium luzonense Broth. = Hymenosry/ium recurvirostrum
var. cylindricum (Bartr.) Zand., see treatment of Hymenosrylium]
[var. minus Broth. in Hall. As4 = Hymenosrylium recurvirostrum
(Hedw.) Dix.fide Sollman in Touw, J. Hattori Bot. Lab. 71: 343,
1992]
Hymenosrylium papillinerve Dix. Afr2
[Hymenosrylium pellucidum Broth. & Yas. As2 = Hymenosrylium
recurvirostrum (Hedw.) Dix.fide Chuang, J. Hattori Bot. Lab. 37:
469, 1973]
Hymenosry/ium recurvirostrum (Hedw.) Dix. Eur As1 As2 As3 As4
AsS Afr1 Afr2 Afr4 Am1 Am2 Am3 Am4 Am6 Austr1 Austr2
[var. aurantiacum (Mitt.) Gangu1ee, Mosses E. India 648, 1972
hom. il/eg. incl. var. prior= Hymenostylium recurvirostrum var.
luzonense (Broth.) Bartr. fide Gangulee, Mosses E. India 3: 648,
1972 = Hymenosrylium recurvirostrum var. cylindricum (Bartr.)
Zand. (based on Hymenostylium g/aucum var. cylindricum Bartr.,
1936), see treatment of Hymenostylium]
var. anoectangioides (TMr.) Wijk & Marg. Eur
var. bicolor (Dix.) Crum As3
var. cataractarum (Schimp.) Podp. Eur As1 As2 As3
var. commutatum (Mitt.) Podp. Eur As1 As2 As3 Am1
var. cylindricum (Bartr.) Zand. in Zander & Eckel, Canad. J. Bot.
60: 1599, 1982 (Hymenosrylium glaucum var. cylindricum Bartr.,
1936) As2 As3 Am2 Am3 Oc
var. insigne (Dix.) Bartr. Eur As2 As3 Am1
[var. latifolium (Zett.) Wijk & Marg. Eur As1 Am1 =Hymenostylium recurvirostrum (Hedw.) Dix. fide Zander in Sharp et a!.,
Moss Fl. Mex.]
[var. luzonense (Broth .) Bartr. As4 = Hymenostylium
recurvirostrum var. cylindricum (Bartr.) Zand., see treatment of
Hymenosrylium]
var. sendtnerianaeforme (Gyorf.) Podp. Eur
fo. crassicoligum Van!k in Smarda, Cas. Morav. Mus. Zemsk. 32:
15, 1948 Eur
fo. laeviuscu/a (Lindb.) Podp., Consp. Muse. Eur. 173, 1954 (Barbuta curvirostris var.) Eur As1
fo. megalosporum (Latz.) Podp., Consp. Muse. Eur. 172, 1954
(Hymenosrylium curvirostre fo.) Eur
fo. microcarpum (Nees & Homsch.) Podp., Consp. Muse. Eur. 172,
1954 (Gymnostomum) Eur As2
fo. minimum (Amann) Podp., Consp. Muse. Eur. 172, 1954
(Hymenostylium curvirostre var.) Eur
fo. pallidisetum (Nees & Homsch.) Podp., Consp. Muse. Eur. 172,
1954 (Gymnostomum) Eur
fo. serrulatum (Roll) Podp., Consp. Muse. Eur. 172, 1954
(Hymenostylium curvirostre fo.) Eur
Hymenostylium rigescens (C. Miill.) Broth. Afr2 Afr4
=
307
Hymenostylium scaturiginosum (C. Miill.) Broth. Afr2 Afr4
[Hymenosrylium secundum C. Miill. Afr2 =Molendoa sendtneriana
(BSG) Limpr., see treatment of Molendoa]
[Hymenosrylium shepheardae Card. & Dix. As3 Anoectangium
shepherdae (Card. & Dix.) Zand., see treatment of Anoectangium]
Hymenostylium sinense Sak. As2
[Hymenostylium sordidum Card. As2 = Gymnostomum
recurivrostrum Hedw. fide Saito, J. Hattori Bot. Lab. 39: 39:
452, 1975 =Hymenostylium recurvirostrum (Hedw.) Dix.]
Hymenostylium subcrispulum Ther. Afr3
Hymenosrylium xanthocarpum (Hook.) Brid. [good species fide
Aziz & Vohra, Bull. Bot. Surv. India 30: 185, 1988] As3
[Hymenostylium validinerve Dix. & P. Yarde As3 = Molendoa
sendtneriana (BSG) Limpr., see treatment of Molendoa]
=
HYOPHILA Brid.
Hyophila acuminata Broth. & P. Yarde Afr2 Afr3
Hyophila acuminata Bartr. hom. illeg. As4
Hyophila acutifolia Saito, J. Hattori Bot. Lab. 39: 470, 1975 As2
[Hyophila acutiuscu/a Broth. Afr2 Trichostomum acutiusculum
(Broth.) Zand., see treatment of Trichostomum]
[Hyophila afrophaea (C. MUll.) Wamst. Afr4 = Trichostomum
brachydontium Bruch ex F. Mii11. fide Magill, Fl. S. Afr. I.
Mosses 1: 260, 1981]
[Hyophila amblyphylla Card. As2 = Merceya ligulata (Spruce)
Schimp. fide Noguchi, J. Jap. Bot. 35(10): 316, 1960 = Scopelophila ligu/ata (Spruce) Spruce]
[Hyophila angustifolia Card. hom. il/eg. As2 = Trichostomum
platyphyllum (Broth. ex lhs.) Chen fide Saito, J. Hattori Bot.
Lab. 39:434, 1975]
Hyophila angustifo/ia Par. & Ren. Afr3
Hyophila angustiuscula Baumg. & Dix. As4
Hyophila anoectangioides C. Miill. ex Dus. Afr2
Hyophila apiculata Fleisch. As4
[Hyophila arborea (Mitt.) Jaeg. Am5
Trichostomum arboreum
(Mitt.) Zand., see treatment of Trichostomum]
Hyophila argentinica Ther. Am6
[Hyophila asanoi Sak. As2 = Barbula indica (Hook.) Spreng. fide
Saito, J. Hattori Bot. Lab. 39: 488, 1975]
Hyophila ascensionis Card. Afr2
Hyophi/a assimilis Broth. Am5
[Hyophila atrovirens (C. MUll.) Broth. Afr2 Afr4 = Hyophila
involuta (Hook.) Jaeg . fide Magill, Fl. S. Afr. I. Mosses 1: 228,
1981 (1982)]
var. oubanguiensis Ther. & P. Yarde Afr2
Hyophila baginsensis C. Miill. Afr2 Afr4
Hyophila bartramiana Steere Am2
[Hyophila basutensis Sim Afr4 = Didymodon certatodonteus (C.
MUll.) Dix. fide Magill, Fl. S. Afr. I. Mosses 1: 235, 1981
(1982)]
Hyophila beruensis Dix. Oc
[Hyophila biloinsularis Sak. As2 = Barbula indica (Hook.) Spreng.
fide Saito, J. Hattori Bot. Lab. 39: 488, 1975 as "bilo-insulare"]
Hyophila bingeri Broth. & Par. Afr2
Hyophila blanda (Hook. & Wils.) Jaeg. Am5
Hyophila brevifolia Hampe Am5
[Hyophila calymperoides Ther. & Nav. Afr2 = Bryoerythrophyllum
campylocarpum (C. Miill.) Crum., see treatment of Bryoerythrophy/lum]
Hyophila combae Broth. Oc
[Hyophila comosa Dix. & P. Yarde As3 = Hyophila rosea Williams
fide Zander in Sharp et a!., Moss Fl. Mex. = Hyophila
=
=
308
GENERA OF THE POTTIACEAE
nymaniana (Aeisch.) Menzel, Willdenowia 22: 198, 1992]
Hyophila compacta (Welw. & dub.) Jaeg. Afr2
Hyophila congo/ens is Ther. & Nav. Afr2
[Hyophila contorta (Kunz.) Jaeg. Afr1 = Trichostomum contortum
(Kunze) Sergio, Portug. Acta Biol. (B) 14: 169, 1985]
[Hyophila crenulata C. MUll. ex Dus. Afr2 = Hyophila involuta
(Hook.) Jaeg.fide Sollman, Lindbergia 10: 55, 1984]
var. brevifolia Biz., Svensk. Bot. Tidsk. 63: 444, 1969 nom. inval.
Afr2
[Hyophila crenulata Mach. hom. illeg. Eur = Hyophila machadoana
Sergio, Rev. Bryol. Lichenol. 36:628, 1969 nom. nov. =Bryoerythrophyllum machadoanum (Sergio) Hill fide Hill, J. Bryol. 11:
601, 1981 (1982) = Bryoerythrophyllum campylocarpum (C.
MUll.) Crumfide Sollman, Lindbergia 16: 22, 1990]
Hyophila crenulatula C. Miill. ex Par. Afr2
Hyophila crispula (Sak.) Sak.
[Hyophila cucullatifolia Gao, Jia & Cao, Bull. Bot. Res. (Harbin)
11(2): 29. 1991 As2 = Weisiopsis cucullatifolia (Gao, Jia & Cao)
Zand., see treatment of Weisiopsis]
Hyophila cuspidatissima Par. & Broth. Afr2
[Hyophila cyathiformis (Dix.) Sim Afr2 Afr4 =Hypodontium dregei
(Homsch.) C. Miill. fide Magill & Schelpe, Mem. Bot. Surv. S.
Afr. 43: 23, 1979]
Hyophila dendroides Dix. As3
Hyophila elliptica Baumg. & Frohl. As4
[Hyophila erosa Sim As4 = Oreoweisia erosa (C. MUll.) Kindb. fide
Magill & Schelpe, Mem. Bot. Surv. S. Afr. 43: 23, 1979]
Hyophila excurrentinervis Par. & Broth. Afr2
[Hyophila flavipes Broth. As2 =Hyophila involuta (Hook.) Jaeg. fide
Saito, J. Hattori Bot. Lab. 39: 468, 1975]
Hyophilafouta-djalloni Par. & Broth. Afr2
Hyophila girodii Ren. & Card. Afr3
Hyophila glaucoviridis Par. & Broth. Afr2
Hyophila grandiretis Sak.. hom. illeg. As2
Hyophila grossidens Broth. Am4
Hyophila guaraya Herz. Am4
Hyophila gymnostomoides (Welw. & dub.) Jaeg. Afr2
Hyophila holstii Broth. Afr2
Hyophila incurva Jaeg. Am2 Am5
Hyophila integrifolia Dix. & Ther. Afr2
Hyophila involuta (Hook.) Jaeg. Eur As2 As3 As4 Afr4 Ami Am2
Am3 Am4 Am5 Oc
fo. circinata (C. MUll.) Chen, Hedwigia 80: 190, 1941 (Pottia) As3
fo. flaccida (Warns!.) Podp., Consp. Muse. Eur. 199, 1954 (Trichostomum warnstorfii var.) Eur
fo. serrata (Amann) Podp., Consp. Muse. Eur. 199, 1954 (Hyophila
riparia var.) Eur
Hyophila involutifolia (C. MUll.) Jaeg. Am4
Hyophilajavanica (Nees & Blum.) Brid. Asl As2 As3 As4
[Hyophila khartoumensis (Pettet) A. Sm. & H. Whiteh., J. Bryol. 8:
14, 1974 (Tortula) Afr2 = Desmatodon bogosicus C. MUll. fide
Corley et al., J. Bryol. 11: 620, 1981 (1982)]
Hyophila kurziana Gangulee, Nov. Hedw. 12:422, 1966 As3
Hyophila laete-virens Broth. Am5
Hyophi/a lanceolata Ren. & Card. Afr3
Hyoplzila latifolia Broth. Afr2
[Hyoplzila lauterbachii Broth. in Schum. & Lauterb. As4 =Hyophila
involuta (Hook.) Jaeg. fide Norris & Koponen, Acta Bot. Fenn.
137: 109, 1987]
Hyophila leikipaiae (C. Miill.) Broth. Afr2
Hyophila leioneura Ren. & Par. Afr3
[Hyophila leprieurii (Mont.) Jaeg. Am5 = Hyophila tortula
(Schwaegr.) Hampe fide Florsch., Mosses Suriname 1: 173, 1964
= Hyophila involuta (Hook.) Jaeg.
[Hyophila ligulaefolia Broth. & Par. Afr2 = Trichostomum
ligulaefolium (Broth. & Par.) Zand., see treatment of Trichostomum]
Hyophila linguaeformis Broth. ex lhs. As2
[Hyophila Ungulata Card. Am2 = Neohyophila sprengelii var.
stomatodonta (Card.) Zand. fide Zander, Bryologist 86: 137,
1983 = Plaubelia sprengelii var. stomatodonta (Card.) Zand.,
see treatment of Plaubelia]
[Hyophila liukiensis Sak. As2 = Hyophila involuta (Hook.) Jaeg.
fide Saito, J. Hattori Bot. Lab. 39: 468, 1975]
[Hyophila /ombokensis Broth. in Hall. As4 = Trichostomum
brachydontium Bruch fidy Sollman in Touw, J. Hattori Bot.
Lab. 71:345, 1992]
Hyophila loxorhyncha C. MUll. ex Anstr. Am5
[Hyophila /usitanica Card. & Dix. Eur = B,ryoerythrophyllum
lusitanicum (Card. & Dix.) Hill, J. Bryol. 11: 600, 1981 (1982)]
[Hyophila machadoana Sergio, Rev. Bryol. Lichenol. 36: 628, 1969
nom. nov. Eur = Bryoerythrophy/lum machadoanum (Sergio)
Hill fide Hill, J. Bryol. 11: 601, 1981 [1982] = Bryoerythrophyllum campylocarpum (C. MUll.) Crum fide Sollman,
Lindbergia 16: 22, 1990]
Hyophila mattogrosensis Broth. Am5
Hyophila melanostoma (Mitt.) Jaeg. Am5
[Hyophila mexicana Ther. Am2 = Trichostomum brachydontium
Bruch ex F. MUll. fide Zander in Sharp et al., Moss A. Mex.]
[Hyophila machadoana Sergio, Rev. Bryol. Lichenol. 36: 628, 1969
nom. nov. for Hyophila crenulata Mach. =Bryoerythrophyllum
machadoanum (Sergio) Hill fide Hill, J. Bryol. 11: 601, 1981
(1982)]
[Hyophila micholitzii Broth. = Hyophila involuta (Hook.) Jaeg.]
var. sterilis Fleisch. As3 As4 Oc
[Hyophila microcarpa (Schimp. ex Besch.) Broth. Am2 Am3 Am4
Am5 = Weissia sinaloensis Bartr. fide Zander in Sharp et al.,
Moss Fl. Mex. Trichostomum sinaloense (Bartr.) Zand., see
treatment of Trichostomum]
[Hyophila minutissima (Mitt.) Jaeg. Am4 Am5
Tisserantiella
minutissima (Mitt.) Zand., see treatment of Tisserantiella
(Rhachitheciaceae), Excluded Taxa]
Hyophila millifolia Dix. & P. Yarde As3
Hyophila mosenii Both. Am5
Hyophila muelleri (Dub.) Jaeg. As3
[Hyophila naganoi Sak. As2 = Hyophila propagulifera Broth. fide
Saito, J. Hattori Bot. Lab. 39: 471, 1975]
[Hyophila nakayamae Sak.. As2 = Barbula indica (Hook.) Spreng.
fide Saito, J. Hattori Bot. Lab. 39: 488, 1975]
[Hyophila nakayamae Sak.. As2 = Barbula indica (Hook.) Spreng.
fide Saito, J. Hattori Bot. Lab. 39: 488, 1975]
Hyophila neocaledonica Broth. & Par. Oc
Hyophila niam-niamiae C. MUll. Afr2
Hyophila novae-guineae Broth. ex C. MUll. As4
Hyophila novae-seelandiae Dix. & Sainsb. Austr2
Hyophila nymaniana (Fleisch.) Menzel, Willdenowia 22: 198, 1992
(Glyphomitrium) As3 As4 Am2
Hyophila obtusifolia (C. MUll.) Jaeg. Am5 [= Gyroweisia sp. cf.
Brotherus 1902]
Hyophila ochracea Broth. Am5
[Hyophila okamurae Broth. As2 = Hyophila propagulifera Broth.
fide Saito, J. Hattori Bot. Lab. 39: 471, 1975]
Hyophila ovalifolia (Hampe) Hampe Am5
Hyophila pampanini Zodda Afrl
Hyophila paraguensis Broth. Am5
Hyophila parieta/is Card. in Grand. Afr3
=
=
309
GENERA, SPECIES AND INFRASPECIAC TAXA
[Hyophila perannulata Ren. & Card. As3 = Trichostomum criorum
Zand. nom. nov., see treatment of Trichostomum]
Hyophila perpendiculata Dix. Afr2
Hyophila perpusilla Ther. & Trab. Afr1
Hyophila potieri Besch., Rev. Bryol. 7(2): 21, 1880 Afr2 Afr3
var. atroviridis Ren. & Card. Afr3
var. denticulta Broth. Afr2
Hyophi/a procera Par. & Broth. Afr2
Hyophi/a propagulifera Broth. As2
[var. e/ara Sak. As2 = Hyophila propagulifera Broth. fide Saito, J.
Hattori Bot. Lab. 39: 471, 1975]
(Hyophila pulchella (Ther. & Hilp.) Wijk & Marg. Afr2 = Tisserantiella pulchella (Ther. & Hilp.) Zand., see treatment of Tisserantiella (Rhachitheciaceae)]
Hyophila punctulata (Mitt.) Kindb. Afr3
Hyophila regnellii C. MUll. ex Angstr. Am5
[Hyophila rosea Williams As2 As3 As4 Am2 =Hyophila nymaniana
(Fleisch.) Menzel, Willdenowia 22: 198, 1992]
Hyophi/a rubiginosa Hampe Am5
Hyophi/a sakalavensis Par. & Ren. Afr3
Hyophila samoana Mitt. Oc
Hyophila setschwanica (Broth.) Hilp. ex Chen As2
Hyophila siamensis Dix. As3
[Hyophila sieboldii Besch. As2 = Barbula unguiculata Hedw. fide
Sollman, Lindbergia 10: 54, 1984]
[Hyophila sinaloensis (Bartr.) Bartr. Am2 Am3 Am4
Weissia
sinaloensis Bartr.fide Zander in Sharp et al., Moss Fl. Mex.]
[Hyophila somaliae C. MUll. Afr2
Torre/la somaliae (C. Miill.)
Zand., see treatment of Torre/la]
Hyophila spathulata (Harv.) Jaeg. As3
[Hyophila sranfordensis (Steere) A. Sm. & Whiteh., J. Bryol. 8: 13,
1974. Eur Am1 = Tortula stanfordensis Steere = Hennediella
stanfordensis (Steere) Blockeel, J. Bryol. 16: 191, 1991]
[Hyophila srenophylla Card. As2 = Trichostomum platyphyllum
(Broth. ex lhs.) Chen fide Saito, J. Hattori Bot. Lab. 39: 434,
1975]
[Hyophila sromatodonta Card. Am2 = Neohyophila sromarodonta
(Card.) Crum = Neohyophila sprengelii var. sromatodonra (Card.)
Zand., Bryo1ogist 86: 138, 1983 = Plaubelia sprengelii var.
sromarodonta (Card.) Zand., see treatment of Plaubelia]
Hyophila streimannii Norris & T. Kop., Acta Bot. Fenn. 137: 111,
1989 As4
[Hyophila styriaca Glow. Eur = Gymnosromum aeruginosum Sm. fide
Corley et al., J. Bryol. 11: 649, 1981 (1982)]
Hyophila subacutiuscula P. Yarde & Ther. Afr2
[Hyophila subangusrifolia Ther. Am2 = Weissia subangustifolia
(Ther.) Zand., Bryo1ogist 86: 156, 1983
Trichostomum
subangusrifolium (Ther.) Zand., see treatment of Trichosromum]
Hyophila subcucullata Williams Am3
Hyophila subflaccida Broth. & Dix. As3
[Hyophila subspathulara Sak. As2 = Trichosromum platyphyllum
(Broth. ex Ihs.) Chen fide Saito, J. Hattori Bot. Lab. 39: 434,
1975]
Hyophi/a tisseranrii P. Yarde Afr2
[Hyophi/a rorrula (Schwaegr.) Hampe Am1 Am2 Am3 Am4 Am5
Am6 =Hyophi/a involuta (Hook.) Jaeg.]
Hyophi/a treleasii Card. Afr1
Hyophila uleana C. Miill. Am5
[Hyophila usambarica Broth. Afr2 = Trichostomum brachydonrium
Bruch, see treatment of Trichosromum]
[Hyophila validinervis Card. & P. Yarde As3 = Hyophila involuta
(Hook.) Jaeg.fide Sollman, Lindbergia 10: 55, 1984]
Hyophila variegara Angstr. Am5
=
=
=
Hyophila vicroriae C. Miill. ex Dus. Afr2
Hyophila viridula Card. & P. Yarde As3
Hyophila vitiana (C. Miill.) Jaeg. Oc
[Hyophi/a walkeri Broth. As3 =Torre/la walkeri (Broth.) Zand., see
treatment of Tortel/a]
Hyophila warmingii Hampe Am5
var. angustifolia Broth. Am5
[Hyophila zeyheri (Hampe) Jaeg. Afr2 Afr4 = Trichostomum
brachydontium Bruch ex F. Miill. fide Magill, Fl. S. Afr. I.
Mosses 1: 260, 1981 (1982)]
var. brevimucronata P. Yarde Afr2
var. lanceolara Sirn Afr4
=
[HYOPHILOPSIS Card. & Dix.
Tortula sect. Hyophilopsis
(Card. & Dix.) Zand., see treatment of Tortula]
[Hyophi/opsis entosthodontacea Card. & Dix. As3 = Torrula
enrosthodontacea (Card. & Dix.) Zander, see treatment of Torrula]
=
[HYOPHILOPSIS Crum non Card. & Dix., 1911 hom. illeg.
Neohyophila Crum = Plaubelia Brid.]
[Hyophilopsis lingulata (Card.) Crum Am2 = Neohyophila
sprengelii var. stomatodonta (Card.) Zand. fide Zander,
Bryologist 86: 137, 1983
Plaubelia sprengelii var.
sromarodonta (Card.) Zand., see treatment of Plaubelia]
[Hyophi/opsis sparhulifolia (Bartr.) Crum Am2 = Neohyophila
sprengelii (Schwaegr.) Crum s. lat. fide Zander, Bryologist 86:
135, 1983 = Plaubelia sprengelii (Schwaegr.) Zand., see treatment of Plaubelia]
[Hyophilopsis sprengelii (Schwaegr.) Crum Am1 Am2 Am3 Am4
Am5 = Neohyophi/a sprengelii (Schwaegr.) Crum = Plaubelia
sprengelii (Schwaegr.) Zand., see treatment of Plaubelia]
[Hyophilopsis sromarodonta (Card.) Crum Am2 Am3 = Neohyophila sprengelii var. stomatodonta (Card.) Zand., Bryologist 86:
135, 1983 = Plaubelia sprengelii var. stomatodonta (Card.)
Zand., see treatment of Plaubelia]
=
HYPODONTIUM C. Miill., Hedwigia 38: 96, 1899
Hypodonrium dregei (Homsch.) C. Miill., Hedwigia 38: 97, 1899
Afr4
Hypodonrium pomiforme (Hook.) C. Miill., Hedwigia 38: 91, 1899
Afr4
[KLEIOWEISIOPSIS Dix. referred to Orthotrichaceae, see treatment of Kleioweisiopsis, Excluded Taxa]
[Kleioweisiopsis involuta Biz., Rev. Bryol. Lichenol. 40: 119, 1974
Afr2 Weissia bizotii Zand. nom. nov.-see treatment of Weissia]
=
LEPTOBARBULA Schimp.
Leprobarbula berica (De Not.) Schirnp. Eur As3 As5 Afrl
var. meridiana/is (Schimp.) Limpr. Eur
var. winteri (Schimp.) Limpr. Eur
LEPTODONTIELLA Zand. & Hegew.
Leptodontiella apicu/ata (Zand.) Zand. & Hegew., Bryologist 79: ·
16, 1976Am4
LEPTODONTIUM (C. Miill.) Hampe
[Leprodonrium abyssinicum Broth . Afr2 = Leprodo ntium
capituligerum C. Miill. fide Sloover, Bull. Jard. Bot. Nat!.
Belgique 57(3/4) : 448, 1987]
[Leptodontium acutifolium Mitt. Am2 Am4 = Leptodontium
310
GENERA OF THE POTTIACEAE
pungens (Mitt.) Kindb.fide Zander, Bryologist 75: 256, 1972]
[var. grimmioides (Britt.) Herz. = Leptodontium pungens (Mitt.)
Kindb. fide Zander, Bryologist 75: 256, 1972]
[Leptodontium acutissimum Bartr. Am4 = Leptodontium pungens
(Mitt.) Kindb. fide Zander, Bryologist 75 : 256, 1972]
Leptodontium aggregatum (C. Miill.) Kindb. As4
[subsp. hyalinum Fleisch., Musci Fl. Buitenz. 1: 369, 1904 As4
Leptodontium aggregatum (C. Miill.) Kindb. fide Norris &
Koponen, Acta Bot. Fenn. 137: 106, 1987]
[var. hyalinum (Fleisch.) Broth., Nat. Pfl. 1(3): 1190, 1909 As4
Leptodontium aggregatum (C. Miill.) Kindh. fide Norris &
Koponen, Acta Bot. Fenn. 137: 106, 1987]
[Leptodontium a/bovaginatum Herz. = Bryoerythrophyllum jamesonii
(Tayl.) Crumfide Zander, Bryologist 75: 278, 1972]
[Leptodontium al/orgei Biz. Afr2 = Cynodontium sp. fide Sloover,
Bull. Jard. Bot. Nat!. Belgique 57(3/4): 449, 1987]
[Leptodontium angustinerve Ther. (= Anoectangium compactum
Schwaegr. fide Zander, Bryologist 75: 278, 1972) =Hymenostylium recurvirostrum (Hedw.) Dix.fide Zander, 80: 243, 1977]
[Leptodontium anoectangiaceum (C. Miill.) Broth. Am5 = Leptodontium stellatifolium (Hampe) Broth. fide Zander, Bryologist 75:
240, 1972]
[Leptodontium anomalum Dix. & Ther. Am4 =Leptodontium wa/lisii
(C. Miill.) Kindb.fide Zander, Bryologist 75: 259, 1972]
[Leptodontium apiculatum Zand., Bryologist 75: 238, 1972 Am4
Leptodontiel/a apicu/ata (Zand.) Zand. & Hegew., Bryologist 79:
16, 1976]
[Leptodontium arachnoideum C. Miill. Am6 = Leptodontium
capituligerum C. Miill.fide Zander, Bryologist 75: 273, 1972]
Leptodontium araucarieti (C. Miill.) Par. Am4 Am5
[Leptodontium arsenii Ther. = Bryoerythrophyllum ferruginascens
(Stirt.) Giac. fide Zander, Bryologist 75 : 278, 1972]
Leptodontium brachyphyllum Broth. & Ther. Am2 Am4 Afr4
[Leptodontium brasiliense Mitt. Am5 = Leptodontium viticulosoides
(P. Beauv.) Wijk & Marg. var. viticulosoides fide Zander,
Bryologist 75: 244, 1972]
[Leptodontium braunioides C. Miill. Am6 = Leptodontium pungens
(Mitt.) Kindb. fide Zander, Bryologist 75 : 256, 1972]
[Leptodontium brevicaule Bartr. Oc = Leptodontium flexifolium
(Dicks.) Hampe in Lindh. fide Zander, Bryologist 75: 231, 1972]
[Leptodontium buesii Williams Am4 = Leptodontium luteum (Tayl.)
Mitt. fide Zander, Bryologist 75: 264, 1972]
[Leptodontium calymperoides Ther. Am4 (= Leptodontium
capituligerum C. Mi.ill. fide Zander, Bryologist 75 : 273, 1972) =
Leptodontium tricolor (Williams) Zand. fide Zander, Bryologist
79: 20, 1976]
Leptodontium capituligerum C. Miill. Am2 Am4 Am5 Am6 Afr2
Afr3
[Leptodontium chrysobaseum (C. Mi.ill.) Broth. Am5 =Leptodontium
stel/atifo/ium (Hampe) Broth. fide Zander, Bryologist 75 : 240,
1972]
[Leptodontium citrinum (Hampe) Hampe Am5 = Leptodontium
viticu/osoides var. panamense (Lor.) Zand., Bryologist 75: 250,
1972 =Leptodontium viticulosoides var. sulphureum (Lor.) Zand.
fide Zander, Bryologist 86: 156, 1983]
Leptodontium debatii (Husn.) Hag. Eur
[Leptodontium densifolium (Mitt.) Mitt. Am4 = Leptodontium
viticu/osoides (P. Beauv.) Wijk & Marg. var. viticulosoides fide
Zander, Bryologist 75: 244, 1972]
[Leptodontium dentatum (Mitt.) Kindb. As3 As4 = Leptodontium
flexifolium (Dicks.) Hampe in Lindh. fide Zander, Bryologist 75 :
231 , 1972 cf. Zander, Bryologist 84: 546, 1972]
[Leptodontium erectifolium Dix. As4 = Leptodontium flexifolium
=
=
=
(Dicks.) Hampe in Lindh. fide Norris & Koponen, Acta Bot.
Fenn: 137: 105, 1987]
Leptodontium erythroneuron Herz. Am4
[Leptodontium exasperatum Card . Am2 = Leptodontium
viticulosoides var. exasperatum (Card.) Zand. fide Zander,
Bryologist 75: 254, 1972]
[Leptodontium excelsum (Sull.) Britt. Ami Am2 = Leptodontium
viticulosoides var. panamense (Lor.) Zand., Bryologist 75: 251,
1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryoloist 86: 156, 1983]
[Leptodontium felipponei Broth. Am6 = Leptodontium
capituligerum C. Mi.ill.fide Zander, Bryologist 75: 273, 1972]
[Leptodontium fernandezianum Broth. in Skottsb. Am6 = Leptodontium longicaule var. microruncinatum (Dus.) Zand. fide
Zander, Bryologist 75: 269, 1972]
[Leptodontium ferrugineum Broth. Am4 = Leptodontium wallisii
(C. Mi.ill.) Kindh.fide Zander, Bryologist 75 : 259, 1972]
[Leptodontium filescens (Hampe) Mitt. Am2 Am4 = Leptodontium
flexifolium (Dicks.) Hampe in Lindh. fide Zander, Bryologist
75: 231 , 1972.
[var. denticulatum Bartr. Am2 = Leptodontium filico/a Herz. fide
Zander, Bryologist 75: 241, 1972]
[Leptodontium filicaule Dix. Afr2 = Leptodontium flexifolium
(Dicks.) Hampe in Lindh. fide Sloover, Bull. Jard. Bot. Nat!.
Belgique 57(3/4): 428, 1987]
Leptodontiumfilicola Herz. Am2 Am4 Am5 Am6
[Leptodontium filiformis (Lor.) Steere Am4 = Leptodontium
flexifolium (Dicks.) Hampe in Lindh. fide Zander, Bryologist
75 : 231, 1972]
Leptodontiumflexifolium (Dicks.) Hampe in Lindh. (see Karttunen,
Taxon 37:156-157, 1988 for discussion of Dickson as authority) Ami Am2 Am4 Afr2 As2 As3 As4
[var. americanum (Grout) Grout Ami Am2 = Leptodontium
flexifolium (Dicks.) Hampe in Lindh. fide Zander, Bryologist
75: 231. 1972]
fo. compacta Hessel. ex Rosevinge, Bot. Iceland 2: 452, 1918 Eur
[fo. gemmifera Monk., Laubm. Eur. 274, 1927 Eur = Leptodontium gemmascens (Mitt. in Hunt) Braithw.]
fo. gemmipara Frahm, Nov. Hedw. 24:418, 1973 [1975] Eur
Leptodontium fuhrmannii Broth. & Irmsch. (type not available for
study fide Zander, Bryologist 75: 278, 1972) Am4
Leptodontium fuscescens Bartr. Am5 = Leptodontium
capituligerum C. Mi.ill:'fide Zander, Bryologist 75: 273, 1972]
Leptodontium gambaragarae Negri Afr2
Leptodontium gemmascens (Mitt. in Hunt) Braithw. [good species
fide Zander, Bryologist 75 : 236, 1972] Eur Afr3
[Leptodontium gemmigerum Broth. in Mildbr. Afr2 =Leptodontium
longicaule Mitt. var. longicaule fide Zander, Bryologist 75:
268, 1972]
[Leptodontium gracile C. Mi.ill. ex Britt. hom. illeg. Am4 = Leptodontium capituligerum C. Mi.ill. fide Zander, Bryologist 75:
273, 1972]
[var. gemmascens Bartr. in Bauer Am5 = Leptodontium
capituligerum C. Mi.ill.fide Zander, Bryologist 75: 273, 1972]
[Leptodontium gracilescens C. Mi.ill . Am4 = Leptodontium
capituligerum C. Mi.ill.fide Zander, Bryologist 75: 273, 1972]
[Leptodontium graci/limum Nog. As2 = Leptodontium flexifolium
(Dicks. ex With.) Hampe in Lindh. fide Zander, Bryologist 75:
232, 1972]
Leptodontium handelii Ther. As2
[Leptodontium humillimum Broth. in Hall. As4 = Leptodontium
flexifolium (Dick. ex With.) Hampe in Lindh. fide Zander,
Bryologist 75: 231, 1972]
GENERA, SPECIES AND INFRASPECIFIC TAXA
=
[Leptodontium hyalinum (Fleisch.) Fleisch. As4
Leptodontium
aggregatum (C. Miill.) Kindb. fide Norris & Koponen, Acta Bot.
Fenn. 137: 106, 1987]
[Leptodontium insolitum Ther. & P. Yarde Afr2 = Barbula eubryum
C. Miill. fide Sloover, Bull. Jard. Bot. Natl. Belgique 57(3/4):
449, 1987]
var. perundulatum Ther. & P. Yarde Afr3
[Leptodontium integrifolium Williams = Barbula integrifolia (Williams) Zand.fide Zander, Bryologist 75: 277, 1972]
Leptodontium interruptum (Mitt.) Broth. Afr4 Austr2
[Leptodontium japonicum Sak. = Didymodon rigidicaulis (C. Miill.)
Saito fide Saito, J. Hattori Bot. Lab. 39: 502, 1975 =Didymodon
ferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11: 599, 1981
(1982)]
[Leptodontium joannis-meyeri C. Miill. Afr2 = Leptodontium pungens
(Mitt.) Kindb. fide Sloover, Bull. Jard. Bot. Natl. Belgique
57(3/4): 434, 1987]
[var. cameruniae Broth. in Mildbr. Afr2 = Leptodontium pungens
(Mitt.) Kindb. fide Sloover, Bull. Jard. Bot. Nat!. Belgique
57(3/4): 434, 1987]
[Leptodontium kinabaluense Dix. As4 = Leptodontium flexifolium
(Dicks.) Hampe in Lindb.fide Zander, Bryologist 75: 231, 1972]
[Leptodontium laevigatum Herz. Am5 = Leptodontium viticulosoides
(P. Beauv.) Wijk & Marg. var. viticulosoides fide Zander,
Bryologist 75: 245, 1972]
[Leptodontium laticuspis Broth. Am4 = Leptodontium syntrichioides
(C. Miill.) Kindb.fide Zander, Bryologist 75: 271, 1972]
Leptodontium latifolium Broth. Afr2
[Leptodontium /axifolium Broth. Am4 =Leptodontium araucarieti (C.
Miill.) Par. fide Zander, Bryologist 75: 265, 1972]
Leptodontium /eptoprion C. Miill. nom. inval. Afr2
[Leptodontium limbatulum Fleisch. As4 = Leptodontium flexifolium
(Dicks.) Hampe in Lindb.fide Zander, Bryologist 75: 231, 1972]
Leptodontium longicaule Mitt. Am2 Am4 Afr2 Afr3 Afr4
subsp. stellatum (Brid.) S1oover, Bull. Jard. Bot. Nat!. Belgique
57(3/4): 444, 1987 (Leptodontium) Afr3
var. microruncinatum (Dus.) Zand., Bryologist 75: 269, 1972 Am4
Am6 Afr2 Afr3 Afr4
Leptodontium luteum (Tayl.) Mitt. Am4 Afr2
[Leptodontium mandonii C. Miill. Am4 = Leptodontium /ongicau/e
Mitt. var.longicaulefide Zander, Bryologist 75: 268, 1972]
[Leptodontium matucamense Besch. Am4 = Leptodontium pungens
(Mitt.) Kindb.fide Zander, Bryologist 75: 256, 1972]
[Leptodontium microruncinatum Dus. Am6
Leptodontium
longicaule var. microruncinatum (Dus.) Zand. fide Zander,
Bryologist 75: 269, 1972]
[Leptodontium nakaii Okam. As2 =Leptodontiumflexifolium (Dicks.)
Hampe in Lindb.fide Saito, J. Hattori Bot. Lab. 39:463, 1975]
[Leptodontium norvegicum Kaal. Eur =Bartramia pomiformis Hedw.
fide Frisvoll, J. Bryol. 12: 186, 1982]
Leptodontium novae-seelandiae C. Miill. Austr2
[Leptodontium orthotrichoides (C. Miill.) Par. Am6 = Zygodon
palmarum C. Miill. (Orthotrichaceae) fide Malta, Gatt. Zygodon
170, 1926 see Zander, Bryologist 75: 278, 1972]
[Leptodontium papil/osum [Hampe] Kindb. Am4 = Leptodontium
longicaule Mitt. var. /ongicaule fide Zander, Bryologist 75: 268,
1972]
Leptodontium paradoxic urn Stone & Scott, J. Bryol. 11: 701, 1981
Austr1
[Leptodontium pergemmascens Broth. As2 = Leptodontium
flexifolium (Dicks.) Hampe in Lindb. fide Zander, Bryologist 75:
231, 1972]
[Leptodontium persquarrosum Broth. in Mildbr. Afr2 =Leptodontium
=
311
wallisii (C. Miill.) Kindb. fide Zander, Bryologist 75: 259,
1972]
Leptodontium planifolium Herz. Am4
[Leptodontium procumbens C. Miill. Am4 = Leptodontium
viticu/osoides var. panamense (Lor.) Zand., Bryologist 75: 251,
1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryologist 86: 156, 1983]
Leptodontium proliferum Herz. Am2 Am4
[Leptodontium pumilum (C. Miill.) Kindb. Afr2 = Leptodontium
pungens (Mitt.) Kindb. fide Sloover, Bull. Jard. Bot. Nat!.
Belgique 57(3/4): 434, 1987]
Leptodontium pungens (Mitt.) Kindb. Afr2 Am2 Am4 Am5 Am6
[Leptodontium quennoae C. Miill. Am6 = Leptodontium
viticu/osoides (P. Beauv.) Wijk & Marg. var. viticu/osoidesfide
Zander, Bryologist 75: 244, 1972]
[Leptodontium ramosum Crum & Richards, J. Bryol. 13: 194, 1984
Am2 = Leptodontium stoloniferum Zand., see treatment of
Leptodontium]
Bryoery[Leptodontium recurvifolium (Tayl.) Lindb. Eur Aml
throphyllum recurvifolium (Tayl.) Zand., Bryologist 75: 277,
1972 = Paraleptodontium recurvifolium (Tayl.) Long, J.
Bryology 12: 181, 1982
Oxystegus recurvifolius (Tayl.)
Zand., Lindbergia 8: 187, 1982 = Trichostomum recurvifolium
(Tayl.) Zand., see treatment of Trichostomum]
Leptodontium repens (C. Miill.) Kindb. Afr2
[Leptodontium rhacomitrioides Lor. & C. Miill. Am4 Am6 = Leptodontium viticu/osoides (P. Beauv.) Wijk & Marg. var.
viticulosoidesfide Zander, Bryologist 75: 244, 1972]
[Leptodontium rhynchophorum Dix. Afr2 = Zygodon sp. fide
Sloover, Bull. Jard. Bot. Nat!. Belgique 57(3/4): 449, 1987]
[Leptodontium rigidum Broth. Am5 = Leptodontium viticulosoides
(P. Beauv.) Wijk & Marg. var. viticulosoides fide Zander,
Bryologist 75: 245, 1972]
[Leptodontium rufescens Broth. in Herz. Am4 = Leptodontium
longicaule var. microruncinatum (Dus.) Zand. fide Zander,
Bryologist 75: 269, 1972]
Leptodontium saxicola [C. Miill.] C. Miill. ex Par. (type not available for study fide Zander, Bryologist 75: 278, 1972) Am5
Leptodontium scaberrimum Broth. As2
[Leptodontium schiffneri Broth. Am5 = Leptodontium wallisii (C.
Miill.) Kindb.fide Zander, Bryologist 75: 259, 1972]
[Leptodontium serrae (C. Miill.) Par. Am5 = Leptodontium
viticulosoides var. panamense (Lor.) Zand., Bryologist 75: 251,
1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryologist 86: 156, 1983]
[Leptodontium serrifolium (C. Miill.) Broth. Afr2 = Leptodontium
pungens (Mitt.) Kindb. fide Sloover, Bull. Jard. Bot. Nat!.
Belgique 57(3/4): 434, 1987]
[Leptodontium setschwanicum Broth. As2 = Didymodon
erosodenticulatus (C. Miill.) Saito fide Saito, J. Hattori Bot.
Lab. 39: 504, 1975]
[Leptodontium sikokianum Sak. in Oti As2 = Dichodontium
pel/ucidum (Hedw.) Schimp. (Dicranaceae) fide Saito, J.
Hattori Bot. Lab. 39: 528, 1975]
[Leptodontium skottsbergii Bartr. Am6 = Leptodontium pungens
(Mitt.) Kindb.fide Zander, Bryologist 75: 256, 1972]
[Leptodontium spinosum Williams Am4 = Zygodon pichinchensis
(Tayl.) Mitt. (Orthotrichaceae) fide Zander, Bryologist 75: 278,
1972]
[Leptodontium spongiosum Herz. Am4 = Leptodontium flexifolium
(Dicks.) Hampe in Lindb. fide Zander, Bryologist 75: 231,
1972]
[Leptodontium squamifolium (C. Miill.) Broth. Am5 = Leptodont-
=
=
GENERA OF THE POTTIACEAE
312
ium stellatifolium (Hampe) Broth. fide Zander, Bryologist 75:
240, 1972]
Leptodontium stellaticuspis Bartr. Am4
Leptodontium stellatifolium (Hampe) Broth. Am5
[Leptodontium stellatum (Brid.) Ren. Afr3 =Leptodontium longicau/e
suhsp. stel/atum (Brid.) Sloover, Bull. Jard. Bot. Nat!. Belgique
57(3/4): 444, 1987]
Leptodontium stoloniferum Zand., Bryologist 75: 239, 1972 Am2
Am4
Leptodontium styriacum (Jur.) Limpr. Eur As1
[Leptodontium subalpinum (De Not.) Lindh. Eur = Dichodontium
pellucidum (Hedw.) Schimp. (Dicranaceae) fide Frahm,
Lindbergia 12: 81, 1987]
[Leptodontium subcirrhifolium (C. Mull.) Kindh. Am4 = Leptodontium viticulosoides var. panamense (Lor.) Zand., Bryologist 75:
251, 1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryologist 86: 156, 1983]
[Leptodontium subfilescens Ther. & Nav. Afr2 = Leptodontium
jlexifolium (Dicks.) Hampe in Lindh. fide Sloover, Bull. Jard.
Bot. Nat!. Belgique 57(3/4): 428, 1987]
[Leptodontium subgracile Ren. & Card. Am2 Am4 = Leptodontium
longicaule Mitt. var. longicau/e fide Zander, Bryo1ogist 75: 268,
1972]
[Leptodontium subgrimmioides Broth. & Ther. Am4 = Leptodontium
pungens (Mitt.) Kindh.fide Zander, Bryo1ogist 75: 256, 1972]
Leptodontium subintegrifolium Ther. ex Herz., Fedde Rep. Spec.
Nov. Reg. Yeg. 45: 45, 1938. Am4
Leptodontium sub/aevifolium Broth. in Mildhr. Afr2
[Leptodontium subplanifolium Ther. Bryoerythrophyllum jamesonii
(Tayl.) Crumfide Zander, Bryologist 75: 278, 1972]
[Leptodontium sulphureum (C. Miill.) Mitt. Am2 Am4 (= Leptodontium viticulosoides var. panamense (Lor.) Zand., Bryologist 75:
250, 1972) =Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryologist 86: 156, 1983]
[var. flagel/aceum Bartr. Am2 Leptodontium viticulosoides var.
jlagellaceum (Bartr.) Zand. fide Zander, Bryologist 75: 255,
1972]
[var. motelayi (Ren. & Card.) Bartr. Am2 = Leptodontium
viticu/osoides var. panamense (Lor.) Zand., Bryologist 75: 251,
1972 =Leptodontium viticulosoides var. sulphureum (Lor.) Zand.
fide Zander, Bryologist 86: 156, 1983]
Leptodontium viticu/osoides var.
[var. panamense Lor. Am2
panamense (Lor.) Zand., Bryologist 75: 250, 1972 = Leptodontium viticulosoides var. sulphureum (Lor.) Zand., Bryologist 86:
156, 1983]
Leptodontium syntrichioides (C. Mull.) Kindh. Am2 Am4
Leptodontium taiwanense Nog. As2
[Leptodontium tenerascens Broth. in Mildhr. Afr2 = Leptodontium
jlexifolium (Dicks.) Hampe in Lindh. fide Zander, Bryologist 75:
231, 1972]
[var. majus Broth. in Mildbr. Afr2 = Leptodontium jlexifolium
(Dicks.) Hampe in Lindb.fide Zander, Bryo1ogist 75: 231, 1972]
var. planifolium P. Yarde & Ther. Afr2
[var. subfilescens (Ther. & Nav.) Ther. Afr2 = Leptodontium
jlexifolium (Dicks.) Hampe in Lindb. fide Sloover, Bull. Jard.
Bot. Nat!. Belgique 57(3/4): 428, 1987]
Leptodontium tricolor (Williams) Zand. in Zand. & Hegew.,
Bryologist 79: 20, 1976 (Williamsiella) Am4
Leptodontium trifarium Broth. [type not available for study fide
Zander, Bryologist 75: 278, 1972] Am5
[Leptodontium turgidum Herz. Am4 = Leptodontium viticulosoides
(P. Beauv.) Wijk & Marg. var. viticulosoides fide Zander,
Bryologist 75: 244, 1972]
=
=
=
[Leptodontium ulocalyx (C. Miill.) Mitt. Am2 Am4 =Leptodontium
viticulosoides var. panamense (Lor.) Zand., Bryologist 75: 250,
1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand.fide Zander, Bryologist 86: 156, 1983]
[var. cirrhifolium (Mitt.) Bartr. Am2 Am4 = Leptodontium
viticulosoides var. panamense (Lor.) Zand., Bryologist 75: 251,
1972 = Leptodontium viticulosoides var. sulphureum (Lor.)
Zand .fide Zander, Bryologist 86: 156, 1983]
[Leptodontium undulatum Herz. Am4 = Leptodontium longicaule
var. microruncinatum (Dus.) Zand. fide Zander, Bryologist 75:
269, 1972]
[Leptodontium vaginatum Herz. Am4 = Leptodontium wallisii (C.
Mull.) Kindb.fide Zander, Bryologist 75: 259, 1972]
[Leptodontium valerianum Bartr.
Gymnostomum valerianum
(Bartr.) Zand., Bryologist 75: 277, 1972
Tuerckheimia
valeriana (Bartr.) Zand., Misc. Bryol. Lichenol. 8: 27, 1978]
[Leptodontium variegatum Herz. Am5 = Leptodontium wallisii (C.
Mull.) Kindb.fide Zander, Bryologist 75: 259, 1972]
Leptodontium viticulosoides (P. Beauv.) Wijk & Marg. Am2 Am4
Am5 Am6 Afr2 Afr3 Afr4 As2 As3 As4
var. abbreviatum (Dix.) Wijk & Marg. As3
var. exasperatum (Card.) Zand., Bryologist 75: 254, 1972 Am2
Am4
var.jlagel/aceum (Bartr.) Zand., Bryologist 75: 255, 1972 Am2
[var. panamense (Lor.) Zand., Bryologist 75: 250, 1972 Ami
Am2 Am3 Am4 Am5 = Leptodontium viticulosoides var.
su/phureum (Lor.) Zand.fide Zander, Bryologist 86: 156, 1983,
I.C.B.N. Art. 57.3]
var. subdenticu/atum (C. Mull.) Wijk & Marg. As4
var. sulphureum (Lor.) Zand., Bryologist 86: 86, 1983 Ami Am2
Am3Am4Am5
[Leptodontium volkensii Broth. Afr2 = Leptodontium viticulosoides
(P. Beauv.) Wijk & Marg. var. viticu/osoides fide Zander,
Bryologist 75: 244, 1972]
Leptodontium wallisii (C. Mull.) Kindb. Am4
[Leptodontium warnstorfii Fleisch. As4 = Leptodontium flexifolium
(Dicks.) Hampe in Lindb. fide Zander, Bryologist 75: 231,
1972]
Leptodontium zygodontoides C. Mull. (type not available for study
fide Zander, Bryologist 75: 278, 1972) Am6
=
=
LUISIERELLA Ther. & P. Yarde
Luisierel/a barbu/a (Schwaegr.) Steere As2 As4 Ami Am2 Am3
Am50c
[LYDIAEA Laz. =Microbryum Schimp.]
[Lydiaea v/assovii (Laz.) Laz. As1 = Phascum vlassovii Laz. fide
Savicz-Ljubitzkaja, L. I. & Z. N. Smirnova, Handh. Mosses
U.S.S.R. Acrocarp. 1970 Microbryum vlassovii (Laz.) Zand.,
see treatment of Microbryum]
=
[MELOPHYLLUM Herz. = Bryomanginia Ther. fide Zander,
Bryologist 75: 333, 1978 = Astomiopsis C. Mtill. (Ditrichaceae)
fide Buck & Zander, Bryologist 83: 255, 1980]
[Melophyllum radiculosum Herz. Am4 (= Bryomanginia saintpierrei Ther. fide Zander, Bryologist 75: 333, 1978) = Astomiopsis radiculosa (Herz.) Buck & Zander (Ditrichaceae) fide
Buck and Zander, Bryologist 83: 255, 1980]
= Scopelophila (Mitt.) Lindb. fide Zander,
Bryologist 70: 406, 1967]
[Merceya difficilis Herz. & Ther., Fedde Rep. Spec. Nov. Reg. Yeg.
45: 46, 1938 Am4 = Scopelophila ligulata (Spruce) Spruce fide
[MERCEYA Schimp.
GENERA, SPECIES AND INFRASPECIFIC TAXA
Zander, Bryologist 70: 412, I967]
[Merceya gedeana (Lac.) Nog. As2 As3 As4 = Scope/ophila
cataractae (Mitt.) Broth. fide Zander, Bryologist 70: 408, I967]
[Merceya latifolia Kindb. Ami
Crumia /atifo/ia (Kindb. ex
Macoun) Schof.]
[Merceya /igu/ata (Spruce) Schimp. Eur As2 As3 As4 AsS Afri Ami
Am2 Am4 = Scopelophila ligu/ata (Spruce) Spruce fide Zander,
Bryologist 70: 4II, I967]
var. acutiuscu/a (Broth.) Chen Eur
[Merceya /ongirostris (Griff.) Wijk & Marg. As3 = Trichostomum
contractum Zand. nom. nov., see treatment of Trichostomum]
Merceya repandu/a Baumg. & Frohl. As4
[Merceya serratinervis Tak. As2 = Encalypta streptocarpa Hedw.
fide lwats. & Nog., J. Hattori Bot. Lab. 37: 368, IC)73]
=
[MERCEYOPSIS Broth. & Dix. ex Dix. = Scope/ophila (Mitt.)
Lindh., see treatment of Scope/ophila]
[Merceyopsis angu/osa Broth. & Dix. ex Dix. As3 Am5 = Weisiopsis
angu/osa (Broth. & Dix. in Dix.) Hilp. Gangu/eea angu/osa
(Broth. & Dix.) Zand., Phytologia 65: 427, 1989]
[Merceyopsis crassinervis (Broth.) Bartr., Bishop Mus. Occ. Pap. IS :
98, I939 Oc = Molendoa crassinervis Broth. fide Nog. in Hoe,
Lyonia I : 28, I974 = Hymenosty/ium recurvirostrum var.
cylindricum (Bartr.) Zand. fide Zander & Eckel, Canad. J. Bot.
60: I599, I982]
[Merceyopsis excavata Sak. As2 = Gymnostomum aeruginosum Sm.
fide Saito, J. Hattori Bot. Lab. 39: 450, I975]
[Merceyopsis hymenostylioides Broth. & Dix. ex Dix. As3 Gymnostomum hymenostylioides (Broth. & Dix.) Zand., see treatment of
Gymnostomum]
[Merceyopsis /ongirostris (Griff.) Broth. & Dix. ex Dix. As3
Merceya /ongirostris (Griff.) Wijk & Marg.
Trichostomum
contractum Zand. nom. nov., see treatment of Trichostomum]
[Merceyopsis robusta Dix. As3 = Tuerckheimia robusta (Dix.) Zand.
comb. nov., see treatment of Tuerckheimia]
[Merceyopsis spathulifolia Dix. & P. Yarde As3
Barbu/a
spathu/ifo/ia (Dix. & P. Yarde) Zand., see treatment of Barbula]
=
=
=
=
=
MICROBRYUM Schimp.
Microbryum brevicau/e (Tayl.) Zand. (Gymnostomum), see treatment
of Microbryum Austr1
Microbryum curvico//e (Hedw.) Zand. (Phascum), see treatment of
Microbryum Eur Afri AsS
Microbryum davallianum (Sm.) Zand. (Gymnostomum), see treatment
of Microbryum Eur AsS Afrl Ami Austrl
var. commutatum (Limpr.) Zand. (Portia) , see treatment of Microbryum Eur AsS Afri
var. conicum (Schleich. ex Schwaegr.) Zand. (Gymnostomum), see
treatment of Microbryum Eur Afri AsS Am2
Microbryum floerkeanum (Web. & Mohr) Schimp. (Phascum), see
treatment of Microbryum Eur Afri Ami AsS
var. arbense (Loitl.) Zand. (Phascum), see treatment of Microbryum Eur
var. badium BSG Eur Ami AsS
Microbryum longipes (Guerra, Martinez & Ros) Zand. (Phascum),
see treatment of Microbryum Eur
Microhryum raddei (Broth.) Zand. (Tortula) As I
Microbryum rectum (With.) Zand. (Phascum), see treatment of
Microbryum Eur Afr1
Microbryum rufochaete (Magill) Zand. (Acau/on), see treatment of
Microbryum Afr4
Microbryum starckeanum (Hedw.) Zand. (Weisia), see treatment of
Microbryum Eur AsS Afri Ami Am2 Austri Austr2
313
var. brachyodus (BSG) Zand. (Anacalypta starckeana var.), see
treatment of Microhryum Eur AsS Afrl
var. hrevidens (Latz.) Zand. (Portia starckeana var.), see treatment of Microbryum Eur
var. fosbergii (Bartr.) Zand. (Portia), see treatment of Microbryum Am I Am2
var. leiostoma (Corb in Corb. & Pitard) Zand. (Portia starckeana
var.) Afri
var. subgymnostoma (De Not.) Zand. (Anacalypta starckeana
var.), see treatment of Microbryum Eur
var. submutica (Latz.) Zand. (Portia starckeana var.), see treatment of Microbryum Eur
fo. brevifolium (Limpr.) Zand. (Portia starckeana fo.), see treatment of Microbryum Eur
fo. dextrorsum (Limpr.) Zand. (Portia starckeana fo.), see treatment of Microbryum Eur
fo. microphy//um (Wamst.) Zand. (Portia starckeana fo.), see
treatment of Microbryum Eur
Microbryum subplanomarginatum (Dix.) Zand. (Pottia), see treatment of Microbryum Afr4
Microbryum tasmanicum (Dix. & Rodw.) Zand. (Phascum), see
treatment of Microbryum Austri
Microbryum v/assovii (Laz.) Zand. (Phascum), see treatment of
Microhryum As1 Ami
Microbryum zeelandiae (R. Br. ter) Zand. (Anacalypta), see treatment of Microhryum Austr2
[MILDEELA Limpr. = Pottia subg. Mildee//a (Kindb.) Broth. =
Tortula sect. Tortula, see treatment of Tortula]
[Mildee//a bryoides (Dicks.) Limpr. =Pottia bryoides (Dicks.) Mitt.
=Tortula protobryoides Zand., see treatment of Tortula]
[fo. /ongiseta Roll, Hedwigia 56: 157, 1915 =Portia bryoides fo.
/ongiseta (Roll) Podp.]
MIRONIA Zand. (nom. nov. for Morinia Card., see treatment of
Mironia)
Mironia crassicuspis (Robins.) Zand., see treatment of Mironia
(Barbu/a) Am2
Mironia ehrenbergiana (C. Miill.) Zand. (Barbula), see treatment
of Mironia nbergiana (C. Miill.) Ther. Am2 Am4
var. e/ongata (Wils. in Mitt.) Zand., see treatment of Mironia
(Barbula) Am4
Mironia stenotheca (Ther.) Zand., see treatment of Mironia (Barbula)Am2
MOLENDOA Lindh.
Mo/endoa andina (Mitt.) Broth. Am4
[Molendoa boliviano Both. in Herz. Am4 = Molendoa sendtneriana
(BSG) Limpr.fide Zander, Bryo1ogist 80: 248, I977]
[var. brevifolia Herz. Am4 = Molendoa sendtneriana (BSG)
Limpr.fide Zander, Bryologist 80: 248, I977]
[Molendoa burmensis Bartr. As3 =Hymenostylium recurvirostrum
var. cylindricum (Bartr.) Zand. in Zand. & Eckel;, see treatment
of Molendoa]
[Molendoa clavuligera Castelli, Rev. Bryol. Lichenol. 34: 716,
1966 (1967)] Eur =Mo/endoa taeniatifolia Herz. (see Corley et
al., J. Bryol. 11 : 623, 1981 (1982)]
[Molendoa crassinervis Broth. Oc = Hymenostylium recurvirostrum
var. cylindricum (Dix .) Bartr. fide Zander & Eckel, Canad. J.
Bot. 60: IS99, I982]
Molendoa cucul/ata (Herz.) Hilp. Am4
Molendoa duthiei (Broth.) Broth. As3
[Molendoa excelsa (C. Mill!.) Broth. Am6 = Mo/endoa
314
GENERA OF THE POTTIACEAE
sendtneriana (BSG) Limpr. fide Zander, Bryologist 80: 247,
1977]
Molendoafuegiana Bartr. Am6
[Molendoa guentheri Broth. Am4 = Molendoa sendtneriana (BSG)
Limpr.fide Menzel, J. Hattori Bot. Lab. 71: 219, 1992]
[Molendoa herzogii Broth. in Herz. Am4 = Molendoa sendtneriana
(BSG) Limpr.fide Zander, Bryo1ogist 80: 248, 1977]
Molendoa hornschuchiana (Hook.) Lindb. ex Limpr. Eur As1 As2
Afr1 Am1
var. minor (Mol.) Limpr. Eur
[Molendoa japonica Broth. As2 = Molendoa sendtneriana var.
japonica (Broth.) lwats. = Hymenostyliella japonica (Broth.)
Saito, J. Jap. Bot. 46: 145, 1971 = Didymodon japonicus (Broth.)
Saito fide Saito, J. Hattori Bot. Lab. 39: 508, 1975]
Molendoa kitaibelana Gytirf. in Bauer Eur
[Molendoa obtusifolia Broth. & Par. Am1 Am2 = Molendoa
sendtneriana (BSG) Limpr. fide Zander, Bryologist 80: 248,
1977]
[var. densissima Ther. Am2 = Molendoa sendtneriana (BSG)
Limpr.fide Zander, Bryo1ogist 80: 248, 1977]
[var. incrassata Ther. Am2 = Molendoa sendtneriana (BSG)
Limpr.fide Zander, Bryologist 80: 248, 1977]
Molendoa ogala/ensis (Smith Merrill) Zand. (Ozobryum), see treatment of Molendoa Am1
Molendoa platyphyllum (Williams) Zand. (Anoectangium), see treatment of Molendoa Am4]
Molendoa roylei (Mitt.) Broth. As2
Molendoa schliephackei (Limpr.) Zand. (Pleuroweisia), see treatment
of Molendoa) Eur As2
Molendoa sendtneriana (BSG) Limpr. Eur As1 As2 As3 Ami Am2
Am4Am5Am6
[var. japonica (Broth.) lwats. As2 = Hymenostyliella japonica
(Broth.) Saito, J. Jap. Bot. 46: 145, 1971 =Didymodonjaponicus
(Broth.) Saito fide Saito, J. Hattori Bot. Lab. 39: 508, 1975]
var. limprichtii Gytirf. Eur
var. sudetica Podp. Eur
[var. tenuinervis (Limpr.) Pilous Eur Am1 = Molendoa
sendtneriana (BSG) Limpr. fide Zander, Bryologist 80: 247,
1977]
var. transcaspica Gytirf. in Fedch. As1
var. yunnanica Gytirf in Ther.
fo. andreaeoides (Limpr.) Pilous, Preslia 30: 168, 1958 (Grimmia)
Eur
fo. propagu/ifera Podp., Cas. Morav. Mus. Zemsk. 13: 42, 1913
Eur
Molendoa seravschanica Broth. & Gyorf. in Fedch. As2
[Molendoa sordida (Mitt.) Steere Am4 = Hymenostylium
recurvirostrum (Hedw.) Dix., see treatment of Molendoa]
Molendoa sublaevis Demar. & P. Yarde Afr2
Molendoa subobtusifolia Broth., Rev. Bryol. 47: 7, 1921 (not in Index
Muscorum) Am4
Molendoa taeniatifolia Herz. Eur
[Molendoa tenuinervis Limpr. Eur = Molendoa sendtneriana var.
tenuinervis (Limpr.) Pilaus = Molendoa sendtneriana (BSG)
Limpr.fide Zander, Bryo1ogist 80: 247, 1977]
[fo. arctica Gytirffy, Bryologist 15: 1912 Am1 = Molendoa
sendtneriana var. tenuinervis Limpr.fide Grout, Moss Fl. N. Am.
1(3): 150, 1933 = Molendoa sendtneriana (BSG) Limpr. fide
Zander, Bryologist 80: 247, 1977]
[Molendoa warburgii (Crundw. & Hill) Zand. (Anoectangium), see
treatment of Molendoa]
[MORINIA Card.= Mironia]
[Morinia crassicuspis (Robins.) Zand., Bryologist 81: 556, 1978
(Barbula) Am2 = Mironia crassicuspis (Robins.) Zand., see
treatment of Mironia]
[Morinia ecuadorensis Bartr. Am4 = Morinia ehrenbergiana var.
elongata (Wils.) in Mitt.) Zand. fide Zander, Bryologist 81:
557, 1978 = Mironia ehrenbergiana var. elongata (Wils. in
Mitt.) Zand., see treatment of Mironia]
[Morinia ehrenbergiana (C. Miill.) Ther. Am2 Am4 = Mironia
ehrenbergiana (C. Miill.) Zand., see treatment of Mironia]
[var. elongata (Wils. in Mitt.) Zand., Bryologist 81: 557, 1973
(Barbula) Am4 = Mironia ehrenbergiana var. elongata (Wils.
in Mitt.) Zand., see treatment of Mironia]
[Morinia saitoana Zand, Delg. & Eckel, Bryologist 83: 510, 1980
(1981) Am2 = Morinia stenotheca (Ther.) Zand., Bryologist 86:
156, 1983 = Mironia stenotheca (Ther.) Zand., see treatment of
Mironia]
[Morinia setschwanica (Broth.) Hilp. As2 = Didymodon erosodenticulatus (C. Miill.) Saito fide Saito, J. Hattori Bot. Lab. 39:
504, 1975]
[Morinia stenotheca (Ther.) Zand., Bryologist 86: 156, 1983 (Barbuta) Am2 = Mironia stenotheca (Ther.) Zand., see treatment of
= Mironia stenotheca (Ther.) Zand., see treatment of Mironia]
[NEOCARDOTIA Ther. & Bartr. = Rhexophyllum Herz. fide Hilpert, Bot. Centralb. Beih. 50(2): 684, 1933]
[Neocardotia subnigra (Mitt.) Ther. & Bartr. Am1 Am2 Am4 =
Rhexophyllum subnigrum (Mitt.) Ther. ex Hilp. fide Hilpert,
Bot. Centralb. Beih. 50(2): 684, 1933]
NEOHYOPHILA Crum
[Neohyophila Ungulata (Card.) Crum Am2 = Neohyophila
sprengelii var. stomatodonta (Card.) Zand., Bryologist 86: 138,
1983 = Plaubelia sprengelii var. stomatodonta (Card.) Zand.,
see treatment of Plaubeliaj
[Neohyophila spathulifolia (Bartr.) Crum Am2 = Neohyophila
sprengelii (Schwaegr.) Crum s. lat. fide Zander, Bryo1ogist 86:
135, 1983 = Plaubelia sprengelii (Schwaegr.) Zand., see treatment of Plaubelia]
[Neohyophila sprengelii (Schwaegr.) Crum Aml Am2 Am3 Am4
Am5 = Plaubelia sprengelii (Schwaegr.) Zand., see treatment
of Plaubelia]
[var. stomatodonta (Card.) Zand., Bryologist 86: 138, 1983 (Hyophila) Am1 Am2 Am3 = Plaubelia sprengelii var.
stomatodonta (Card.) Zand., see treatment of Plaubelia]
[Neohyophila stomatodonta (Card.) Crum Ami Am2 Am3 = Neohyophila sprengelii var. stomatodonta (Card.) Zand., Bryologist
86: 138, 1983]
[OXYSTEGUS (Limpr.) Hilp. =Trichostomum Bruch]
[Oxystegus burmensis (Bartr.) Gangulee, M. E. India 653, 1972
"burmense" (Trichostomum) As3 = Trichostomum tenuirostre
(Hook. & Tayl.) Lindb.fide Sollman, Lindbergia 10: 55, 1984]
Oxystegus crassicostatus Norris & T. Kop., Acta Bot. Fenn. 137:
90, 1989 As4
[Oxystegus cuspidatus (Doz. & Molk.) Chen As2 As4 = Trichostomum cuspidatum (Doz. & Molk.) Doz. & Molk. = Trichostomum tenuirostre (Hook. & Tayl.) Lindb. fide Sollman,
Lindbergia 10: 55, 1984]
[Oxystegus cylindricus (Brid.) Hilp. Eur As1 As3 Afr1 Afr2 Afr3
Afr4 Am1 Am2 Am4 = Oxystegus tenuirostris (Hook. & Tayl.)
GENERA, SPECIES AND INFRASPECIFIC TAXA
A. Sm. fide Smith, J. Bryol. 9: 393, 1977 = Trichostomum
tenuirostre (Hook. & Tayl.) Lindh.]
[subsp. hibernie us (Mitt.) Podp. Eur = Oxystegus hibernicus (Mitt.)
Hilp.fide Corley et al., J. Bryol. 11: 623, 1981 (1982) = Trichostomum hibernicum (Mitt.) Dix.]
var. daldinianus (De Not.) Podp Eur
[var. holtii (Braithw.) Podp. Eur = Oxystegus tenuirostris var. holtii
(Braithw.) A. Sm. fide Smith, J. Bryol. 9: 393, 1977 = Trichostomum tenuirostre var. holtii (Braithw.) Dix.]
var. irriguus (Limpr.) Podp. Eur
var. norvegicus (Hag.) Podp. Eur
var. robustus (Schimp.) Podp. Eur
[fo. asper (Podp.) Podp., Consp. Muse. Eur. 176, 1954 (Trichostomum cylindricum var.) Eur = Trichostomum cylindricum var.
asperum Podp.]
fo. filifer (Herz.) Podp., Consp. Muse. Eur. 176, 1954 (Oxystegus
cylindric us var.) Eur
fo. longifolius Podp., Consp. Muse. Eur. 176, 1954 (Trichostomum
cylindricum fo. longifolius Latz. hom. illeg.) Eur
[Oxystegus cylindrothecus (Mitt.) Gangulee, Nov. Hedw. 12: 430,
1966 (Tortula) As3 = Trichostomum cylindrothecum (Mitt.)
Broth. = Trichostomum bombayense C. Miill. fide Townsend, J.
Bryol. 12: 561, 1983]
[Oxystegusfallax (Herz.) Hilp. Am4 = Trichostomumfallax Herz.]
[Oxystegus hibernicus (Mitt.) Hilp. Eur = Trichostomum hibernicum
(Mitt.) Dix.]
[Oxystegus indicus (Dix. & P. Yarde) Hilp. As3 = Pseudosymblepharis indica Dix. & P. Yarde]
[Oxystegus khasianus (Mitt.) Gangulee, Nov. Hedw. 8: 149, 1964
(Tortula) As3 = Psuedosymblepharis khasiana (Mitt.) Zand., see
treatment of Pseudosymblepharis]
[Oxysregus lignicola (Herz.) Hilp. Am5 = Trichostomum lignicola
Herz.]
[Oxystegus linearis (Web. & Mohr) Hilp. Am3 Am5 = Tortella
linearis (Web. & Mohr.) Zand., see treatment of Tortella]
Oxystegus melanostoma (Mitt.) Hilp. Am5
[Oxystegus recurvifolius (Tayl.) Zand., Lindbergia 8: 187, 1982
(Bryum) Eur Am1 = Trichostomum recurvifolium (Tayl.) Zand.,
see treatment of Trichostomum]
[Oxystegus rhodesiae (Broth.) Hilp. Afr2 = Trichostomum rhodesiae
Broth.]
[Oxystegus soulae (C. Miill. in Ren. & Card.) Wijk & Marg. Afr3 =
Trichostomum soulae (C. Miill. in Ren. & Card.) Zand., see treatment of Trichostomum]
[var. corticicola (Ren. & Card.) Wijk & Marg. Afr3 = Trichostomum soulae var. corticicola (Ren. & Card.) Zand., see treatment
of Trichostomum]
[Oxystegus spiralis (Grout) Crum & Anderson, Bryologist 92: 533,
1989 = Trichostomum spirale Grout, see treatment of Trichostomum]
[Oxystegus stenophyllus (Mitt.) Gangulee, Nov. Hedw. 12: 429, 1966
As3 = Oxystegus cylindricus (Brid.) Hilp. fide Saito, J. Hattori
Bot. Lab. 39: 437, 1975 =Oxystegus tenuirostris (Hook. & Tayl.)
A. Sm. fide Smith, J. Bryol. 9: 393, 1977 = Trichostomum
tenuirostre (Hook. & Tayl.) Lindh.]
[Oxystegus svihlae (Bartr.) Gangulee, M. E. India 654, 1972 (Triclwstomum) As2 = Tuerckheimia svihlae (Saito) Zand., see treatment
of Tuerckheimia]
[Oxystegus syrrhopodontoides (Herz.) Hilp. Am4 = Trichostomum
syrrhopodontoides Herz. = Trichostomum tenuirostre (Hook. &
Tayl.) Lindb.fide W. Steere, in !itt., having seen type at JE]
[Oxystegus verrucosus (Broth. & Par.) Hilp. Oc = Pseudosymblepharis verrucosa (Broth. & Par.) Zand., see treatment of Pseudo-
315
symblepharis]
[Oxystegus tenuirostris (Hook. & Tayl.) A. Sm., J. Bryol. 9: 393,
1977 Eur As1 As3 Afr1 Afr2 Afr3 Afr4 Am1 Am2 Am4 =
Trichostomum tenuirostre (Hook. & Tayl.) Lindh.]
[var. gemmiparus (Schimp.) Zand., Lindbergia 4: 285, 1978
(Didymodon cylindricus var.) Eur As3 Am2 Am3 Am5 =
Trichostomum tenuirostre var. gemmiparum (Schimp.) Zand.,
see treatment of Trichostomum]
[var. holtii (Braithw.) A. Sm., J. Bryol. 9: 393, 1977 Eur Am1 =
Trichostomum tenuirostre var. holtii (Braithw.) Dix.]
[var. stenocarpus (Ther.) Zand., Misc. Bryol. Lichenol. 9: 73,
1982 (Weisiopsis) As2 Am1 Am2 = Trichostomum spirale
Grout, see treatment of Trichostomum]
[OZOBRYUM Smith Merrill, Novon 2: 255, 1992 = Molendoa
Lindh., see treatment of Molendoa]
[Ozobryum ogalalense Smith Merrill, Novon 2: 255, 1992 Ami =
Molendoa ogalalensis (Smith Merrill) Zand., see treatment of
Molendoa]
PACHYNEUROPSIS H. Mill.
Pachyneuropsis bartlettii (Bartr.) H. Mill., Taxon 19: 822, 1970
(Pachyneurum Bartr. hom. illeg.) As4
[PARALEPTODONTIUM Hill., J. Bryology 12: 181, 1982 =
Oxystegus (Limpr.) Hilp. fide Zander, Lindbergia 8: 187, 1982
= Trichostomum Hedw., see treatment ofTrichostomumJ
[Paraleptodontium recurvifolium (Tayl.) Long, J. Bryology 12:
181, 1982 = Oxystegus recurvifolius (Tayl.) Zand., Lindbergia
8: 187, 1982 = Trichostomum recurvifolium (Tayl.) Zand., see
treatment of Trichostomum]
[PHASCONICA C. Miill. = Weissia subg. Phasconica (C. Miill.)
Zand., see treatment of WeissiaJ
[Phasconica balansae C. Miill. Austr1 Oc = Weissia balansae (C.
Miill.) Zand., see treatment of Weissia]
[Phasconica lorentzii C. Miill. Am6 = Weissia lorentzii (C. Miill.)
Zand., see treatment of Weissia]
[Phasconica tisserantii P. Yarde Afr2 Afr4 (good species of
Phasconica fide Magill, Fl. S. Afr. 1. Mosses 1: 251, 1981
[1982]) = Weissia unguiculara (Mitt.) Crundw. & Nyh., J.
Bryol. 8: 69, 1974 = Trichosromum unguiculatum (Mitt.) Zand.,
see treatment of Trichostomum]
PHASCOPSIS Stone, J. Bryol. 11: 17, 1980
Phascopsis rubicunda Stone, J. Bryol. 11 : 17, 1980 Austr1
[PHASCUM Hedw. =Tortula Hedw., see treatment of Tortula]
Phascum beaugleholei Stone, J. Bryol. 15: 766, 1989 Austr1
[Phascum brittoniae Crum & Steere Am3 = Uleobryum
peruvianum Broth., see treatment of Uleobryum]
Phascum calodictyon C. Miill. Am6
[Phascum carinatum Hampe Am5 = Bruchia carinata (Hampe)
Zand. (Ditrichaceae) see Excluded Taxa]
[Phascum crispum Hedw. = Astomum crispum (Hedw.) Hampe =
Weissia longlfolia Mitt. fide Crundwell & Nyholm, J. Bryol. 7:
13, 1972]
var. minus De Not. Eur
[Phascum curvicolle Hedw. Eur Afrl AsS= Microbryum curvicolle
(Hedw.) Zand., see treatment of Microbryum]
[Phascum cuspidatum Hedw. Eur As1 As2 As5 Afr1 Am1 Am2
Am4 = Tortula atherodes Zand. nom. nov., see treatment of
Tortula]
GENERA OF THE POTIIACEAE
316
=
[subsp. e/atum (Brid.) C. Jens. Eur Tortula atherodes var. elata
(Brid.) Zand., see treatment of Tortula]
[subsp. mitraeforme (Limpr.) C. Jens. Eur Tortula atherodes var.
mitraeformis (Limpr.) Zand., see treatment of Tortula]
[subsp. papillosum (Lindh.) Guerra & Ross in Guerra, Jimenez, Ros
& Carri6n, Cryptogamic Bryol. Lichenol. 12: 391, 1991
Tortula atherodes var. papillosa (Lindh.)
(Phascum p.) Eur
Zand., see treatment of Tortula]
[subsp. piliferum (Hedw.) Amann Eur As1 As5 Afr2 Am1 =Tortula atherodes var. pi/ifera (Hedw.) Zand., see treatment of Tortula]
[var. affine (Nees & Homsch.) Hampe Eur Tortula atherodes var.
affinis (Nees & Homsch.) Zand., see treatment of Tortula]
[var. arcuatum Herrnstadt & Heyn, Bryologist 94: 175, 1991 As5
Tortula atherodes var. arcuata (Herrnstadt & Heyn) Zand., see
treatment of Tortula)
[var. curvisetum (Dicks.) Nees & Homsch. Eur = Tortula atherodes
var. curviseta (Dicks.) Zand., see treatment of Tortula]
Tortula atherodes var.
[var. diaphorum (Hag.) C. Jens. Eur
diaphora (Hag.) C. Jens., see treatment of Tortula]
[var. elatum (Brid.) Drumm. Eur
Tortula atherodes var. e/ata
(Brid.) Zand., see treatment of Tortula]
[var.flagel/aceum Ruthe in Warnst. Eur Phascum cuspidatum fo.
flagel/a ceum (Ruthe in Warnst.) Podp., Consp. Muse. Eur. 222,
1954]
[var. grandiuscu/um (Brid.) Roll, Hedwigia 56(1-3): 156, 1915
(Phascum grandiusculum) Eur = Phascum cuspidatum var.
Tortula
elatum (Brid.) Drumm. fide Web. & Mohr 1807
atherodes var. e/ata (Brid.) Zand., see treatment of Tortula]
[var. henrici (Ren. & Card.) Wijk & Marg. Am1 = Phascum
cuspidatum fide Crum & Anders., Mosses E. N. Amer. 1: 350,
1981 Tortula atherodes Zand., see treatment of Tortula]
[var. intertextum (Brid.) Brid. Eur
Tortula atherodes var.
intertexta (Brid.) Zand., see treatment of Tortula]
[var. marginatum Hemnstadt & Heyn, Bryologist 94: 175, 1991
As5 = Tortula atherodes var. marginata (Hemnstadt & Heyn)
Zand., see treatment of Tortula]
[var. mitraeforme Limpr. Eur =Tortula atherodes var. mitraeformis
(Limpr.) Zand., see treatment of Tortula]
[var. papi/losum (Lindh.) C. Hartm. in Hartm., Handb. Skand. Fl.
ed. 10, 2: 119, 1871 (Phascum papillosum) Eur = Tortula
atherodes var. papillosa (Lindh.) Zand., see treatment of Tortulal
[var. piliferum (Hedw.) Hook. & Tayl. Eur As1 As5 Afr2 Aml
Tortula atherodes var. pilifera (Hedw.) Zand., see treatment of
Tortula]
[var. retortifolium Guerra & Ros in Guerra, Jimenez, Ros &
Carri6n, Cryptogamic Bryol. Lichenol. 12: 390, 1991 Eur =Tortula atherodes var. retortifolia (Guerra & Ros in Guerra, Jimenez,
Ros & Carri6n) Zand., see treatment of Tortula]
[var. schreberianum (Dicks.) Brid. Eur Am1 = Tortula atherodes
var. schreberiana (Dicks.) Zand., see treatment of Tortula]
[var. varium Steud. Eur = Phascum cuspidatum var. schreberianum
Tortula
(Dicks.) Brid. fide Brid., Mant. Muse. 8, 1819
atherodes var. schreberiana (Dicks.) Zand., see treatment of
Tortulal
fo. affine (Nees & Homsch.) Casares Gil, Fl. Iber. Musg. 256, 1932
(Phascum affine) Eur
fo. flagellaceum (Ruthe in Warnst.) Podp., Consp. Muse. Eur. 222,
I954 (Phasum cuspidatum var.) Eur
Phascum cuynettii Biz. & Pier. ex Guerra, Jimenez, Ros & Carri6n,
Cryptogamic Bryol. Lichenol. 12: 401, 1991 (Phascum cuynettii
Biz. & Pier., Rev. Bryol. Lichenol. 36: 506, 1969 nom . inval., single element not cited as holotypefide Crosby eta!. 1992) Eur
=
=
=
=
=
=
=
=
=
=
=
=
[Phascumfloerkeanum Web. & Mohr Eur Afri Ami As5 =Microbryum floerkeanum (Web. & Mohr) Schimp., see treatment of
Microbryum]
[var. arbense (Loitl.) Podp. Eur Microbryum floerkeanum var.
arbense (Loitl.) Zand., see treatment of Microbryum]
[var. badium (BSG) Schimp. Eur Aml As5 = Microbryum
floerkeanum var. badium BSG]
[Phascum fosbergii (Bartr.) Guerra in Guerra, Martinez & Ros, J.
Bryol. I6: 59, I990 Ami Am2 = Microbryum starckeanum var.
fosbergii (Bartr.) Zand.]
Phascum galilaeum Hemnstadt & Heyn, Bryologist 94: 175, 1991
As5
[Phascum halophi/um Smarda Eur = Phascum cuspidatum Hedw.
fide Corley et a!., J. Bryol. 11 : 62I, 1981 [1982] = Tortula
atherodes Zand., see treatment of Tortula]
[Phascum hyalinotrichum Card. & Ther. Am1 Am2 = Stegonia
hyalinotrichum (Card. & Ther.) Zand., see treatment of Stegonia]
[Phascum kilimandscharicum (C. Mtill.) Roth Afr2 = Bryum
kilimandscharicum (C. Mtill.) Zand. (Bryaceae) see Excluded
Taxa]
Phascum laticostum Stone, J. Bryol. 15: 753, 1989 Austrl
[Phascum /eptophyllum C. MUll. Afr2 Afr4 = Chenia leptophyl/a
(C. Mtill.) Zand., see treatment of Chenia]
[Phascum longipes Guerra, Martinez & Ros, J. Bryol. 16: 55, I990
Eur ["longipedis" typographical error cf. A. Smith, J. Bryol.
16: 335, 1991] = Microbryum longipes (Guerra, Martinez &
Ros) Zand., see treatment of Microbryum]
Phascum longipi/um Stone, J. Bryol. 15: 755, 1989 Austr1
Phascum nepalense Brid. As3
[Phascum nitidum Hedw. = Pseudephemerum nitidum (Hedw.)
Reim.]
var. bulbiferum Husn. Eur
[var. strictum (Dicks.) Wils., Bryol. Brit. 35, 1855 nom. illeg.
incl. Phascum nitidum var. minus Bayrh. = Pseudephemerum
nitidum var. minimum (Rabenh.) Wijk & Marg. cf. Wilson
1855]
Phascum peraristatum C. MUll. Afr4
[Phascum pi/iferum Schreb. ex Hedw. = Phascum cuspidatum var.
piliferum (Hedw.) Hook. & Tayl. = Tortula atherodes var.
pilifera (Hedw.) Zand., see treatment of Tortula]
var. subacaule Schwaegr. ex Steud. Eur
[var. imberbe Hartm., Handb. Skand. Fl. ed. 3: 261, 1838 =Phascumfloerkeanum Web. & Mohr. 1807]
Phascum piptocarpum Dur. & Mont. in Mont. Afr1
Phascum readeranum Stone, J. Bryol. 15: 757, 1989 Austr1
[Phascum rectum With. =Portia recta (With.) Mitt. = Microbryum
rectum (With.) Zand. , see treatment of Microbryum]
var. luxurians De Not. Eur
[Phascum robustum (Broth.) ex Roth) Stone, J. Bryol. 15: 747,
1989 Austr I Acaulon robustum Broth. ex Roth]
var. crassinervium (C. MUll.) Stone, J. Bryol. 15: 751, 1989
Austr1
[Phascum schimperianum Sull. in A. Gray, Man. Bot. E. N. Am.
ed. 2: 615, 1856 = Acaulon schimperianum (Sull. in A. Gray)
Sull.]
Microbryum
[Phascum tasmanicum Dix. & Rodw. Austri
tasmanicum (Dix. & Rodw.) Zand., see treatment of Microbryum]
var. unguiculatum Stone, J. Bryol. 15: 761 , 1989 Austri
[Phascum vlassovii Laz. As1 Am1 = Microbryum vlassovii (Laz.)
Zand., see treatment of Microbryum]
[Phascum vogesiacum Moug. & Nest!., Stirp. Crypt. Vog.-Rhen.
=
=
=
317
GENERA, SPECIES AND INFRASPECIFIC TAXA
fasc. 7: n. 706, 1823 inval. Eur =Bruchia vogsesiaca Schwaegr.]
PLAUBELIA Brid.
Plaubelia involuta (Magill) Zand. (Weisiopsis), see treatment of
Plaubelia Afr4
Plaubelia perinvoluta Zand. (nom . nov. for Desmatodon involutus
Bartr. hom. illeg.), see treatment of Plaubelia As3
Plaubelia sprengelii (Schwaegr.) Zand. (Barbula), see treatment of
Plaubelia Ami Am2 Am3 Am4 Am5
var. stomatodonta (Card.) Zand. (Hyophila), see treatment of
Plaubelia Am 1 Am2 Am3
PLEUROCHAETE Lindh.
Pleurochaete beccarii Vent. Afr2
Pleurochaete ecuadoriensis Broth. Am4 Am6
[Pleurochaete luteola (Besch.) Ther. Ami Am2 Am3 Am4 Pleurochaete squarrosa var. luteola (Besch.) Zand., see treatment of
Pleurochaete]
Pleurochaete malacophylla (C. Mtill.) Broth. As5
Pleurochaete squarrosa (Brid.) Lindh. Eur Asl As2 As3 AsS Afrl
Afr2 Am 1 Am2 Am3 Am4
[var. crispifolia Nog. As2 = Pseudosymblepharis angustata (Mitt.)
Chenfide Saito, J. Hattori Bot. Lab. 39: 439, 1975]
var. densifolia Amann Eur
var. luteo/a (Besch.) Zand., see treatment of Pleurochaete (Pleurochaete luteola) Ami Am2 Am3 Am4
var. nitida (Fam.) Podp. Eur
fo. brevifolia (Amann) Podp., Consp. Muse. Eur. 183, 19S4
(Pleurochaete squarrosa var.) Eur
fo. subintegra Latz., Bot. Centralb. Beih. 48:477, 1931 Eur
=
= Molendoa Lindh. (see treatment of
Molendoa)]
[Pleuroweisia schliephackei Limpr. Eur = Molendoa schliephackei
(Limpr.) Zand., see treatment of Molendoa]
[PLEUROWEISIA Limpr.
= Tortula Hedw., see treatment of Tortula]
[Portia acutidentata Card. & Ther. Afr4 Hennediella acutidentata
(Card. & Ther.) Zand., see treatment of Hennediella]
[Portia affinis (Hook. & Tayl.) Herrnstadt & Heyn, Bryologist 94:
17S, 1991 (Weissia) AsS =Portia mutica Vent. in De Not. Eur
AsS Afr1 =Portia starckeana var. brachyoda (BSG) C. Miill . fide
Moenkemeyer, Laub. Eur. 327, 1927 Microbryum starckeanum
var. brachyodus (BSG) Zand., see treatment of Microbryum]
Portia alpicola Dix. As3
Portia altipes Broth. AmS
Portia antarctica (Angstr.) C. Mtill. Am6
Portia appertii Wamst. Afrl
Portia areo/ata (Knight) R. Br. ter Austr2
[Portia arizonica Wareh. in Grout Ami = Microbryum starckeanum
(Hedw.) Zand. var. starckeanum, see treatment of Microbryum]
[var. mucronulata Wareh. in Gn;ut Am1 = Microbryum
starckeanum var. brachyodus (BSG) Zand., see treatment of
Microbryum]
[Portia aurantiaca (Mitt.) C. Mtill. in Chen, Hedwigia 80: 62, 1941
inval. = Hymenostylium aurantiacum Mitt. = Hymenostylium
recurvirostrum var. luzonense (Broth.) Bartr. fide Gangulee,
Mosses E. lndia 3: 648, 1972 = Hymenostylium recurvirostrum
var. cylindricum (Bartr.) Zand., see treatment of Hymenostylium]
[Portia austrogeorgica Card. (good species fide Matteri 1977) Am6
Hennediella austrogeorgica (Card.) Blockeel, J. Bryol. 16: 191,
1991]
[POTTIA (Reichenb.) Ehrh. ex Ftirnr.
=
=
=
[Portia brevicaulis (Tayl.) C. Mtill. (see Catcheside, Mosses S.
Austr. 128, 1980) Austr1 = Microbryum brevicaule (Tayl.)
Zand., see treatment of Microbryum]
[Portia bryoides (Dicks.) Mitt. Eur AsS Am1 Am2 = Tortula
protobryoides Zand. (nom . nov. for Phascum bryoides Dicks.),
see treatment ofTortula]
[var. brevifolia (De Not.) Wijk & Marg. Eur
Tortula
protobryoides var. brevifolia (De Not.) Zand., see treatment of
Tortula]
[var. thornhillii (Wils.) Braithw. Eur = Tortula protobryoides var.
rhornhillii (Wils.) Zand., see treatment of Tortula]
fo. atroviridis (Schirnp.) Podp., Consp. Muse. Eur. 224, 19S4
(Phascum bryoides var.) Eur
fo. brachycarpa Monk., Laubm. Eur. 326, 1927 (Phascum
bryoides var.) Eur
fo. curviseta (Monk.) BSG, Laub. Eur. 326, 1927 (Phascum
bryoides var.) Eur
fo. longiseta (Roll) Podp., Consp. Muse. Eur. 224, 19S4
(Mildeella bryoides fo.) Eur
fo. minor (Nees & Homsch.) Podp., Consp. Muse. Eur. 224, 19S4
(Phascum bryoides var.) Eur
fo. pilifera (Schultz) Monk., Bryol. Eur. 326, 1927 (Phascum
bryoides var.) Eur
[Portia caespitosa (Bruch ex Brid.) C. Mtill. cf. Corley et al., J.
Bryol. 11 : 620, 1981 [1982] Eur = Trichostomum caespitosum
(Bruch ex Brid.) Jur. , see treatment of Trichostomum]
Portia caucasica (Lindh.) Par. Eur
[Portia charcotii Card. Ant = Portia heimii var. charcotii (Card.)
Sav. & Smim., Novosti Sist. Niz. Rast. 196S: 2S8, 196S =
Portia heimei (Hedw.) Ftimr. in Hampe var. heimei fide
Matteri, Brit. Antarct. Surv. Bull. 46: 141 , 1977]
Portia chucutensis Card. & Broth. Am6
[Portia ciliatiseta Sak. = Rhabdoweisia crispata (With.) Lindb. fide
Saito, J. Hattori Bot. Lab. 39: S28, 197S]
[Portia conica (Schleich. ex Schwaegr.) Nyh., 111. Fl. Nord. Mosses
2: 82, 1989 Eur As5 Afrl = Microbryum davallianum var.
conicum (Schleich. ex Schwaegr.) Zand., see treatment of
Microbryum]
Portia cucullata (Hampe) Jaeg. Am4
[Portia cuneifolia Solms ex Schimp. Eur = Tortula zoddae Zand.
comb. nov., see treatment of Tortula]
[Portia davalliana (Sm.) C. Jens. Eur AsS Afr1 Am1 Austrl (=
Portia starckeana subsp. minutulum (Schleich. ex Schwaegr.)
Chamber!. fide Chamberlain, Roy. Bot. Gard. Edinburg Not.
29: 403, 1969) Microbryum davallianum (Sm.) Zand., see
treatment of Microbryum]
[subsp. commutata (Limpr.) Podp. Eur AsS Afrl = Microbryum
davallianum var. commutatum (Limpr.) Zand., see treatment of
Microbryum]
var. angustifolia (Par.) Podp. Eur
[var. charcotii (Card.) Sav. & Smim., Novosti Sist. Niz. Rast.
196S: 2S8, 196S =Portia heimei (Hedw.) Ftimr. in Hampe var.
heimeifide Matteri, Brit. Antarct. Surv. Bull. 46: 141, 1977]
[var. conica (Schleich. ex Schwaegr.) Podp. Eur As5 Afr1 =
Portia starckeana subsp. conica (Schleich. ex Schwaegr.)
Chamber!. fide Chamberlain, Roy. Bot. Gard. Edinburg Not.
29: 403, 1969 = Microbryum da vallianum var. conicum
(Schleich. ex Schwaegr.) Zand., see treatment of Mit;robryum]
var. microphylla (Wamst.) Podp. Eur
var. oblonga (B.&S. in BSG) Podp. Eur
[fo. conica (Schleich. ex Schwaegr.) Pilous, Preslia 10: 130, 1969
Eur As5 Afrl nom. inval. basion. non cit.
Microbryum
daval/ianum var. conicum (Schleich. ex Schwaegr.) Zand., see
=
=
=
318
GENERA OF THE POTTIACEAE
treatment of Microbryum]
[Portia davallii Wareham in Grout, Moss Fl. N. Am. 1(4): 201, 1929
nom. illeg. incl. typon . Gymnostomum davallianum Sm. Portia
davalliana (Sm.) C. Jens. (= Portia starckeana subsp. minutulum
(Schleich. ex Schwaegr.) Chamber!. fide Chamberlain, Roy. Bot.
Gard. Edinburg Not. 29: 403, 1969) = Microbryum davallianum
(Sm.) Zand., see treatment of Microbryum)
[Pottia denticulata Dix. & P. Yarde hom. il/eg. As3 = Chenia
/eptophylla (C. MUll.) Zand., see treatment of Chenia and vide
Arts & Sollman, Lindbergia 17:22, 1991]
[Pottia drummondii (Wils.) Willis Austr1 = Tortula willisiana Zand.
nom. nov., see treatment of Tortula]
[var. obscura Willis Austr1 = Tortula willisiana var. obscura (Willis) Zand., see treatment of Tortula]
Portiaflavipes Mont. Am6
[Pottia fosbergii Bartr. Am1 Am2 = Microbryum starckeanum var.
fosbergii (Bartr.) Zand., see treatment of Microbryum]
Pottia globosa Catcheside, Mosses S. Austr. 124, 1980 Austr1
Pottia groenlandica (Kindb.) Par. Am1
[Pottia heimii (Hedw.) Ftirnr. in Hampe Eur As1 As2 Am1 Am6
Austr1 Austr2 Ant= Hennediel/a heimii (Hedw.) Zand., see treatment of Hennediella]
[var. alpina Amann Eur = Hennediella heimii var. a/pina (Amann)
Zand., see treatment of Hennediella]
[var. arctica Lindb. Eur As1 Am1 = Hennediella heimii var. arctica
(Lindb.) Zand., see treatment of Hennediella]
[var. brachyphylla Warnst. Eur = Hennedie/la heimii var.
brachyphylla (Warnst.) Zand., see treatment of Hennediella]
[var. brevicuspis Warnst. Eur = Hennediella heimii var. brevicuspis
(Warnst.) Zand., see treatment of Hennediella]
var. brevinervis Sav.-Ljubits., Bot. Mater. Otd. Sporov. Rast. Bot.
Inst. Korarova Akad. Nauk SSSR 16: 193, 1963 Ant
[var. breviseta Warnst. Aml = Hennediella heimii var. breviseta
(Warnst.) Zand., see treatment of Hennediella]
[var. drummondii Warnst. Am1 = Hennediel/a heimii var.
drummondii (Warnst.) Zand., see treatment of Hennediella]
[var. eurystoma Card. & Broth. Am6 = Hennediella heimii var.
eurystoma (Card. & Broth.) Zand., see treatment of Hennediella]
[var. guessfeldtii (Schlieph.) Warnst. Eur = Hennediella heimii var.
guessfeldtii (Schlieph.) Zand., see treatment of Hennediella]
[var. krausei (Warnst.) Warnst. Eur = Pottia heimii (Hedw.) Hampe
fide Podp~ra, Consp. Muse. Eur. 1954 = Hennediella heimii
(Hedw.) Zand.]
[var. lanceolata Warnst. Eur Am1 = Hennediella heimii var.
lanceo/ata (Warnst.) Zand., see treatment of Hennediel/a]
[var. magellanica Warnst. Am6 = Hennediella heimii var.
magellanica (Warnst.) Zand., see treatment of Hennediella]
[var. maxima Card. Am6 = Hennediella heimii var. maxima (Card.)
Zand., see treatment of Hennediel/a]
[var. spegazzinii (C. MUll.) Warnst. Am6 = Hennediel/a heimii var.
spegazzinii (C. MUll.) Zand., see treatment of Hennediella]
[var. thaxteri Card. & Ther. Am6 = Hennediella heimii var. thaxteri
(Card. & Ther.) Zand., see treatment of Hennediella]
fo. cylindrica (BSG) Podp., Consp. Muse. Eur. 232, 1954 (Pottia
heimii var.) Eur
fo. longiseta (Arnell) Podp., Consp. Muse. Eur. 232, 1954 (Portia
heimii var.) Eur
fo. ryanii (Philib.) Warnst., Hedwigia 58: 101, 1916 Eur = Pottia
heimii var. arctica Lindb. cf. Par., Ind. Bryol. ed. 2, 4: 90, 1905]
fo. tschuctschica Warnst., Hedwigia 58: 102, 1916 (Pottia
tschuctschica C. MUll. syn. inval.) As1
Pottia humillima (Angstr.) Par. Am5
[Pottia inconspicua (Griff.) C. MUll. ex Chen, Hedwigia 80: 63, 1941
=
syn. inval. As2 As3 As4 = Reimersia inconspicua (Griff.)
Chen.]
[Portia intermedia (Turn.) FUrnr. Eur As2 Afr1 Am1 = Tortula
modica Zand. nom. nov., see treatment of Tortula]
[var. corsa Fleisch. & Warnst. Eur = Tortula modica var. corsa
(Fleisch. & Warnst.) Zand., see treatment of Tortula]
[var. gymnandra Schiffn. Eur = Tortula modica var. gymnandra
(Schiffn.) Zand., see treatment of Tortula]
[var. gymnogyna Schiffn. Eur = Tortula modica var. gymnogyna
(Schiffn.) Zand., see treatment of Tortula]
[var. revoluta Schiffn. Eur = Tortula modica var. revoluta
(Schiffn.) Zand., see treatment of Tortula]
[var. stenocarpa Velen. Eur = Tortula modica var. stenocarpa
(Velen.) Zand., see treatment of Tortula]
[var. tenuis Vent. Eur = Tortula modica var. tenuis (Vent.) Zand.,
see treatment of Tortula]
fo. cylindrica Roll, Hedwigia 56: 158, 1915 Eirr
fo. robusta Podp., Vslyedky 11. V. Kl. Pfirod. Prost~jov 8: 32,
1906 Eur
[Pottia lanceo/ata (Hedw.) C. MUll. Eur As2 As5 Afr1 Am1 =
Tortula /anceo/a Zand. nom. nov., see treatment of Tortula]
[subsp. leucodonta (Schimp.) Boul. Eur = Pottia lanceo/ata var.
angustata (B.&S. in BSG) C. MUll. fide Husnot, Muse. Gall.
1(3): 76, 1885 = Tortula lanceola var. angustata (B.&S. in
BSG) Zand., see treatment ofTortula]
[var. albidens Corb. Eur = Tortula lanceo/a var. albidens (Corb.)
Zand., see treatment of Tortula]
[var. angustata (B.&S. in BSG) C. MUll. Eur As5 Afr1 = Tortula
lanceola var. angustata (B.&S. in BSG) Zand., see treatment of
Tortula]
[var. lejolisii Corb. Eur Afr1 = Tortula /anceo/a var. /ejolisii
(Corb.) Zand., see treatment of Tortu/a]
var. leucocephala Schimp. in Paques nom. nud.? fide Index
Muscorum 4: 195, 1967 Eur
[var. leucodonta Schimp. Eur = Tortula lanceola var. leucodonta
(Schimp.) Zand., see treatment of Tortula]
[var. macrophylla Warnst. Eur = Tortula lanceola var.
macrophylla (Warnst.) Zand., see treatment of Tortula]
[var. microphylla Warnst. Eur = Tortula lanceola var.
microphylla (Warnst.) Zand., see treatment of Tortula]
[var. mucronata Amann Eur = Tortula lanceola var. mucronata
(Amann) Zand., see treatment of Tortula]
[var. ova/ifolia Warnst. Eur = Tortula lanceo/a var. ova/ifolia
(Warnst.) Zand., see treatment of Tortula]
[var. papillosa Corb. Eur = Tortula /anceola var. papillosa
(Corb.) Zand., see treatment of Tortula]
[var. rigidior (Schwaegr.) Wijk & Marg. Eur = Tortula lanceo/a
var. rigidior (Schwaegr.) Zand., see treatment of Tortula]
var. trabecu/ata Podp. Eur
fo. breviseta (Roll) Podp. , Consp. Muse. Eur. 229, 1954 (Pottia
/anceolata var.) Eur
fo. conica Roll, Hedwigia 56: 159, 1915 Eur
fo. major Roll, Hedwigia 56: 158, 1915 Eur
[fo. minor Geh. in Warnst., Hedwigia 58: 134, 1916 syn. inval.
Eur = Pottia /anceo/ata var. microphylla Warnst. fide Warnstorf 1916]
[fo. ovata Roll, Hedwigia 56(1-3): 159, 1915 Eur = Pottia
lanceolata subfo. ovata (Roll) Podp.]
fo. pilifera Roll, Hedwigia 56: 159, 1915 Eur
fo. stenocarpa Podp., Vy'sledky VII. Sborn. Klubu Pfir. v Brno
5: 13, 1923 Eur
fo. tortuosa Roll, Hedwigia 56: 158, 1915 Eur
subfo. ovata (Roll) Podp., Consp. Muse. Eur. 229, 1956
GENERA, SPECIES AND INFRASPECIFIC TAXA
subfo. conica (Roll) Podp., Consp. Muse. Eur. 229, 1954 (Pottia
lanceolata fo.) nom. inval. Eur
[Portia latifolia (Schwaegr.) C. Miill. = Stegonia latifolia (Schwaegr.)
Vent.]
[fo. pilifera (Brid.) Monk., Laubm. Eur. 333, 1927 Eur Asl Ami=
Stegonia latifolia var. pilifera (Brid.) Broth.]
Portia latzii Catcheside, Mosses S. Austr. 126, 1980 Austrl
Pottia ligularifolia C. Miill., Hedwigia 34: 123, 1895 Am5
Pottia longifolia R. Br. ter Austr2
[Pottia longirostris Hampe ex C. Miill. Afr2 = Hennediella
longirostris (Hampe ex C. Miill.) Zand., see treatment of Hennediella]
[Portia macowaniana C. Miill. Afr4 [see Magill, Fl. S. Afr. I. Mosses
1: 207, 1981 (1982)] =Portia starckeana subsp. conica (Schleich.
ex Schwaegr.) Chamber!., Roy. Bot. Gard. Edinburg Not. 29:
403, 1969 = Microbryum davallianum var. conicum (Schleich. ex
Schwaegr.) Zand., see treatment of Microbryum]
[Pottia macrocarpa Schimp. Am6 = Funaria macrocarpa (Schimp.)
Zand. (Funariaceae) see Excluded Taxa]
[Portia macrophylla (R. Br. ter) Sainsb. Am6 Austr2 = Hennediella
macrophylla (R. Br. ter) Par., see treatment of Hennediella]
[Pottia maritima (R. Br. ter) Broth. Austr2 = Tortula maritima (R. Br.
ter) Zand, see treatment of Dendia]
[Pottia minor (C. Miill.) Wijk & Marg. As3 Afr2 = Tortula minor (C.
Miill.) Zand., see treatment of Tortula]
[var. elatior (C. Miill.) Wijk & Marg. As3 Afr2 = Tortula minor
var. elatior (C. Miill.) Zand., see treatment of Tortula]
[Pottia mirabilis Broth. & Par. Afr2 = Physcomitrium mirabile
(Broth. & Par.) Zand. (Funariaceae) see Excluded Taxa]
[Pottia mutica Vent. in De Not. Eur As5 Afr1 = Pottia starckeana
var. brachyoda (BSG) C. Miill. fide Monkemeyer, Laub. Eur.
327, 1927 = Microbryum starckeanum var. brachyodus (BSG)
Zand., see treatment of Microbryum]
var. brachyphylla Wamst. Austrl
var. gymnostoma Corb. Afr1
var.leucodonta Corb. Afr1
var. parvifolia Warns!. Eur
[var. salina (Warns!.) Podp. Eur =Portia starckeana subsp. conica
(Schleich. ex Schwaegr.) Chamber!., Roy. Bot. Gard. Edinburg
Not. 29: 403, 1969 = Microbryum davallianum var. conicum
(Schleich. ex Schwaegr.) Zand., see treatment of Microbryum]
Portia namaquensis Magill, Fl. S. Afr. I. Mosses 1: 206, 1981 [1982]
Afr4
Pottia naumanii C. Miill. Afr4
Portia neocaledonia Ther. nom. inval. Oc
[Pottia nevadensis Card. & Th~r. Am1 = Tortula nevadensis (Card. &
Th~r.) Zand., see treatment of Tortu/a]
Pottia notarisii Schimp. Eur
var. cyclopica Zodda, Malphighia 24: 274, 1911 Eur
Pottia obliqua R. Br. ter Austr2
[Portia obtusifolia (C. Miill.) C. Miill. Am5 = Pottia gardneriana C.
Miill., Syn. 2(9-10): 754, 772, 1851 Hyophila obtusifolia (C.
Miill.) Jaeg.]
[Pottia oedipodioides C. Miill. Afr4 = Hennediel/a oedipodioides (C.
Miill.) Zand., see treatment of Hennediella]
[Pottia pal/ida Lindb. Eur Afr1 As5 = Tortula pal/ida (Lindb.) Zand.,
see treatment of Tortula]
[var. longicuspis Warnst. Eur As5 Afr1
Tortula pal/ida var.
longicuspis (Warns!.) Zand., see treatment of Tortula]
Pottia pellata (Schimp.) Broth. Am2
Pottia propagulifera Herz. Eur As2
[Portia pusilla Warnst. Afr2 hom. illeg. = Pottia minor (C. Miill.)
Wijk & Marg. = Tortula minor (C. Miill.) Zand., see treatment of
=
=
319
Tortula]
[Pottia randii Kenn. Eur Am1 = Tortula randii (Kenn.) Zand., see
treatment of Tortula]
[Pottia recta (Sm.) Mitt. Eur Afr1 = Microbryum rectum (With.)
Zand., see treatment of Microbryum]
Portia recurvifolia Warnst. Afr1
Pottia sampaiana Mach. Eur
Portia scabrifolia Bartr. Austr1
Pottia serrata R. Br. ter Austr2
[Portia splachnobryoides C. Miill. As2 = Chenia leptophylla (C.
Miill.) Zand., see treatment of Chenia and vide Arts & Sollman,
Lindbergia 17:22, 1991]
[Portia splachnoides (Homsch.) Broth. Afr4 =Tortula splachnoides
(Homsch.) Zand., see treatment ofTortula]
[Pottia starckeana (Hedw.) C. Miill. Eur As5 Afr1 Am1 Am2
Austr1 Austr2 = Microbryum starckeanum (Hedw.) Zand., see
treatment of Microbryum]
[subsp. conica (Schleich. ex Schwaegr.) Chamber!., Roy. Bot.
Gard. Edinburg Not. 29: 403, 1969 Eur Afr1 As5 Am2 =
Microbryum dava//ianum var. conicum (Schleich. ex
Schwaegr.) Zand., see treatment of Microbryum]
[subsp. minutula (Schleich. ex Schwaegr.) Bouvet, Bull. Soc.
Etudes Sci. Angers 26: 87, 1896 (Gymnostomum) Eur Afr1
Austr1 = Microbryum davallianum (Hedw.) Zand. var.
davallianum, see treatment of Microbryum]
[subsp. minutula (Schleich. ex Schwaegr.) Chamber!., Roy. Bot.
Gard. Edinburg Not. 29: 403, 1969 (Gymnostomum) hom. illeg.
Eur Afr1 Austr1 =Portia starckeana subsp. minutula (Schleich.
ex Schwaegr.) Bouvet = Microbryum davallianum (Hedw.)
Zand. var. davallianum, see treatment of Microbryum
[var. brachyodus (BSG) C. Miill. Eur As5 Afr1 = Microbryum
starckeanum var. brachyodus (BSG) Zand., see treatment of
Microbryum]
[var. brevidens Latz. Eur = Microbryum starckeanum var.
brevidens (Latz.) Zand. (Portia starckeana var.), see treatment
of Microbryum]
[var. fosbergii (Bartr.) Zand., Novon 3: 92, 1993 (Pottia) =
Microbryum starckeanum var. fosbergii (Bartr.) Zand., see
treatment of Microbryum]
[var. leiostoma Corb. in Corb. & Pitard Afr1 = Microbryum
starckeanum var. leiostoma (Corb in Corb. & Pitard) Zand., see
treatment of Microbryum]
[var. subgymnostoma (De Not.) Grav. Eur
Microbryum
starckeanum var. subgymnostoma (De Not.) Zand., see treatment of Microbryum]
[var. submutica Latz. Eur = Microbryum starckeanum var.
submutica (Latz.) Zand., see treatment of Microbryum]
[fo. brevifolia Limpr., Laubm. Deutsch. 1: 536, 1888 Eur =
Microbryum starckeanum fo. brevifolium (Limpr.) Zand., see
treatment of Microbryum]
[fo. dextrorsa Limpr., Laubm. Deutsch. 1: 536, 1888 Eur =
Microbryum starckeanum fo. dextrorsum (Limpr.) Zand., see
treatment of Microbryum]
[fo. microphylla Warns!., Hedwigia 58: 144, 1917 Eur =Microbryum starckeanum fo. microphyllum (Warnst.) Zand., see
treatment of Microbryum]
[fo. minima Latz., Bot. Centralbl. Beih. 2: 483, 1931 Eur = Pottia
starckeana (Hedw.) C. Miill. = Microbryum starckeanum
(Hedw.) Zand., see treatment of Microbryum]
Pottia stenocarpa P. Yarde Afr2
[Pottia stevensii R. Br. ter Austr2 = Tortula sainsburyana Zand.
nom. nov., see treatment of Tortula]
[Pottia subplanomarginata Dix. Afr4 = Microbryum
=
GENERA OF THE POTTIACEAE
320
subplanomarginatum (Dix.) Zand., see treatment of Microbryum]
Portia tasmanica Broth. Austr1
[Portia texana Wareh. in Grout Am1 Am2 =Portia starckeana subsp.
conica (Schleich. ex Schwaegr.) Chamberl. fide Chamberl., Roy.
Bot. Gard. Edinburg Not. 29: 403, 1969 = Microbryum
davallianum var. conicum (Schleich. ex Schwaegr.) Zand., see
treatment of Microbryum]
Portia thraustophyl/a (Angstr.) Par. Am5
[Portia truncata (Hedw.) B.&S. Eur As1 As2 As3 Afr1 Am1 Am6
Austr1 Austr2 = Tortula truncata (Hedw.) Mitt. in Godm., see
treatment of Tortula]
[subsp. intermedia (Turn.) Bouvet, Bull. Soc. Etudes Sci. Angers
26: 86, 1896 (Gymnostomum) Eur As2 Afr1 =Portia intermedia
(Tum.) Fiirnr. =Tortula modica Zand. nom. nov., see treatment of
Tortula]
[subsp. lirtoralis (Mitt.) Giac. Eur Afr1
Tortula truncata var.
littoralis (Mitt.) Zand., see treatment of Tortula]
[var. brevirostris (Lisa) De Not. Eur = Tortula truncata var.
brevirostris (Lisa) Zand., see treatment of Tortula]
[var. illyrica (Latz.) Podp. Eur = Tortula truncata var. illyrica
(Latz.) Zand., see treatment of Tortula]
Tortula truncata var.
[var. littoralis (Mitt.) Wamst. Eur Afr1
lirtoralis (Mitt.) Zand., see treatment of Tortula]
[var. major (Web. & Mohr) B.&S. = Portia intermedia (Tum.)
Fiirnr. = Tortula modica Zand. nom. nov., see treatment of Tortula]
[var. minutissima Wamst. Eur = Tortula truncata var. minutissima
(Wamst.) Zand., see treatment of Tortula]
fo. angustata Wamst., Hedwigia 58: 116, 1916 Eur
fo. angustifolia Wanst., Hedwigia 58: 122, 1916 Eur
fo. brevicuspis Wamst., Hedwigia 58: 119, 1916 Eur
fo. compacta Roll, Hedwigia 56(1-3): 158, 1915 Eur
fo. elongata Hammerschtn., Mitt. Bayer. Bot. Ges. 2: 184, 1908 Eur
fo . latifolia Wamst., Hedwigia 58: 120, 1916 Eur
fo. longicuspis Wamst., Hedwigia 58: 118, 1916 Eur
fo. longifo/ia Wamst., Hedwigia 58: 121, 1916 Am2
fo. major Roll, Hedwigia 56(1-3): 158, 1915 nom. illeg. non Portia
truncata var. major (Web. & Mohr) B.&S. Eur
fo. pusi/la Roll, Hedwigia 56(1-3): 158, 1915 Eur
fo. serrulata Roll, Hedwigia 56(1-3): 158, 1915 Eur
fo. spathulata (Wamst.) Wamst., Hedwigia 58: 115, 1916 (Portia
truncatula var.) Eur
[Portia truncatula (With.) Bus. = Portia truncata (Hedw.) BSG
Tortula truncata (Hedw.) Mitt. in Godm.]
var. macrocarpa Bus. Eur
Portia u/eana Par. Am5
[Portia verrucosa Wamst. ex Rehm. Afr4 = Portia starckeana var.
conica (Schleich. ex Schwaegr.) Chamberl., Bot. Gard. Edinburg
Not. 29: 403, 1969 = Microbryum davallianum var. conicum
(Schleich. ex Schwaegr.) Zand., see treatment of Microbryum]
[Pottia viridifolia Mitt. Eur =Portia crinita Wils. ex BSG fide Corley
et al., J. Bryol. 11: 620, 1981 [ 1982] =Portia wilsonii var. crinita
(Wils. ex BSG) Wamst. = Tortula wilsonii var. crinita (Wils. ex
B.&S.) Zand., see treatment of Tortula]
[var. flavescens (Corb.) Podp. Eur = Tortula wilsonii var. crinita
(Wils. ex B.&S.) Zand.]
[Portia watsonii R. S. Chopra, Taxon, Indian Mosses 155, 1975
(nom . nov. for Portia denticu/ata Dix. & P. Yarde hom. il/eg.) =
Chenia /eptophylla (C. Miill.) Zand., see treatment of Chenia]
[Portia willisiana Sainsb., Rev. Bryol. Lichenol. 25: 237, 1956 Austr1
= Weissia willisiana (Sainsb.) Catcheside, Mosses S. Austr. 194,
1980]
[Portia wilsonii (Hook.) B.&S. Eur As1 As5 Afr1 Am1 = Tortula
=
=
=
wilsonii (Hook.) Zand., see treatment of Tortula]
[subsp. asperula (Mitt.) Kindb. Eur = Tortula wilsonii var.
asperula (Mitt.) Zand., see treatment of Portia]
Tortula
[var. crinita (Wils. ex B.&S.) Wamst. Eur As5 Afr1
wilsonii var. crinita (Wils. ex B.&S.) Zand., see treatment of
Portia]
[var. mucronifolia (Bruch) Wamst. As1 As5 = Tortula wilsonii
var. mucronifolia (Bruch in F. Miill.) Zand., see treatment of
Portia]
fo. pilifera (Limpr.) Podp., Consp. Muse. Eur. 228, 1954 (Portia
wilsonii var.) Eur
[Portia zealandiae (R. Br. ter) Par. Austr2 = Microbryum
zee/andiae (R. Br. ter) Zand., see treatment of Microbryum]
[Portia zea/andioides Dix. & Sainsb. Austr2 = Portia starckeana
var. conica (Schleich. ex Schwaegr.) Chamberl., Roy. Bot.
Gard. Edinburg Not. 29: 403, 1969 Microbryum davallianum
var. conicum (Schleich. ex Schwaegr.) Zand., see treatment of
Microbryum]
=
=
[PRIONIDIUM Hilp. = Didymodon Hedw. fide Saito, J. Hattori
Bot. Lab. 39: 500, 1975]
[Prionidium erosodenticulatum (C. Miill.) Chen As2 = Didymodon
erosodenticulatus (C. Miill.) Saito fide Saito, J. Hattori Bot.
Lab.39: 504, 1975]
[Prionidium setschwanicum (Broth.) Hilp. As2 = Didymodon
erosodenticu/atus (C. Miill.) Saito fide Saito, J. Hattori Bot.
Lab. 39: 504, 1975]
[PSEUDALOINA Delgad., Bryologist 85: 401, 1982 (1983) =
Crossidium Jur., see treatment of Crossidium]
[Pseuda/oina woodii Delgad., Bryologist 85: 401, 1982 [1983] As5
Crossidium woodii (De1gad.) Zand., see treatment of Crossidium]
=
PSEUDOCROSSIDIUM Williams
Pseudocrossidium apiculatum Williams Am4 Am6
[Pseudocrossidium aureum (Bartr.) Zand., Phytologia 44: 207,
1979 (Barbula) Am1 Am2 = Pseudocrossidium crinitum
(Schultz) Zand., see treatment of Pseudocrossidium]
Pseudocrossidium austrorevolutum (Besch. in Britt.) Zand. (Barbula), see treatment of Pseudocrossidium Am4
Pseudocrossidium carinatum (Gill. ex Grev.) Zand. (Tortu/a), see
treatment of Pseudocrossidium Am6
Pseudocrossidium chilense Williams Am6
Pseudocrossidium crinitum (Schultz) Zand., see treatment of
Pseudocrossidium (Barbula) Afr2 Afr4 Am1 Am2 Am6 Austr1
Austr2
Pseudocrossidium elatum (Williams) Delg., Bryologist 78: 278,
1975 (Crossidium) Am4
Pseudocrossidium excavatum (Mitt.) Williams Am4
Pseudocrossidium hornschuchianum (Schultz) Zand., Phyto1ogia
44: 205, 1979 (Barbula) Eur As5 Afr1 Afr4 Am1 Austr1
Pseudocrossidium leucocalyx (Mont.) Ther. Am6
[Pseudocrossidium obtusulum (Lindb.) Crum & Anderson,
Bryologist 92: 533, 1989 Am1 Pseudocrossidium revolutum
var. obtusulum (Lindb.) Tan, Zand. & T. Tayl., see treatment of
Pseudocrossidium]
Pseudocrossidium mendozense (Mitt.) Zand. (Tortula), see treatment of Pseudocrossidium Am6
Pseudocrossidium pachygastrellum (Herz.) Broth. Am4
Pseudocrossidium perrevolutum (C. Miill.) Zand. (Barbula), see
treatment of Pseudocrossidium Am6
var. acutifolium (C. Miill.) Zand. (Barbula perrevoluta var.), see
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
treatment of Pseudocrossidium Am6
var. linearifolium (C. Mtill.) Zand. (Barbula perrevoluta var.), see
treatment of Pseudocrossidium Am6
Pseudocrossidium porphyreoneurum (C. Miill. ex Vent.) Zand.
(Barbula), see treatment of Pseudocrossidium Afr2 Afr4
Pseudocrossidium replicatum (Tayl.) Zand., Phytologia 44: 206, 1979
(Barbula) Afr2 Afr4 Aml Am2 Am4 Am6
var. longicuspidatum Zand., Willdenowia 16: 258, 1986 Am4
Pseudocrossidium revolutum (Brid. in Schrad.) Zand., Phytologia 44:
204, 1979 (Barbula) Eur As5 Afrl Afr4
var. obtusulum (Lindh.) Tan, Zand. & T. Tayl., Lindbergia 7: 41,
1981 (Barbula hornschuchiana var.) Eur Am1
Psuedocrossidium steerei Churchill, Bryo1ogist 93: 353, 1990 Am4
PSEUDOSYMBLEPHARIS Broth.
Pseudosymblepharis angustata (Mitt.) Hilp. As2 As3 As4
Pseudosymblepharis angustifolia (Crum & Steere) Zand. (Trichostomum), see treatment of Pseudosymblepharis Am3
[Pseudosymblepharis bartramii Ther. ex Bartr. Am2 = Pseudosymblepharis schimperiana (Par.) Crum, see treatment of Pseudosymblepharis]
Pseudosymblepharis cavernarum (Broth.) Zand. (Trichostomum), see
treatment of Pseudosymblepharis Am5
Pseudosymblepharis circinnatula (Broth. in Voltzk) Zand. (Trichostomum), see treatment of Pseudosymblepharis Afr3
Pseudosymblepharis duriuscula (Mitt.) Chen As2 As3
Pseudosymblepharis indica Dix. & P. Yarde As3
Psuedosymblepharis khasiana (Mitt.) Zand. (Tortula), see treatment
of Pseudosymblepharis As3
[Pseudosymblepharis mauiensis (C. MUll.) Broth. As4 Oc = Pseudosymblepharis angustata (Mitt.) Hilp. fide Norris & Koponen,
Acta Bot. Fenn. 137: 94, 1987]
[Pseudosymblepharis pallidens Dix. As3 Trichostomum pallidens
(Dix.) Zand., see treatment of Trichostomum]
[Pseudosymblepharis papillosula (Card. & Ther.) Broth. As2 As3 =
Pseudosymblepharis angustata (Mitt.) Hilp. fide Saito, J. Hattori
Bot. Lab. 39: 439, 1975]
var. robusta Tix., Rev. Bryol. Lichenol. 34: 134, 1966 As3
Pseudosymblepharis perlongifolia (Frohl.) Zand. (Trichostomum), see
treatment of Trichostomum As4
Pseudosymblepharis schimperiana (Par.) Crum Am2 Am3 Am4 Am5
[Pseudosymblepharis socotrana (Mitt.) Ther. Afr2 = Weissia
artocosana Zand. nom. nov., see treatment of Weissia]
[Pseudosymblepharis subduriuscula (C. Miill.) Chen As2 As3 As4
Pseudosymblepharis angustata (Mitt.) Hilp. fide Norris &
Koponen, Acta Bot. Fenn. 137: 94, 1987]
Pseudosymblepharis syrrhopodontoides (Dix.) Zand., see treatment
of Pseudosymblepharis Afr2
Pseudosymblepharis verrucosa (Broth. & Par.) Zand. (Trichostomum), see treatment of Pseudosymblepharis Oc
=
=
[PSEUDOTIMMIELLA Biz., Cryptogamie Bryol. Lichenol. 1:425,
1980 = Diphyscium Mohr (Diphysciaceae) see Excluded Taxa]
[Pseudotimmiella pocsii Biz., Cryptogamie Bryol. Lichenol. 1: 425,
1980 Afr2 =Diphyscium poscii (Biz.) Zand. (Diphysciaceae) see
Excluded Taxa]
PTERYGONEURUM Jur.
[Pterygoneurum arcticum Steere, Bryologist 62: 217, 1960 Aml
Pterygoneurum lamellatum (Lindh.) Jur.fide Steere, 1976]
Pterygoneurum californicum Crum, Madroiio 19: 92, 1967 Am1
[Pterygoneurum cavifolium Jur.
Pterygoneurum ovatum (Hedw.)
Dix.]
=
321
var. muticum Schiffn. As5
[fo. crossidioides Sapegin, Bot. Jahrb. Syst. 46(Beibl. 105): 13,
1911 Eur = Pterygoneurum ovatum fo. crossidioides (Sapegin)
Podp., Consp. Muse. Eur. 235, 1954]
[fo. /ongiseta Amann, Bull. Soc. Vaudoise Sci. Nat. 57: 123,
Pterygoneurum ovatum fo. longiseta (Amann)
1929 Eur
Podp.]
Pterygoneurum chotticum (Trab.) Broth. Afrl
Pterygoneurum crossidioides Frey, Herrnstadt & Kiirschner, Nov.
Hedw. 50: 239-244, 1990 As5
Pterygoneurum kemsleyi Willis Austrl
Pterygoneurum kozlovii Laz., Vizn. List. Moch. Ukr. S.R.R. 104,
1936 nom . inval. sin. descr. lat. Eur Pterygoneurum kozlovii
Laz. ex Laz.
Pterygoneurum kozlovii Laz. ex Laz., Bot. Zhum. Ak. Nauk Uk.
RSS 3: 61, 1946 Eur Am1
Pterygoneurum lamellatum (Lindh.) Jur. Eur Asl As2 Am1
Pterygoneurum macleanum Wamst. Afr4
Pterygoneurum medium (Salm.) Broth. As1 As5
Pterygoneurum ovatum (Hedw.) Dix. Eur As1 As5 Afrl Am1 Am6
Austrl
var. humile (Amann) Podp. Eur
var. perraldieri (Besch.) Wijk & Marg. Afrl
fo. crossidioides (Sapegin) Podp., Consp. Muse. Eur. 235, 1954
Eur
fo. epilosum (Brid.) Podp., Consp. Muse. Eur. 235, 1954 (Gymnostomum ovatum var.) Eur
fo. incanum (Nees & Homsch.) Podp., Consp. Muse. Eur. 235,
1954 (Gymnostomum ovatum var.) Eur As5 Afr1 Aml
fo. /ongisetum (Amann) Podp., Consp. Muse. Eur. 235, 1954
(Pterygoneurum cavifolium fo.) Eur
fo. majus Roll, Hedwigia 56: 160, 1915 Eur
fo. minimum Roll, Hedwigia 56: 160, 1915 Eur
fo. polycarpum (Gyorf.) Podp., Consp. Muse. Eur. 235, 1954
(Pterygoneurum cavifolium var.) Eur
[Pterygoneurum rosei Williams Am4
Aloina rosei (Williams)
Delgadillo, Bryologist 76: 273, 1973]
[Pterygoneurum smardaeanum Vanek Eur (= Phascum cuspidatum
Hedw. but see discussion by Corley et al., J. Bryol. 11: 64 7,
1981 [ 1982]) = Pterygoneurum koslovii Laz. fide Abramova et
al., Notyl. Syst. Plant non Vascular. Acad. Sci. URSS lnst. Bot.
V. L. Komarovii 10: 314, 1973]
Pterygoneurum subsessile (Brid.) Jur. Eur As1 Afr1 Aml Am6
var. henrici (Rau) Wareh. in Grout Am1
var. kieneri Hab. Am1
fo. robustum Roll, Hedwigia 56: 160, 1915 Eur
fo. tenellum Roll, Hedwigia 56: 160, 1915 Eur
Pterygoneurum subsessile (Brid.) Jur. xP. ovatum (Hedw.) Dix.fide
Podp., Consp. Muse. Eur. 235, 1954 As1
fo. apterum Lazar., Opr. Listv. Mchov Ukr. 181, 1955 nom. inval.
sin. descr. lat. Eru
=
=
=
QUAESTICULA Zand. (see treatment of Quaesticula)
Quaesticula navicularis (Mitt.) Zand. (Weissia), see treatment of
Quaesticula Am2 Am3
[RECHINGERELLA Frohl., Ann. Naturh. Mus. Wien 66: 36,
1962 (1963) hom. illeg. non Petrak. = Weissia Hedw. fide H.
Robinson, Bryologist 69: 112, 1966]
[Rechingerella macedonica Frohl., Ann. Naturh. Mus. Wien 66: 36,
1962 (1963) = Weissia controversa Hedw. fide H. Robinson,
Bryologist 69: 112, 1966]
322
GENERA OF THE POTTIACEAE
REIMERSIA Chen
Reimersia inconspicua (Griff.) Chen As2 As3 As4
[RHAMPHIDIUM Mitt. referred to Ditrichaceae in Excluded Taxa)
RHEXOPHYLLUM Herz.
[Rhexophyllum laciniatum Herz. = Rhexophyllum subnigrum (Mitt.)
TMr. ex Hilp.fide Zander in Sharp et al., Mo. Fl. Mexico]
Rhexophyllum subnigrum (Mitt.) TMr. ex Hilp. Am1 Am2 Am4
SAGENOTORTULA Zand.
Sagenotortula quitoensis (Tayl. in Hook.) Zand., Phytologia 6S: 430,
1989 (Tortula) Am2 Am4 Am6
[SAITOA Zand. = Saitoel/a Menzel, J. Hattori Bot. Lab. 71: 239,
1992)
[Saitoa peruviana (Williams) Zand., Phytologia 6S: 431, 1989
(Globulina) Am2 Am4 = Saitoel/a peruviana (Williams) Menzel,
J. Hattori Bot. Lab. 71: 240, 1992)
SAITOELLA Menzel, J. Hattori Bot. Lab. 71: 239, 1992
Saitoel/a peruviana (Williams) Menzel, J. Hattori Bot. Lab. 71: 240,
1992 (Giobulina) Am2 Am4
SARCONEURUM Bryhn
[Sarconeurum antarcticum Bryhn, Nyt Mag. Naturvidensk. 40: 20S,
1902 =Sarconeurum glaciale (C. Mi.ill.) Card. & Bryhnfide Robinson in Llano, Antarctic Terrestrial Biology, Antarctic Res. Ser.
20: 171, 1972)
Sarconeurum glaciale (C. Mi.ill.) Card. & Bryhn Am6 Ant
[Sarconeurum tortel/oides Greene, Sci. Rep. Brit. Antarct. Surv. 64:
38, 1970 Ant= Tortel/a tortel/oides (Greene) Robinson fide Robinson in Llano, Antarctic Terrestrial Biology, Antarctic Res. Ser.
20: 171, 1972 =Tortella alpico/a Dix.)
SCOPELOPHILA (Mitt.) Lindb.
Scopelophila cataractae (Mitt.) Broth. Eur As2 As3 As4 AsS Afr2
Am1Am2Am4
Scopelophila infericola Hoe, Bryologist 76: 192, 1973 Oc
Scopelophila ligulata (Spruce) Spruce Aml Am2 Am4 Eur Mrl As2
As3 As4 AsS
[Scopelophila norrisii Zand., Bryologist 88: 3S3, 1986 Am2 = Weisiopsis norrisii (Zand.) Zand., see treatment of Weisiopsis]
Semibarbula ranuii Gangulee, Nova Hedw. 8: 149, 1964 As3
[Semibarbula rufipes (Schimp. ex Besch.) Hilp. Am2 = Barbula
indica (Hook.) Spreng. fide Zander, Phytologia 44: 18S, 1979)
[Semibarbula scaberrima (Broth. & Par.) Hilp. As3 = Barbula
scaberrima Broth. & Par.)
[Semibarbula stulhmannii (Broth.) Hilp. Mr2 Afr4 = Barbula
stulhmannii (Broth.) Broth.)
[Semibarbula trachyphylla (Broth.) Laz. in Lazaret al., Biul. Mosk.
Obsh. lspyt. Prir. Otd. Bioi. 7S(3): 147, 1970 (Tortula) comb.
inval. basion. non cit. As1 = Tortula trachyphylla Broth.]
[SERPOTORTELLA Dix. referred to Serpotortellaceae, see
Excluded Taxa]
[SPLACHNOBRYUM C. Mi.ill. referred to Splachnobryaceae, see
Excluded Taxa]
STEGONIA Vent.
Stegonia hyalinotrichum (Card. & Ther.) Zand. (Phascum), see
treatment of Stegonia Am1 Am2
Stegonia /atifolia (Schwaegr.) Vent. ex Broth. Eur As1 As2 AsS
Aml
var. pilifera (Brid.) Broth. Eur Asl Am1 [= Stegonia pilifera
(Brid.) Crum & Anderson, Bryologist 92: S33, 1989)
[fo. pilifera (Brid.) Nyh., Ill. Moss Fl. Fenno. 98, 19S6 nom.
inval. ref incompl. (Coscinodon) Eur As1 Am1 = Stegonia
pilifera (Brid.) Crum & Anderson, Bryologist 92: S33, 1989 =
Stegonia latifolia var. pilifera (Brid.) Broth.)
Stegonia mouretii (Corb.) Broth. Mr1
var. crinita Corb. ex Jelenc. Mrl
[Stegonia pilifera (Brid.) Crum & Anderson, Bryologist 92: S33,
1989 Eur As1 Am1 = Stegonia /atifolia var. pilifera (Brid.)
Broth.)
[STEPHANODICTYON Dix. = Trichostomum Bruch, see treatment of Trichostomum)
[Stephanodictyon angustinerve Fri:ihl. As4 = Trichostomum
finukamactum Zand. nom. nov., see treatment of Trichostomum)
[Stephanodictyon borneense Dix. As4 (= Pseudosymblepharis
subduriuscu/a (C. Mi.ill.) Chen fide Eddy, Handb. Males.
Mosses 2: 1S3, 1991) = Trichostomum borneense (Dix.) Zand.,
see treatment ofTrichostomumJ
[Stephanodictyon obscurirete Dix. As4 (= Pseudosymblepharis
angustata (Mitt.) Chen fide Eddy, Handb. Males. Mosses 2:
1S3, 1991) = Trichostomum brachydontium Bruch, see treatment ofTrichostomumJ
[SEBILLEA Biz., Rev. Bryol. Lichenol. 40: 120, 1974 nom. inval.
fide Crosby et al. 1992, probably referable to Dicranaceae, see
Excluded Taxa)
[Sebillea brasiliensis Biz., Rev. Bryol. Lichenol. 40: 120, 1974 nom.
inva/. fide Crosby et al. 1992 Am5 = Dicranaceae? see Excluded
Taxa)
STONEA Zand.
Stonea o/eaginosa (Stone) Zand., Phytologia 6S: 431, 1989 (Tortula) Austrl
[SEMIBARBULA Herz. ex Hilp. = Barbula Hedw.)
[Semibarbu/a congoana (Ther.) Biz., Rev. Bryol. Lichenol. 38: S49,
1971-72 (1973) Afr2 = Barbula congoana Ther.)
[Semibarbula e/ongata Hilp. Mr2 = Barbula zambesiaca Magill in
Magill & Schelpe, Mem. Bot. Surv. S. Afr. 43: 21, 1979 nom.
nov.)
[Semibarbula lambarenensis (P. Yarde) Biz., Rev. Bryol. Lichenol.
38: S94, 1971-72 (1973) Mr2 = Barbula lambarenensis P.
Yarde)
[Semibarbula orienta/is (Web.) Wijk & Marg. As2 As3 As4 Mr2
Afr3 Afr4 Oc = Barbula indica (Hook.) Spreng. fide Saito, J.
Hattori Bot. Lab. 39:488, 197S)
[STREBLOTRICHUM P. Beauv. = Barbula Hedw. fide Saito, J.
Hattori Bot. Lab. 39: 499, 197S)
[Streblotrichum bico/or (BSG) Loeske Eur = Barbula bicolor
(BSG) Lindb.)
[Streblotrichum canaliculata (Dix.) R. S, Chopra, Taxon. Indian
Mosses 138, 197S As3 = Didymodon canaliculatus Dix.)
[Streblotrichum convolutum (Hedw.) P. Beauv. Eur As1 As2 AsS
Mr1 Am1 Am2 Austr2 =Barbula convoluta Hedw.)
[subsp. commutatum (Jur.) Giac. Eur AsS Mr1 = Barbula
convolutum subsp. commutata (Jur.) Boul.)
var. commutatum (Jur. Amann Eur AsS Mr1
var. girodii (Ther.) Podp. Eur
323
GENERA, SPECIES AND INFRASPECIFIC TAXA
=
[var. sardoa (BSG) Podp. Eur Barbula convoluta var. sardoa
BSG]
fo.? angustifolium (Warns!.) Podp., Consp. Muse. Eur. 216, 1954
nom. inval. dispon. non cit. (Barbula convoluta var.) Eur
[fo.? brevifolium (Podp.) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon. non cit. (Barbula convoluta fo.) Eur = Barbula
convoluta fo. brevifolia Podp.]
fo.? filiformis (Hag.) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon . non cit. (Barbula convoluta var.) Eur
[fo.? insolatum (Latz.) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon . non cit. (Barbula convoluta fo.)
Barbula
convoluta Co. insolata Latz.]
Co.? /atifolium (Amann) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon. non cit. (Streblotrichum convolutum var.) Eur
fo.? obtusatum (Artr. & Loeske in Bauer) Podp., Consp. Muse. Eur.
216, 1954 nom. inval. dispon . non cit. (Barbula convoluta var.)
Eur
fo.? propaguliferum (Glow.) Podp., Consp. Muse. Eur. 216, 1954
nom. inval. dispon. non cit. (Barbula convoluta var.) Eur
[fo.? rufescens (Loeske & Quelle) Podp., Consp. Muse. Eur. 216,
1954 nom. inval. dispon. non cit. (Barbula convoluta fo.) =Barbula convo/uta fo. rufescens Loeske & QueUe]
[fo.? rufipes (Bauer) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon. non cit. (Barbula convoluta fo.) Eur (Barbula
convoluta fo. rufipes Bauer]
Co.? stockumii (Warns!.) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon . non cit. (Barbula convoluta var.) Eur
fo.? turfaceum (Warnst.) Podp., Consp. Muse. Eur. 216, 1954 nom.
inval. dispon . non cit. (Barbula convoluta var.) Eur
fo.? uliginosum (Limpr.) Podp., Consp. Muse. Eur. 216, 1954 nom.
Barbula
inval. dispon . non cit. (Barbula convoluta var.)
convoluta Co. uliginosa (Limpr.) Monk.]
[Streblotrichum croceum (Brid.) Loeske Eur As2 Afr1 = Barbula
crocea (Brid.) Web. & Mohr]
var. carpaticum Pilous, Preslia 41: 120, 1969 Eur
[var. coreense (Card.) Wijk & Marg. = Barbula coreense (Card.)
Saito, J. Hattori Bot. Lab. 39: 484, 1975]
var. dichodontioides Pilous, Prestia 41 : 121 , 1969 Eur
var.funckianum (Schultz) Podp. Eur
var. grimmioides Pilous, Prestia 41 : 121, 1969 Eur
var. lingulatum Pilous, Prestia 41 : 121, 1969 Eur
fo. cataractarum (Loeske) Podp., Consp. Muse. Eur. 217, 1954
(Streblotrichum croceum var.) Eur
[Streblotrichum enderesii (Garov.) Loeske Eur As1
Barbula
enderesii Garov.]
Streblotrichum gracillimum Herz. As2
[Streblotrichum hypse/ostegium (Card.) Hilp. Am2 = Barbula indica
(Hook.) Spreng. fide Zander, Phytologia 44: 196, 1979]
Streblotrichum obtusifolium (Hilp.) Chen As2
[Streblotrichum pringlei (Card.) Hilp. Am2 = Barbula indica (Hook.)
Spreng. fide Zander, Phytologia 44: 186, 1979]
Streblotrichum propaguliferum Li & Zhang, Acta Bot. Yunnan. 5(4):
385, 1983 As2
=
=
=
=
STREPTOCAL YPT A C. Mi.ill.
Streptoca/ypta lorentziana C. Mi.ill. Am6
Streptocalypta santosii (Bartr.) Zand., Lindbergia 8: 165, 1982 [1983]
(Tarte/la) Am2
Streptocalypta tortelloides (Card.) Zand., Lindbergia 8: 163, 1982
[1983] (Barnesia) Am2
STREPTOPOGON Wils. in Mitt.
[Streptopogon australis Mitt. Afr4 [(= Didymodon sp. fide Salmon
1903: 109) = Daltonia cf. angustifolia Dozy & Molk. (Daltoniaeeae), see treatment of Streptopogon]
Streptopogon calymperes C. Mi.ill. ex Geh. Afr3 Am2 Am3 Am4
Streptopogon calymperoides Demar. & P. Yarde Afr2
Streptopogon cavifolius Mitt. Am2 Am4
Streptopogon clavipes (Spruce) Spruce ex Mitt. Am4
Streptopogon elimbatus Card. in P. Yarde Afr3
Streptopogon erythrodontus (Tayl.) Wils. Afr3 Am2 Am3 Am4 Oe
var. intermedius Salm. Am4
var. rutenbergii (Geh.) Salm. Afr2 Afr3
Streptopogon heterophy/lus Herz. Am4
Streptopogonjuarezii Sharp, Phytologia 61: 372, 1986 Am2
Streptopogon lindigii Hampe Am4
Streptopogon matudianus Crum Am2
Streptopogon peruvianus Broth. ex Herz. nom. inval. sin. descr.
Am4
[Streptopogon rzedowskii Cardenas, Phytologia 61: 297, 1986 Am2
= Leptodontium proliferum Herz. fide Cardenas, Anales Inst.
Biol. Univ. Nac. Auton. Mexico 58, Ser. Bot. 1: 94, 1987
(1989)]
Streptopogon spathulatus Herz. Am4
Streptopogon stenophy/lus P. Yarde & Tber. Afr2
Streptopogon subelimbatus Card. in Grand. Afr3
STREPTOTRICHUM Herz.
Streptotrichum ramico/a Herz. Am4
SYNTRICHIA Brid.
[Syntrichia abranchesii (Luis.) Ochyra, Fragm. Aorist. Geobot. 37:
Syntrichia caninervis var.
212 , 1992 (Tortula) Eur
abranchesii (Luis.) Zand., see treatment of Syntrichia]
Syntrichia abruptinervis (Dix.) Zand. (Tortula) see treatment of
Syntrichia Austr2
[Syntrichia aciphylla (B.&S.) Jur. = Tortula ruralis var. alpina
Wahlenb.]
[var. calva Amann Eur = Syntrichia norvegica var. calva
(Amann) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
Syntrichia aculeata (Wils.) Zand. (Barbu/a), see treatment of Syntrichia Am4
Syntrichia a/pestris (Dix. in Herz.) Zand. (Tortula) see treatment of
Syntrichia Afr2
Syntrichia ammonsiana (Crum & Anders.) Ochyra, Fragm. Aorist.
Geobot. 37: 212, 1992 (Tortula) Am1 Afr4
Syntrichia amplexa (Lesq.) Zand. (Barbula), see treatment of Syntrichia Eur Am1
Syntrichia amphidiacea (C. Miill.) Zand. (Barbula), see treatment
of Syntrichia Afr2 Am1 Am2 Am3
Syntrichia anderssonii (Angstr.) Zand. (Tortula) , see treatment of
Syntrichia Am6 Austr2 Ant
var. fagico/a (C. Mi.ill.) Zand. (Barbu/a conotricha var.), see
treatment of Syntrichia Am6
Syntrichia andico/a (Mont.) Ochyra, Fragm. Florist. Geobot. 37:
212, 1992 (Tortula) Am2 Am4
Syntrichia antarctica (C. Miill. & Hampe) Zand. (Barbula), see
treatment of Syntrichia Afr4 Am6 Austr1 Austr2
Syntrichia baileyi (Broth.) Zand. (Tortula), see treatment of Syntrichia Austr1
Syntrichia bartramii (Steere in Grout) Zand. (Tortula), see treatment of Syntrichia Am 1 Am2
Syntrichia bipedicel/ata (Britt.) Zand. (Tortula), see treatment of
Syntrichia Am4
Syntrichia bogotensis (Hampe) Mitt. (Barbula), see treatment of
Syntrichia [l.M. citation of Am3 is incorrect as Mitten's (1869)
=
324
GENERA OF THE POTTIACEAE
"Guadelupe" refers to an area in Colombia, not the West Indies]
Am4
Syntrichia bolanderi (Lesq. & James) Zand. (Barbula), see treatment
of Syntrichia Eur Afr1 Am1
Syntrichia brachyclada (Card.) Zand. (Tortula), see treatment of Syntrichia Am6
Syntrichia brandisii (C. Miill.) Zand. (Barbula), see treatment of Syntrichia As3
Syntrichia brevisetacea (F. Miill.) Zand. (Barbula), see treatment of
Syntrichia Austr1
Syntrichia cainii (Crum & Anders.) Zand. (Tortula), see treatment of
Syntrichia Am1
Syntrichia ca/cicola Amann Eur
Syntrichia campestris (Dus.) Zand. (Tortula), see treatment of Syntrichia Am6
Syntrichia caninervis Mitt. Eur As2 Am1
var. abranchesii (Luis.) Zand. (Tortula), see treatment of Syntrichia
Eur
var. gypsophila (Amann ex Roth) Ochyra, Fragm. Florist. Geobot.
37: 212, 1992 (Tortula ruralis var.) Eur As2
var. spuria (Amann) Zand. (Tortula), see treatment of Syntrichia
Eur
Syntrichia cava/Iii (Negri) Ochyra, Fragm. Florist. Geobot. 37: 212,
1992 (Tortula) Afr2
Syntrichia chisosa (Magill, Delg. & Stark) Zand. (Tortula), see treatment of Syntrichia Am1 Am2 Afr4
Syntrichia ciliata (Broth.) Zand. (Tortula), see treatment of Syntrichia
Am4
Syntrichia conferta (Bartr.) Zand. (Tortula), see treatment of Syntrichia Ant
Syntrichia costesii (Ther.) Zand. (Tortula), see treatment of SyntrichiaAm6
Syntrichia didymodontoides (Broth. in Dryg.) Zand. (Tortula), see
treatment of Syntrichia Afr4
[Syntrichia echinata (Schiffn.) Herrnstadt & Ben-Sasson, Bryologist
85: 216, 1982 (Tortu/a) Eur As5 Tortula princeps subsp.
echinata (Schiffn.) Kramer, Bryophyt. Biblioth. 21: 86, 1980
Syntrichia princeps var. echinata (Shiffn.) Zand. (Tortula), see
treatment of Syntrichia]
Syntrichia epilosa (Broth. ex Dus.) Zand. (Tortula), see treatment of
Syntrichia Am6
var. pilifera (Ther.) Zand. (Tortula epilosa var.), see treatment of
Syntrichia Am6
[Syntrichia erythroneura (C. Miill.) Sim Afr4 = Tortula antarctica
(Hampe in C. Miill.) Wils. in Hook. f. fide Kramer, J. Hattori Bot.
Lab. 65: 89, 1988 = Syntrichia antarctica (Hampe in C. Miill.)
Zand., see treatment of Syntrichia]
Syntrichia filaris (C. MUll. in Neum.) Zand. (Barbula), see treatment
of Syntrichia Am6 Ant
Syntrichia fiagellaris (Schimp.) Zand. (Barbula), see treatment of
Syntrichia Am6
var. densiretis (Ther.) Zand. (Tortula fiagel/aris var.), see treatment
of Syntrichia] Am6
Syntrichia fontana (C. Miill. in Neum.) Zand. (Barbula), see treatment of Syntrichia Am6 Ant
Syntrichia fragilis (Tayl.) Ochyra, Fragm. Florist. Geobot. 37: 212,
1992 (Tortula) Am1 Am2 Am3 Am4 As3 Afr3 Afr4
Syntrichia fuscoviridis (Car.) Zand. (Tortula), see treatment of Syntrichia Am6
Syntrichia geheebiaeopsis (C. Miill.) Zand. (Barbula) Afr4 Ant
Syntrichia gemmascens (Chen) Zand. (Desmatodon)see treatment of
Syntrichia As2
Syntrichia gemmascens var. hopeiensis (Chen) Zand. (Desmatodon
=
gemmascens var.), see treatment of Syntrichia As2
Syntrichia g/acialis (Kunze ex C. Miill.) Zand. (Barbula), see treatment of Syntrichia Am4 Am6
Syntrichia gromschii (Ther.) Zand. (Tortula), see treatment of Syntrichia Am6
[Syntrichia hadacii Vondr., Bull. Soc. Amis Sci. Lett. Poznon, ser.
D, 6: 121, 1965 As2 = Tortula caninervis var. gypsophila
(Roth) Kramer, Bryoph. Biblioth. 21: 106, 1980 Syntrichia
caninervis var. gypsophila (Roth.) Ochyra, Fragm. Florist.
Geobot. 37: 212, 1992]
Syntrichia handelii (Schiffn.) Bach. Eur As5
var. ferganensis (Laz.) Ochyra, Fragm. Florist. Geobot. 37: 212,
1992 (Tortula) As1
[var. pseudodesertorum Vondr., Bull. Soc. Amis Sci. Lett.
Poznon, ser. D, 6: 121, 1965 As5 = Syntrichia pseudohandelii
Frohl.fide Agnew & Vondra~ek, Feddes Rep. 86: 401, 1975]
Syntrichia inermis (Brid.) Bruch in Hueb. Eur As1 As2 As3 As5
Afr1 Am1 Am2
var. submarginata (Schiffn.) Podp. Eur
Syntrichia intermedia Brid. Eur As1 As3 As5 Afr1 Afr4 Am1 Am2
var. calva (Dur. & Sag.) Delogn. Eur
Syntrichiajaffuelii (Ther.) Zand. (Tortula), see treatment of SyntrichiaAm6
Syntrichia /acerifolia (Williams) Zand. (Tortula), see treatment of
Syntrichia Am4
Syntrichia laevipila Brid. Eur As 1 As3 As5 Afr 1 Am 1 Am6
var. meridiana/is (Schimp.) Jur. Eur
Syntrichia /atifolia (Hartm.) Hiib. Eur Am1
var. propagu/ifera (Milde) Amann Eur
fo. perfragilis Amann, Fl. Mousses Suisse Add. 3: 36, 1933 Eur
[Syntrichia /eptopyxis (C. Miill.) Lazar., J. Bot. Acad. Sci. Ukraine
1(3-4): 95, 1940 (Barbula) As2 =Tortula /eptopyxis (C. Miill.)
Lindb. & Am.]
Syntrichia /eucostega (C. Miill.) Zand. (Barbula), see treatment of
Syntrichia Afr4 Austr1
var. trachyneura (Dix.) Zand. (Tortula) Afr4
Syntrichia limensis (Williams) Zand. (Tortula) see treatment of
Syntrichia Am4
Syntrichia linguifolia (Herz.) Zand. (Tortula), see treatment of Syntrichia Am4
Syntrichia longimucronata (X.-j. Li) Zand. (Tortula), see treatment
of Syntrichia As2
Syntrichia magellanica (Mont. in Gay) Zand. (Tortula), see 'treatment of Syntrichia Am6 Ant
[Syntrichia mniadelphus (C. Miill.) Herz.
Tortula mniadelphus
(C. Miill.) Broth. = Tortula quitoensis Tayl. in Hook. fide
Mishler in Sharp et al., Moss Fl. Mex. = Sagenotortula
quitoensis (Tayl. in Hook.) Zand., Phytologia 65: 430, 1989]
var. coch/earifolia Herz. Am6?
Syntrichia mol/is (B.&S. ex C. Miill.) Zand. (Barbula), see treatment of Syntrichia Afr2
[Syntrichia mongolica Boros, Trans. Brit. Bryol. Soc. 6: 70, 1970
As1 = Syntrichia submontana (Broth.) Ochyra, Fragm. Florist.
Geobot. 37: 212, 1992]
[Syntrichia montana Nees in Raab = Tortula intermedia (Brid.)
Berk.]
var. brevifolia C. Jens. Eur Am1
[var. gelida Podp. Eur Tortula intermedia var. gelida (Amann)
Wijk & Marg.]
var. nivalis Amann Eur
var. rufipila Amann Eur
[fo. brevifolia (Am.) Podp., Consp. Muse. Eur. 258, 1954 (Tortuta ruralis fo.) Eur = Tortula ruralis fo. brevifolia Am. in
=
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
Rabenh.]
Tortula mucronifolia
[Syntrichia mucronifolia (Sehwaegr.) Brid.
Sehwaegr.]
var. systilia Amann Eur
fo. perpusilla (Warnst.) Podp., Consp. Muse. Eur. 25I, I954 (Tortuta mucronifolia var.) Eur
fo. pygmaea Mart. ex Warnst., Hedwigia 52: 74, I9I2 Eur
Syntrichia norvegica Web. Ami Eur Afri Afr4 As I As2 As3
var. calva (Amann) Ochyra, Fragm. Florist. Geobot. 37: 2I2, I992
Eur
Syntrichia obtusissima (C. MUll.) Zand. (Barbula), see treatment of
Syntrichia Ami Am2
Syntrichia pagorum (Milde) Amann, see treatment of Syntrichia Eur
Afr4 Am 1 Am2 Austr 1
Syntrichia papil/osa (Wils.) Jur. Eur Afr4 Ami Am2 Am4 Am5 Am6
Austri Austr2
var. chilensis (Ther.) Zand. (Tortula papillosa var.), see treatment
of Syntrichia Am4
var. meridiana/is (Warns!.) Zand. Eur
fo. saxatilis (Warnst.) Podp., Consp. Muse. Eur. 253, 1954 (Tortula
papillosa var.) Eur
Syntrichia percarnosa (C. Milll.) Zand. (Barbula), see treatment of
Syntrichia Am4 Am6
Syntrichia perichaetialis Herz. nom. inval. Am6
Syntrichia phaea (Hook. f. & Wils.) Zand. (Trichostomum) see treatment of Syntrichia Austr2
Syntrichia pichinchensis (Tayl.) Zand. (Tortula), see treatment of
Syntrichia Am3 Am4
Syntrichia princeps (De Not.) Mitt. Eur As1 As3 AsS Afr1 Afr4 Am1
Am2 Am4 Am6 Austr1 Austr2 Oe Ant
var. brachycarpa (De Not.) Zand. (Tortula princeps var.), see treatment of Syntrichia Eur
var. parnassica (Sehiffn.) Podp. Eur
Syntrichia prostrata (Mont.) Zand. (Tortula), see treatment of Syntrichia Am4 Am6
[Syntrichia pseudodesertorum (Frohl.) Agnew & Vondr., Feddes Rep.
86(6--8): 402, 1975 (Tortula) As3
Syntrichia pseudohandelii (Frohl.), Agnew & Vondr., Feddes Rep.
86(6--8): 401, 1975 (Tortula) As3
Syntrichia pseudorobusta (Dus.) Zand. (Tortula), see treatment of
Syntrichia Am6
[Syntrichia pulvinata (Jur.) Jur. Eur Afr1 Am1 = Tortula pulvinata
(Jur.) Limpr. = Tortula virescens (De Not.) De Not. fide Kramer,
Bryoph. Biblioth. 21: 99, 1980 = Syntrichia virescens (De Not.)
Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
fo. macrophylla (Warns!.) Podp., Consp. Muse. Eur. 255, 1954
(Tortula pulvinata var.) Eur
fo. microphylla (Warnst.) Podp., Consp. Muse. Eur. 255, 1954
(Tortula pulvinata var.) Eur
fo. versispora (Warnst.) Podp., Consp. Muse. Eur. 255, 1954 (Tortuta pulvinata var.) Eur
Syntrichia pygmaea (Dus.) Zand. (Tortula) see treatment of Syntrichia Am6 Austr2
Syntrichia ramosissima (Ther.) Zand. (Tortula), see treatment of Syn trichia Am4
Syntrichia reflexa Zand. (nom. nov. for Tortula rejlexa Li hom.
illeg.), see treatment of Syntrichia As2
Syntrichia rigescens (Broth. & Geh. in Broth.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992 As5
Syntrichia robusta (Hook. & Grev.) Zand. (Tortula) see treatment of
Syntrichia Am5 Am6 Austr1 Ant
var. recurva (Lightowlers) Zand. (Tortula robusta var.) see treatment of Syntrichia Ant
=
325
Syntrichia rubella (Hook. f. & Wils.) Zand. (Tortula), see treatment
of Syntrichia Afr4 Austr1 Austr2
Syntrichia rubra (Mitt. in Hook. f.) Zand. (Tortula) see treatment of
Syntrichia Am6 Austr1 Austr2 Ant
var. subantarctica (Sainsb.) Zand. (Tortula) see treatment of Syntrichia Austr2
[Syntrichia ruraliformis var. subpapil/osissima (Biz. & Pierro! in
Biz.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992 Afr1
Syntrichia ruralis var. subpapil/osissima (Biz. & Pierr.) Zand.
(Tortula ruralis var.), see treatment of Syntrichia]
Syntrichia ruralis (Hedw.) Web. & Mohr Eur As1 As2 As3 AsS
Afr1 Afr2 Afr4 Am1 Am2 Am4 Am5 Austr1 Oe
var. aetnensis Reim. Eur
var. arenicola (Braithw.) Amann Eur As5 Afr1 Am1
[var. calcicola (Amann) Moenk. Eur = Syntrichia calcicola
Amann]
var. gigantea (Lesq.) Zand. (Barbula ruralis var.), see treatment
of Syntrichia Am1
var. gracilis (C. Jens.) Zand. (Tortula ruralis var.), see treatment
of Syntrichia Am1
var. hirsuta (Vent.) Podp. Eur As2 Am1
var. pontresinae (Geh. & Warns!.) Podp. Eur
var. pseudodesertorum Podp. Eur
var. spiralis (Herz.) Zand. (Tortula ruralis var.), see treatment of
Syntrichia Am4
var. submamillosa (Kramer) Zand. (Tortula ruralis var.), see
treatment of Syntrichia As5
var. subpapil/osissima (Biz. & Pierr.) Zand. (Tortula ruralis var.),
see treatment of Syntrichia Afr1
var. substereidosa (Kramer) Zand. (Tortula ruralis var.), see
treatment of Syntrichia As5
fo. brevipila (Warns!.) Podp., Consp. Muse. Eur. 255, 1954 (Tortuta ruralis var.) Eur
[fo. calva (Amann) Podp., Consp. Muse. Eur. 257, 1954 (Syntrichia aciphylla var.) Eur = Syntrichia norvegica var. calva
(Amann) Zand., see treatment of Syntrichia]
fo. compacta (Glow.) Podp., Consp. Muse. Eur. 257, 1954 (Tortuta aciphylla var.) Eur
Lfo. contorta (Podp.) Podp., Consp. Muse. Eur. 255, 1954 (Tortuta ruralis fo.) Eur = Tortula ruralis fo. contorta Podp.]
fo. fallax (Herz.) Podp., Consp. Muse. Eur. 255, 1954 (Tortula
ruralis fo.) Eur
fo. gigantea (Fam.) Podp., Consp. Muse. Eur. 256, 1954 (Barbula
ruraliformis fo.) Eur
fo. gracilis (Meyl.) Podp., Consp. Muse. Eur. 255, 1954 (Tortula
ruralis fo.) Eur
fo./ongipila Papp, Bul. Grlid. Bot. Univ. Cluj 26: 13, 1946 Eur
fo. planifolia (Wamst.) Podp., Consp. Muse. Eur. 256, 1954
(Tortula ruralis var.) Eur
fo. rubicundula (Kindb.) Podp., Consp. Muse. Eur. 256, 1954
(Tortula ruralis subsp.) Eur
fo. rufipila Herz. ex RUbel, Bot. Jahrb. Syst. 47: 478, I912 Eur
fo. rufoneura (Podp.) Podp., Consp. Muse. Eur. 256, 1954 (Tortuta ruralis fo.) Eur
fo. subrufa (Podp.) Podp., Consp. Muse. Eur. 256, 1954 (Tortula
ruralis fo.) Eur
fo. viridis (Matous.) Podp., Consp. Muse. Eur. 256, 1954 (Tortula
ruralis fo.) Eur
Syntrichia saxicola (Card.) Zand. (Tortula) , see treatment of Syntrichia Am6
Syntrichia scabrella (Dus.) Zand. (Tortula), see treatment of Syntrichia Am6
Syntrichia scabrinervis (C. MUll.) Zand. (Barbula), see treatment of
=
326
GENERA OF THE POTTIACEAE
Syntrichia Am4 Am6
Syntrichia serrata (Dix.) Zand. (Tortula serrata Dix. nom. legit. contrary to Index Muscorum; cf. Lightowlers, J. Bryol. 13: 373,
1985) Austr2
Syntrichia serripungens (Lor. & C. Mtill.) Zand. (Barbula) Am4 Am6
var. excesa (C. Mtill.) Zand. (Barbula serripungens var.), see treatment of Syntrichia Am6
Syntrichia sinensis (C. Mtill.) Ochyra, Fragm. Aorist. Geobot. 37:
212, 1992 (Barbula) Eur As1 As2 As3 As5 Afr1
Syntrichia socialis (Dus.) Zand. (Tortula), see treatment of Syntrichia
Am6
Syntrichia subaristata (B.&S. ex C. Mtill.) Zand. (Barbula), see treatment of Syntrichia Afr2
Syntrichia submontana (Broth.) Ochyra, Fragm. Aorist. Geobot. 37:
212, 1992 (Tortula)As1
[Syntrichia subulata (Hedw.) Web. & Mohr. = Tortula subulata
Hedw.]
var. major Htib. Eur
fo. brevifolia Herz., Kryptog. Forsch. 4: 280, 1919 Eur
fo. calcarea (Velen.) Podp., Consp. Muse. Eur. 250, 1954 (Tortula
subulata var.) Eur
fo. compacta (Schiffn.) Podp., Consp. Muse. Eur. 250, 1954 (Tortuta subulata var.) Eur
fo. dentata (Boul.) Podp., Consp. Muse. Eur. 250, 1954 (Barbula
subulata var.) Eur Afr1
[fo. denticulata (Latz.) Podp., Consp. Muse. Eur. 250, 1954 (Tortula subulata fo.) Eur =Tortula subulata fo. denticulata Latz.]
[fo. graeffii (Warnst.) Podp., Consp. Muse. Eur. 250, 1954 (Barbula) Eur =Tortula subulata var. graeffii Warnst.]
fo. inframarginata (Glow.) Podp., Consp. Muse. Eur. 250, 1954
(Tortula subu/ata var.) Eur
fo. lapidico/a (C. Jens.) Podp., Consp. Muse. Eur. 250, 1954
"lapicidicola" (Tortula subulata var.) Eur
fo. mucronata (Farn.) Podp., Consp. Muse. Eur. 250, 1954 (Barbula subulata var.) Eur
fo. recurvomarginata (Breidl. ex Limpr.) Podp., Consp. Muse. Eur.
250, 1954 (Tortula subulata var.) Eur
fo. robusta (Warnst.) Podp., Consp. Muse. Eur. 250, 1954 (Tortula
subulata var.) Eur
fo. saxicola (Lamy) Podp., Consp. Muse. Eur. 250, 1954 (Barbula)
Eur
fo. transiens (Velen.) Podp., Consp. Muse. Eur. 250, 1954 (Tortula
subulata var.) Eur
Syntrichia virescens (De Not.) Ochyra, Fragm. Florist. Geobot. 37:
212, 1992 (Tortula ruralis var.) (cf. Syntrichia virescens (De
Not.) Boros in So6, Magyar Fl. 1: 425, 1964 inval. basion. non
cit.) Eur As5 Afr1 Am1
var. bizotiana (Kramer) Zand. (Tortula virescens subsp.), see treatment of Syntrichia As5
var. iranica (Kramer) Zand., see treatment of Syntrichia As5
var. minor (Biz.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992
(Tortula laevipila var.) As5
Syntrichia viridula (C. Mtill.) Zand. (Barbula) , see treatment of Syntrichia Am4
TENIOLOPHORA Reese
Teniolophorajluviatile (Williams) Reese Am3
TETRACOSCINODON R. Br. ter
Tetracoscinodon irroratus (Mitt.) Zand., see treatment of Tetracoscinodon Austr2
TETRAPTERUM Hampe
Tetrapterum brachype/ma (C. Mtill.) Broth. Austr1
Tetrapterum cylindricum (Tayl.) Jaeg. Austr1
Tetrapterum lamprocarpum (C. Mtill.) Broth. Am6
Tetrapterum lamprothecium (C. Mtill.) Broth. Am6
[Tetrapterum lilliputanum (C. Mtill. ex Roth) Broth. Am6 =Torte//a lilliputana (C. Mtill. ex Roth) Zand., see treatment of Tortella]
[Tetrapterum recurvirostre (C. Mtill.) Broth. Am5 Am6 = Weissia
diffidentia Zand. (nom . nov. for Phascum recurvirostre C.
Mtill.), see treatment of Weissia]
Tetrapterum sullivanii (C. Mtill.) Broth. Austr1
Tetrapterum tetragonum (Hook.) Andrews Afr4
Tetrrapterum vernicosum (Roth) Broth. Am5 = Trichostomum
exulatum Zand. nom. nov. , see treatment of Trichostomum]
Tetrapterum weymouthii (C. Mtill.) Broth. Austr1 .
TIMMIELLA (De Not.) Limpr.
Timmie/la acaulon (C. Mtill.) Zand., see treatment of Timmiella
Am6
[Timmiella alata Herz. Am5 = Hymenostyliel/a alata (Herz.) Robins.]
Timmie/la anomala (BSG) Limpr. Eur As1 As2 As3 As4 As5 Afr1
Am1Am2
[Timmiella argentinica Broth. Am4 =Timmie/la acaulon (C. Mtill.)
Zand., see treatment of Timmie/la]
[Timmie/la barbula Limpr. = Timmiella barbuloides (Brid.)
Moenk.]
var. minor Schimp. ex Luis. Afr1
Timmie/la barbu/oides (Brid.) Monk. Eur As1 As3 As5 Afr1 Afr2
Am4
var. /ongiseta (Brid.) Wijk & Marg. Eur
Timmie/la brevidens Dix. Afr2
Timmie/la cameruniae Broth. Afr2
Timmie/la cornicu/ata (Wahlenb.) Broth. As1
Timmie/la crassinervis (Hampe) Koch Am1
Timmie/la diminuta (C. Mtill.) Chen As2
Timmie//aflexiseta (Bruch) Limpr. Eur Afr1
[var. vancouveriensis (Broth.) Grout Am1 = Timmiella
crassinervis (Hampe) Koch fide Crum, Steere & Anderson,
Bryologist 68: 431, 1973]
Timmiella grosseserrata Schiffn. As5
[Timmie/la japonica Iwas. As2 =Timmiella anomala (BSG) Limpr.
fide lwatsuki & Noguchi, J. Hattori Bot. Lab. 37: 412, 1973]
Timmie/la pelindaba Magill in Magill & Schelpe, Mem. Bot. Surv.
S. Afr. 43 : 3, 1979 Afr4
[Timmie/la subanomala (Besch.) Broth. Am2 =Timmiel/a anomala
(BSG) Limpr.fide Zander in Sharp et al., Moss Fl. Mex.]
Timmie/la subintegra Dix. As3
Timmie/la umbrosa (C. Mtill.) Broth. Am6
TORTELLA (Lindh.) Limpr.
[Tortella acaulon (C. Mtill.) Broth. Am6 = Timmiel/a acaulon (C.
Mtill.) Zand., see treatment of Timmie/la]
Tortella alpicola Dix. Am1 As3 Oc Ant
Tortella aprica (C. Mull:) Broth. Afr2
[Torre/la arctica (Am.) Crundw. & Nyh., Trans. Brit. Bryol. Soc. 4:
187, 1963 (1964) Am1 As1 As3 =Torre/la tortuosa var. arctica
(Am.) Broth. in Fedch.]
Tortella brotheri (Broth.) Broth. Eur
Tortel/a tetrapteroides (C. MUll.) Broth. Am5
[Tortella bryotropica Zand. in Schultze-Motel & Menzel, Beih.
GENERA, SPECIES AND INFRASPECIFIC TAXA
Nov. Hedw. 88: 21, 1987 Am4
[Tortella caespitosa (Schwaegr.) Limpr. Tortella humilis (Hedw.)
Jenn.]
var. /ongirostris Papp Eur
[Torre/la calycina (Schwaegr.) Dix. As3 Afr4 Am6 Austrl Austr2 =
Barbula calycina Schwaegr. fide Magill, Fl. S. Afr. I. Mosses 1:
241, 1981 (1982)]
[Tarte/la ceylonensis Reisch. ex Dix. As3 =Tarte/la humilis (Hedw.)
Jenn.fide Sollman, Lindbergia 16: 24, 1990]
[Tarte/la cirrhata Broth. Austrl = Trichostomum eckelianum Zand.
nom. nov. see treatment ofTrichostomum]
Tarte/la cirrifolia (Mitt.) Broth. Afrl
Tortella contortifolia (Mitt.) Broth. in Par. Am4
Tortella cryptocarpa (Broth.) Zand. (Astomum), see treatment of Tortel/a Am5
[Tarte/la cylindrica (Brid.) Loeske = Trichostomum tenuirostre
(Hook. & Tayl.) Lindh.]
var. perpapillosa (Ihs.) Nog. As2
[fo. irrigua (Limpr.) Monk. Eur = Oxystegus cylindricus var.
irriguus (Limpr.) Podp., Consp. Muse. Eur. 176, 1954]
Tarte/la cyrtobasis Dix. As3
Tarte/la dakinii Willis Austrl
Tortella densa (Lor. & Mol.) Crundw. & Nyh. Eur
Tarte/la eckendorffii (P. Yarde) Zand. (Hymenostomum), see treatment of Tort ella Afr2
Tarte/la e/kantarensis Ther. & Trab. Afrl
Tarte/la erosodentata Sak. As2
[Tarte/la eutrichostomum (C. Miill.) Broth. Afr4 = Tarte/la humilis
(Hedw.) Jenn. fide Magill, Fl. S. Afr. I. Mosses 1: 256, 1981
(1982)]
Torte//aflavovirens (Bruch) Broth. Eur As5 Afrl Am1
subsp. esterelensis (Roth) Giac. Eur
subsp. limose//a (Stirt.) Podp. Eur
subsp. viridiflava (De Not.) Giac. Eur
var. glareicola (T. Christ.) Crundw. & Nyh. Eur
var. /aevis Luis. Afrl
var. papi/losissima Sergio & Cas. de Puig, Portugaliae Acta Bioi.,
ser. B., Sist. 13: 116, 1981 ("papilosissima") Eur
var. viridiflava (De Not.) Cas. Gil. Eur
fo. brevifolia (Schiffn.) Podp., Consp. Muse. Eur. 178, 1954
(Trichostomumflavovirens var.) Eur
fo. nitidicostatum (Zodda) Podp., Consp. Muse. Eur. 178, 1954
(Trichostomumflavovirens var.) Eur
Tortella flavovirens (Bruch) Broth. x Weissia crispa (Hedw.) Mitt.
fide Nicholson, Rev. Bryol. 37: 23, 1910 Eur
Torte//afragilis (Drumm.) Limpr. Eur Asl As2 As3 Afr4 Ami
var. moravica Podp. Eur
var. setacea (Fam.) Par. Eur
[var. tortelloides (Greene) Zand. & Hoe, Bryologist 82: 84, 1979
Ami As3 Oc Ant Ami As3 Oc Ant (Sarconeurum) Tarte/la
torte//oides (Greene) Robins. in Llano= Tarte/la a/pica/a Dix.]
fo. pa/udosa Amann, Bull. Soc. Yaudoise Sci. Nat. 57: 123, 1929
Eur
fo. riparia (Amann) Podp., Consp. Muse. Eur. 182, 1954 (Tortella
tortuosa var.) Eur
Tortellafragillima P. Yarde Afr2
Tortel/afristedtii (C. Miill.) Broth. Austr2
Tarte/la fruchartii (C. Miill.) Zand. (Phascum), see treatment of Tortel/a Am5 Am6
Tarte/la germainii (C. Miill.) Broth. Am4
Tarte/la goniospora (C. Miill.) Zand. (Barbula), see treatment of Tortel/a Oc
Tarte/la goughii Dix. As3
=
=
327
[Tarte/la grossiretis Bartr. Am5 = Pseudosymblepharis schimperiana (Par.) Crum, see treatment of Pseudosymb/epharis]
[Tarte/la guatema/ensis Bartr. Am2 = Tarte/la tortuosa (Hedw.)
Limpr.fide Zander in Sharp et al., Moss Fl. Mex.]
Tarte/la hosseusii Herz. Am6
Tarte/la humilis (Hedw.) Jenn. Eur As2 As5 Afrl Afr2 Afr3 Afr4
Am1Am2Am3Am4Am5Am60c
Tarte/la inclinata (Hedw. f.) Limpr. Eur As5 Afr1 Afr4 Ami Am4
Austrl
var. brachypoda (Besch.) Par. Afrl
var. /eptotheca (Brid.) Par. Afr4
fo. acuminata (Fam.) Par., Index Bryol. ed. 2, 30, 1906 (Barbu/a
inclinata fo.) Eur
[fo. actutifolia Gand. in Dalla Torre, Moose v. Tirol 212, 1904
Tarte/la inc/inata fo. acuminata (Fam.) Par. fide
Eur
Podp~ra, Consp. Muse. Eur. 179, 1954]
fo. alpina Tosco, Webbia 28: 284, 1973 Eur
fo.fragilifolia Herz., Wiener Bot. Z. 93: 37, 1944 Eur
fo. sa/tans Loeske, Herbarium 62: 130, 1922 Eur
fo. mutica (Latz.) Podp., Consp. Muse. Eur. 179, 1954 (Tarte/la
inclinata var.) Eur
Tarte/la inflexa (Bruch) Broth. Eur As2 As5 Afrl
fo. elata (Glow.) Podp., Consp. Muse. Eur. 177, 1954 Eur
Tarte/la involutifolia Dix. Afr2
Tartella japonica (Besch.) Broth. As2 Am2
Torte//ajugicola (Dub.) Par. Am5
Tarte/la kmetiana Pilous Eur
Tarte/la knightii (Mitt.) Broth. Austrl Austr2
Tarte/la lilliputana (C. Miill. ex Roth) Zand. (Phascum), see treatment of Tarte/la Am6
Tarte/la limbata (Schiffn.) Geh. & Herz. Afrl
Tarte/la lindmaniana Broth. Am5
Tarte/la linearis (Web. & Mohr) Zand. (Barbula), see treatment of
Tarte/la Am3 Am5
[Tarte/la mollissima Broth. ex Bartr. Ami Am2 Am3 Am5
Trichostomum mo//issimum (Broth. ex Bartr.) Crum, Bryologist
72: 245, 1969 = Psuedosymblepharis schimperiana (Par.) Crum
fide Zander in Sharp et al., Moss A. Mex.]
Tortella mooreae Sainsb. Austr2
Tarte/la nitida (Lindh.) Broth. Eur As5 Afrl
var. irrigata Winter, Hedwigia 55: 82, 1914 Afrl
var. media (Boul.) Corb. Eur Afrl
var. obtusa (Boul.) Jelenc Eur Afr1
var. subtortuosa (Boul.) Jelenc Eur Afr1
fo. alpigena Grom, Acta Bot. Croat. 26-27: 249, 1967-68 [1969]
Eur
fo. brachyphy//a Latz., Bot. Centralbl. Beih. 48: 476, 1931 Eur
fo. media (Boul.) Podp., Consp. Muse. Eur. 177, 1954 (Trichostomum nitidum var.) Eur
fo. obtusa (Boul.) Podp., Consp. Muse. Eur. 177, 1954 (Trichostomum nitidum var.) Eur
Torre/la novae-valesiae Broth. Austrl
[Tarte/la obtusifolia Dix. Afr2 = Barbula umtaliensis Magill nom.
nov. fide Magill in Magill & Schelpe, Mem. Bot. Surv. S. Afr.
43: 5, 1979]
[Tortella opaca Dix. Afr2 = Weissia opaca (Dix.) Magill in Magill
& Schelpe, Mem. Bot. Surv. S. Afr. 43: 7, 1979]
Torte /Ia perrufula (C. Miill.) Broth. Am6
[Tarte/la petrieana Sim Afr4 = Tarte/la humilis (Hedw.) Jenn. fide
Magill, Fl. S. Afr. l. Mosses 1: 256, 1981 (1982)]
Torre/la pileomayica Herz. Am4
Torte /Ia pseudocaespirosa (C. Miill.) Broth. Am6
var. brachybasis (C. Miill.) Par. Am6
=
=
328
GENERA OF THE POTTIACEAE
var. pungens (C. Mull.) Par. Am6
[Tortella richardsii Bartr. Am2 Am3 = Pseudosymblepharis
schimperiana (Par.) Crum fide Zander in Sharp et al., Moss A.
Mex.]
Tortella rigens Alb. Eur Am1
Tortella rubripes (Mitt.) Broth. Austr2
[Tortella rufiseta (C. Miill.) Broth. Afr4 = Tortella xanthocarpa (C.
Mill!.) Broth. fide Magill, Fl. So. Afr.l. Mosses 1: 255, 1982]
[Torrella santosii Bartr. Am2 = Streptocalypta santosii (Bartr.) Zand. ,
Lindbergia 8: 165, 1982 (1983)]
Tortella satoi Sak. As2
Tortella simplex Robins. in Robinson, Holm-Nielsen & L~jtnant,
Lindbergia 4: 109, 1977 [1978] Am4
Tortella smithii Towns., Trans. Brit. Bryol. Soc. 6: 791, 1969 Afr2
Tortella somaliae (C. Mull.) Zand. (Hyophila), see treatment of Tortel/a Afr2
[Tortella spathulata Sak. As2 =Trichostomum brachydontium Bruch
fide Saito, J. Hattori Bot. Lab. 39: 432, 1975]
Tortella subflavovirens Broth. & Watts Austr1
[Tortella subfragilis Crum & Steere Am3 = Pseudosymblepharis
schimperiana (Par.) Crum fide Zander in Sharp et al., Moss Fl.
Mex.]
[Tortella syrrhopodontoides Dix. Afr2 = Pseudosymblepharis
syrrhopodontoides (Dix.) Zand., see treatment of Pseudosymblepharis]
[Tortella theriotii Broth. & P. Yarde Afr2 = Tortella humilis (Hedw.)
Jenn .fide Sollman, Lindbergia 16: 24, 1990]
[var. angustata Dix. & P. Yarde Afr2 = Tortella humilis (Hedw.)
Jenn. fide Sollman, Lindbergia 16: 24, 1990]
[Tortella tortelloides (Greene) Robins. in Llano, Antarctic Terrestrial
Biology, Antarctic Res. Ser. 20: 170, 1972 Am1 As3 Oc Ant[=
Tortella fragilis var. tortelloides (Greene) Zand. & Hoe,
Bryologist 82: 84, 1979 = Tortella alpicola Dix.]
Tortella tortuosa (Hedw.) Limpr. Eur As1 As2 As3 As5 Afr1 Am1
As2Am6
subsp.fasciculata Culm Eur
var. angustifolia (Jur.) Limpr. Eur
var. arctica (Am.) Broth. in Fedch. As1 As3 Am1
var. brevifolia Breidl. in Limpr. Eur
var. brevirostris Papp Eur
var. cucullata Amann Eur
var. curvula (Hartm.) Hag. Eur
var. dicranoidea (Ferg.) Limpr. Eur
var. fleischeri (Bauer) Latz. Eur
[var. gracilescens (Zett.) Podp. Eur = Tortella tortuosa var. tenella
(Walth. & Mol.) Limpr. fide Podpl!ra, Consp. Muse. Eur. 181,
1954]
var. hamifolia Herz. Eur
var. longifolia (Ren.) Par. Eur
var. nitida Pilous Eur
var. rauligera Latz. Eur
var. rigida (Boul.) Limpr. Eur
var. riparia Amann Eur
var. robusta (Pfeff.) Limpr. Eur
var. rotaeana (De Not.) Limpr. Eur
var. setifera Velen. Eur
var. tenella (Walth. & Mol.) Limpr. Eur
var. viridis (Delogn.) Wild. Eur
fo. alpina Giac., Ann. Bryol.11: 72, 1938 Eur
fo. alpina (Boul.) Podp., Consp. Muse. Eur. 180, 1954 hom. illeg.
(Barbula tortuosa fo.) Eur
fo. atrovirens Latz., Hedwigia 66: 138, 1926 Eur
[fo. brevifolia (Breidl. in Limpr.) Herz. in Riibel, Bot. Jahrb. Syst.
47: 476, 1912 Eur = Torre/la tortuosa var. brevifolia Breid. in
Limpr. fide Podpl!ra, Consp. Muse. Eur. 180, 1954)
fo. dentata (Fam.) Podp., Consp. Muse. Eur. 180, 1954 (Barbula
tortuosa fo.) Eur
[fo. gracilescens (Zett.) Par., Index Bryol. ed 2, 32, 1906 (Barbuta tortuosa var.) Eur =Tortella tortuosa var. tenella (Walth. &
Mol.) Limpr.fide Podpl!ra, Consp. Eur. 181 , 1954]
fo. latifolia Herz., Weiner Bot. Z. 93: 38, 1944 Eur
fo. spinidens (Roth ex Zodd.) Bauer, Fedde Repert. 50: 341, 1941
(Tortella) Eur
fo. subrecurva Latz., Magyar Bot. Lapok 33: 174, 1934 Eur
fo. supera(pina Podp., Consp. Muse. Eur. 181, 1954 nom. illeg.
Eur
Tortella undulatifolia Dix., Anniv. Vol. Bot. Gard. Calcutta 180,
1942 As3
Tortella vernicosa (Ren. & Card.) Broth. Afr3
Tortella walkeri (Broth.) Zand. (Hyophila), see treatment of TortellaAs3
Tortella xanthocarpa (C. Miill.) Broth. Afr4
TORTULA Hedw.
[Tortula abranchesii Luis. Eur = Tortula caninervis var.
abranchesii (Luis.) Kramer, Bryophyt. Biblioth. 21: 108, 1980
= Syntrichia caninervis var. abranchesii (Luis.) Zand., see
treatment of Syntrichia]
[Tortula abruptinervis Dix. Austr2 = Syntrichia pygmaea (Dus.)
Zand. see treatment of Syntrichia]
Tortula abyssinica De Not. Afr2
[Tortula aciphyl/a (B.&S. in BSG) Hartm. = Tortula norvegica
Web.) Lindb. fide Kramer, Bryoph. Biblioth. 21 : 128, 1980 =
Syntrichia norvegica Web.]
var. mucronata (Sendtn.) Limpr. Eur
[Tortula acletoi Robins., Phytologia 12: 390, 1971 Am4 = Hennediella acletoi (Robins.) Zand., see treatment of Hennediella]
[Tortula aculeata (Wils.) Mitt. Am4 = Syntrichia aculeata (Wils.)
Zand., see treatment of Syntrichia]
Tortula aculeonervis (C. Mull.) Broth. Am6
[Tortu/a adusta (Mitt.) Mitt. Austr1 = Barbula hornschuchiana
Schultz fide Catcheside, Mosses S. Austr. 178, 1980] =Pseudocrossidium hornschuchianum (Schultz) Zand., Phytologia 44:
205, 1979)
[Tortula aestiva (Hedw.) P. Beauv. = Tortula muralis var. aestiva
Hedw.]
[fo. terrestris Podp., Sbom. Klubu Pfir. v. Bmo 5: 13, 1923 Eur =
Tortula muralis fo. terrestris (Podp.) Podp., Consp. Muse. Eur.
248, 1954]
Tortula afanassievii Laz. As2
[Tortula afroruralis (C. Mull.) Broth.Afr4 = Tortula leucostega (C.
Mull.) Broth. fide Kramer, J. I-fattori Bot. Lab. 65: 84, 1988 =
Syntrichia leucostega (C. Miill.) Zand., see treatment of Syntrichia]
[Tortula alpestris Dix. in Herz. Afr2 = Syntrichia alpestris (Dix. in
Herz.) Zand. see treatment of Syntrichia]
[Tortula alpina (BSG) Bruch in Breut. = Tortula sinensis (C. Mull.)
Broth. = Syntrichia sinensis (C. Mull.) Ochyra, Frag. Florist.
Geobot. 37: 212, 1992)
[var. inermis (Mild.) De Not. Eur As2 Afr1 Ami Oc = Tortula
fragilis Tayl. fide Corley et al., J. Bryol. 11 : 619, 1981 (1982)]
= Syntrichia fragilis (Tayl.) Ochyra, Fragm. Florist. Geobot.
37: 212, 1992]
[fo. pagorum (Milde) Squiv., Rev. Bryol. Licbenol. 30: 215, 1962
(1963) Eur Ami Am2 Austr1 = Tortula pagorum (Milde) De
Not. = Syntrichia pagorum (Milde) Amann, see treatment of
329
GENERA, SPECIES AND INFRASPECIFIC TAXA
Syntrichia]
Tortula altipes (Broth.) Zand. (Desmatodon), see treatment of Tortula
As1
[Tortula amphidiacea (C. Mull.) Broth. Afr2 Am1 Am2 Am3 Syntrichia amphidiacea (C. MUll.) Zand., see treatment of Syntrichia]
[Tortula ammonsiana Crum & Anders., Bryologist 82: 469, 1979
Am1 Afr4 Syntrichia ammonsiana (Crum & Anders.) Ochyra]
Tortula amphidiifolia (C. Mull.) Broth. Am6
[Tortula amplexa (Lesq.) Steere in Grout Eur Am1 = Syntrichia
amplexa (Lesq.) Zand., see treatment of Syntrichia]
Tortula ampliretis Crundwell & Long, J. Bryol. 10: 104, 1978 Afr1
Tortula anacamptophylla (C. Mull.) Broth. AmS Am6 = Tortula
robusta Hook. & Grev. fide Lightowlers, Brit. Antarct. Surv.
Bull. 67: 67, 198S = Syntrichia robusta (Hook. & Grev.) Zand.
see treatment of Syntrichia]
Syntrichia
[Tortula anderssonii Angstr. Am6 Austr2 Ant
anderssonii (Aongstr.) Zand., see treatment of Syntrichia]
[var. fagicola (C. Mull.) Card. Am4 Syntrichia anderssonii var.
fagicola (C. Mull.) Zand., see treatment of Syntrichia]
[Tortula angustifolia (Herz.) Herz. nom. illeg. Am4
Tortula
herzogii Zand. nom. nov. Bryologist 82: 631, 1979 =Hennediella
angustifolia (Herz.) Zand., see treatment of Hennediella]
[Tortula andicola Mont. Am2 Am4
Syntrichia andicola (Mont.)
Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
[Tortula androgyna (C. Mull.) Broth. Austr1 = Tortula antarctica
(Hampe in C. MUll.) Wils. in Hook. f. fide Kramer, J. Hattori Bot.
Lab. 6S: 84, 1988 = Syntrichia antarctica (C. Mull. & Hampe)
Zand., see treatment of Syntrichia]
[Tortula antarctica (Hampe in C. Mull.) Wils. in Hook. f. Am6 (good
species fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988) Syntrichia antarctica (Hampe in C. MUll.) Zand., see treatment of Syntrichia]
Tortula appressa Mitt. Am6
[Tortula arenae (Besch.) Broth. Am6 Ant
Hennediella arenae
(Besch.) Zand., see treatment of Hennediella]
[subsp. petriei (Broth.) Lightowlers, J. Bryol. 13: 371, 1985 (Tortuta petriei) Austr2 = Hennediella arenae var. petriei (Broth. ex
Beck.) Zand., see treatment of Hennediella]
Tortula argentinica (Broth.) Zand. (Desmatodon), see treatment of
Tortula Am6
Tortu/a astoma Schiffn. AsS
Tortula atherodes Zand. (nom. nov. for Phascum cuspidatum Schreb.
ex Hedw.), see treatment of Tortula Eur As1 As2 AsS Afr1 Ami
Am2Am4
var. affinis (Nees & Hornsch.) Zand. (Phascum affine), see treatment of Tortula Eur
var. atherodes var. arcuata (Herrnstadt & Heyn) Zand. (Phascum
cuspidatum var.) Eur
var. curviseta (Dicks.) Zand. (Phascum curvisetum), see treatment
of Tortula Eur
var. diaphora (Hag.) Zand. (Phascum acaulon var.), see treatment
of Tortula Eur
var. elatum (Brid.) Zand. (Phascum elatum), see treatment of Tortula Eur
var. intertexta (Brid.) Zand. (Phascum intertextum), see treatment
of Tortula Eur
var. mitraeformis (Limpr.) Zand. (Phascum cuspidatum var.), see
treatment of Tortula Eur
var. papillosa (Lindh.) Zand. (Phascum papillosum), see treatment
of Tortula Eur
var. pilifera (Hedw.) Zand. (Phascum piliferum), see treatment of
Tortula Eur As1 AsS Afr2 Am1
=
=
=
=
=
=
=
=
var. retortifolia (Guerra & Ros in Guerra, Jimenez, Ros &
Carri6n) Zand. (Phascum cuspidatum var.), see treatment of
Tortula Eur
var. schreberiana (Dicks.) Zand. (Phascum schreberianum), see
treatment of Tortula Eur Am1
Tortula atrata Ther. Am4
Tortula atrovirens (Sm.) Lindh. Eur As4 AsS Am1 Am2 Am3 Am4
Am6 Austr1 Austr2 Oc Afrl Afr2 Afr3 Afr4
var. brevifolia Ther. Am6
var. edentula (BSG) Schimp. Eur Afr4
var. gasilienii (Vent.) Limpr. Eur
var. leucodonta (Corb.) Zand. (Barbula atrovirens var.), see treatment ofTortu/a Eur
var. subrevolvens Amann Eur
[Tortula aurea Bartr. Am1 Am2 Barbula aurea (Bartr.) Zand. in
Zand. & Steere, Bryologist 81 : 466, 1978 = Pseudocrossidium
aureum (Bartr.) Zand., Phytologia 44: 207, 1979 = Barbula
crinita Schultz fide Sollman, Lindbergia 16: 22, 1990
Pseudocrossidium crinitum (Schultz) Zand., see treatment of
Pseudocrossidium and cf. Sollman, Lindbergia 16: 22, 1990]
[Tortula austroafricana Kramer, J. Hattori Bot. Lab. 6S: 92, 1988
Afr4 Syntrichia austroafricana (Kramer) Zand., see treatment
of Syntrichia]
Tortula austroruralis (C. Mull.) Broth. Am6 Austr1
Tortula baetica (Casas & Oliva) Guerra & Ros in Guerra, Ros &
Carri6n, J. Bryol. 17: 281, 1992 (Tortula muralis var.) Eur
[Tortula baileyi Broth. Austr1 = Syntrichia baileyi (Broth.) Zand.,
see treatment of Syntrichia]
[Tortula bartramii Steere in Grout Am1 Am2 = Syntrichia
bartramii (Steere in Grout) Zand., see treatment of Tortula]
Tortula bauriana Warnst. ex Baur Eur
[Tortula bealeyensis R. Br. ter Austr2 =Tortula anderssoniiAngstr.
fide Lightowlers, Brit. Antarct. Surv. Bull. 67: 4S, 198S & J.
Bryol. 13: 369, 198S Syntrichia anderssonii (Aongstr.) Zand.,
see treatment of Syntrichia]
Tortula berthoana Ther. Am6
[Tortula bipedicel/ata Britt. Am4 Syntrichia bipedicellata (Britt.)
Zand., see treatment of Syntrichia]
[Tortula bistratosa Flow. Ami = Tortula caninervis (Mitt.) Broth.
fide Corley eta!., J. Bryol. 11: 619, 1981 (1982)] = Syntrichia
caninervis Mitt.]
[Tortula bizotii Laz., Vopr. Evol. Biogeogr. Genet. Sel. 14S, 1960
nom. inval. typon . non cit. As5 = Tortula virescens subsp.
bizotii (Laz.) Kramer, Bryophyt. Biblioth. 21: 102, 1980 comb.
inval. basion. inval.]
Tortula bogosica (C. Mull.) Zand. (Desmatodon), see treatment of
Tortula Afr1 Afr2 Afr4
[Tortula bogotensis (Hampe) Mitt. Am4 (l.M. citation of Am3 is
incorrect as Mitten's [1869] "Guadelupe" refers to an area in
Syntrichia bogotensis
Colombia, not the West Indies)
(Hampe) Mitt., see treatment of Syntrichia]
[Tortula bolanderi (Lesq. & James) Howe Eur Afr1 Am1 = Syntrichia bolanderi (Lesq. & James) Zand., see treatment of SyntrichiaJ
[Tortula brachychaete Dus. Am6 = Willia brachychaete (Dus.)
Zand., see treatment of Willia]
[Tortula brachyclada Card. Am6 = Syntrichia brachyclada (Card.)
Zand., see treatment of Syntrichia]
[Tortula brachydontia (Bruch in F. Mull.) Mitt., J. Linn. Soc. Bot.
12: 148, 1869 Am4 = Trichostomum brachydontium Bruch in
F. MUll.]
[Tortula brachypelma Des. Am6 = Tortula magel/anica Mont. in
Gay fide Kramer, J. Hattori Bot. Lab. 6S: Ill, 1988 = Syntric-
=
=
=
=
=
=
330
GENERA OF THE POTTIACEAE
hia magellanica (Mont.) Zand.]
[Tortula brachytricha (C. MUll.) Broth. Austr1 = Tortula antarctica
(Hampe in C. MUll.) Wils. in Hook. f. fide Kramer, J. Hattori Bot.
Lab. 65: 89, 1988 = Syntrichia antarctica (Hampe in C. MUll.)
Zand.]
[Tortula brandisii (C. MUll.) Broth. As3 = Syntrichia brandisii (C.
Miill.) Zand., see treatment of Syntrichia]
Tortula brevimucronata (C. MUll.) Broth. Afr4
Tortula brevipes (Lesq.) Broth. Am1 Am2
[Tortula brevis Whitehouse & Newton, J. Bryol. 15: 83, 1988 Eur =
Hennediella brevis (Whitehouse & Newton) Blockeel, J. Bryol.
16: 191, 1991]
Tortula breviseta Mont. Am6
[Tortula brevisetacea (F. MUll.) Ther. Austr1 = Syntrichia
brevisetacea (F. MUll.) Zand., see treatment of Syntrichia]
Tortula brevissima Schiffn. Eur As5
Tortula brunnea (C. MUll.) Broth. Am4
Tortula brunnea Broth. & Watts hom. il/eg. Austr1
Tortula buchtienii Herz. Am4
Tortula bullata Herz. Am6 hom. il/eg.
Tortula bushii (Card. & Th~r.) Steere nom. inval. in syn. Am1
Tortula buyssonii (Philib.) Broth. Eur
[Tortula cabulica J. Frohl., Mitt. Thiir. Bot. Ges. 1: 61, 1955 As5 =
Trichostomopsis aaronis (Lor.) Agnew & Towns., Israel J. Bot.
19: 258, 1970 = Didymodon aaronis (Lor.) Guerra in Guerra &
Ros, Cryptogamie Bryol. Lichenol. 8: 55, 1987]
[Tortula cainii Crum & Anders. Am1 = Syntrichia cainii (Crum &
Anders.) Zand., see treatment of Syntrichia]
[Tortula calcico/ens Kramer, Bryophyt. Biblioth. 21: 90, 1980 (Tortula ca/cico/a Greebe hom. illeg., Tortula ruralis subsp. calcico/a
(Amann) Giac.) Eur (= Tortula ruralis (Hedw.) Gaertn., Meyer &
Scherb. fide Corley et al., J. Bryol. 11: 619, 1981 [1982]) = Syntrichia ca/cico/a Amann]
Tortula ca/ifornica Bartr. Ami Am2
[Tortula ca/obo/ax (C. MUll.) Broth. Afr4 = Willia calobolax (C.
MUll.) Lightowlers, J. Bryol. 13: 370, 1985]
[var. angustinervia (C. Miill.) Par. Afr4 = Willia calobolax var.
angustinervia (C. Miill.) Lightowlers, J. Bryol. 18: 370, 1985]
[Tortula campestris Dus. Am6 = Syntrichia campestris (Dus.) Zand.,
see treatment of Syntrichia]
Tortula canescens Mont. Eur As1 As5 Afr1
var. /ongipila (Carb.) Jelenc Afrl
[Tortula caninervis (Mitt.) Broth. Eur As2 Ami = Syntrichia
caninervis Mitt.]
[subsp. spuria (Amann) Kramer, Bryophyt. Biblioth. 21: 106, 1980
(Tortula) Eur = Syntrichia caninervis var. spuria (Amann) Zand.,
see treatment of Syntrichia]
[var. abranchesii (Luis.) Kramer, Bryophyt. Biblioth. 21: 108, 1980
(Tortula) Eur = Syntrichia caninervis var. abranchesii (Luis.)
Zand.]
[var. gypsophila (G. Roth) Kramer, Bryophyt. Biblioth. 21: 108,
1980 (Tortula ruralis var.) Eur = Syntrichia caninervis var.
gypsophila (Roth.) Ochyra, Fragm. Florist. Geobot. 37: 212,
1992]
Tortula capillaris (Chen) Zand. (Desmatodon), see treatment of Tortula As2
Tortula cardotii Ther. & Nav. Afr2
[Tortula carinata Gill. ex Grev. Am6 = Pseudocrossidium carinatum
(Gill. ex Grev.) Zand., see treatment of Pseudocrossidium]
[Tortula caroliniana Andrews Am1 Am2 Am3 = Tortula
amphidiacea (C. MUll.) Broth. fide Mishler in Sharp et al., Moss
Fl. Mex. = Syntrichia amphideacea (C. Miill.) Zand., see treatment of Syntrichia]
[Tortula cava/Iii Negri Afr2 = Syntrichia cavallii (Negri) Ochyra,
Fragm. Florist. Geobot. 37: 212, 1992]
Tortula cernua (Hueb.) Lindh. Eur As1 As2 Am1
var. xanthopus (Kindb.) Zand. (Desmatodon cernuus var.), see
treatment of Tortula Am 1
Tortula characodonta (C. MUll.) Broth. Am6
[Tortula chisosa Magill, Delg. & Stark, Ann. Missouri Bot. Gard.
70: 200, 1983 Am1 Am2 Afr4 = Syntrichia chisosa (Magill,
Delg. & Stark) Zand., see treatment of Syntrichia]
Tortula chrysopila (C. MUll.) Par. Am6
Tortula chubutensis Dus. Am6
Tortula chubutensis Broth. Am6 hom. illeg.
(nom. nov. for Desmatodon
Tortula chungtienia Zand.
yuennanensis Broth.), see treatment of Tortula As2
[Tortula ciliata Broth. Am4 = Syntrichia ciliata (Broth.) Zand.]
Tortula coch/earifolia P. Yarde Afr2
[Tortula conferta Bartr. Ant (= Tortula grossiretis Card. fide Robins., Antarct. Res. Ser. 20: 172, 1972) = Tortula princeps var.
conferta (Bartr.) Lightowlers, Brit. Antarct. Surv. Bull. 67: 61,
1985 (= Tortula princeps De Not. fide Lightowlers, J. Bryol.
14: 290, 1986 [1987] = Syntrichia princeps (De Not.) Mitt.)=
Syntrichia conferta (Bartr.) Zand., see treatment of Syntrichia
Ant]
[Tortula conica Turton, Gen. Syst. Nat. Veg. Kingd. 2: 1720, 1806
(Barbula conica Brid.) Am1 = Barbula agraria Hedw. fide
Brid., Bryol. Univ. 1: 532, 1827]
Tortula contorta (C. MUll.) Mont. in Gay Am6
[Tortula convoluta (Hedw.) Gaertn., Meyer & Scherb. = Barbula
convoluta Hedw.]
var. major De Not. Eur
[Tortula costesii Ther. Am6 = Syntrichia costesii (Ther.) Zand., see
treatment of Syntrichia]
Tortula crawfordii (Par.) Watts in Watts & White!. Austr1
Tortula crenata Mitt. Am4
[Tortula crenulata Warnst = Tortula subulata var. serrulata
Warnst.]
var. /atifolia (Warnst.) Warns!. Eur
Tortula cucullifolia Frohl., Mitt. Thiirin. Bot. Ges. 1(2-3): 62, 1955
Eur
Tortula cuneifolia (Dicks.) Turn. Eur As5 Afr1 [for discussion of
Dickson as authority see Karttunen, Taxon 37: 156-157, 1988]
subsp.freibergii (Dix. & Loeske) Giac. Eur
var. blissii Zand. var. nov., see treatment of Tortula Am1
var. luteomarginata Hohn. Eur
var. marginata Fleisch. Eur
var. piligera Latz. Eur
fo. brevifolia Fleisch. in E. Bauer, Muse. Eur. Exs. n. 178, 1906
Eur
[Tortula cuspidata (Schultz) Chev. Eur Afr1 = Barbula unguiculata
fo. cuspidata (Schultz) Monk., Laub. Eur. 286, 1927]
var. elongata Chev. Eur
[Tortula cuspidata Hook. f. & Wils. Austrl = Tortula
anderssoniiAngstr. fide Kramer, J. Hattori Bot. Lab. 65: 84,
1988 = Syntrichia anderssonii (Aongstr.) Zand., see treatment
of Syntrichia J
Tortula cuspidatissima (C. MUll.) Broth. Afr2
Tortula deciduidentata (Sharp & lwats.) Zand. (Crumia), see treatment of Tortula Am1
Tortula demawendica Schiffn. As5
[Tortula densifolia (Hook. f. & Wils.) Hook. f. & Wils. Am6 Ant=
Hennediella densifolia (Hook. f. & Wils.) Zand., see treatment
of Hennediella]
[Tortula denticulata (Wils. in Mitt.) Mitt. Am3 Am4 = Hennediella
331
GENERA, SPECIES AND INFRASPECIFIC TAXA
denticulata (Wils. in Mitt.) Zand., see treatment of Hennediella]
[Tortula deserta (C. Miill.) Broth. Afr4 = Desmarodon
longipedunculatus (C. Miill.) Magill, Fl. S. Afr. I. Mosses I: 210,
1981 (1982) = Hennediella /ongipedunculata (C. Miill.) Zand.,
see treatment of Hennediella]
[Tortula desertorum Broth. Eur As! As3 As5 Ami = Tortula
caninervis (Mitt.) Broth. fide Corley et al., J. Bryol. 11: 619, 1981
(1982)] = Syntrichia caninervis Mitt.]
[Tortu/a dicksoniana (Schultz) Podp. = Tortu/a cuneifolia (Dicks.)
Tum.]
[fo. piligera (Latz.) Podp., Consp. Muse. Eur. 245, 1954 (Tortu/a
cuneifolia var.) Eur = Tortula cunelfolia var. pi/igera Latz.]
[fo. brevifolia (Fleisch. in Bauer) Podp., Consp. Muse. Eur. 245,
1954 Eur = Tortula cuneifolia fo. brevifolia Fleisch. in Bauer]
[Tortula didymodontoides Broth. in Dryg. Afr4 = Syntrichia didymodontoides (Broth. in Dryg.) Zand., see treatment of Syntrichia]
[Tortula domingensis Ther. Am3 = Brachymenium domingensis
(Ther.) Zand. ·see Excluded Taxa]
[Tortula eckeliae Zand., Bryologist 88 : 354, 1985 (1986) Am2 =Hennediella heteroloma var. eckeliae (Zand.) Zand., see treatment of
Hennediella]
[Tortu/a e/ongata (Wils. in Mitt.) Mitt. hom. il/eg. Am2 Am4 Morinia ehrenbergiana var. elongata (Wils . in Mitt.) Zand.,
Bryologist 81 : 557, 1978 (1979) = Mironia ehrenbergiana var.
e/ongata (Wils. in Mitt.) Zand., see treatment of Mironia]
Tortula entosthodontacea (Card. & Dix.) Zander (Hyophi/opsis), see
treatment of Tortula As3
Tortula epi/osa Broth. ex Dus. Am6
var. pi/ifera Ther. Am6
[Tortula erythrodonta (Tayl.) Wils. = Streptopogon erythrodontus
(Tayl.) Wils.]
[var. c/avipes Spruce Am4
Streptopogon clavipes (Spruce)
Spruce ex Mitt.]
[Tortu/a erythroneura (C. Miill.) Broth. Afr4 = Tortu/a antarctica
(Hampe in C. Miill.) Wils. in Hook. f. fide Kramer, J. Hattori Bot.
Lab. 65: 89, 1988 Syntrichia antarctica (Hampe in C. Miill.)
Zand., see treatment of Syntrichia]
Barbula
[Tortula eubryum (C. Miill.) Broth. in Engl. Afr2 Afr4
eubryum C. Miill. fide Magill, Fl. S. Afr. I. Mosses I: 245, 1981
(1982)]
Tortula euryphylla Zand. (nom. nov. for Dicranum latlfolium
Hedw.), see treatment of Tortula Eur As! As2 As3 As5 Afrl
Ami
subsp. brevifolia (Kindb.) Zand. (Tortula latifolia (Hedw.) Lindb.
subsp. brevifolia Kindb.), see treatment of Tortula Eur
var. euca/yptrata (Lindb.) Zand. (Tortula) Eur
var. flavescens (Brid.) Zand . (Dicranum latifolium var.), see treatment of Tortu/a Eur
var. spelaea (Amann) Zand. (Desmatodon), see treatment of Tortuta Eur
var. subobliqua (Lindb.) Zand. (Desmatodon latifolius var.), see
treatment of Tortula Eur
Tortula evanescens Broth. Austrl
[Tortula exce/sa Card. Ant = Tortu/a filaris (C. Miill. in Newm.)
Broth. fide Lightowlers, Brit. Antarct. Surv. Bull. 67: 49, 1985 =
Syntrichia filaris (C. Miill. in Neum.) Zand., see treatment of
Tortula]
[Tortulafallax (Hedw.) Schrad. ex Tum.= Barbulafallax Hedw.]
var. brevifolia Chev. Eur
Tortulafelipponei Ther. in Felipp. Am6
[Tortula ferganensis Laz. As1
Torrula handelii var. ferganensis
(Laz.) Kramer, Bryologist 81: 385, 1978
Syntrichia handelii
var. ferganensis (Laz.) Ochyra, Fragm. Florist. Geobot. 37: 212,
=
=
=
=
=
=
1992 (Torrula) As!]
Tortu/a ferruginea Bartr., Rev. Bryol. Lichenol. 33: 325, 1964-65
[1965] Am4
[Tortula filaris (C. Miill. in Neum.) Broth. Am6 Ant = Syntrichia
filaris (C. Miill. in Neum.) Zand., see treatment of Tortula]
[Tortula flagellaris (Schimp.) Mont. in Gay Am6 = Syntrichia
flagellaris (Schimp.) Zand., see treatment of Syntrichia]
[var. densiretis Ther. Am6 Syntrichiaflagellaris var. densiretis
(Ther.) Zand., see treatment of Syntrichia]
[Tortula flavinervis Dix. Austr1 Austr2 = Barbu/a crinita Schultz
fide Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
[var. gigantea Dix. & Sainsb. Austr2 = Barbula crinita Schultz
fide Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
[var. obscura Dix. Austr2 = Barbula crinita Schultz fide Weber,
Lindbergia 1: 216, 1972 = Pseudocrossidium crinitum (Schultz)
Zand., see treatment of Pseudocrossidium]
[var. parviretis Sainsb. Austrl = Barbula crinita Schultz fide
Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium crinitum
(Schultz) Zand., see treatment of Pseudocrossidium]
Tortu/aflavipes Broth. hom . il/eg. Am6
[Torru/a flexomarginata (C. Miill. & Hampe) Mitt. Austr1 = Tortuta antarctica (Hampe in C. Miill.) Wils. in Hook. f. fide
Kramer, J. Hattori Bot. Lab. 65: 84, 1988 = Syntrichia
antarctica (C. Miill. & Hampe) Zand., see treatment of Syntrichia]
[Tortula flexuosa Brid. Am4 = Macromitrium sp. fide C. Miill.,
Syn. 1(5): 645, 1849]
[Tortula flexuosa Hook. hom. il/eg. Afr4 = Barbula calycina
Schwaegr. fide Magill, Fl. S. Afr. I. Mosses I: 240, 1981
(1982)]
[Tortula fontana (C. Miill.) Broth. Am6 Ant= Syntrichia fontana
(C. Miill. in Neum.) Zand., see treatment of Syntrichia]
[Tortu/a fragilis Tayl. Ami Am2 Am3 Am4 Syntrichia fragi/is
(Tayl.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992 (Tortula)
Ami Am2 Am3 Am4 As3 Afr3 Afr4]
Tortulafragillima Herz. Am4
Tortula fuegiana (Mitt.) Mitt. (= Tortula princeps var. magellanica
(Mont. in Gay) Lightowlers, Brit. Antarct. Surv. Bull. 67: 61,
1985 = Syntrichia magellanica (Mont. in Gay) Zand., see treatment of Syntrichia) a good species fide cf. Kramer, J.' Hattori
Bot. Lab. 65: 84, 1988 Afr4 Am6 Ant
[Tortula fuscomucronata (C. Miill.) Broth. Afr2 = Pseudocrossidium replicatum C. Miill. fide Townsend, Lindbergia 10: 178,
1985]
[Torru/a fuscoviridis Card. Am6
Syntrichia fuscoviridis (Car.)
Zand., see treatment of Syntrichia]
Syntrichia
[Tortula geheebiaeopsis (C. Miill.) Broth. Afr4 Ant
geheebiaeopsis (C. Miill.) Zand., see treatment of Synrrichia]
[Tortula g/acialis (Kunze ex C. Miill.) Bomt. in Gay Am4 Am6
Syntrichia g/acialis (Kunze ex C. Miill.) Zand., see treatment of
Syntrichia]
Tortula goudotii (Hampe) Mitt. Am4
var. boliviano Broth. in Herz. Am4
[Tortula graeffii (Warnst.) Warnst. = Syntrichia subu/ata fo.
graeffii (Warnst.) Podp., Consp. Muse. Eur. 250, 1954 =Tortuta subulata var. graeffii Warnst.]
[var. angustifolia Warns!. Eur Tortula subu/ata fo. angustifolia
(Wanst.) Sav.-Ljub., Novosti Sis!. Niz. Rast. 6: 248, 1969
(1970)]
var. /atifo/ia Warns!.
fo. denticulata Warns!., Hedwigia 52: 77, 1912 Eur
=
=
=
=
=
=
GENERA OF THE POTI'IACEAE
332
=
[Tortula gregaria Mitt. As 3 Am1 Am2
Barbula indica var.
gregaria (Mitt.) Zand.fide Zander, Phytologia 44: 185, 1979]
Tortula grandiretis Broth. Eur As1
[Tortula gromschii Ther. Am6 = Syntrichia gromschii (Ther.) Zand.,
see treatment of Syntrichia]
[Tortula grossiretis Card. Am6 Ant = Tortula princeps De Not. fide
Lightowlers, Brit. Antarct. Surv. Bull. 67: 58, 1985) Syntrichia
princeps (De Not.) Mitt.]
[var. atrata Card. Am6 = Tortula princeps De Not. fide
Lightowlers, Brit. Antarct. Surv. Bull. 67: 58, 1985 Syntrichia
princeps (De Not.) Mitt.]
[Tortula guatemalensis Bartr. Am2 = Tortula bogotensis (Hampe)
Mitt. fide Mishler in Sharp eta!., Moss Fl. Mex.]
Tortula guepinii (B.&S. in BSG) Broth. Eur Am1 Am2
[Tortula handelii Schiffn. Eur AsS = Syntrichia handelii (Schiffn.)
Bach.]
[var. ferganensis (Laz.) Kramer, Bryologist 81: 385, 1978 As1 =
Syntrichia handelii var. ferganensis (Laz.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992 (Tortula) As1]
[Tortula hellenica Schiffn. & Baumg. Eur =Didymodon australasiae
(Hook. & Grev.) Zand.fide Sollman, Lindbergia 10: 54, 1984]
[var. brevifolia Schiffn. & Baumg. Eur = Didymodon australasiae
(Hook. & Grev.) Zand. fide Sollman, Lindbergia 10: 54, 1984]
[Tortula herzogii Zand. Bryologist 82: 631 , 1979 (Calyptopogon
angustifolius Herz.; nom. nov. for Tortula angustifolia Herz. hom.
illeg.) Am4 =Syntrichia angustifolia (Herz.) Zand., see treatment
of Syntrichia]
[Tortula heteroloma Card. Am2 = Hennediella heteroloma (Card.)
Zand., see treatment of Hennediella]
[Tortula heteroneura Card. Ant = Tortula princeps De Not. fide
Lightowlers, Brit. Antarct. Surv. Bull. 67: 58, 1985 Syntrichia
princeps (De Not.) Mitt.]
[Tortula hildebrandtii (C. Miill.) Broth. Afr2 Afr3 Afr4 = Tortula
schmidii (C. MUll.) Broth. fide Magill, Fl. S. Afr. I. Mosses 1:
215, 1981 (1982) = Tortulafragilis Tayl.fide Frey & Ktirschner,
Nova Hedw. 46: 96, 1988 Syntrichia fragilis (Tayl.) Ochyra,
Fragm. Florist. Geobot. 37: 212, 1992]
var. papillosa (Broth.) Wijk & Marg. Afr2
[Tortula hirsuta (Vent.) Laz. in Laz., Visotsk., & Mamatk., Biul.
Mosk Obsh. lspyt Prir. Otd. Bioi. 73(3): 145, 1968 Eur Syntrichia ruralis var. hirsuta (Vent.) Podp.]
[Tortula humida Mitt. Am4 =Didymodon humidus (Mitt.) Zand., see
treatment of Didymodon]
Tortula humillima Card. & Copp. Afr1
[Tortula husnotii (Besch.) Broth. Am3 = Barbula agraria Hedw. fide
Zander, Phytologia 44: 202, 1979]
[Tortula hyalinotricha (C. MUll.) Broth. Afr4 (= Tortula princeps De
Not. fide Lightowlers, J. Bryol. 14: 290, 1986 (1987) Syntrichia
princeps (De Not.) Mitt.) = Tortula magellanica Mont. in Gay
Syntrichia
fide Kramer, J. Hattori Bot. Lab. 65 : 111, 1988
magellanica (Mont.) Zand.]
[Tortula incerta (Mitt.) Broth. Afr3 Trichostomum incertum (Mitt.)
Zand., see treatment of Trichostomum]
[Tortula inermis (Brid.) Mont. Eur As1 As2 As3 AsS Afr1 Am1 Am2
=Syntrichia inermis (Brid.) Bruch in Hueb.]
Syntrichia inermis var.
[var. submarginata Schiffn. Eur
submarginata (Schiffn.) Podp.]
[Tortula intermedia (Brid.) Berk., Handb. Brit. Mosses 134, 1855 Eur
As1 As3 AsS Afr1 Afr4 Am1 Am2 =Syntrichia intermedia Brid.]
[subsp. handelii (Schiffn.) Wijk & Marg. Eur AsS
Tortula
handelii Schiffn.fide Kramer, Bryoph. Biblioth. 21: 109, 1980]
[var. calva (Dur. & Sag.) Wijk & Marg. Eur = Syntrichia
intermedia var. calva (Dur. & Sag.) Delogn.]
=
=
=
=
=
=
=
=
=
=
var. gelida (Amann) Wijk & Marg. Eur
var. planifolia (Fleisch. & Wamst.) Wijk & Marg. Eur
var. subcalcicola (Giac.) Wijk & Marg. Eur
[Tortula irregularis Sim Afr4 = Tortula hildebrandtii (C. MUll.)
Broth. fide Magill & Schelpe, Mem. Bot. Surv. S. Afr. 43: 25,
1979 = Tortula schmidii (C. MUll.) Broth. fide Magill, Fl. S.
Afr. I. Mosses 1: 215, 1981 (1982) = Tortulafragilis Tayl.fide
Frey & KUrschner, Nova Hedw. 46: 96, 1988 = Syntrichia
fragilis (Tayl.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
[Tortula jaffuelii Ther. Am6 Syntrichia jaffuelii (Ther.) Zand., see
treatment of Syntrichia]
Tortula kabir-khanii (Broth.) Zand. (Desmatodon), see treatment of
Tortula As3
[Tortu/a khartoumensis Pettet, Trans. Brit. Bryol. Soc. 5: 322, 1966
(1967) Afr2 Hyophila khartoumensis (Pettet) A. Sm. & H.
Whiteh., J. Bryol. 8: 14, 1974 = Desmatodon bogosicus C.
MUll. fide Townsend & Whitehouse, J. Bryol. 10: 475, 1979
and Corley et al., J. Bryol. 11 : 620, 1981 (1982)]
[Tortula kingii Robins., Bryologist 70: 24, 1967 Am4 = Tortula
percarnosa (C. Miill.) Broth. = Syntrichia percarnosa (C.
MUll.) Zand., see treatment of Tortula]
[Tortula kunzeana (C. MUll.) Mont. in Gay Afr4 Am6 Hennediella kunzeana (C. Miill.) Zand., see treatment of Hennediella]
Syntrichia lacerifolia (Wil[Tortula /acerifolia Williams Am4
liams) Zand., see treatment of Syntrichia]
Tortu/a /aevinervis Broth. ex Dus. Am6
[Tortula laevipila (Brid.) Schwaegr. Eur As1 As3 AsS Afr1 Am1
Arn6 Ant Austr2 = Syntrichia laevipila Brid., see treatment of
Syntrichia]
var. gemmifera Squiv. Eur
[var. meridiana/is (Schimp.) Wijk & Marg. As1 As3 AsS Afr1
Am1 Am6
Syntrichia /aevipila Brid. var. meridiana/is
(Schimp.) Jur.]
[var. minor (Biz.) Biz. AsS = Syntrichia virescens var. minor
(Biz.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
var. notarisii Barkm. Eur
var. saccardoana (De Not.) Barkm. Eur
var. wachteri Barkm. Eur
Tortula lanceola Zand. (nom . nov. for Encalypta lanceolata Hedw.
non Tortula lanceolata (Hedw.) P. Beauv.), see treatment of
Tortula Eur As2 AsS Afrl Am1
var. albidens (Corb.) Zand. (Pottia /anceo/ata var.), see treatment
ofTortula Eur
var. angustata (B.&S. in BSG) Zand. (Anaca/ypta /anceo/ata
var.), see treatment of Tortula Eur AsS Afr1
var. lejolisii (Corb.) Zand. (Portia lanceo/ata var.), see treatment
of Tortula Eur Afrl
var. /eucodonta (Schimp.) Zand. (Pottia lanceolata var.), see
treatment of Tortula Eur
var. macrophyl/a (Wamst.) Zand. (Pottia /anceolata var.), see
treatment of Tortula Eur
var. microphylla (Wamst.) Zand. (Pottia lanceolata var.), see
treatment of Tortula Eur
var. mucronata (Amann) Zand. (Pottia lanceolata var.), see treatment of Tortula Eur
var. ovalifolia (Wamst.) Zand. (Pottia /anceolata var.), see treatment of Tortula Eur
var. papillosa (Corb.) Zand. (Pottia lanceolata var.), see treatment of Tortula]
var. rigidior (Schwaegr.) Zand. (Enca/ypta lanceolata var.), see
treatment of Tortula Eur
[Tortula latifolia Bruch ex Hartm. Eur Am1 = Syntrichia latifo/ia
(Hartm.) HUb.]
=
=
=
=
=
333
GENERA, SPECIES AND INFRASPECIFIC TAXA
[var. propagulifera (Milde) Limpr., Laubm. Deutsch!. 1(11): 678,
1888 Eur = Syntrichia latifolia var. propagulifera (Milde)
Amann]
[fo. propagulifera (Milde) Baumgartn. in Degen, Fl. Veleb. 3: 415,
1938 Eur Tortula latifolia var. propagulifera (Milde) Limpr.
Syntrichia latifolia var. propagulifera (Milde) Amann]
[Tortula latrobeana (C. Miill.) Mitt. Austr1 = Tortula antarctica
(Hampe in C. Miill.) Wils. in Hook. f. fide Kramer, J. Hattori Bot.
Lab. 65: 84, 1988 = Syntrichia antarctica (C. Miill. & Hampe)
Zand., see treatment of Syntrichia]
Tortula laureri (Schultz) Lindb. Eur As1 As2 Afr4 Am1
var. setschwanica (Broth.) Zand. (Desmatodon), see treatment of
Tortula As2
Tortula lazarenkoi Sav. in Sav.-Ljub. & Smirn., Handb. Mosses
U.S.S.R. 330, 1970 (nom . nov. for Tortula scabrinervis Laz.)
As1
[Tortula /echleri (C. Miill.) Broth. Am6 =Torrula magellanica Mont.
in Gay fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988 = Syntrichia
mage/lanica (Mont. in Gay) Zand., see treatment of Syntrichia]
[Tortula leiostoma Herz. Am2 Am4 = Hennedie/la limbata (Mitt.)
Zand., see treatment of Hennediella and Long, J. Bryol. 10: 381,
1979]
Tortula leiostomoides P. Yarde Afr2
[Tortula lemniscata Zand., Willdenowia 16: 253, 1986 Am4 = Syntrichia aculeata (Wils.) Zand., see treatment of Tortula]
Tortula leptopyxis (C. Miill.) Lindb. & Am. As!
[Tortula leptosyntrichia (C. Mtill.) Broth. Am6 = Tortula robusta
Hook. & Grev.fide Lightowlers, Brit. Antarct. Surv. Bull. 67: 67,
1985 Syntrichia robusta (Hook. & Grev.) Zand. see treatment
of Syntrichia]
Tortula leptotheca (Broth.) Chen As2
Tortula le-testui P. Yarde Afr2
Tortula leucochlora (C. Miill.) Broth. Afr2
[Tortula leucostega (C. Miill.) Broth. Afr4 Syntrichia leucostega
(C. Miill.) Zand., see treatment of Syntrichia]
[var. trachyneura (Dix.) Kramer, J. Hattori Bot. Lab. 65: 107, 1988
Syntrichia leucostega var. trachyneura (Dix.) Zand. (Tortula),
see treatment of Syntrichia]
Tortula leucostoma (R. Br.) Hook. & Grev. Eur Am1 As! As2
Torrula ligulata Herz. Am4
[Tortula limensis Williams Am4
Syntrichia limensis (Williams)
Zand. see treatment of Syntrichia]
[Tortula linearifolia P. Beauv., Prodr. 92, 1805 (1806) (Bryum
acuminatum Sw. nom. inval.; Barbula acuminata Brid. nom.
inval.) = Barbula agraria Hedw.fide C. Miill., Syn. 1: 604, 1849]
[Tortula linguifolia Herz. Am4 = Syntrichia linguifolia (Herz.) Zand.,
see treatment of Syntrichia]
[Tortula lingulaefolia Card. & Broth. Am6 = Tortula arenae (Besch.)
Broth. fide Lightowlers, Brit. Antarct. Surv. Bull. 67: 46, 1985]
Tortula Ungulata Lindb. Eur
subsp. montenegrina (Breidl. & Szysz.) Podp. Eur
[Tortula lithophila Dus. Am6 = Sarconeurum glaciale (C. Miill.)
Card. & Bryhnfide Green, Brit. Ant~>Tct. Surv. Bull. 41-42: 187,
1975 and Lightowlers, Bryologist 88: 365, 1985]
Tortula litorea Card. in Card. & Broth. Am6
[Tortula longimucronata X.-j. Li, Acta Bot. Yunnan. 3: 107, 1981
As2 = Syntrichia longimucronata (X.-j. Li) Zand., see treatment
of Syntrichia]
Desmatodon
[Tortula longipedunculata (C. Miill.) Broth. Afr4
longipedunculatus (C. Miill.) Magill, Fl. S. Afr. I. Mosses 1: 210,
Hennediella longipedunculata (C. Miill.) Zand.,
1981 [1982]
see treatment of Hennediella]
[Tortula lorentzii (C. Miill.) Broth. Am6 Chenia lorentzii (C. Miill.)
=
=
=
=
=
=
=
=
=
Zand., see treatment of Chenia]
[Tortula luteola Mitt. Austrl =Barbula crinita Schultz fide Weber,
Lindbergia 1: 216, 1972 = Pseudocrossidium crinitum (Schultz)
Zand., see treatment of Pseudocrossidium]
Tortula madagassa Dix. Afr3
[Tortula magellanica Mont. in Gay Am6 = Tortula princeps var.
magellanica (Mont.) Lightowlers, Brit. Antarct. Surv. Bull. 67:
61, 1985 = Syntrichia magellanica (Mont.) Zand., see treatment
of Syntrichia]
Tortula mairei Ther. & Trab. Afr1
Tortula marginata (B.&S. in BSG) Spruce Eur As3 As5 Afr1 Ami
Am3
subsp. limbata (Lindb.) Podp. Eur Afrl
fo. ragusina Latz., Bot. Centralbl. Beih. 48(2): 484, 1931 Eur
Tortula maritima (R. Br. ter) Zand. (Dendia), see treatment of Tortula Austr2
[Tortula maudii R. Brown ter Austr2 = Tortula rubella Hook. f. &
Wils . fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988 Syntrichia rubella (Hook. f. & Wils.) Zand., see treatment of Syntrichia]
Tortula meridiana/is (Luis.) Mach. Eur
Tortula minima Herz. Am4
Tortula minor (C. Miill.) Zand. (Portia), see treatment of Tortula
As3 Afr2
var. elatior (C. Mtill.) Zand. (Portia minor var.), see treatment of
Tortula As3 Afr2
[Tortula minuscula Williams Am4 = Didymodon minuscula (Williams) Zand., see treatment of Didymodon]
Tortula minutirosula (C. Miill.) Broth. Am6
[Tortula mniadelphus (C. Miill.) Broth. Am2 Am4 Am6 = Tortula
quitoensis Tayl. in Hook. fide Mishler in Sharp et al., Moss Fl.
Sagenotortula quitoensis (Tayl. in Hook.) Zand.,
Mex.
Phytologia 65: 430, 1989]
[Torrula mniifolia (Sull.) Mitt. Am2 Am3 Am4 = Dolotortula
mniifolia (Sull.) Zand., Phytologia 65: 426, 1989]
Tortula modica Zand. (nom. nov. for Gymnostomum intermedium
Tum.), see treatment ofTortula Eur As2 Afl Am1
var. corsa (Fleisch. & Wamst.) Zand. (Portia intermedia var.),
see treatment ofTortula Eur
var. gymnandra (Schiffn.) Zand. (Portia intermedia var.), see
treatment of Tortula Eur
var. gymnogyna (Schiffn.) Zand. (Portia intermedia var.), see
treatment of Tortula Eur
var. revoluta (Schiffn.) Zand. (Portia intermedia var.), see treatment ofTortula Eur
var. stenocarpa (Velen.) Zand. (Portia intermedia var.), see treatment of Tortula Eur
var. tenuis (Vent.) Zand. (Pottia intermedia var.), see treatment of
Tortula Eur
[Tortula mollis (B.&S . ex C. Miill.) Broth. Afr2 = Syntrichia mollis
(B.&S. ex C. Miill.) Zand., see treatment of Syntrichia]
[Torrula mongolica (Boros) Ochyra & Pacyna, Fragm. Florist. &
Geobot. 26: 73, 1980 (Syntrichia) Asl = Tortula submontana
Broth. fide Kramer, Bryoph. Biblioth. 21: 89, 1980) = Syntrichia submontana (Broth.) Ochyra, Fragm. Florist. Geobot. 37:
212, 1992]
[Tortula monoica Card. Am6 Ant = Tortula princeps var.
magellanica (Mont.) Lightowlers, Brit. Antarct. Surv. Bull. 67:
61, 1985 also see Kramer, J. Hattori Bot. Lab. 65: 84, 1988
Syntrichia magellanica (Mont.) Zand., see treatment of Syntrichia]
[Tortula montana (Nees) Lindb. hom. illeg. = Tortula intermedia
(Brid.) Berk.]
=
=
=
334
GENERA OF THE POTTIACEAE
var. patudosa Roll, Hedwigia 56(1-3): 164, 1915 Eur
fo. brevifolia Arnold in Rabenh. Bryoth. Eur. 22: n. 1069, 1869 Eur
Tortuta mucronifolia Schwaegr. Eur As1 As2 Afrl Aml
var. arctica Hook. & Grev. Ami Am6
var. aristata (Warnst.) C. Miill. ex Warnst. Eur
var. brevifolia (Warnst.) Warnst. Asl
var. emucronata Am. Asl
var. hyperborea (Hag.) C. Jens. Eur
var. mucronata (Warns!.) Warns!. Eur
fo. angustifolia Warnst., Hedwigia 52: 79, 1912 Eur
fo. breviseta Warnst., Hedwigia 52: 79, 1912 Eur
fo. tatifolia Warnst., Hedwigia 52: 79, 1912 Eur
fo. tongifolia Warns!., Hedwigia 52: 79, 1912 Eur
fo. perpusil/a Warns!., Hedwigia 52: 80, 1912 Eur
fo. recurvata Warns!., Hedwigia 52: 79, 1912 Am1
subfo. crassiseta Warns!., Hedwigia 52: 80, 1912 Eur
Tortuta muralis Hedw. Cosm
subsp. aestiva (Brid. ex Hedw.) Meyl. Eur As2 Afrl Ami Am6
var. aestiva Brid. ex Hedw. Eur As2 Afrl Ami Am6
[var. atbida Podp. Eur = Tortuta muralis fo. atbida (Podp.) Podp.,
Consp. Muse. Eur. 247, 1954]
[var. baetica Cas. & Oliva, Acta Bot. Malacitana 7: 104, 1932 Eur
= Tortuta baetica (Casas & Oliva) Guerra & Ros in Guerra, Ros
& Carri6n, J. Bryol. 17: 281, 1992]
var. brevipeduncutata Rehrn. ex Sirn Afr4
var. emarginata Broth. Eur
var. heribaudii (Corb.) Corb. in Culm. Eur
var. humilis Papp. Eur
var. israelis (Biz. & Bil.) Biz. As5
var. tongipila Dus. Am6
var. madagascariensis Ther. Afr3
var. obcordata (Schimp.) Limpr. (good taxon fide Iwatsuki &
Noguchi, J. Hattori Bot. Lab. 37: 413, 1973) Eur As2 As5 Afrl
var. subtaevis Latz. Eur
var. vulcanicola (Schiffn.) Podp. Eur
fo. aestiva (Brid. ex Hedw.) Koppe, Mitth. Thiiring. Bot. Vereins
N.F. 50: 137, 1943 (Tortula muralis var.) Eur As2 Afrl Aml
Am6
fo. albida (Podp.) Podp., Consp. Muse. Eur. 247, 1954 (Tortula
muralis var.) Eur
fo. brachyrhyncha (Warnst.) Podp., Consp. Muse. Eur. 248, 1954
(Tortula aestiva var.) Eur
fo. brevifolia (Schiffn.) Podp., Consp. Muse. Eur. 248, 1954 (Tortuta aestiva var.) AsS
fo.? calcarea Podp., Consp. Muse. Eur. 247, 1954 nom. invat.
dispon . non cit. (Tortula muralis var.) Eur
fo. dentata Loeske, Moosfl. Harz 178, 1903 Eur
fo.? heterophylla Podp., Consp. Muse. Eur. 247, 1954 nom. inval.
dispon. non cit. (Barbuta muralis var.) Eur
fo. incana Sapegen, Bot. Jahrb. System. 46(Beibl. 105): 14, 1911
(Tortuta muralis var.) Eur Afr1 Afr4
fo. metanocarpa Podp., Sitzungsber. Kong!. Bohrn. Ges. Wiss. Prag
1899(46): 16, 1899 Eur
fo. obcordata (Schimp.) Monk., Laubm. Eur. 304, 1927 (Tortula
muralis var.) Eur As2 As5 Afrl
fo.? ovata Podp., Consp. Muse. Eur. 248, 1954 nom. inval. dispon.
non cit. (Tortula muralis var.) Eur
fo. rupestris (Schultz) SapC!hin, Bot. Jahrb. 46(Beibl. 104): 14, 1911
(Tortula muralis var.) Eur
fo. terrestris (Podp.) Podp., Consp. Muse. Eur. 248, 1954 (Tortula
aestiva fo.) Eur
Tortula murina (C. Miill.) Broth. Austrl
[Tortula nankomontana Nog. = Desmatodon latifolius (Hedw.) Brid.
fide Saito, J. Hattori Bot. Lab. 39: 71, 1975 = Tortuta
euryphylla Zand.]
Tortula napoana De Not. Am4
[Tortula navicularis (Mitt.) Broth. Am3 = Quaesticula navicularis
(Mitt.) Zand., see treatment of Quaesticula]
Tortula nevadensis (Card. & Ther.) Zand. (Pottia), see treatment of
Tortula Am!
[Tortula nigra Zand., Willdenowia 16: 255, 1986 = Syntrichia
percarnosa (C. Miill.) Zand., see treatment of Tortuta]
[Tortuta nigrescens Griff. in Gangulee, Mosses E. India 3: 700,
1972 nom. invat. sin descr. As3 = Barbula nigrescens Mitt. =
Didymodon nigrescens (Mitt.) Saito]
[Tortula norvegica (Web. & Mohr) Wahlenb. Aml Eur Afrl Asl
As2 =Syntrichia norvegica Web.]
[var. catva (Amonn) Kramer, Bryoph. Biblioth. 21: 132, 1980
(Syntrichia aciphylla var.) = Syntrichia norvegica var. catva
(Amann) Zand., see treatment of Syntrichia]
[Tortuta novoguinensis Bartr. As4 = Tortuta caroliniana Andrews
fide Eddy, Handb. Males. Mosses 2: 246, 1991 = Tortuta
amphidiacea (C. Miill.) Broth. fide Mishler in Sharp et al.,
Moss Fl. Mex. Syntrichia amphideacea (C. Miill.) Zand., see
treatment of Syntrichia]
Tortuta oamaruana R. Br. ter Austr2
Tortuta obscuriretis Ther. Am6
Tortuta obtusifolia (Schwaegr.) Math. Eur Asl As2 AsS Afrl Am1
[Tortuta obtusissima (C. Miill.) Mitt. Aml Am2
Syntrichia
obtusissima (C. Miill.) Zand. (Barbuta), see treatment of Syntrichia]
[var. connectens (Card.) Ther. Am2 = Tortula obtusissima (C.
Miill.) Mitt. fide Crum, Appalachian-Ozarkian Element Moss
Fl. Mexico (dissertation, Univ. Microfilms) 165, 1951 Syntrichia obtusissima (C. Miill.) Zand.]
[Tortuta oteaginosa Stone, J. Bryol. 10: 117, 1978 Austrl = Stonea
oteaginosa (Stone) Zand., Phytologia 65: 432, 1989]
[Tortuta ovata (Hedw.) Dix., Rep. Brit. Bryol. Soc. 4: 118, 1939
(Gymnostomum) Eur Asl AsS Afrl Am1 Am6 Austrl = Pterygoneurum ovatum (Hedw.) Dix.
[Tortuta pachyneura Dix. & Sainsb. Austr2 = Willia catobotax (C.
Mlill.) Lightowlers, J. Bryol. 13: 370, 1985
[Tortuta pagorum (Milde) De Not. Eur Afr4 Aml Am2 Austr1 (=
Tortuta atpina fo. pagorum (Milde) Squiv. fide Squivet de
Carondelet, Rev. Bryol. Lichenol. 30: 215, 1962 [1963]) =Syntrichia pagorum (Milde) Amann, see treatment of Syntrichia]
Tortuta pal/ida (Lindb.) Zand. (Portia), see treatment of Tortuta
Eur Afrl As5
var. tongicuspis (Warnst.) Zand. (Pottia pal/ida var. see treatment
of Tortuta Eur As5 Afrl
[Tortula panduraefo/ia (C. Miill. & Hampe) Broth. Austrl = Tortuta antarctica (Hampe in C. Miill.) Wils. in Hook. f. fide
Kramer, J. Hattori Bot. Lab. 65: 84, 1988
Syntrichia
antarctica (C. Miill. & Hampe) Zand., see treatment of Syntrichia]
[Tortula panduriformis R. Brown ter Austr2 = Tortula rubella
Hook. f. & Wils.fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988
Syntrichia rubella (Hook. f. & Wils.) Zand., see treatment of
Syntrichia]
[Tortula papillosa Wils. in Spruce Eur Afr4 Aml Am2 Am4 Am5
Am6 Austrl Austr2 = Syntrichia papillosa (Wils. in Spruce)
Jur., see treatment of Syntrichia]
[var. chilensis Ther. Am4
Syntrichia papillosa var. chilensis
(Ther.) Zand., see treatment of Syntrichia]
[var. meridiana/is Warnst. Eur
Syntrichia papillosa var.
meridiana/is (Warns!.) Zand.]
=
=
=
=
=
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
[Tortula papillosissima (Copp.) Broth. Ami Eur (= Tortula ruralis
(Hedw.) Gaertn., Meyer & Scherb. fide Corley et al., J. Bryol. 11:
619, 1981 [1982]) = Tortula ruralis subsp. hirsuta (Vent.) W.
Kramer= Syntrichia ruralis var. hirsuta (Vent.) Podp.]
Tortula parramattana Mitt. Austrl
Tortula parva Card. Am2
[var. latifolia Ther. Am2 = Tortula fragilis Tayl. fide Crum,
Appalachian-Ozarkian Element Moss Fl. Mexico (dissertation,
Univ. Microfilms) 166, 1951 = Syntrichiafragilis (Tayl.) Ochyra,
Fragm. Florist. Geobot. 37: 212, 1992)
Tortula pau/senii Broth. Asl
Tortula perarmata Broth. Am6
[Tortula percarnosa (C. Miill.) Broth. Am4 Am6 = Syntrichia
percarnosa (C. Miill.) Zand., see treatment of Tortula)
[Tortula perlimbata Geh. ex Card. in Luis. hom. il/eg. Afrl = Tortula
so/msii fo. perlimbata Diill, Cryptogamie Bryol. Lichenol. 1: 179,
1980 (Tortula perlimbata Geh. ex Herz. non Geh. ex Card. in
Luis. see Crosby & Bauer 1983) = Hennediella solmsii fo.
perlimbata (Diill) Zand., see treatment of Hennediella]
Tortula perpusilla (C. Miill.) Broth. Am6
[Tortula peruviana Mitt. Am4 = Barbula peruviana (Mitt.) Jaeg., see
treatment of Barbula]
[Tortula petriei Broth. ex Beck. Austr2 = Tortula arenae subsp.
petriei (Broth.) Lightowlers, J. Bryol. 13: 371, 1985 = Syntrichia
arenae subsp. petriei (Broth. ex Beck.) Zand., see treatment of
Syntrichia]
[Tortula phaea (Hook. f. & Wils.) Dix. Austr2 = Syntrichia phaea
(Hook. f. & Wils.) Zand. see treatment of Syntrichia]
[Tortula pichinchensis Tayl. Am3 Am4 = Syntrichia pichinchensis
(Tayl.) Zand., see treatment of Syntrichia]
Tortula pierrotii Biz., Acta Bot. Acad. Sci. Hung. 18: 21, 1973
"pierrqtti" Afr2
[Tortula pilifera Hook. As4? Afr2 Afr4 Am6 = Barbula crinita
Schultz fide Magill, Fl. S. Afr. I. Mosses 1: 237, 1981 (1982) =
Pseudocrossidium crinitum (Schultz) Zand., see treatment of
Pseudocrossidium]
[var. denticulata (Dus.) Wijk & Marg. Am6 = Barbula crinita
Schultz fide Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium)
[var. gracilis (Dus.) Wijk & Marg. Am6 = Barbula crinita Schultz
fide Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
[var. longifolia Sim Afr4 = Barbula crinita Schultz fide Weber,
Lindbergia 1: 216, 1972 andfide Magill, Fl. S. Afr. I. Mosses 1:
237, 1981 (1982))
[var. oliviensis (Card.) Wijk & Marg. Am6 = Barbula crinita
Schultz fide Weber, Lindbergia 1: 216, 1972 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
Tortula planicosta Herz. Am6
Tortula planifo/ia X.-j. Li, Acta Bot. Yunnan. 3: 109, 1981 As2
Tortula platyphylla Mitt. Am6
Tortula plinthobia (Sull. & Lesq.) Aust., Musci Appalach. Suppl.
1:10, 1878 cf. Zander, Bryologist 82: 551, 1979 Ami
Tortula podocarpi (C. Miill.) Broth. Am6
Tortula po/ycarpa Dus. Am6
Tortula polylepidis Herz. Am4
[Tortula polyseta (C. Miill.) Warnst. Am2 Am4 = Hennediella
polyseta (C. Miill.) Zand., see treatment of Hennediella]
[Tortula porphyreoneura (C. Miill.) Townsend, J. Bryol. 10: 576,
1979 (Barbula) Afr2 Afr4 = Psuedocrossidium porphyreoneurum
(C. Miill.) Zand., see treatment of Pseudocrossidium]
Tortula porteri (Jam. in Aust.) Zand., see treatment of Tortula Ami
[Tortula preissiana (C. Miill.) Broth. Austr1 = Tortula antarctica
335
(Hampe in C. Miill.) Wils. in Hook. f. fide Kramer, J. Hattori
Bot. Lab. 65: 84, 1988 = Syntrichia antarctica (Hampe in C.
Miill.) Zand., see treatment of Syntrichia]
[Tortula princeps De Not. Eur As1 As3 As5 Afrl Afr4 Ami Am2
Am4 Am6 Austrl Austr2 Oc Ant = Syntrichia princeps (De
Not.) Mitt., see treatment of Syntrichia]
[subsp. echinata (Schiffn.) Kramer, Bryophyt. Biblioth. 21: 86,
1980 (Tortula) Eur As5 (= Tortula princeps De Not. fide
Corley et al., J. Bryol. 11: 618, 1981 [1982]) = Tortula princeps
subsp. echinata (Schiffn.) Kramer, Bryophyt. Biblioth. 21: 86,
1980 = Syntrichia princeps var. echinata (Shiffn.) Zand., see
treatment of Syntrichia]
[subsp. parnassica (Schiffn.) Kramer, Bryophyt. Biblioth. 21: 87,
1980 (Tortula muelleri var.) Eur = Syntrichia princeps var. '
parnassica (Schiffn.) Podp.]
[var. brachycarpa De Not. Eur = Syntrichia princeps var.
brachycarpa (Zand.) De Not., see treatment of Syntrichia]
[var. conferta (Bartr.) Lightowlers, Brit. Antarct. Surv. Bull. 67:
61, 1985 (Tortula) Ant (= Tortula princeps De Not. fide
Lightowlers, J. Bryol. 14: 290, 1986 [1987] = Syntrichia
princeps (De Not.) Mitt., see treatment of Syntrichia) = Syntrichia conferta (Bartr.) Zand., see treatment of Syntrichia]
[var. echinata (Schiffn.) Biz. Eur As5 (= Tortula princeps De
Not. fide Corley et al., J. Bryol. 11: 619, 1981 [1982]) =Tortula princeps subsp. echinata (Schiffn.) Kramer, Bryophyt.
Biblioth. 21: 86, 1980 = Syntrichia princeps var. echinata
(Shiffn.) Zand., see treatment of Syntrichia]
[var. magel/anica (Mont.) Lightowlers, Brit. Antarct. Surv. Bull.
67: 61, 1985 (Tortula) Am6 Ant = Syntrichia magellanica
(Mont.) Zand., see treatment of Syntrichia]
[var. parnassica (Schiffn.) Wijk & Marg. Eur = Syntrichia
princeps var. parnassica (Schiffn.) Podp.]
[Tortula propagulata Laz., J. Bot. Acad. Sci. Ukr. 3(3-4): 61, 1946
As1 = Tortula laevipila (Brid.) Schwaegr. var. propagulifera
Lindh. fide Sav.-Ljub. & Smim., Handb. Mosses USSR 352,
1970]
[Tortula propagulosa Sharp Am1 = Rhachithecium perpusil/um
(Thwait. & Mitt.) Broth. fide Zander, Bryologist 81: 458, 1978]
Tortula propinqua (C. Miill.) Broth. Austrl
Tortula protobryoides Zand. (nom . nov. for Phascum bryoides
Dicks.), see treatment of Tortula Eur AsS Ami Am2
var. brevifolia (De Not.) Zand. (Phascum bryoides var.), see
treatment of Tortula Eur
var. thornhillii (Wils.) Zand. (Phascum bryoides var.), see treatment of Tortula] Eur
[Tortula prostrata Mont. Am4 Am6 = Syntrichia prostrata (Mont.)
Zand. see treatment Syntrichia]
fo. angustofolia Herz., Arch. Es. Farm. Fac. Ci. Med. C6rdoba
(Secc. Ci.) 7: 41, 1938 nom. inval. descr. hisp. Am6
fo. latifolia Herz., Arch. Es. Farm. Fac. Ci. Med. C6rdoba (Secc.
Ci.) 7: 41, 1938 nom. inval. descr. hisp. Am6
[Tortula pseudaciphylla (Kindb.) Broth. Ami = Tortula intermedia
(Brid.) Berk. fide Steere in Grout, Moss Fl. N. Am. 1: 246,
1939]
[Tortula pseudodesertorum Frohl., Ann. Naturhist. Mus. Wien 67:
155, 1964 As3 = Syntrichia pseudohandelii (Frohl.), Agnew &
Vondr., Feddes Rep. 86(6-8): 401, 1975]
[Tortula pseudohandelii Frohl., Ann. Naturhist. Mus. Wien 67: 155,
1964 As3 = Syntrichia pseudohandelii (Frohl.), Agnew &
Vondr., Feddes Rep. 86(6-8): 401, 1975]
Tortula pseudolatifolia Card. Am6
Tortula pseudoprinceps Dix. As3
[Tortula pseudorobusta Dus. Am6 = Syntrichia pseudorobusta
336
GENERA OF THE POTIIACEAE
(Dus.) Zand., see treatment of Syntrichia]
Tortula pugionata (C. Miill.) Broth. As2
[Tortula pulvinata (Jur.) Limpr. Eur Afr1 Am1 = Tortula virescens
(De Not.) De Not. fide Kramer, Bryoph. Biblioth. 21: 99, 1980 =
Syntrichia virescens (De Not.) Ochyra, Fragm. Aorist. Geobot.
37: 212, 1992]
subsp. papillinervis (C. Mill!. & Kindb.) Par. nom. dub. Ami
Tortula pulvinatula Dus. Am6
[Tortula purpureovelutina Herz. Am6 = Barbula crinita Schultz fide
Sollman, Lindbergia 16: 22, 1990 = Pseudocrossidium aureum
fide Zander, Phytologia 44: 207, 1979 = Pseudocrossidium
crinitum (Schultz) Zand., see treatment of Pseudocrossidium]
[Tortula pusilla Angst. hom. illeg. Am6 = Tortula princeps var.
magel/anica (Mont.) Lightowlers, Brit. Antarct. Surv. Bull. 67:
61, 1985 = Syntrichia magellanica (Mont.) Zand., see treatment
of Syntrichia]
[Tortula pygmaea Dus. Am6 = Syntrichia pygmaea (Dus.) Zand. see
treatment of Syntrichia]
[Tortula quitoensis Tayl. Am2 Am4 Am6 = Sagenotortula quitoensis
(Tayl. in Hook.) Zand., Phytologia 65: 430, 1989]
[Tortula raddei Broth. As1 = Microbryum raddei (Broth.) Zand., see
treatment of MicrobryumJ
Tortula rallieri Card. Afr4
[Tortula ramosissima Ther. Am4 = Syntrichia ramosissima (Ther.)
Zand., see treatment of Syntrichia]
Tortula randii (Kenn.) Zand. (Portia), see treatment of Tortula Eur
Aml
Tortula raucopapillosa (X.-j. Li) Zand. (Desmatodon), see treatment
of Tortula As2
Tortula readeri (C. Miill.) Broth. Austr1
Tortula recurvata Hook. (good species fide Catcheside, Mosses S.
Austr. 150, 1980 as Desmatodon) Afr4 Austrl
[Tortula recurvifolia (Schimp.) Aust., Musci Appalach. Suppl. 1: 10,
496, 1878 hom. il/eg. (Barbula) Eur As2 As3 Am1 = Barbula
reflexa (Brid.) Brid.fide Limpricht 1888 = Didymodonfal/ax var.
reflexus (Brid.) Zand., Bryologist 83: 230, 1980 = Didymodon
rigidicaulis (C. MUll.) Saito fide Saito, J. Hattori Bot. Lab. 39:
502, 1975 =Didymodon ferrugineus (Schimp. ex Besch.) Hill, J.
Bryol. 11: 599, 1981 (1982)]
[Tortula reflexa X .-j. Li, Acta Bot. Yunnan. 3: 109, 1981 hom. illeg.
non Brid. As2 = Syntrichia reflexa Zand. nom. nov., see treatment
of Syntrichia]
[Tortula rehmannii (C. Miill.) Sim Afr2 Afr4 = Barbula rehmannii C.
MUll. fide Magill, Fl. S. Afr. I. Mosses 1: 245, 1981 (1982)]
[Tortula remotifolia Tak. As2 = Tortula norvegica (Web. & Mohr)
Wahlenb.fide Kramer, Bryoph. Biblioth. 21: 128, 1980 = Syntrichia norvegica Web.]
[Tortula reticularia (C. MUll.) Broth. Afr4 = Tortula papillosa Wils.
in Spruce fide Magill & Schelpe, Mem. Bot. Surv. S. Afr. 43: 25,
1979 and see Magill, Fl. S. Afr. I. Mosses 1: 218, 1981 (1982)]
Tortula revolutifolia Laz. As1
Tortula revolvens (Schimp.) Roth Eur Afr1
var. obtusata Reim. Eur As5
[Tortula rhizophylla (Sak.) Iwats. & Saito Eur As2Am1 Am2 Austr1
= Chenia rhizophylla (Sak.) Zand., Phytologia 65: 425, 1989 =
Chenia leptophylla (C. MUll.) Zand., see treatment of Chenia]
Tortula rhodonia Zand. (nom. nov. for Desmatodon wilczekii Meyl.),
see treatment of Tortula Eur
[Tortula rigescens Broth. & Geh. in Broth. As5 = Syntrichia
rigescens (Broth. & Geh. in Broth.) Ochyra, Fragm. Florist.
Geobot. 37: 212, 1992]
Tortula ripico/a Ther. Am2
[Tortula rivularis Dus. Am6 = Tortula fontana (C. MUll.) Broth. fide
Lightowlers, Brit. Antarct. Surv. Bull. 67: 53, 1985 = Syntrichia fontana (C. MUll. in Neum.) Zand., see treatment of Syntrichia]
[Tortula robusta Hook. & Grev. Am5 Am6 Austr1 Ant= Syntrichia
robusta (Hook. & Grev.) Zand. see treatment of Syntrichia]
[var. taxa Bartr. Am6 = Tortula robusta Hook. & Grev. fide
Lightowlers, Brit. Antarct. Surv. Bull. 67: 67, 1985 = Syntrichia robusta (Hook. & Grev.) Zand. see treatment of Syntrichia]
[var. recurva Lightowlers, Brit. Antarct. Surv. Bull. 64: 64, 1984
Ant = Syntrichia robusta var. recurva (Lightowlers) Zand.
(Tortula robusta var.) see treatment of Syntrichia]
[var. runcinata (C. MUll.) Broth. Am4 Am6 = Tortula robusta
Hook. & Grev. fide Lightowlers, Brit. Antarct. Surv. Bull. 67:
67, 1985 = Syntrichia robusta (Hook. & Grev.) Zand. see treatment of Syntrichia]
[Tortula robustula Card. Am6 = Tortula filaris (C. Miill.) Broth.
fide Lightowlers, Brit. Antarct. Surv. Bull. 67: 49, 1985 = Syntrichia filaris (C. Miill. in Neum.) Zand., see treatment of Tortuta]
[Tortuta rubella Hook. f. & Wils. good species fide Kramer, J.
Hattori Bot. Lab. 65: 84, 1988 Afr4 Austr1 Austr2 = Syntrichia
rubella (Hook. f. & Wils.) Zand., see treatment of Syntrichia]
[Tortula rubra Mitt. in Hook. f. Am6 Austrl Austr2 Ant= Syntrichia rubra (Mitt. in Hook. f.) Zand. see treatment of Syntrichia]
[var. subantarctica (Sainsb.) Lightowlers, J. Bryol. 13: 373, 1985
(Tortula subantarctica) inval. basion. non cit. Austr2 = Syntrichia rubra var. subantarctica (Sainsb.) Zand.]
[Tortula rubripila Dix. As3 = Tortula norvegica (Web. Lindh. fide
Kramer, Bryoph. Biblioth. 21 : 128, 1980 = Syntrichia
norvegica Web.]
Tortula rufa (Besch.) Broth. hom. illeg. Afr3
[Tortula ruraliformis (Besch.) Grout, Mosses Hand!. Micr. 167,
1903 Eur As5 Afrl Am1 = Tortula ruralis var. ruraliformis
(Besch.) Wild. = Syntrichia ruralis var. arenicota (Braithw.)
Amann, cf. Kramer, Bryoph. Biblioth. 21: 116, 1980]
[Tortuta ruraliformis (Besch.) lngh. (inval. fide Crundwell, Trans.
Brit. Bryol. Soc. 6: 325, 1971) Eur As5 Afr1 Am1 = Tortula
ruralis var. ruraliformis (Besch.) Wild. = Syntrichia ruralis
var. arenico/a (Braithw.) Amann, cf. Kramer, Bryoph. Biblioth.
21: 116, 1980]
[var. subpapillosissima (Biz. & Pierr.) Kramer, Bryophyt.
Biblioth. 21: 120, 1980 Afr1 = Syntrichia ruralis var.
subpapil/osissima (Biz. & Pierr.) Zand., see treatment of Syntrichia]
[Tortuta ruralis (Hedw.) Gaertn., Meyer & Scherb. Eur As1 As2
As3 As5 Afr1 Afr2 Afr4 Am1 Am2 Am4 Am5 Austr1 Oc =
Syntrichia ruralis (Hedw.) Web. & Mohr]
[subsp. calcicota (Amann) Giac. Eur = Tortuta calcicotens
Kramer, Bryophyt. Biblioth. 21: 90, 1980 (= Tortuta ruralis
(Hedw.) Gaertn., Meyer & Scherb. fide Corley et a!., J. Bryol.
11: 619, 1981 [1982]) = Syntrichia calcicola Amann]
[subsp. hirsuta (Vent.) Kramer, Bryophyt. Biblioth. 21: 126, 1980
Eur As2 Am1 = Syntrichia ruralis var. hirsuta (Vent.) Podp.]
[subsp. rura/iformis (Besch.) Dix. Eur As5 Afr1 Am1 = Syntrichia ruralis var. arenicota (Braithw.) Amann cf. Kramer,
Bryoph. Biblioth. 21: 116, 1980]
[var. aciphyl/oides Podp. Eur = Tortuta ruratiformis (Besch.)
lngh.fide Kramer, Bryoph. Biblioth. 21: 116, 1980 = Syntrichia
ruralis var. arenicota (Braithw.) Amann cf. Kramer, Bryoph.
Biblioth. 21: 116, 1980]
[var. atpina Wahlenb. Eur As! Afrl Ami = Tortula norvegica
(Web.) Wahlenb. ex Lindb.fide Corley et al., J. Bryol. 11 : 619,
1981 (1982)]
337
GENERA, SPECIES AND INFRASPECIFIC TAXA
var. alpina De Not., Mem. R. Ace. Sc. Torino Cl. Sc. Fis. Mat. 40:
29I , I838 hom. illeg. Eur
var. brevifolia Wamst., Krypt. Fl. Brandenburg 2: 277, I904 hom.
illeg. (Tortula ruralis fo. brevifolia Am. in Rabenh., I869) Eur
[var. calcicola (Amann) Barkm. Eur = Tortula calcicolens Kramer,
Bryophyt. Biblioth. 2i: 90, i980 (= Tortula ruralis (Hedw.)
Gaertn., Meyer & Scherb. fide Corley et al., J. Bryol. ii: 6i9,
i98I [I982]) = Syntrichia ruralis var. calcicola (Ammann)
Monk. = Syntrichia calcicola Amann]
[var. densa Velen. Eur = Tortula ruralis (Hedw.) Gaertn., Meyer &
Scherb. var. ruralis fide Kramer, Bryoph. Biblioth. 2i: I22, i980
= Syntrichia ruralis (Hedw.) Web. & Mohr]
[var. gigantea (Lesq.) Koch Ami= Syntrichia ruralis var. gigantea
(Lesq.) Zand., see treatment of Syntrichia]
[var. gracilis C. Jens. Ami = Syntrichia ruralis var. gracilis (C.
Jens.) Zand., see treatment of Syntrichia]
[var. gypsophila Amann ex Roth Eur = Tortula caninervis var.
gypsophila (G. Roth) Kramer, Bryophyt. Biblioth. 2i: 108, I980
= Syntrichia caninervis var. gypsophila (Roth.) Ochyra, Fragm.
Florist. Geobot. 37: 2i2, i992]
[var. hirsuta (Vent.) Par. Eur As2 Ami = Syntrichia ruralis var.
hirsuta (Vent.) Podp.]
[var. latifolia Amott, Disp. Meth. Esp. Mouss. 38, I825 [I826] Eur
= Tortula latifolia (Amott) Hartm.]
[var. pontresinae (Geh. & Wamst.) Wijk & Marg. Eur = Syntrichia
ruralis var. pontresinae (Geh. & Wamst.) Podp.]
[var. pseudoaciphylla Wint. Eur = Tortula norvegica (Web.) Lindh.
var. norvegica fide Kramer, Bryoph. Biblioth. 2i: i28, i980 =
Syntrichia norvegica Web.] '
[var. rufipila Herz. Eur = Tortula norvegica (Web.) Lindh. var.
norvegicafide Kramer, J. Hattori Bot. Lab. 65 : 84, I988]
[var. ruraliformis (Besch.) Wild. Eur As5 Afrl Ami = Syntrichia
ruralis var. arenicola (Braithw.) Amann cf. Kramer, Bryoph.
Biblioth. 2I: ll6, i980]
[var. spiralis Herz. Am4 = Syntrichia ruralis var. spiralis (Herz.)
Zand., see treatment of Syntrichia]
[var. submamillosa Kramer, Bryophyt. Biblioth. 2I: i27, i980 As5
= Syntrichia ruralis var. submamillosa (Kramer) Zand., see treatment of Syntrichia]
[var. subpapillosissima Biz. & Pierr., Acta Bot. Acad. Sci.
Hungaricae I8: ll, I973 Afri = Tortula ruraliformis var.
subpapillosissima (Biz. & Pierr.) Kramer, Bryoph. Biblioth. 2I:
I20, i980 = Syntrichia ruralis var. subpapillosissima (Biz. &
Pierr.) Zand., see treatment of Syntrichia]
[var. substereidosa Kramer, Bryophyt. Biblioth. 2I: i25, I980 As5
= Syntrichia ruralis var. substereidosa (Kramer) Zand., see treatment of Syntrichia]
fo. brevifolia Am. in Rabenh., Bryoth. Eur. 1069, I869 Eur
fo. contorta Podp., Sitzungsber. Kong!. Bohm. Ges. Wiss. Prag
I899(46): i6, I899 Eur
[fo. fallax Herz., Bot. Z. 93: 4i, i944 Eur = Syntrichia ruralis fo.
fallax (Herz.) Podp.]
[fo.fuscipila Herz., Kryptog. Forsch. 4: 280, i920 Eur = Syntrichia
ruralis fo. rufipila Herz. ex Ri.ibel fide Podp., Consp. Muse. Eur.
256, I954]
[fo. gracilis Meyl., Rev. Bryol. 52: 54, I925 Eur = Syntrichia
ruralis fo. gracilis (Meyl.) Podp.]
fo. pseudoruraliformis Tosco, Webbia 28: 286, i973 Eur
[fo. rufoneura Podp., Sitzungsber. Kong!. Bohm. Ges. Wiss. Prag
I899(46): I6, 1899 Eur = Syntrichia ruralis fo. rufoneura (Podp.)
Podp.]
[fo. subrufa Podp., Veroff. Geobot. lnst. Ri.ibel Zi.irich I: 244, I924
Eur = Syntrichia ruralis fo. subrufa (Podp.) Podp.j
[fo. viridis Matous., Hedwigia 44: 31, I904 Eur = Syntrichia
ruralis fo. viridis (Matous.) Podp.]
Tortula sabinae Townsend, Lindbergia 10: 176, 1985 Afr2
[Tortula saharae Trab. Afr1 =Tortula caninervis (Mitt.) Broth. fide
Kramer, Bryoph. Biblioth. 2i: 103, i980 = Syntrichia
caninervis Mitt.]
Tortula sainsburyana Zand. (nom . nov. for Portia stevensii) Austr2
Tortula santiagensis Broth. Am6
Tortula santorinensis Schiffn. Eur
fo. typica Schiffn., Verh. Zool.-Bot. Ges. Wien 69: 329, I920
nom. illeg. = Tortula santorinensis Schiffn. fo. santorinensis]
var. apiculata Schiffn. Eur
Tortula satoi Sak. As2
Tortula savatieri (Besch.) Broth. Am4
[Tortula saxicola Card. Am6 = Syntrichia saxicola (Card.) Zand.,
see treatment of Syntrichia]
[Tortula scabrella Dus. Am6 = Syntrichia scabrella (Dus.) Zand.,
see treatment of Syntrichia]
[Tortula scabrinervis (C. Mi.ill.) Mont. in Gay Am4 Am6 = Syntrichia scabrinervis (C. Mi.ill.) Zand., see treatment of Syntrichia]
[Tortula scabrinervis Laz. Asi hom. illeg. = Tortula lazarenkoi L.
Sav. in Sav.-Ljub. & Smim., Handb. Mosses U.S.S.R. 330,
1970 nom. nov.]
[Tortula schmidii (C. Mi.ill.) Broth. As3 Afr3 Afr4 = Tortula
fragilis Tayl. fide Frey & Ki.irschner, Nova Hedw. 46: 96,
1988) = Syntrichia fragilis (Tayl.) Ochyra, Fragm. Florist.
Geobot. 37: 2I2, I992]
[Tortula schnyderi (C. Mi.ill.) Broth. Am6 = Syntrichia schnyderi
(C. Mi.ill.) Zand., see treatment of Syntrichia]
[Tortula scotteri Zand. & Steere, Bryologist 8I : 463, I978 Ami =
Hilpertia scotteri (Zand. & Steere) Zand., Phytologia 65: 428,
I989J
[Tortula searlii R. Brown ter Austr2 = Tortula rubella Hook. f. &
Wils . fide Kramer, J. Hattori Bot. Lab. 65 : 84, I988 = Syntrichia rubella (Hook. f. & Wils.) Zand., see treatment of Syntrichia]
[Tortula semirubra (C. Mi.ill.) Broth. Afr4 = Tortula princeps De
Not. fide Lightowlers, J. Bryol. i4: 290, I986 (I987) Syntrichia princeps (De Not.) Mitt.]
[Tortula serrata Dix. (nom . legit. contrary to Index Muscorum; cf.
Lightowlers, J. Bryol. 13: 373, I985) Austr2 = Syntrichia
serrata (Dix.) Zand., see treatment of Syntrichia]
[Tortula serripungens (Lor. & C. Mi.ill.) Broth. Am4 Am6 Syntrichia serripungens (Lor. & C. Mi.ill.) Zand., see treatment of
Syntrichia]
[var. exesa (C. Mi.ill.) Herz. Am6 = Syntrichia serripungens var.
excesa (C. Mi.ill.) Zand., see treatment of Syntrichia]
[Tortula serrulata Hook. & Grev. (see Lightowlers, J. Bryol. I3,
373, I985) Am6 = Hennediella serrulata (Hook. & Grev.)
Zand., see treatment of Hennediella]
[Tortula serrulata Wamst. hom. illeg. = Tortula subulata var.
serrulata Wamst.]
fo. tenuiseta Warnst., Hedwigia 52: 76, i9I2 Eur
fo. minor (A. Braun) Warnst., Hedwigia 52: 76, i912 (Syntrichia
subulata var.) Eur
[Tortula sinensis (C. Mi.ill.) Broth. in Levier Eur Asl As2 As3 As5
Afri = Syntrichia sinensis (C. Mi.ill.) Ochyra, Fragm. Florist.
Geobot. 37: 2i2, i992]
Tortula sinuata Bartr., Rev . Bryol. Lichenol. 33: 325; I964-65
[i965] Am6
[Tortula socialis Dus. Am6 = Syntrichia socialis (Dus.) Zand., see
treatment of Tortula]
Tortula solmsii (Schimp.) Limpr. Eur Afri
=
=
GENERA OF THE POTTIACEAE
338
var. minor Roth Afr1
fo. perlimbata DUll, Cryptogamic Bryol. Lichenol. 1: 179, 1980
(Tortula perlimbata Geh. ex Herz. non Geh. ex Card. in Luis. see
Crosby & Bauer 1983) Afr1
Tortula solomensis (Broth.) Zand. (Desmatodon), see treatment of
Tortula As2
Tortula sordida Herz. Am4
Tortula splachnoides (Homsch.) Zand. (Phascum), see treatment of
Tortula Afr4
[Tortula spuria Amann Eur
Tortula caininervis subsp. spuria
(Amann) Kramer, Bryophyt. Biblioth. 21: 106, 1980 Syntrichia
caninervis var. spuria (Amann) Zand., see treatment of Syntrichia]
Tortula squarripila Ther. Am6
[Tortula stanfordensis Steere Eur Am1 Austr1 = Hyophila
Hennediella
stanfordensis (Steere) A. Sm. & Whiteh.
stanfordensis (Steere) Blockeel, J. Bryol. 16: 191, 1991]
[Tortula stenophylla Mitt. As2 = Oxystegus tenuirostris (Hook. &
Grev.) A. Sm. (as Oxystegus cylindricus (Brid.) Hilp.) fide Saito,
J. Hattori Bot. Lab. 39: 437, 1975 = Trichostomum tenuirostre
(Hook. & Tayl.) Lindb.]
Tortula stenophylla Card. & Broth. hom. illeg. Am6
Tortula streptopogoniacea (C. Mii11.) Broth. Austr1
[Tortula subantarctica Sainsb. Austr2 = Tortula rubra var.
subantarctica (Sainsb.) Lightow1ers, J. Bryol. 13: 373, 1985]
[Tortula subaristata (B.&S. ex C. Miill.) Broth. Afr2 Syntrichia
subaristata (B.&S. ex C. MUll.) Zand., see treatment of Syntrichia]
Tortula subbrunnea Broth. & Watts Austr1
Tortula subcaroliniana Biz., Svensk Bot. Tidskr. 63: 446, 1969 Afr2
Tortula sublimbata (Mitt.) Broth. As2
[Tortula submontana Broth. As1 = Syntrichia submontana (Broth.)
Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
[Tortula subobliqua Williams Am4 = Chenia subobliqua (Williams)
Zand., Phyto1ogia 65: 425, 1989]
Tortula subrufa Card. in Grand. Afr3
[Tortula subspathulata (C. MUll.) Broth. Afr2 =Tortulafragilis Tayl.
fide Townsend, Lindbergia 10: 177, 1985 = Syntrichia fragilis
(Tayl.) Ochyra, Fragm. Florist. Geobot. 37: 212, 1992]
[Tortula subspiralis (Hampe) Broth. Austr1 = Barbula hornschuchiana Schultz fide Catcheside, Mosses S. Austr. 178, 1980]
Psuedocrossidium hornschuchianum (Schultz) Zand. Phytologia
44: 205, 1979]
[Tortula subtorquatifolia Dix. Afr4 = Desmatodon convolutus (Brid.)
Grout fide Magill, Fl. S. Afr. I. Mosses 1: 210, 1981 [1982] =
Tortula atrovirens (Sm.) Lindb.]
Tortula subtranscaspica Frohl., Mitt. ThUring. Bot. Ges. 1(2-3): 63,
1955 Eur?
Tortula subulata Hedw. Eur As1 As2 As3 As5 Afr1 Am1 Am3
subsp. angustata (Schimp.) Kindb. Eur Afr1 Am1
subsp. serrulata (Warns!.) Giac. Eur
var. angustata (Schimp.) Limpr. Eur Afr1 Am1
var. angustifolia (Schimp. ex Milde) Roll, Hedwigia 56(1-3): 162,
1915 Eur
var. bifrons Hag. Eur
var. buergeneri Warnst. Eur
var. elongata (Lang.) Par. Eur Am1
var.flaviseta Warnst. Eur
var. graeffii Warns!. Eur
var.laxifolia Warnst. Eur
var. microphylla Warnst. Eur
var. minor Wahlenb. Eur
var. mutica (Schimp.) Limpr. Eur Afr1 Am1
=
=
=
=
=
var. serrulata Warnst. Eur
var. subinermis (B.&S. in BSG) Wils. Eur As5 Afr1 Am1
fo. angustifolia (Wanst.) Sav.-Ljub., Novosti Sist. Niz. Rast. 6:
248, 1969 [1970] (Tortula graeffii var.) Eur
fo. denticulata Latz., Magyar Bot. Lapok 29: 119, 1930 Eur
fo. pygmaea Mart. ex Warnst., Hedwigia 52: 74, 1912 Eur
[Tortula synecia R. Brown ter Austr2 = Tortula rubella Hook. f. &
Wils. fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988 = Syntrichia rubella (Hook. f. & Wils.) Zand., see treatment of Syntrichia]
Tortula systylia (Schimp.) Lindb. Eur As1 As3 Am1
[Tortula tanganyikae Dix. Afr2 = Syntrichia amphidiaceus (C.
MUll.) Zand. see treatment of Syntrichia]
[Tortula tenella Broth. Afr4 Austr1 Austr2 = Tortula rubella Hook.
f. & Wils.fide Kramer, J. Hattori Bot. Lab. 65: 84, 1988 = Syntrichia rubella (Hook. f. & Wils.) Zand., see treatment of Syntrichia]
Tortula thianschanica Broth. As1
Tortula thompsonii (C. MUll.) Zand. (Trichostomum), see treatment
of Tortula As2 As3
Tortula tonkinensis (Besch.) Zand. (Desmatodon), see treatment of
Tortula As3
[Tortula tortuosa Hedw. = Tortella tortuosa (Hedw.) Limpr.]
var. microcarpa Hartm. Eur
Tortula toutonii Biz., Acta Bot. Acad. Sci. Hung. 18: 26, 1973 Afr2
[Tortula trachyneura Dix. Afr4 = Tortula ruralis fo. trachyneura
(Dix.) Sim = Tortula ruralis (Hedw.) Gaetm., Meyer & Scherb.
fide Magill, Fl. S. Afr. I. Mosses 1: 219, 1981 (1982)]
Tortula trachyphylla Broth. As1
Tortula transcaspica Broth. As1
Tortula truncata (Hedw.) Mitt. in Godm. Eur As1 As2 As3 Afr1
Am1 Am6 Austr1 Austr2
var. littoralis (Mitt.) Zand. (Portia littoralis), see treatment of
Tortula Eur Afr1
var. brevirostris (Lisa) Zand. (Gymnostomum truncatum var.), see
treatment of Tortula Eur
var. illyrica (Latz.) Zand. (Pottia) , see treatment of Tortula Eur
var. minutissima (Warns!.) Zand. (Pottia truncata var.), see treatment of Tortula Eur
[Tortula tutigae Sak. As2 = Pottia intermedia (Tum.) FUmr. fide
· Saito, J. Hattori Bot. Lab. 39: 522, 1975
Tortula modica
Zand. nom. nov., see treatment of Tortula]
Tortula ucrainica (Laz.) Zand. (Desmatodon), see treatment of
Tortula Eur
Tortula umbrosa Dus. Am6
[Tortula unguiculata (Hedw.) A. Roth ex P. Beauv. hom. illeg.
Barbula unguicu/ata Hedw.]
var. compacta De Not. Eur
var. robusta Lindb. in Hartm., Handb. Skand. Fl. ed. 10, 2: 91,
1871 Eur
Tortula vahliana (Schultz) Mont. in Gay Eur AsS Afr1 Am1 Am6
var. minor (Husn.) Par. Afr1
[Tortula vectensis Warb. & Crundw., Trans. Brit. Bryol. Soc. 4:
763, 1965 Eur Am1 = Tortula rhizophyl/a (Sak.) Jwats. &
Saito, Misc. Bryol. Lichenol. 6: 59, 1972 = Chenia leptophyl/a
(C. MUll.) Zand., see treatment of Chenia]
[Tortula velenovskyi Schiffn. Eur Am6 = Hilpertia velenovskyi
(Schiffn.) Zand., see treatment of Hilpertia ]
Tortula vesicu/osa (C. Miill.) Broth. Austr1
var. involucrata (C. MUll.) Par. Austr1
[Tortula virescens (De Not.) De Not. Eur Afr1 Am1 = Syntrichia
virescens (De Not.) Ochyra, Fragm. Florist. Geobot. 37: 212,
1992]
=
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
[subsp. bizotii (Laz.) Kramer, Bryophyt. Biblioth. 21: 102, 1980
comb. inval. basion. inval. (Tortula) As5 = Tortula virescens
subsp. bizotiana Kramer, J. Hattori Bot. Lab. 65: 123, 1988 =
Syntrichia virescens var. bizotiana (Kramer) Zand., see treatment
of Syntrichia]
[var. iranica Kramer, Bryophyt. Biblioth. 21: 101, 1980 AsS= Syntrichia virescens var. iranica (Kramer) Zand., see treatment of
Syntrichia]
Tortula viridipila Dix. & Sainsb. Austr2
[Tortula viridula (C. Miill.) Broth. Am4 = Syntrichia viridula (C.
MUll.) Zand., see treatment of Syntrichia]
Tortula websteri Robins., Bryo1ogist 68: 316, 1965 As3
Tortula wilczekii Meyl. Afr1
[Tortu/a williamsii Bartr. Am1 = Scopelophila cataractae (Mitt.)
Broth. fide Zander, Bryologist 70: 408, 1967 (1968)]
Tortula willisiana Zand. (nom. nov. for Phascum drummondii Wils.),
see treatment of Tortula Austr1
var. obscura (Willis) Zand. (Phascum drummondii var.), see treatment of Tortula Austr1
Tortula wilsonii (Hook.) Zand. (Gymnostomum) Eur As1 AsS Afr1
var. asperula (Mitt.) Zand. (Pottia), see treatment of Pottia Eur
var. crinita (Wils. ex B.&S.) Zand. (Pottia), see treatment of Pottia
Eur As5 Afr1
var. mucronifolia (Bruch in F. Miill.) Zand. (Entosthymenium), see
treatment of Pottia As1 AsS
Tortula xerophila Hen. Arn4
Tortula yuennanensis Chen As2
Tortula zoddae Zand. (nom . nov. for Portia cuneifolia Solms ex
Schimp.), see treatment of Tortula Eur
TRACHYCARPIDIUM Broth.
Trachycarpidium brisbanicum (C. Miill.) Stone, Muelleria 3: 122,
1975 (Acaulon) As4 Austr1
Trachycarpidium echinatum Dix. As4
Trachycarpidium lonchophyllum (Roth) Zand. (Astomum), see treatment of Trachycarpidium Am5
[Trachycarpidium novae-valesiae Broth. ex Roth. Austr1 = Bryobartramia novae-valesiae (Broth.) Stone & Scott, J. Bryol. 7: 604,
1973, Encalyptaceae]
Trachycarpidium tisserantii Dix. & P. Yarde Afr2
Trachycarpidium verrucosum (Besch.) Broth. Oc
TRACHYODONTIUM Steere, Bryologist 89: 17, 1986
Trachyodontium zanderi Steere, Bryologist 89: 17, 1986 Am4
[TRICHOSTOMOPSIS Card. = Didymodon Hedw. fide Zander,
Phyto1ogia 41: 14, 1978]
[Trichostomopsis aaronis (Lor.) Agnew & Towns., Israel J. Bot. 19:
258, 1970 Eur As5 Afr1 = Didymodon aaronis (Lor.) Guerra in
Guerra & Ros, Cryptogamic Bryol. Lichenol. 8: 55, 1987]
[Trichostomopsis australasiae (Hook & Grev.) Robins., Phytologia
20: 187, 1970 (Tortula) Eur Afr4 Am1 Am2 Am4 Am6 Austr1
Austr2 = Didymodon australasiae (Hook. & Grev.) Zand. fide
Zander, Phytologia 41: 21, 1978]
[Trithostomopsis brevifolia Bartr. Afr1 Am1 Am2 = Trichostomopsis
australasiae (Hook. & Grev.) Robins., Phytologia 20: 187, 1970
= Didymodon australasiae (Hook. & Grev.) Zand. fide Zander,
Phytologia 41: 21, 1978]
[Trichostomopsis crispifolia Card. Am2 = Trichostomopsis umbrosus
(Card.) Robins., Phytologia 20: 185, 1970 = Didymodon
umbrosus (C. Miill.) Zand., Phytologia 41: 22, 1978]
fo. crassiretisTher., Smiths. Misc. Coll. 85(4): 9, 1931 Am1
Trichostomopsis curvipes (C. Miill.) Robins., Phytologia 20: 186,
339
1970 (Barbula) Am6
[Trichostomopsis diaphanobasis (Card.) Grout Am1 Am2 =
Trichostomopsis australasiae (Hook. & Grev.) Robins.,
Phytologia 20: 187, 1970 = Didymodon australasiae (Hook. &
Grev.) Zand.fide Zander, Phyto1ogia 41: 21, 1978]
[Trichostomopsis fayae Grout Am1 Am2 = Didymodon
australasiae (Hook. & Grev.) Zand.fide Zander, Phytologia 41:
21, 1978]
[Trichostomopsis haussknechtii (Jur. & Milde) Agnew & Towns.,
Israel J. Bot. 19: 258, 1970 (Barbula) As5 = Didymodon
haussknechtii (Jur. & Mild.) Broth.]
[Trichostomopsis trivia/is (C. Miill.) Robins., Phytologia 20: 187,
1970 (Barbula) Eur Afr4 = Didymodon trivia/is (C. Miill.)
Guerra in Guerra & Ros, Cryptogamic Bryol. Lichenol. 8: 64,
1987]
[Trichostomopsis umbrosa (C. Miill.) Robins., f>hytologia 20: 185,
1970 (Barbula) Am2 = Didymodon austra/asiae var. umbrosus
(C. Miill.) Zand. fide Zander, Cryptogamic Bryol. Lichenol. 2:
400, 1981 (1982) = Didymodon umbrosus (C. Miill.) Zand.]
fo. propagulifera Brogues, Acta Phytotax. Barcinon. 21: 15, 1976
[1977] nom. inval. sin. descr. lat. et holotyp. non cit. Eur
TRICHOSTOMUM Bruch
Trichostomum abyssinicum (Ther.) Zand. (Weissia), see treatment
of Trichostomum Afr2
Trichostomum acutiuscu/um (Broth.) Zand. (Hyophila), see treatment of Trichostomum Afr2
Trichostomum acutum Hen. Am4
Trichostomum aduncum Par. Oc
Trichostomum aequatoriale Spruce ex Dix. Afr4 Am4
Trichostomum affine Warnst. hom. il/eg. Eur
Trichostomum alpinum Kindb. Am1
[Trichostomum andinum Sull. Am4 = Erythrophyllopsis andina
(Sull.) Zand., Bryologist 80: 159, 1977 Erythrophyllastrum
andinum (Sull.) Zand., see treatment of Erythrophyllastrum]
[Trichostomum angustifolium Crum & Steere Am3 = Pseudosymblepharis angustifolia (Crum & Steere) Zand., see treatment of
Pseudosymblepharis]
Trichostomum apophysatulum Hen. Am4
Trichostomum arboreum (Mitt.) Zand. (Weissia), see treatment of
Trichostomum Am5
Trichostomum arcticum Kaal. Eur As1 Am1
[Trichostomum ardjunense Fleisch. As4 =Trichostomum crispulum
Bruch in F. Miill . fide Norris & Koponen, Acta Bot. Fenn. 137:
96, 1989]
[Trichostomum aristatulum Broth. Austr1 = Trichostomum
brachydontium Bruch in F. Miill. fide Norris & Koponen, Acta
Bot. Fenn. 137: 96, 1989]
Trichostomum aristatulum (Broth.) Hi1p. ex Chen hom. illeg. As2
Trichostomum atrocaule (Saito) Zand. (Weissia), see treatment of
Trichostomum As2
Trichostomum austrocrispum (Beck.) Zand. (Phascum), see treatment of Trichostomum Austr1 Austr2
var. longifolium (R. Br. ter) Zand. (Phascum longifolium), see
treatment of Trichostomum Austr2
[Trichostomum atrorubens Besch. As2 As3 = Bryoerythrophyllum
wallichii (Mitt.) Chenfide Saito, J. Hattori Bot. Lab. 39: 479,
1975]
Trichostomum ayresianum Schimp. ex Besch. Afr3
Trichostomum barbuloides (Broth.) Chen hom. illeg. As2
[Trichostomum bartramii H. Miller, J. Hattori Bot. Lab. 30: 272,
1967 (Trichostomum mauiense Broth. hom. illeg.) Oc = Trichostomum crispulum Bruch in F. Miill. fide Norris & Koponen,
=
GENERA OF THE POTTIACEAE
340
Acta Bot. Fenn. 137: 96, 1987]
[Trichostomum bartramii lwats. & Tan, Misc. Bryol. Lichenol. 7:
152, 1977 hom. illeg. As4 = Trichostomum brevifolium Bartr.
hom. il/eg.) = Trichostomum philippenense Iwats. & Tan,
Kalikasan 8: 195, 1979]
[Trichostomum baurianum Warnst. ex Amann Eur = Barbula
spadicea (Mitt.) Braithw. fide Geissler, Candollea 40: 198, 1985
=Didymodon spadiceus (Mitt.) Limpr.]
Trichostomum be/Iii Bartr. Am4
[Trichostomum bermudanum (Mitt.) Par. Am1 = Weissiajamaicensis
(Mitt.) Grout, see treatment of Weissia]
Trichostomum bombayense C. Miill. As3
Trichostomum borbonicum Een, Lindbergia 3: 217, 1976 (nom. nov.
for Trichostomum glaucoviride Ren. & Card. hom. illeg. non C.
Miill. 1882) nom. il/eg. incl. spec. prior., Trichostomum cardotii
Biz. [= Trichostomum cardotii Biz. nom. inval. basion. non cit.]
Trichostomum borneense (Dix.) Zand. (Stephanodictyon), see treatment of Trichostomum Dix. As4
Trichostomum brachydontium Bruch in F. Miill. Eur As1 As2 As3
As4 As5 Afr1 Afr2 Afr3 Afr4 Am1 Am2 Am3 Am4 Am6
suhsp. cuspidatum (Braithw.) Giac. Eur
suhsp. densum (B.&S . in BSG) Giac. Eur Afr1
suhsp. mutabile (Bruch) Giac. Eur As1 As2 As5 Afr1 Am2 Am3
Am4
var. angustifolium (Lindh.) Wijk & Marg. Eur
var. antillarum Bartr. in Orcutt Am3
var. cophocarpum (Schimp.) P. Cout. Eur
var. cuspidatum (Braithw.) Sav. Eur
var. cylindricum (Schimp.) P. Cout. Eur
var. devonicum (Podp.) Wijk & Marg. Eur
var. esquirolii (Ther.) Chen As2
var. littorale (Mitt.) C. Jens. in Weim. Eur Afrl
var. longifolium (Warns!.) Wijk & Marg. Eur
var. lutescens (Lindh.) Wijk & Marg. Eur
var. nigroviride (Ren. & Card.) Luis. Afrl
var. robustum (Ren. & Card.) During, Lindbergia 7: 121, 1981 Afr2
var. sibiricum (Podp.) Laz. As1
var. unguiculatum (Philih.) Corh. & Jah. Eur
[Trichostomum brevifolium Bartr. As4 hom. il/eg.
Trichostomum
bartramii lwats. & Tan, Misc. Bryol. Lichenol. 7: 152, 1977 hom.
illeg. = Trichostomum philippenense Iwats. & Tan, Kalikasan 8:
195, 1979]
Trichostomum brittonianum Zand. (nom . nov. for Hymenostomum
flavescens E. Britt. in N. Britt. & Millsp.), see treatment of
Trichostomum Am3
[Trichostomum bulbilliferum Herz. Am5 = Oxystegus tenuirostris var.
gemmiparus (Schimp.) Zand., Lindbergia 4: 285, 1978 = Trichostomum tenuirostre var. gemmiparum (Schimp.) Zand., see treatment of Trichostomum]
[Trichostomum burmense Bartr. As4 Oxystegus burmensis (Bartr.)
Gangulee, Moss. E. India 653 , 1972 = Trichostomum tenuirostre
(Hook. & Tayl.) Lindh. fide Sollman, Lindbergia 10: 55, 1984]
Trichostomum caespitosum (Bruch ex Brid.) Jur. [= Pottia caespitosa
(Bruch ex Brid.) C . Miill. fide Corley et al., J. Bryol. 11: 620,
1981 (1982)], see treatment of Trichostomum Eur
var. mucronatum Pilous Eur
Trichostomum calymperaceum Broth. & Par. Afr2
[Trichostomum cardotii Biz., Rev. Bryol. Lichenol. 40: 120, 1974
nom. inval. basion. non cit. (nom . nov. for Trichostomum
glaucoviride Ren. & Card.) Afr3 = Trichostomum borbonicum
Een, Lindbergia 3: 217, 1976]
Trichostomum carinatum Bartr. As4
Trichostomum castaneum (Crum & Steere) Zand. (Hymenostomum),
=
=
see treatment of Trichostomum Am3
[Trichostomum cavernarum Broth. Am5 = Pseudosymblepharis
cavernarum (Broth.) Zand., see treatment of Pseudosymblepharis]
[Trichostomum chlorphyllum C. Miill. Am2 = Weissia jamaicensis
(Mitt.) Grout fide Zander in Sharp et al., Moss Fl. Mex.]
[var. brevifolium Ther. Am2 = Trichostomum crispulum Bruch in
F. Miill.fide Zander in Sharp et al., Moss Fl. Mex.]
[Trichostomum circinnatulum Broth. in Voltzk. Afr3 = Pseudosymblepharis circinnatula (Broth. in Voltzk) Zand., see treatment
of Pseudosymblepharis]
Trichostomum clavinerve (Card. & P. Yarde) H. 0 . Whitt. in H. 0.
Whitt. & B. Whitt., Bryologist 77: 433, 1974 (Weissia) Oc
[Trichostomum clintonii C. Miill. Am2 = Oxystegus tenuirostris
(Hook. & Tayl.) A. Sm. fide Zander in Sharp et al., Moss Fl.
Mex. = Trichostomum tenuirostre (Hook. & Tayl.) Lindh.]
Trichostomum compactulum C. Miill. Am6
Trichostomum connivens (Broth.) Par. Eur
Trichostomum contortum (Kunze) Sergio, Portug. Acta Bioi. (B)
14: 169, 1985 (Hymenostomum) Afrl
Trichostomum contractum Zand., see treatment of Trichostomum
(nom . nov. for Gymnostomum longirostre Griff.) As2
Trichostomum criotum Zand. (nom. nov. for Hyophila perannulata
Ren. & Card.), see treatment of Trichostomum As3
Trichostomum crispulum Bruch in F. Miill. Eur As1 As2 As5 Afr1
Afr2Am1 Am2
suhsp. brevifolium (B .&S. in BSG) Giac. Eur Afr1
suhsp. hammerschmidii (Loeske & Paul) Amann Eur
var. acuminatum Meyl. Eur
var. algarvicum Schimp. Eur Afr1
var. armatum (Ther. & Trah.) Biz. Afrl
var. brevifolium B.&S. Eur Afr1
var. cucul/atum (Roth) Podp. Eur Am1
var. elatum Schimp. Eur As5 Afr1
var. hammerschmidii (Loeske & Paul) Blumr. Eur
var. mucronatulum (Card.) Podp. Eur Afrl
var. muticum Podp. Eur
var. nigroviride (Braithw.) Dix. Eur
var. pseudoweisia Schimp. Eur Afr1
var. sudeticum Velen. Eur
fo. brevissimum Reim., Hedwigia 79: 261, 1940 Eur
fo. longifolium Bouvet, Bull. Soc. Et. Sc. Angers 26: 84, 1896
(Trichostomum crispulum var. longifolium Schimp. nom. illeg.)
Eur
fo. tophaceum Latz., Bot. Centralhl. Beih. 48(2): 242, 1931 Eur
[Trichostomum crispulum Besch. Am2 hom. il/eg. = Oxystegus
tenuirostris (Hook. & Tayl.) A. Sm. (as Oxystegus cylindricus
(Brid.) Hilp.) fide Crum, Appalachian-Ozarkian Element Moss
Fl. Mexico (dissertation, Univ. Microfilms) 137, 1951 _
Trichostomum tenuirostre (Hook. & Tayl.) Lindh.]
[Trichostomum cuspidatissimum Card. & Ther. Eur As1 Am1 =
Trichostomum arcticum Kaal. fide Frisvoll, Bryologist 81: 156,
1978]
[Trichostomum cuspidatum (Doz. & Molk.) Doz. & Molk. As2 As4
= Trichostomum tenuirostre (Hook. & Tayl.) Lindh. fide
Sollman, Lindbergia 10: 55, 1984]
[Trichostomum cylindricum (Brid.) C. Miill. nom. inval. =
Oxystegus tenuirostris (Hook. & Tayl.) A. Sm., J. Bryol. 9:
393, 1977 Trichostomum tenuirostre (Hook. & Tayl.) Lindh.]
var. asperum Podp., Shorn. Kluh. Ptirod. Bmo 5: 11 , 1923 Eur
var. cataractarum (Loeske) Culm. (Streblotrichum croceum var.)
Eur
[fo. longifolia Latz., Hedwigia 66: 138, 1926 hom. il/eg. Eur
=
=
341
GENERA, SPECIES AND INFRASPECIFIC TAXA
Oxystegus cylindricus fo. longifolia Podp.]
[Trichostomum cylindrotheca (Mitt.) Broth. As3 = Oxystegus
cylindrothecus (Mitt.) Gangulee, Nov. Hedw. 12: 430, 1966 =
Trichostomum bombayense C. Miill.fide Townsend, J. Bryol. 12:
561, 1983]
Trichostomum deciduaefolium (Saito) Zand. (Weissia), see treatment
of Trichostomum As2
Trichostomum decurvifolium Dix. Afr2
Trichostomum distans Hampe in C. Miill. Afr2
Pseudosymblepharis angustata
[Trichostomum dubium Ther. Oc
(Mitt.) Hilp. fide Norris & Koponen, Acta Bot. Fenn. 137: 94,
1989]
Trichostomum duidense Bartr. Am5 [= Trichostomum brachydontium
Bruch in F. Muell.fide Sollman, Lindbergia 10: 55, 1984 but recognized here as a good species]
Trichostomum edentulum Broth. in Herz. Am4
[Trichostomum ehrenbergii Lor. = Barbula ehrenbergii (Lor.)
Fleisch.]
var. denticuspis Broth., Bot. Jahrb. 30: 261, 1901loc?
[Trichostomum ekmanii Ther. Am3 = Weissia jamaicensis (Mitt.)
Grout, see treatment of Trichostomum]
Trichostomum el/iottii Broth. ex Dus. Am6
[Trichostomum etessei Broth. & Par. Oc. = Trichostomum crispulum
Bruch in F. Miill. fide Norris & Koponen, Acta Bot. Fenn. 137:
96, 1989]
Trichostomum exulatum (C. Miill. ex Roth) Zand. (Tetrapterum), see
treatment of Trichostomum Am5
Trichostomumfal/aciosum Welch & Crum Am3
Trichostomumfal/ax Herz. Am4
fo. minutum Herz., Biblioth. Bot. 87: 92, 1916 Am4
Trichostomumflavescens Dix. Afr2
Trichostomum flavisetum DC. in Lam. & DC. Eur [= Ditrichum sp.
fide W. Margadant in !itt. DC]
Trichostomumfragilifolium Dix. Afr2
[Trichostomum fuscomucronatum C. Miill. Afr2 = Pseudocrossidium
rep/icatum (Tayl.) Zand. fide Townsend, Lindbergia 10: 178,
1985]
[Trichostomum glaucoviride Ren. & Card. Afr3 hom. illeg. Trichostomum cardotii Biz., Rev. Bryol. Lichenol. 40: 120, 1974 nom.
inval. basion. non cit. (nom . nov. for Trichostomum glaucoviride
Ren. & Card.) = Trichostomum borbonicum Een, Lindbergia 3:
217, 1976]
Trichostomum gracillimum C. Miill. Am6
Trichostomum hibernicum (Mitt.) Dix. Eur
Trichostomum hyalinoblastum (Broth.) Broth. As3
Trichostomum imshaugii (Vitt) Zand., see treatment of Trichostomum
(Barbula) Austr2
Trichostomum incertum (Mitt.) Zand. (Weissia), see treatment of
Trichostomum Afr3
[Trichostomum inclinans Schimp. ex Besch., Mem. Soc. Nat. Sci.
Natl. 16: 176, 1872 = Rhamphidium dicranoides (C. Miill.) Par.
fide Zander in Sharp et al., Moss Fl. Mex.]
Trichostomum insulare (Besch.) Broth. Oc
Trichostomum involutum Sull. Am3 Am4
var. minus Sull. Am3
Trichostomum involutum Broth. As2 hom. illeg.
[Trichostomum involvens Card. Am2 = Trichostomum crispulum
Bruch in F. Miill.fide Zander in Sharp et al., Moss Fl. Mex.]
[Trichostomum jamaicense (Mitt.) Jaeg. Am1 Am2 Am3 = Weissia
jamaicensis (Mitt.) Grout fide Zander in Sharp et aL, Moss Fl.
Mex.]
Trichostomum kanieriense R. Br. ter Austr2
[Trichostomum khasianum (Mitt.) Broth. As3 Oxystegus khasianus
=
=
=
(Mitt.) Gangulee, Nov. Hedw. 8: 149, 1964 =
Psuedosymblepharis khasiana (Mitt.) Zand., see treatment of
Pseudosymblepharis]
Trichostomum knightii Hampe ex C. Miill. Am2
Trichostomum lambii Bartr., Rev. Bryol. Lichenol. 33: 324,
1964-65 [1965] Am5
[Trichostomum lamprothecium C. Miill. Am2 = Trichostomum
crispulum Bruch in F. Miill. fide Zander in Sharp et al., Moss
Fl. Mex.]
[Trichostomum lanuginosum Hedw. Rhacomitrium lanuginosum
(Hedw.) Brid.]
var. incanum Hornsch. Am1
Trichostomum /aticostatum Ther. Oc
Trichostomum /eptocylindricum C. Miill. Am5
[Trichostomum leiodontium C. Miill. = (Torre/la xanthocarpa (C.
Miill.) Broth. fide Broth, Nat. Pfl. 1(3): 397, 1902) =Torre/la
humi/is (Hedw.) Jenn.fide Sollman, Lindbergia 16: 22, 1990]
Trichostomum leptotheca C. Miill. Austr1
Trichostomum lignicola Herz. Am5
Trichostomum ligulaefolium (Broth. & Par.) Zand. (Hyophila), see
treatment of Trichostomum Afr2
Trichostomum lil/ei Dix. As2
Trichostomum lindigii (Hampe) Zand. (Systegium), see treatment of
Trichostomum Am4
[Trichostomum linea/ifolium C. Miill. Am4 = Weissia jamaicensis
(Mitt.) Grout, see treatment of Weissia]
[Trichostomum lingulatum Hook. f. & Wils., Fl. N. Zeal. 2: 71,
1854 = Didymodon tophaceus (Brid.) Lisa, see treatment of
Didymodon]
Trichostomum lorifolium Broth. & Par. Afr2
[Trichostomum makanuiense R. Br. ter, Trans. N. Zeal. lnst. 35:
332, 1903 nom. inval. in Robinson, Phytologia 20: 187, 1970 =
Trichostomopsis australasiae (Hook. & Grev.) Robins. loc. cit.
Didymodon australasiae (Hook. & Grev.) Zand. var.
australasiae]
[Trichostomum marginatum Robins., Phytologia 12: 389, 1971
Arn4 = Didymodon marginatum (Robins.) Zand., see treatment
of Didymodon]
[Trichostomum mauiense Broth. hom. illeg. Oc
Trichostomum
bartramii H. Miller, J. Hattori Bot. Lab. 30: 272, 1967 =
Trichostomum crispulum Bruch in F. Miill. fide Norris &
Koponen, Acta Bot. Fenn. 137: 96, 1989]
Trichostomum melanostomum (Mitt.) Zand. (Weissia) Am5
Trichostomum mildeanum Jur. As5
Trichostomum minusculum Dix. & P. Yarde As3
Trichostomum minutissimum Sak. Oc
Trichostomum mitteneanum Zand. nom. nov. (Weissia umbrosa
Mitt.), see treatment of Trichostomum Am4
[Trichostomum molariforme Zand., Bryologist 85: 126, 1982 Am1
Am2 Am3 = Trichostomum portoricense Crum & Steere, see
treatment of Trichostomum]
[Trichostomum mol/issimum (Broth. ex Bartr.) Crum Am1 Am2
Am3 Am5 = Pseudosymblepharis schimperiana (Par.) Crum
fide Zander in Sharp et al., Moss Fl. Mex.]
Trichostomum mouense Broth. & Par. Oc
[Trichostomum mutabile Bruch in De Not. - Trichostomum
brachydontium subsp. mutabile (Bruch) Giac.]
var. alpinum Amann Eur
var. brevifolium Schiffn. Eur
var. majus Podp. Eur
[var. robustum Ren . & Card. Afrl = Trichostomum
brachydontium var. robustum (Ren. & Card.) During,
Lindbergia 7: 121, 1981]
=
=
=
342
GENERA OF THE POTTIACEAE
var. subalpinum BSG nom. inval. prov. Eur
fo.litorale-mutabile Herz., Abh. Leop.-Car. Ak. Naturf. 73(3): 471,
1907 Eur
fo. mutabile-cuspidatum Herz., Abh. Leop.-Car. Ak. Naturf. 73(3):
476, 1907 Eur
subfo.? crispulomutabile Herz., Abh. Leop.-Car. Ak. Naturf. 73(3):
471, 1907 Eur
subfo.? flaccidolitorale Herz., Abh. Leop.-Car. Ak. Naturf. 73(3):
471, 1907 Eur
subfo.? strictolitorale Herz., Abh. Leop.-Car. Ak. Naturf. 73(3):
470, 1907 Eur
Trichostomum muticum Par. Afr3
[Trichostomum nigrescens P. Beauv., Prodr. 47, 1805 (Bryum
nervosum Vill. 1789 nom. inval.) Eur = p.p. Racomitrium
aquaticum (Schrad.) Brid. (Grimmiaceae) cf. Brid., Muse. Rec.
2(3): 51, 1803)]
[Trichostomum nitidum (Lindb.) Schirnp. = Tortella nitida (lindb.)
Broth.]
var. irrigatum Wint. Afrl
Trichostomum nordenskioeldii Schimp. in Heer, A. Foss. Arct. 2(3):
88, 1870 Eur
Trichostomum novogranatense Broth. & lrmsch. Am4
[Trichostomum oblongifolium Bartr. Oc = Trichostomum
brachydontium Bruch in F. Muell. fide Sollman, Lindbergia 10:
55, 1984]
[Trichostomum obtusifolium (Schwaegr.) Boul., Fl. Crypt. Est.
Muscin. 491, 1872 hom. illeg. Eur Asl As2 As5 Afr1 Am1
Desmatodon obtusifolius (Schwaegr.) Schimpr.
Tortula
obtusifolia (Schwaegr.) Math. fide Corley et al., J. Bryol. 11: 620,
1981 (1982)]
[Trichostomum orientale Web.= Barbula indica (Hook.) Spreng.]
fo. sterile Aeisch., Musci Fl. Buitenz. 1: 346, 1902 [1904] As4
Trichostomum orthodontum (C. Miill.) Broth. As3
Trichostomum ovatifolium Zand. (nom. nov. for Hymenostomum
anomalum Broth. in Herz.), see treatment of Trichostomum Am4
Trichostomum pallidens (Dix.) Zand. (Pseudosymblepharis), see
treatment of Trichostomum As3
Trichostomum paludicola (Broth.) Hilp. Am4
Trichostomum eckelianum Zand., see treatment of Trichostomum
(nom. nov. for Trichostomum cirrhatum Hampe hom. illeg.)
Austrl
Trichostomum pennequinii Ren. & Par. Afr3
Trichostomum perangustum Besch. Afr4
Trichostomum perannulatum Dix. & P. Yarde As3
Trichostomum perinvolutum Tix., Rev. Bryol. Lichenol. 34: 132,
1966 As3
Trichostomum perligulatum (Flow. ex Crum) Zand. (Weissia), see
treatment of Trichostomum Am1
[Trichostomum perlongifolium J. Frohl. fide As4 = Pseudosymblepharis perlongifolia (Frohl.) Zand., see treatment of Pseudosymblepharis]
Trichostomum perplexum P. Yarde Afr2
Trichostomum perpusillum C. Miill. ex Wamst. Am2
Trichostomum perrieri Ther. Afr2
Trichostomum persicum Jur. & Milde As5
[Trichostomum perviride Broth. Am3 Am4 = Weissia veviridis Zand.
nom. nov., see treatment of Weissia]
Trichostomum philippenense Iwats. & Tan, Kalikasan 8: 195, 1979
(nom. nov. for Tricho.<tomum brevlfolium Bartr.) As4
Trichostomum planifolium (Dix.) Zand. (Weissia), see treatment of
Trichostomum As1 As2 Am2
Trichostomum platyphyllum (lhs.) Chen As2
Trichostomum plicatum C. Miill. Am6
=
=
Trichostomum pomangium Broth. & Herz. in Herz. Am4
Trichostomum portoricense Crum & Steere Aml Am2 Am3
Trichostomum prionodon C. Miill. Am5
Trichostomum pulicare (Besch.) Zand. (Hymenostomum) , see treatment of Trichostomum Afr3
Trichostomum pygmaeum Bartr. Am2
[Trichostomum quitense Hampe in C. Miill. nom. nud. = Tortula
pichinchensis Tayl.]
var. longifolium Herz. comb. inval. Am4
Trichostomum recurvifolium (Tayl.) Zand., see treatment of Trichostomum Eur Aml
[Trichostomum rehmannii Sim Afr4 = Trichostomum tortuloides
Sull. & Lesq. fide Magill & Schelpe, Mem. Bot. Surv. S. Afr.
43: 25, 1979 = Tortella xanthocarpa (C. Miill.) Broth. fide
Magill, Fl. S. Afr. I. Mosses 1: 255, 1981]
Trichostomum rhodesiae Broth. Afr2
Trichostomum rigidulum (Hedw.) Turn. = Didymodon rigidulus
Hedw.]
var. flaccidum Roll nom. inval. prov. Eur
var. rigidum Roll nom. inval. prov. Eur
Trichostomum robustum Broth. ex lhs. As2
Trichostomum ruvenzorense (Broth.) Broth. Afr2
Trichostomum sarawakense Dix. As4
[Trichostomum schlimii C. Miill. Am4 (= ?Oxystegus tenuirostris
(Hook. & Tayl.) A. Sm. (as Oxystegus cylindricus (Brid.) Hilp.)
fide Crum, Appalachian-Ozarkian Element Moss Fl. Mexico
Trichostomum
(dissertation, Univ. Microfilms) 137, 1951)
tenuirostre (Hook. & Tayl.) Lindb., see treatment of Trichostomum]
Trichostomum siamense Broth. in Hosseus, Beih. Bot. Centralbl.
28: 362, 1911 As3
Trichostomum sinaloense (Bartr.) Zand. (Weissia), see treatment of
Trichostomum Am2 Am3
[Trichostomum sitkanum Card. & Ther. Aml = Tortella tortuosa
var. arctica (Amell) Broth. in Fedch., see treatment of Tortella]
Trichostomum soulae (C. Miill. in Ren. & Card.) Zand.
(Ptychomitrium), see treatment of Trichostomum Afr3
var. corticicola (Ren. & Card.) Zand. (Barbula), see treatment of
Trichostomum Afr3
Trichostomum sparsifolium (Ren. & Card.) Card. in Grand. Afr3
Trichostomum spirale Grout Am1 Am2 As2 (= Oxystegus
tenuirostris (Hook. & Tayl.) A. Sm. var. stenocarpus (Ther.)
Zand., Misc. Bryol. Lichenol. 9: 73, 1982) = Oxystegus spiralis
(Grout) Crum & Anderson, Bryologist 92: 533, 1989)
Trichostomum sporaphyllum (Ren. & Card.) Card. ln Grand. Afr3
Trichostomum stanilandsii R. Br. ter Austr2
[Trichostomum stenocarpum Wils. in Seem., Bot. Voyage Herald
344, 1857 Am2 =Atractylocarpus stenocarpus (Wils. in Seem.)
Zand., Bryologist 85: 128, 1982]
[Trichostomum stenocarpum (Ther.) Crum, Bryologist 84: 390,
1981 hom. illeg. (Weisiopsis) Am2 = Oxystegus tenuirostris
var. stenocarpus (Ther.) Zand., Misc. Bryol. Lichenol. 9: 73,
1982 = Trichostomum spirale Grout, see treatment of Trichostomum]
Trichostomum stenophyllum (Mitt.) Broth. [see discussion of
Townsend, J. Bryol. 12: 563, 1983] As2
[Trichostomum striatum (P. Beauv.) Am. Aml = p.p. Racomitrium
patens (Hedw.) Hiib. cf. Amott foe . cit.]
Trichostomum subangustifolium (Ther.) Zand. (Hyophila), see treatment of Trichostomum Am2
Trichostomum subcirrhatum Hampe Am5
Trichostomum subconnivens Ther. Am3
Trichostomum subdenticulatum Aust. Aml hom. illeg.
=
343
GENERA, SPECIES AND INFRASPECIFIC TAXA
Trichostomum subintegrum (Broth.) Broth. Afr2
Trichostomum sublamprothecium Par. Am3
Trichostomum subminusculum Dix. & P. Yarde As3
[Trichostomum subulifolium Bartr. As4
Pseudosymblepharis
angustata (Mitt.) Hilp. fide Norris & Koponen, Acta Bot. Fenn.
137: 94, 1989]
Trichostomum sumatranum Baumg. in J. Fri:ihl. As4
[Trichostomum svihlae Bartr. As4 = Oxystegus svihlae (Bartr.)
Gangulee, M. E. India 6S4, 1972 = Tuerckheimia svih/ae (Bartr.)
Zand., see treatment of Tuerckheimia]
[Trichostomum syrrhopodontoides Herz. Am4 = Trichostomum
tenuirostre (Hook. & Tayl.) Lindh. fide W. Steere, in litt. having
seen type at JE]
Trichostomum tenuirostre (Hook. & Tayl.) Lindh. Eur Asl As3 Afrl
=
Afr2Afr3Af~Am1Am2Am40c
var. gemmiparum (Schimp.) Zand., see treatment of Trichostomum
Eur As3 Am2 Am3 AmS
var. holtii (Braithw.) Dix. Eur Aml
Trichostomum termitarum (C. Miill.) Zand. (Weissia), see treatment
of Trichostomum AmS
[Trichostomum tophaceum Brid.
Didymodon tophaceus (Brid.)
Lisa]
var. brevifo/ium Lindh. hom. il/eg. Eur
var. leucotrichion Brid. Eur
[fo. acutifolium Boul., Mouss. France 449, 1884 Eur Didymodon
tophaceus fo. acutifolius (Boul.) Zodda, Malphighia 22: S09,
1908]
fo. brevicau/e (Schimp.) Boul., Muscin. France 449, 1884 (TricJwstomum tophaceum var.) Eur
[fo. cylindricum Boul., Muscin. France 449, 1884 Eur =Didymodon
tophaceus fo. cylindrica (Boul.) Limpr., Laubm. Eur. 1: SS4,
1888]
[fo. e/atum Boul., Muscin. France 449, 1884 Eur Didymodon
tophaceus fo. elatus (Boul.) Artaria in E. Bauer, Musci Eur. Am.
Exsicc. 37: n. 1823, 192S]
[fo. lingulatum Boul., Muscin. France 449, 1884 Eur = Didymodon
tophaceus fo. /ingulatus (Boul.) Monk., Siisswasserfl. 14: 67,
1914]
[fo. recurvifolium Boul., Muscin. France 449, 1884 Eur = Didymodon tophaceus fo. recurvifolius (Boul.) De Willd., Prodr. 434,
1899]
[fo. truncatum Boul., Muscin. France 449, 1884 Eur = Didymodon
tophaceus fo. truncatus (Boul.) Limpr., Laubm. Deutsch. 1: SS4,
1888]
Trichostomum tortelloides (Broth. & Dix.) Zand. (Calymperes), see
treatment of Trichostomum As3
[Trichostomum tortuloides Sull. & Lesq. Afr4 =Tarte/la xanthocarpa
(C. Miill.) Broth. fide Magill, Fl. S. Afr. I. Mosses 1: 2SS, 1981
(1982)]
[Trichostomum trifarium (Hedw.) Sm.
Didymodon trifarius
(Hedw.) Ri:ihl. nom. dub. fide Zander, Cryptogamic Bryol.
Lichenol. 2: 414, 1981 (1982)]
var. sterile Mazz. Eur
Trichostomum trirete Dix. Afr2
[Trichostomum triumphans De Not. ex Schimp. Eur AsS Afr1 = Weissia triumphans (De Not.) Hill, J. Bryol. 11: 600, 1981 (1982)]
[subsp. monspeliense (Schimp.) Boul. Eur Afr1
Weissia
triumphans var. monspeliensis (Schimp.) Zand., see treatment of
Weissia]
subsp. pallidisetum (H. Miill.) Giac. Eur AsS Afrl
subsp. philibertii (Schimp.) Boul. Eur Afrl
var. azoricum (Card.) Podp. Eur Afrl
var. brachyodon (Spindl.) Podp. Eur
=
=
=
=
=
=
[var. monspeliense (Schimp.) Boul. Eur Afr1
Weissia
triumphans var. monspeliensis (Schimp.) Zand., see treatment
of Weissia]
var. pallidisetum (H. Miill.) Husn. Eur AsS Afr1
var. philibertii (Schimp.) Husn. Eur Afrl
Trichostomum tucumanense Bartr., Rev. Bryol. Lichenol. 33: 324,
1964-6S [ 196S] AsS
Trichostomum unguiculatum (Mitt.) Zand., see treatment of Trichostomum Afr2 Afr4
Trichostomum uncifolium Dix., Anniv. Vol. Bot. Gard. Calcutta
180, 1942 As3
Trichostomum urceolare (Hampe) Zand. (Hyophila), see treatment
of Trichostomum AmS
Trichostomum usambaricum (Broth.) Broth. Afr2
Trichostomum villaumei Ther. Afr3
[Trichostomum viridulum Bruch in F. Miill. .Eur AsS Afr1 [=
Trichostomum crispulum fo. longifolium Bouvet] = Trichostomum crispulum Bruch in F. Miill. fide Corley et al., J. Bryol.
11:623, 1981 (1982)]
Trichostomum wagneri (C. Miill.) Broth. Am4
Trichostomum weisioides C. Miill. AmS
Trichostomum whittonii R. Br. ter Austr2
Trichostomum williamsii Zand. (nom. nov. for Astomum chilense
Williams), see treatment of Trichostomum Am4
[TRIDONTIUM Hook. f. in Hook. referred to Grimmiaceae, see
Excluded Taxa]
[Tridontium tasmanicum Hook. f. in Hook. As4 Austr1 Austr2 =
Grimmiaceae, see Excluded Taxa]
TRIQUETRELLA C. Miill.
Triquetrella arapilensis Luis. Eur
Triquetrella californica (Lesq.) Grout Am1
[Triquetrella ferruginea (Besch.) Ther. Am2 = Didymodon fa/lax
var. rejlexus (Brid.) Zand. fide Zander, Bryologist 83: 230,
1980 = Didymodon rigidicaulis (C. Miill.) Saito fide Saito, J.
Hattori Bot. Lab. 39: S02, 197S = Didymodon ferrugineus
(Schimp. ex Besch.) Hill, J. Bryol. 11: S99, 1981 (1982)]
[Triquetrel/a filicaulis Dus. Am6 Afr4 = Triquetrel/a patagonica
fo.filicaulis (Dus.) Herz.]
Triquetrellafragilis C. Miill. Austr1
[Triquetrella japonica Dix. in Sak. As2 nom. inval. descr. angl. =
Didymodon rigidicaulis (C. Miill.) Saito fide Saito, J. Hattori
Bot. Lab. 39: S02, 197S = Didymodonferrugineus (Schimp. ex
Besch.) Hill, J. Bryol. 11: S99, 1981 (1982)]
[Triquetrella nipponensis Sak. As2 = Dichodontium pellucidum
(Hedw.) Schimp. (Dicranaceae) fide Saito, J. Hattori Bot. Lab.
39: S02, 197S]
Triquetrella papillata (Hook. f. & Wils.) Broth. Austrl Austr2
Triquetrella patagonica C. Miill. Am6
fo. filicaulis (Dus.) Herz., Arch. Esc. Farm. Fac. Ci. Med. C6rdoba (Secc. Ci.) 7: 40, 1938 Am6 Afr4
[Triquetrella preissiana (Hampe) C. Miill. Austrl = Triquetrella
papillata (Hook. f. & Wils.) Broth. fide Watts & Whitelegge,
Proc. Linn. Soc. N.S.W. 27: 1-90, 1902 cf. Stone & Scott, Mo.
So. Australia 220, 1976, and, see treatment of Triquetrella]
[Triquetrella recurvifolia Dix. & Sak. = Didymodon rigidicaulis (C.
Miill.) Saito fide Saito, J. Hattori Bot. Lab. 39: S02, 197S =
Didymodonferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11:
S99, 1981 (1982)]
Triquetrella richardsiae C. Miill. Austrl
Triquetrella spiculosa Ther. Am4
Triquetrel/a tasmanica (Broth.) Granzow-de la Cerda, Bryologist
344
GENERA OF THE POTTIACEAE
92: 383, 1989 Austr1
[Triquetrella tenuicaulis Sak. As2 = Didymodon rigidicaulis (C.
MUll.) Saito fide Saito, J. Hattori Bot. Lab. 39: 502, 1975 = Didymodon ferrugineus (Schimp. ex Besch.) Hill, J. Bryol. 11 : 599,
1981 (1982)]
Triquetrel/a tristicha (C. MUll.) C. MUll. Afr4
TUERCKHEIMIA Broth.
[Tuerckheimia angustifolia (Saito) Zand., Misc. Bryol. Lichenol. 8:
27, 1978 nom. inval. (Gymnostomum) Am1 Am2 As2 = Tuerckheimia svihlae (Bartr.) Zand., see treatment ofTuerckheimia]
[Tuerckheimia angustinervis (Card.) Broth. Am1 Am2 Am3 Am5 =
Weissia jamaicensis (Mitt.) Grout fide Zander in Sharp et al.,
Moss A. Mex.]
[Tuerckheimia calculosa Zand. & Eckel, Mem. New York Bot. Gard.
45: 293, 1987 Am2 = Quaesticula navicularis (Mitt.) Zand., see
treatment of Quaesticula]
Tuerckheimia guatemalensis Broth. Am2
[Tuerckheimia linearis (Web. & Mohr) Britt. Am3 Am5 = Oxystegus
linearis (Web. & Mohr) Hilp. fide Zander, Bryologist 85: 128,
1982 = Tortel/a linearis (Web. & Mohr) Zand., see treatment of
Tortella]
[Tuerckheimia longifolia Ther. Am3 = Oxystegus tenuirostris var.
gemmiparus (Schimp.) Zand., Lindbergia 4: 285, 1978 = Trichostomum tenuirostre var. gemmiparum (Schimp.) Zand., see treatment of Trichostomum]
Tuerckheimia robusta (Dix.) Zand. comb. nov., see treatment of
Tuerckheimia (Merceyopsis) As3
Tuerckheimia svihlae (Bartr.) Zand. (see treatment of Tuerckheimia)
Ther. Afr2 = Trichostomum abyssinicum (Ther.) Zand., see treatment of Trichostomum]
Tuerckheimia valeriana (Bartr.) Zand., Misc. Bryol. Lichenol. 8: 27,
1978 (Leptodontium) Am2
ULEOBRYUM Broth.
Uleobryum curtisii Stone, J. Bryol. 13: 19, 1984 Austr1
Uleobryum peruvianum Broth. Am2 Am3 Am4 Austr1
Uleobryum occultum (Roth) Zand. (Aschisma), see treatment of UleobryumAm3
WEISIOPSIS Broth.
[Weisiopsis angulosa (Broth. & Dix.) Hilp. As3
Ganguleea
angulosa (Broth. & Dix.) Zand., Phytologia 65: 427, 1989]
Weisiopsis anomala Broth. & Par. As2
Weisiopsis bahiensis (C. MUll.) Broth. Am5
[Weisiopsis hol/andii Bartr. Oc = Luisierel/a barbula (Schwaegr.)
Steere, see treatment of Luisierel/a]
Weisiopsis cucullatifolia (Gao, Jia & Cao) Zand. (Hyophila), see
treatment of Weisiopsis As2
[Weisiopsis hyophiloides Dix. & Ther. As2 = Trichostomum
platyphyllum (lhs.) Chen p.p. & Hyophila involuta (Hook.) Jaeg.
p.p.fide Saito, J. Jap. Bot. 47: 17, 1972]
[Weisiopsis involuta Magill, A. S. Afr. I. Mosses 1: 225, 1981 (1982)
Afr4
Plaubelia involuta (Magill) Zand., see treatment of
Plaubelia]
Weisiopsis nigeriana (Egun. & Olar.) Zand. (Gyroweisia), see treatment of Weisiopsis Afr2
Weisiopsis norrisii (Zand.) Zand. (Scopelophila), see treatment of
Weisiopsis Am2
Weisiopsis oblonga Ther. Am2
Weisiopsis plicata (Mitt.) Broth. Afr2 Afr3 Afr4
[Weisiopsis pulchriretis Dix. Afr4 = Streptocalypta pulchriretis (Dix.)
Zand., see treatment of Streptocalypta]
=
=
[Weisiopsis spathulifolius Crum & Bartr. Am3 = Neohyophila
sprengelii (Schwaegr.) Crum s. lat. fide Zander, Bryologist 86:
137, 1983 = Plaubelia sprengelii (Schwaegr.) Zand., see treatment of Plaubelia]
[Weisiopsis stenocarpa Ther. Am2 = Trichostomum stenocarpum
(Ther.) Crum, Bryo1ogist 84: 390, 1981 = Oxystegus
tenuirostris var. stenocarpus (Ther.) Zand., Misc. Bryol.
Lichenol. 9: 73, 1982 = Trichostomum spirale Grout, see treatment of Trichostomum]
WEISSIA Hedw.
Weissia abbreviata (Thwait. & Mitt.) Zand. (Systegium), see treatment of Weissia As3 As4
[Weissia abyssinica Ther. Afr2 = Trichostomum abyssinicum
(Ther.) Zand., see treatment of Trichostomum]
[Weissia acuta Hedw. 8/india acuta (Hedw.) BSG]
var. curvu/a Hartm. Eur
Weissia alianuda Tan, Mem. New York Bot. Gard. 68 : 5, 1992
(nom. nov. for Weissia nuda Mitt.) Austr1
[Weissia amblyphylla (Dix.) Dix. As3 hom . illeg. non Zett 1869
Weissia norketii R. S. Chopra, Taxon. Indian Mosses 125,
1975]
Weissia andersoniana Zand., Monogr. Syst. Bot. Missouri Bot.
Gard. 11: 195, 1985 (nom. nov. for Weissia glauca Bartr. hom.
i/leg.) Am1 Am2
[Weissia andrewsii Bartr. Am1 Am2 = Weissia controversa Hedw.
fide Stoneburner & Wyatt, Monogr. Syst. Bot. Missouri Bot.
Gard. 11: 184, 1985 and, Bryologist 88: 302, 1985]
Weissia argentinica C. MUll. Am6
Weissia artocosana (Mitt.) Zand. (nom. nov. for Pseudosymblepharis socotrana (Mitt.) Ther.), see treatment of Weissia Afr2
[Weissia atrocaulis Saito, J. Hattori Bot. Lab. 39: 425, 1975 As2 =
Trichostomum atrocaule (Saito) Zand., see treatment of Trichostomum]
Weissia austrocrispa (Beckett) Stone, J. Bryol. 11: 236, 1980
Austr2
Weissia ayresii Schimp. in Besch. Afr3
Weissia balansae (C. MUll.) Zand. (Phasconica), see treatment of
Weissia Austr1 Oc
Weissia balansaeana (Besch.) C. MUll. Am5 Am6
Weissia bizotii Zand. (nom. nov. for Kleioweisiopsis involuta Biz.),
see treatment of Weissia Afr2
Weissia brachycarpa (Nees & Hornsch.) Jur. (replaces Weissia
microstoma (Hedw.) C. MUll. hom. illeg. fide Koponen et al.,
Flora Fenn. 6: 61, 1977)
var. obliqua (Nees & Hornsch.) Hill, J. Bryol. 11: 601, 1981
[1982]
Weissia brachypelma C. MUll. Afr2
Weissia brachypoma Townsend, J. Bryol. 11 : 695, 1981 [1982]
Afr2
Weissia brevitheca Chen nom. inval. descr. germ. As2
Weissia breutelii C. MUll. Am2 Am3 Am4 Am5
Weissia canaliculata Hampe Am5
[Weissia cirrata Hedw. =Dicranoweisia cirrata (Hedw.) Lindh.]
var. atra Brid. Eur
var. ovatum HUb. Eur
var. porrigens Brid. Eur
[Weissia clavinervis Card. & P. Yarde Oc
Trichostomum
clavinerve (Card. & P. Yarde) Whitt. in Whitt. & Whitt.,
Bryologist 77: 433, 1974]
Weissia condensa (Yoit) Lindh. Eur As1 As3 As5 Afr1 Am1
Austr2 (replaces Weissia tortilis (Schwaegr.) C. MUll. hom.
illeg.fide Corley et al., J. Bryol. 11 : 609, 1981 [1982])
=
=
=
345
GENERA, SPECIES AND INFRASPECIFIC TAXA
Weissia controversa Hedw. Eur As1 As2 As3 As4 As5 Afr1 Afr2
Afr4 Am1 Am2 Am3 Am4 Am5 Am6 Austr1 Austr2
subsp. perssonii (Kindb.) Podp. Eur
var. amblyodon (Brid.) Sendtn. Eur As2 Afr1 Am1 Am3
var. amoene-viridis Nees. & Hornsch. Eur
var. arenico/a (Limpr.) Podp. Eur
[var. australis (Aust.) Schornh. Am1 Am3 = Weissia controversa
var. longiseta (Lesq. & James) Crum, Steere & Anders. fide
Crum et al., Bryolgist 67: 164, I973 = Weissia controversa
Hedw ..fide Stoneburner, Bryo1ogist 88: 302, 1985]
[var. brachycarpa Nees. & Hornsch. Eur Weissia brachycarpa
(Nees & Hornsch.) Jur. fide Koponen et al., Flora Penn. 6: 6I,
1977
[var. crispata (Nees & Hornsch.) Nyholm, Ill. Moss Fl. Penn. II.
Musci 775, 1969 (Hymenostomum) nom. inval.]
var. densifolia (BSG) Demaret in Demaret & Castagne, Fl. Gen.
Belgique, Bryoph. 2: 242, I964 Eur Afr1 Am1
var. exigua Schultz ex Nees & Hornsch. Eur
var. gibbosula (Amann) Podp. Eur
var. gymnostoma (Dix.) Sainsb. Austr1 Austr2
var. hioramii Tber. ex Biz, Bull. Soc. Linn. Lyon 34: 311, 1965
nom. inval. Am2
var. /atifolia (Yelen.) Podp. Eur
[var./ongiseta (Lesq. & James) Crum, Steere & Anders., Bryologist
67: I64, I967, il/eg. hom. (Weissia) Am1 Am3 = Weissia
controversa Hedw.fide Stoneburner, Bryologist 88: 302, 1985]
var. macrophylla (Ther. & P. Yarde) Wijk & Marg. Afr2
var. microdonta (Hedw.) Rtihl. Eur Afr1 Ami
var. microstoma HUb. Eur
var. minutissima (Par.) Wijk & Marg. As2
var. nitidifolia (Pam.) Podp. Eur
var. pi/lansii Schelpe, Trans. Roy. Soc. South Africa 44: Il3, I979
nom. nov. (Weissia viridula var. brachycarpa Dix. non Nees &
Hornsch.) Afr4
var. rutilans Hiib. Eur
var. stenocarpa Nees & Hornsch. Eur Afri Ami
var. subglobosa (Limpr.) Podp. Eur
var. tenuiseta (Besch.) Nog. Eur
var. turfosa (Podp.) Podp. Eur
[var. wo/ffii (Lesq. & James) Crum, Steere & Anders., Bryologist
67: 164, 1967 "wolfii" (Weissia) Ami = Weissia controversa
Hedw.fide Stoneburner, Bryologist 88: 302, 1985]
fo.fascicu/ata (Latz.) Podp., Consp. Muse. Eur. I89, I954 (Weissia
viridula fo.) Eur
fo. protonematica (Latz.) Podp., Consp. Muse. Eur. I89, I954
(Weissia viridula fo.) Eur
fo. longiseta (Latz.) Podp., Consp. Muse. Eur. I89, 1954 (Weissia
viridula fo.) Eur
fo. ticinensis (Pam.) Podp., Consp. Muse. Eur. 190, 1954 (Weissia
viridula fo.) Eur
[Weissia crispa (Hedw.) Mitt. hom. il/eg. Eur As I As2 As3 As5 Afr1
= Weissia longifolia Mitt. fide Corley et al., J. Bryol. 11: 623,
198I (1982)]
[subsp. sterilis (Nich.) Dix. Weissia sterilis Nich. fide Crundwell
& Nyholm, J. Bryol. 7: 11, 1972]
[var. aciculata (Mitt.) Dix. Eur = Weissia /ongifolia Mitt. fide
Crundwell & Nyholm, J. Bryol. 7: I3, I972]
Weissia crispa (Hedw.) Mitt. [= Weissia longifolia Mitt.] x W.
crispata (Nees & Hornsch.) C. Miill. fide Nicholson, Rev. Bryol.
32: 20, I905 Eur
Weissia crispa (Hedw.) Mitt. [= Weissia longifolia Mitt.] x W.
microstoma (Hedw.) C. Miill. fide Nicholson, Rev. Bryol. 33: I,
1906 Eur
=
=
Weissia crispata (Nees & Hornsch.) C. Miill. = Weissia fa/lax
Sehlm.
Weissia crispata (Nees & Hornsch.) C. Miill. [= Weissia fa/lax
Sehlm.] x W. crispa (Hedw.) Mitt. fide Nicholson, Rev. Bryol.
32: 22, I905 Eur
[Weissia crispula Hedw. Dicranoweisia crispula (Hedw.) Mild.]
var. ambigua Hook. f. Austr2
var. longirostris Nees & Hornsch. Eur
var. subulata Nees & Hornsch. Eur
Weissia cucullata C. MUll. Afr4
Weissia cucullata P. Yarde Afr2 hom. i/leg.
[Weissia cucullifolia Dix. & Sak. in Sak. As2 = Weissia planifolia
Dix.fide Saito, J. Hattori Bot. Lab. 39: 427, 1975 = Trichostomum planifolium (Dix.) Zand., see treatment of Trichostomum]
[Weissia curvirostris C. Miill. hom. il/eg .
Hymenostylium
recurvirostrum (Hedw.) Dix.]
var. curvipes Kindb. Ami
[Weissia deciduaefolia Saito, J. Hattori Bot. Lab. 39: 429, I975 As2
Trichostomum deciduaefolium (Saito) Zand., see treatment of
Trichostomum]
Weissia dieterlenii Ther. Afr4
Weissia diffidentia Zand. (nom . nov. for Phascum recurvirostre C.
MUll.), see treatment of Weissia Am5 Am6
Weissia edentula Mitt. As3 As4 Oc
Weissia erythrogona Brid. Am1
[Weissia euteiches Zand., Monogr. Syst. Bot. Missouri Bot. Gard.
11: 195, 1985 (nom. nov. for Weissia tortilis (Schwaegr.) C.
MUll.) = Weissia condensa (Yoit) Lindh. (replaces Weissia
tortilis (Schwaegr.) C. MUll. hom. il/eg. fide Corley et al., J.
Bryol. li: 609, 1981 (1982)]
Weissia exserta (Broth.) Chen As2
Weissiafal/ax Sehlm. Eur As2 As5 Afri Ami
var. alpina (Schimp.) Podp. Eur
var. subgymnostoma (Podp.) Podp. Eur
[Weissiaflavescens (Britt. inN. Britt. & Millsp.) Reese, Bryologist
94: 54, I99I Ami Am3 = Trichostomum brittonianum nom.
nov., see treatment of Trichostomum]
Weissia felipponei Ther. in Felipp. Am6
Weissiafornicata Brid. Eur
Weissia ghatensis Dix. & P. Yarde As3
[Weissia glauca Bartr. Ami Am2 hom. illeg.
Weissia
andersoniana Zand., Monogr. Syst. Bot. Missouri Bot. Gard.
11: 195, I985]
Weissia glaziouii Zand. (nom . nov. for Hymenostomum striatum
Geh. & Hampe), see treatment of Weissia Am5
[Weissia graeca Schiffn. Eur = Weissia controversa Hedw. fide
Corley et al., J. Bryol. II: 623, I98I (1982)]
[Weissia hedwigii Crum, Bryologist 74: I69, I97I Eur Asi As2
As3 As5 Afri Aml (nom . nov. for Weissia microstoma
(Heow.) C. Miill. hom. illeg.) = Weissia brachycarpa (Nees &
Hornsch.) Jur. var. brachycarpafide Corley et al., J. Bryol. II:
650, I98I (I982) and Koponen et al., Flora Penn. 6: 61, I977]
Weissia humicola C. MUll. Afr2 Afr4
Weissia inoperculata (Crum) Crum, Steere & Anders., Bryologist
67: 164, I973 Ami (Hymenostomum)
Weissia jamaicensis (Mitt.) Grout Am I Am2 Am3
Weissia jamesonii Tayl. Am4
Weissia kaikouraensis R. Br. ter Austr2
[Weissia krassavinii (Lazar.) Lazar. in Lazar. et al., Biul. Mosk.
Obsh. Ispyt. Prir, Otd. Biol. 73(2): I42, I968 comb. inval.
basion. non cit. (Hymenostomum) Asi = Weissia krassavinii
(Lazar.) Lazar. ex Ochyra]
Weissia krassavinii (Lazar.) Lazar. ex Ochyra, J. Hattori Bot. Lab.
=
=
=
=
346
GENERA OF THE POTTIACEAE
64: 343, 1988 As1
Weissia kunzeana C. Miill. Am6
[Weissia lancifolia (R. Br. ter) Wijk & Marg. Austr2 = Tridontium
tasmanicum Hook. f. in Hook., see treatment of Tridontium]
[var. waymouthii (R. Br. ter) Wijk & Marg. Austr2 = Didymodon
waymouthii (R. Br. ter) Zand., see treatment of Didymodon]
Weissia laticuscula C. Miill. Afr2 Afr4 (good species fide Magill, Fl.
S. Afr.l. Mosses 1: 267, 1981 [1982])
Weissia leptocarpa Schimp. ex Besch. in Ler. & Levier Eur Afrl
hom. illeg.
Weissia levieri (Limpr.) Kindb. cf. Crundwell & Nyholm, J. Bryol. 7:
16, 1972 Eur
Weissia ligulaefolia (Bartr.) Grout Aml
Weissia lineaefolia C. Miill. Afr2
Weissia longidens Card. As2 (good species fide Saito, J. Hattori Bot.
Lab. 39: 429, 1975)
Weissia longifolia Mitt. Eur Asl As2 As3 AsS Afr1
var. angustifolia (Baumg. in Giznb.) Crundw. & Nyh., J. Bryol. 7:
14, 1972 (Astomum crispum var.) Eur
Weissia lorentzii (C. Miill.) Zand. (Phasconica), see treatment of
Weissia Am6
Weissia ludoviciana (Sull.) Reese & Lemmon, Bryologist 68: 282,
1965 (Astomum) Aml
Weissia ludoviciana (Sull.) Reese & Lemmon X W. controversa
Hedw.fide Reese & Lemmon, Bryologist 68: 280, 1965 Arn1
Weissia macrospora Card. & P. Yarde As3
[Weissia malayensis (Fleisch.) Manuel, Fed. Mus. Jour. 26: 161, 1981
(Hymenostomum) As2 As4 = Barbula indica (Hook.) Spreng. fide
lwatsuki & Noguchi, J. Hattori Bot. Lab. 37: 355, 1973 as Barbuta cruegeri]
Weissia melanostoma Mitt. Am5
Weissia micacea (Schlecht.) C. Miill. Am3 Am5 ·
[Weissia microstoma (Hedw.) C. Miill. Eur Asl As2 As3 AsS Afrl
Aml hom. illeg. = Weissia brachycarpa (Nees & Hornsch.) Jur.
var. brachycarpafide Hill, J. Bryol. 11: 601, 1981 (1982) and
Koponen eta!., Flora Fenn. 6: 61, 1977]
[Weissia minycarpa Zand., Monogr. Syst. Bot. Missouri Bot. Gard.
11: 196, 1985 (nom . nov. for Trichostomum microcarpum
Schimp. ex Besch. non Weissia microcarpa Hook. f. & Wils.)
Am2 Am3 Am4 = Weissia sinaloensis Bartr.fide Zander in Sharp
et a!., Moss Fl. Mex. = Trichostomum sinaloense (Bartr.) Zand.,
see treatment of Trichostomum]
Weissia mittenii (BSG) Mitt. Eur
Weissia muehlenbergiana (Sw.) Reese & Lemmon, Bryologist 68:
282, 1965 (Astomum) As2 Am1
Weissia multicapsularis (Sm.) Mitt. Eur As1
Weissia neocaledonica (Ther.) Zand. (Aschisma), see treatment of
Weissia Oc
Weissia newcomeri (Bartr.) Saito, J. Hattori Bot. Lab. 39: 423, 1975
(Hymenostomum) As2
Weissia nitida Reinw. & Hornsch. As4
Weissia norketii R. S. Chopra, Taxon. Indian Mosses 125, 1975
(nom. nov. for Weissia amblyphylla) (Dix.) Dix. As3
[Weissia nuda Mitt. Austr1 hom. illeg. = Weissia alianuda Tan, Mem.
New York Bot. Gard. 68: 5, 1992, nom. nov.]
[Weissia nudiflora C. Miill. & Hampe Austr1 = Weissia controversa
Hedw.fide Catcheside, Mosses S. Austr. 196, 1980]
Weissia obtusata C. Miill. Am5
Weissia obtusifolia C. Miill. Am5
Weissia occidentalis (Flow. ex Crum) Stoneburner, Bryologist 88:
310, 1985 (Astomum) Am1
Weissia occulta Wallr. Eur
Weissia opaca (Dix.) Magill in Magill & Schelpe, Mem. Bot. Surv. S.
Afr. 43: 7, 1979 (Tarte/la) Afr2
Weissia ova/is (Williams) Bartr. Oc
Weissia papillosa Dix. & Nav. Afr2
Weissia papillosissima Laz., Dopov. Akad. Nauk Ukr. R.S.R. Ser.
B 8: 752, 1967 As1
Weissia patagonica Card. & Broth. Am6
[Weissia perligulata Flow. ex Crum, Bryologist 76: 291, 1973 Am1
(= Trichostomum crispulum Bruch ex F. Miill.fide Stoneburner
& Wyatt, Monogr. Syst. Bot. Missouri Bot. Gard. 11: 184,
1985) Trichostomum perligulatum (Flow. ex Crum) Zand.,
see treatment of Trichostomum]
Weissia perpusilla (C. Miill.) Stone, J. Bryol. 11: 231, 1980
(Hymenostomum) Austr1
Weissia phascopsis Zand.
(nom. nov. for Gymnostomum
phascoides Hook. ex Drumm. non Weissia phascoides (BSG)
C. Miill.) Eur Am1
[Weissia planifolia Dix. As 1 As2 Am2 = Trichostomum planifolium
(Dix.) Zand., see treatment of Trichostomum]
[Weissia platyphylloides Card. As2 As3 = Weissia edentula Mitt.
fide Saito, J. Hattori Bot. Lab. 39: 421, 1975]
Weissia platystegia (Dix.) Eddy, Handb. Males. Mosses 2: 165,
1991 (Astomum) As4
Weissia riograndensis (Broth.) Zand. (Hymenostomum), see treatment of Weissia Am5
[Weissia rigescens Broth. As2 hom. illeg. = Weissia controversa
Hedw.fide Saito, J. Hattori Bot. Lab. 39: 426, 1975]
[Weissia riparia Hampe Austr1 = Weissia controversa Hedw. fide
Catcheside, Mosses S. Austr. 196, 1980]
Weissia rostel/ata (Brid.) Lindh. Eur Aml
[var. phascoides (Hook.) Reese & Lemmon, Bryologist 68: 283,
1965 (Astomum) Eur Am1 = Weissia phascopsis Zand. nom.
nov., Monogr. Syst. Bot. Missouri Bot. Gard. 11: 196, 1985 non
Weissia phascoides (BSG) C. Miill.J
Weissia rutilans (Hedw.) Lindh. Eur As1 Afrl Afr2 Am1 Austrl
subsp. ganderi (Jur.) Kindb. Eur
var. ganderi (Jur.) Macoun Eur
var. hillieri Meyl. Eur
var. himalayana Broth. As3
var. subgymnostomum Limpr.
Weissia semidiaphana (Ther.) Zand, Monogr. Syst. Bot. Missouri
Bot. Gard. 11: 197, 1985 (Hymenostomum) Am2
Weissia semiinvoluta C. Miill. Am6
Weissia semipal/ida C. Miill. As2
Weissia sharpii Anders. & Lemmon, Bryologist 76: 133, 1973 Am1
Weissia simplex Brid. ignot.
[Weissia sinaloensis Bartr. Am2 Am3 Am4 (see Zander in Sharp et
al., Moss Fl. Mex.) = Trichostomum sinaloensis (Bartr.) Zand.,
see treatment of Trichostomum]
Weissia socotrana Mitt. Afr2
Weissia splachnum Garov. Eur
Weissia sterilis Nich. see Crundwell & Nyholm, J. Bryol. 7: 11,
1972 Eur
Weissia squarrosa (Nees & Hornsch.) C. Mtill. Eur
Weissia subacaulis (Mitt.) Par. Am4
[Weissia subangustifolia (Ther.) Zand., Bryologist 86: 156, 1983
(Hyophila) Am2 = Trichostomum subangustifolium (Ther.)
Zand., see treatment of Trichostomum]
Weissia submicacea C. Mtill. Am5
Weissia sweetii Bartr. Am1 l= Trichostomum crispulum Bruch in F.
Miill. fide Stoneburner & Wyatt, Monogr. Syst. Bot. Missouri
Bot. Gard. 11: 184, 1985 and Stoneburner, Bryologist 88: 1985,
but retained here because leaf margins narrowly inflexed and
monoicous, possibly synonymous with Weissia perligulata]
=
GENERA, SPECIES AND INFRASPECIFIC TAXA
=
[Weissia termitarum C. Mtill. Am5 Trichostomum termitarum (C.
Mtill.) Zand., see treatment of Trichostomum]
Weissia termitidarum C. Mtill. Afr2 Am5
[Weissia tortilis (Schwaegr.) C. Mill!. hom. illeg. Eur As1 As3 As5
Afr1 Am1 Austr2 = Weissia condensa (Voit) Lindh. fide Corley
et al., J. Bryol. ll: 623, 1981 (1982)]
[subsp. papillosissima Lazar., Dopov. Akad. Nauk Ukr. R.S.R. Ser.
B 8: 752, 1967 nom. inval. in syn. Eur = Weissia papillosissima
Lazar.]
var. intermedia Monk. Eur
var. subcylindrica (BSG) Dix. Eur
[Weissia tortivelata Williams Am4 Trichostomum tenuirostre var.
gemmiparum (Schimp.) Zand., see treatment of Trichostomum]
Weissia triumphans (De Not.) Hill, J. Bryol. ll: 600, 1981 [1982]
(Trichostomum) Eur As5 Afrl
var. monspeliensis (Schimp.) Zand. (Trichostomum), see treatment
of Weissia Eur Afr1
Weissia tyrrhena Fleisch. Eur As1
[Weissia umbrosa Mitt., J. Linn. Soc. Bot. 12: 133, 1869 Am4
Trichostomum mitteneanum Zand. nom. nov., see treatment of
Trichostomum]
Weissia unguiculata (Mitt.) Crundw. & Nyholm, J. Bryol. 8: 69, 1974
(Astomum) Afr2
[Weissia vallis-gratiae C. Mtill., Hedwigia 38: lll, 1899 = Weissia
controversa Hedw.fide Magill, Fl. S. Afr. I. Mosses 1: 265, 1981
(1982)]
[Weissia verticillata Brid. Eucladium verticillatum (Brid.) BSGJ
var. sterilis Mazz. Eur
Weissia veviridis Zand. (nom. nov. for Trichostomum perviride
Broth.), see treatment of Weissia Am3 Am4
[Weissia viridula Hedw. ex Brid. illeg. = Weissia controversa Hedw.]
subsp. longirostris Kindb. in Roll Am I
var. canaliculata Hampe Am4
var. cylindrica Schimp. in Par. Afr1
[var. gymnostoma Dix., New Zealand lnst. Bull. 3: 112, 1923 =
Trichostomum brachydontium Bruchfide Stoneburner, Bryologist
88: 312, 1985]
[var. hioramii Ther. nom. inval. descr. gall. Am3 = Weissia
controversa var. hioramii Ther. ex Biz, Bull. Soc. Linn. Lyon 34:
311, 1965]
var. longidens De Not. Eur
var. longifolia Broth. & Wag. Afr4
var. polycarpa Chen As2
var. rugeliana Ren. & Card. Am1
var. seligerioides Amann, Fl. Mouss. Suisse 2(Add. Rectific.): 1,
=
=
=
347
1991 Eur
fo.fasciculata Latz., Beih. Bot. Centralb. 48: II. 472, 1931 Eur
fo. longiseta Latz., Beih. Bot. Centralb. 48: II. 472, 1931 hom.
illeg. Eur
fo. protonematica Latz., Magyar Bot. Lapok 29: 16, 1930 Eur
fo. ticinensis Fam., Muschi Prov. Pavia, Atti lstituto Bot. Univ.
Pavia 3: 9, 1891 Eur
[Weissia waymouthii R. Br. ter Austr2 Didymodon waymouthii
(R. Br. ter) Zand., see treatment of Didymodon]
Weissia welwitchii Schimp. Eur
Weissia willisiana (Sainsb.) Catcheside, Mosses S. Austr. 194, 1980
(12 Dec. 1980) (Pottia) (comb. also by Stone, J. Bryol. 11: 235,
1980 (27 Mar. 1981) Austr1
Weissia wimmeriana (Sendtn.) BSG Eur As3 Am1
subsp. pallescens (Besch.) Giac. Eur Afrl
var. dalmatica (Latz.) Podp. Eur
var. linderi Broth. & Geh. Eur
var. muralis (Spruce) Breidl. Eur
var. pal/escens (Besch.) Bott. Eur Afr1
[fo. hymenostomoides Spruce= Weissia wimmeriana var. muralis
(Spruce) Breidl.fide Podp., Consp. Muse. Eur. 190, 1954]
[fo. subgymnostoma Limpr., Laubm. Deutsch. 259, 1886 = Weissia wimmeriana var. muralis (Spruce) Breidl. fide Podp.,
Consp. Muse. Eur. 191, 1954]
=
WILLIA C. Mtill. in Neum.
Willia austroleucophaea (Besch.) Broth. Am6 Ant
Willia brachychaete (Dus.) Zand. (Tortula), see treatment of Willia
Am6
Willia calobolax (C. Mtill.) Lightowlers, J. Bryol. 13: 370, 1985
(Barbula) Afr4
var. angustinervia (C. Miill.) Lightowlers, J. Bryol. 13: 370, 1985
(Barbula calobolax var.) Afr4
[Wil/ia grimmioides C. Miill. in Neum. Am6 = Willia austroleucophaea (Besch.) Broth. fide Bell, Brit. Antarct. Surv. Bull.
38: 73, 1974]
[Willia marginata (Hook. f. & Wils.) C. Miill. Afr4 = Hennediel/a
marginata (Hook. f. & Wils.) Zand., see treatment of Hennediella]
[WILLIAMSIELLA Britt.= Leptodontium]
[Williamsiella tricolor (Williams) Britt. Am4 = Leptodontium tricolor (Williams) Zand. in Zand. & Hegew., Bryologist 79: 20,
1976]
BIBLIOGRAPHY
This literature list includes selected taxonomic treatments and morphological or physiological discussions of the Pottiaceae in addition to the citations in the text. Published notices of simple range extensions and other minor works are largely eliminated. Readers
are directed to a far larger bibliography published electronically (Zander 1993b), which is more suitable for ad hoc searches.
A
Abrarnova, A. L., L. S. Blagodatskich & L. A. Czerepanova. 1973.
Conspectus generis Pterygoneurum Jur. (Musci) in URSS. Nov.
Syst. Pl. non Vase., Ac. Sc. URSS, Inst. Bot. V. L. Komarovii 10:
305-316.
Abrarnova, A. L. & U. K. Marnatkulov. 1967. [De proprietatibus
biologicis Barbula johansenii Williams.] Nov. Syst. Plant. non
Vase., Acad. Sci. USSR, Komarov lnst. 1967: 364-371.
Abramova, A. L. & Ts. Tsegmed. 1987. De positione taxonomica
Trichostomopsis arronis (Lor.) Agnew et Townsend notula.
Novosti Sistematiki Nizshikh Rastenii 24: 169-178. [In Russian.]
Agnew, S. & M . Vondnil!ek. 1975. A moss flora of Iraq. Feddes
Repertorium 86(1Hl): 341-489.
Akiyama, H. & N. Nishimura. 1993. The Climacium-type branch
development on the Bartramiaceae and its taxonomic significance. Bryologist 96: 185-191.
Albert, V. A. & B. D. Mishler. 1992. On the rationale and utility of
weighting nucleotide sequence data. Cladistics 8: 73-83.
Allen, B. H. 1992. Teratolgical axillary "hairs" in Didymodon
tophaceus. Bryologist 95: 97-99.
Allen, B. H. & R. E. Magill. 1987. In support of a distinct terminology for bryophyte sexuality. Taxon 36: 57-58.
Allen, B. H. & R. A. Pursell. 1991. A reconsideration of the systematic position of Jonesiobryum. Bryologist 94: 438-442.
Allorge, P. 1938. Observations sur Tortula desertorum Broth.,
mousse aralo-caspienne des plateaux castillans. Rev. Bryol.
Lichenol. 11: 110-112.
Anderson, L. E. 1943. The distribution of Tortula pagorum (Mild.)
De Not. in North America. Bryologist 46: 47--{)6.
Anderson, L. E. 1951. The mosses of North Carolina. V1. Encalyptaceae to Pottiaceae. Bryologist 54: 156-161.
Anderson, L. E . 1954. Hoyer's Solution as a rapid mounting medium
for bryophytes. Bryologist 57: 242-244.
Anderson, L. E. & B. E. Lemmon. 1972. Cytological studies of natural intergeneric hybrids and their parental species in the moss
genera Astomum and Weissia. Ann. Missouri Bot. Gard. 59:
382-416.
Anderson, L. E. & B. E. Lemmon. 1973. Weissia sharpii, a new species from North America, with cytological and and ecological
correlations. Bryologist 76: 131-143.
Anderson, L. E . & B. E. Lemmon. 1974. Gene flow distances in the
moss Weissia controversa Hedw. J. Hattori Bot. Lab. 38: 67-90.
Anderson, L. E. & R. H. Zander. 1973. The mosses of the Southern
Blue Ridge Province and their phytogeographic relationship. J.
Elisha Mitchell Scientific Soc. 89: 15--{)0.
Anderson, L. E. & R. H. Zander. 1986. False anisospory in the moss
Leptodontium viticulosoides (P.-Beauv.) Wijk & Marg. Am. J.
Bot. 73(5): 603 (Abstr.).
Andrews, A. L. 1915. Bryological notes. I. Aschisma kansanum, a
new species, with remarks upon the genus. Torreya 15: 63--{)7.
Andrews, A. L. 1920. Hymenostomum in North America. I. Delimitation of the genus. Bryologist 23: 28-31.
Andrews, A. L. 1922a. Hymenostomum in North America. II. The
case of Astomum sullivantii. Bryologist 25: 66-71.
Andrews, A. L. 1922b [1923]. The status of Gyroweisia in North
America. Bryologist 25: 97-100.
Andrews, A. L. 1922c [1923]. The status of Gyroweisia in North
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Lindb. is an Oxystegus. Lindbergia 8: 185-187.
Zander, R. H. 1982c. Trichostomum molariforme sp. nov. and
Atractylocarpus stenocarpus (Wils.) in Seem.) comb. nov.
Bryologist 85: 126-128.
Zander, R. H. 1982d. Practical species concepts. Beih. Nov. Hedw.
71: 7-11.
Zander, R. H. 1982e. Various thoughts on characters in moss taxonomy. Beih. Nov. Hedw. 71 : 81-85.
Zander, R. H. 1982f. Herbarium and cultivation methods in mosses.
Beih. Nov. Hedw. 71 : 127-130.
371
Zander, R. H. 1982g. Aspects of the taxonomy of the Pottiaceae.
Beih. Nov. Hedw. 71: 225-227.
Zander, R. H. 1982h. The quality of floristic and taxonomic information. Beih. Nov. Hedw. 71: 291-296.
Zander, R. H. 1983a. The genus Streptocalypta C. Muell. (=
Barnesia Card.) Lindbergia 8: 161-165.
Zander, R. H. 1983b. A rapid microscopic mounting medium for
delicate bryophytes. Taxon 32: 618-620.
Zander, R. H. 1983c. A reevealuation of Neohyophila Crum (Pottiaceae). Bryologist 86: 134-139.
Zander, R. H. 1983d. Nomenclatural changes in Hyophila, Leptodontium and Morinia (Pottiaceae). Bryologist 86: 156-157.
Zander, R. H. 1983e. Pottiaceae (Musci) of Cofre de Perote, Veracruz, Mexico. Am. Philos. Soc. Grantees' Rep. 1982: 12-13.
Zander, R. H. 1984. Bryophyte sexual systems: -oicous versus
-oecious. Bryol. Beitr. 3: 46-51.
Zander, R H. 1985a. Nomenclatural transfers · in Weissia (Pottiaceae, Musci). Monogr. Syst. Bot. Missouri Bot. Gard. 11 :
195-197.
Zander, R. H. 1985b. An essay on species concepts developed during revisionary studies. Bryologist 88: 215- 220.
Zander, R. H. 1986a. Notes on Bryoerythrophyllum (Musci).
Bryologist 89: 13-16.
Zander, R. H. 1986b. Two new species of Pottiaceae from Mexico.
Bryologist 88: 353-356.
Zander, R. H. 1986c. Three new taxa of bryophytes from Peru.
Willdenowia 16: 253-258.
Zander, R. H. 1987a. Pottiaceae of North America North of Mexico. Flora Online (Taxacom, Buffalo Museum of Science) Issue
7, Ver. 1.0, 40,853 bytes, Jan. 1987; Ver. 1.1 , 40,048 bytes,
Mar. 1987.
Zander, R. H. 1987b. A bibliography of the Pottiaceae (Musci).
Flora Online (Taxacom, Buffalo Museum of Science) Issue 8,
Ver. 1.0., 150,651 bytes.
Zander, R. H. 1988. Sporophyte reduction series in Pottiaceae
(Musci). Am. J. Bot. 75(6, Part 2): 12 (Abstr.).
Zander, R. H. 1989. Seven new genera in Pottiaceae (Musci) and a
lectotype for Syntrichia. Phytologia 65: 424-436.
Zander, R. H. 1991. [Abstract:] A phylogenetic evaluation of the
genera of the Pottiaceae (Musci). Am. J. Bot. 78(6, suppl.):
11-12.
Zander, R. H. 1993a. A new combination in Pottia (Musci). Novon
3: 92, 1993.
Zander, R. H. 1993b. A bibliography of the Pottiaceae (Musci).
Flora Online (Taxacom, Buffalo Museum of Science) Issue 8,
Ver. 2.0.
Zander, R. H. & H. Crum. 1977 [1978]. Desmatodon steereanus, a
new species of Pottiaceae from Peru. Bryologist 80: 638-640.
Zander, R. H. & A.M. Cleef. 1982 [1983]. Studies on Colombian
Cryptogams. XVI. Taxonomy and ecology of Kingiobryum
paramicola (Dicranaceae, Musci). Proc. Kon. Ned. Akad.
Wetensch., Ser.C 85: 627-634.
Zander, R. H. & C. Delgadillo M. 1984 [1985]. The mosses of the
Tehuacan Valley, Mexico, and notes on their distribution.
Bryologist 87: 319-322.
Zander, R. H., C. Delgadillo M. & P. M. Eckel. 1980 [1981]. Morinia saitoana, a new species of Pottiaceae (Bryopsida) from
Mexico. Bryologist 83: 508-511.
Zander, R. H. & P. M. Eckel. 1980. Tortula cainii: additional
Ontario records and behavior in a common garden. Bryologist
83: 209-211.
Zander, R. H. & P. M. Eckel. 1982. Hymenostylium recurvirostrum
var. insigne and Barbula amplexifolia in British Columbia,
372
GENERA OF THE POTTIACEAE
Canada. Canad. J. Bot. 60: 1596-1600.
Zander, R. H. & P. M. Eckel. 1987. A new species of Tuerckheimia
from Mexico. Mem. New York Bot. Gard. 45: 293-295.
Zander, R. H. & E. Hegewald. 1976. Leptodontiella, gen. nov. and
Leptodontium from Peru. Bryologist 79: 16-21.
Zander, R. H. & F. J. Hermann. 1986 [1987]. Gyroweisia monterreia,
a new moss species from Mexico. Bryo1ogist 89: 227-229.
Zander, R. H. & W. J. Hoe. 1979. Geographic disjunction and
heterophylly in Torre/la fragilis var. tortelloides (= Sarconeurum
tortelloides). Bryologist 82: 84-87.
Zander, R. H. & A. J. Sharp. 1981 [1982] . Two species of
Bryoerythrophyllum new to Mexico. Bryologist 84: 543-547.
Zander, R. H. & W. C. Steere. 1978. Tortula scotteri sp. nov. from
the Northwest Territories of Canada. Bryologist 81: 463-467.
Zander, R. H. & D. H. Yitt. 1979. Gametophytic distinctions of
Zygodon (Orthotrichaceae) and Anoectangium and Leptodontium (Pottiaceae) and the status of Anoectangium rubrigemmium
of Hawaii. Canad. J. Bot. 57: 292-296.
Zijlstra, G. 1990. On conserved types. Bryol. Times 53: 8-9.
Zuttere, P. de, A. Gohimont, R. Schumacker, A. Sotiaux & J. Werner. 1984. Weissia rostellata (Brid.) Lindb. (Musci), nouveau
pour Ia Belgique, Le Grande-Duche de Luxembourg et le
departement du Nord (France). Dumortiera 29/30: 15-19.
GLOSSARY OF SPECIAL TERMS
This glossary includes only those terms especially appropriate to study of the Pottiaceae. See Wyatt (1985) and Magill (1990) for
extensive definitions of bryological terms.
abaxial: away from the axis; the surface of the leaf facing away
from the stem, i.e. the dorsal surface.
adaxial: towards the axis; the surface of the leaf facing towards
the stem, i.e. the ventral surface.
anisospory: spores of two size classes occurring in a single capsule.
autoicous: antheridia, surrounded by perigonial leaves, and
archegonia, surrounded by perichaetial leaves, borne in separate locations on the same plant; here usually indicating the
gonioautoicous condition, perigonia borne laterally on very
short branches in the axils of leaves.
bulliform: bubble-shaped, e.g. !aminal cells of Quaesticula or
Tuerckheimia.
carinate: keeled, with upper halves of the lamina rather appressed,
diverging at a small angle at the costa and vee-shaped in
transverse section.
cee: the letter "c".
cleistocarpous: a capsule lacking a differentiated operculum when
mature, usually dehiscing irregularly.
clockwise: twisted (setae, peristomes) in a clockwise direction,
being seen as twisted in a clockwise direction upwards
towards an observer above the plant. The few taxa with clockwise twisted peristomes (species of Timmiella and to a lesser
extent Leptodontiella) would have the appearance of the
thread on a left-handed screw. Synonym: dextrorse.
collenchymatous: !aminal cells thickened at the corners, the
thickenings having the appearance of small, bright triangles.
colliculate: covered with crowded, small, rounded protrusions,
used here for laminae with strongly bulging upper lamina!
cells, these usually smooth individually but sometimes also
papillose.
correct: as used here, that most recently set right.
counterclockwise: the usual direction of twist of a peristome in
the Pottiaceae is counterclockwise as seen from above with
the direction of twist rising upwards. Laterally, a
counterclockwise-twisted peristome would have the appearance of the thread on a standard, right-handed screw (pers.
comm. D. Wagner). Synonym: sinistrorse. Jackson's (1960)
Glossary of Botanic Terms (Appendix) discusses past confusion in use of terms for direction of twisting.
distal: most distant, usually from the center or body of the plant.
dorsal: as used here meaning the abaxial surface of a leaf.
emergent: a sporophyte with a portion of its capsule extending
just beyond the tips of the perichaetialleaves.
eperistomate: lacking a peristome, used for capsules with a
differentiated operculum but no peristome; this is the latinized form but also spelled aperistomate. See cleistocarpous.
exerted: a sporophyte with the entire capsule visible above the
tips of the perichaetialleaves.
flat: In a treatment of Grimmia (Grimmiaceae) Sayre (1952)
defmed this for the leaf as when the costa is in the same
plane of focus as the !aminal cells at lOOx, or when leaves
on a microscope slide lie mostly open rather than folded
(''keeled"). In practice, "flat" may also refer to leaves that
are broadly channeled across the leaf but which lack a narrow groove ventrally along the costa.
gametoecia: a moss inflorescence, consisting of several leaves
usually somewhat modified in size and shape terminating a
stem or branch, the leaves enclosing archegonia or
antheridia or both, often also paraphyses.
guide cells: large, wide-lumened, thin-walled costal cells, usually 2-4 in number, just ventral to dorsal stereid band (and
sometimes an associated hydroid strand). Synonyms:
eurycysts, Deuters, socii (cf. Ligrone 1985).
hyalocyst: enlarged basal !aminal cells sharply differentiated
from upper cells by larger size and lack of chlorophyll.
immersed: a sporophyte (capsule) entirely hidden by the perichaetialleaves.
keeled: leaves with lateral portions of the upper lamina inserted
on the costa at right angles or less (being vee-shaped in
transverse section) such that the leaves generally lie on their
sides (at least above midleaf) when moist.
ligulate: in bryological usage, strap-shaped or narrowly elliptical, as opposed to spathulate, which usually means obovate.
lumen: the interior of a cell, of various shapes in outline. Plural:
either lumina or lumens is acceptable.
lower: usually refering to the portion of a structure proximal to
the stem or to the base of a plant, e.g. the differentiated
basal lamina! cells in the lower portion of the leaf.
mamilla (or mammilla): a smooth, broad bulge; generally the
whole cell surface, when mamillose, bulges to produce collectively a colliculate lamina! surface, but sometimes (not
GLOSSARY
here) simple, hollow papillae are called mamillae.
molariform: like a millstone; a papilla with a broad, rounded or
flattened surface that is often tuberculated.
multifid: having many branches (or, in the case of papillae, points
or salients of various shapes).
oh: the letter "o".
optical section: the shallow focal plane of a high power microscope produces an image of a section of the tissue observed.
Hollow, hemisphericallaminal papillae thus may appear to be
oh- or cee-shaped, when in fact only a section of the hemisphere is in focus. An upright dome gives an oh-shaped optical section, a tilted dome a cee-shaped section.
p.p.: pro parte, in part.
prorulose: paired papillae formed by superficial projecting ends of
contiguous cells, e.g. as in Barbula indica (see also
"scindulae" fide Weber & Simone 1977).
proximal: most near, generaJly to the center or body of the plant.
salients: branches or points on the papillose surface of a cell.
rhexolytic: rupturing irregularly; in this case, a capsule dehiscing
373
without the aid of an operculum.
stereid: longitudinaJly elongate cells with much-thickened waJls
and smaJI lumina, usuaJly evident in transverse sections of
the costa and stem. Substereid ceJls have rather wide
lumina.
trifarious: leaves distinctly arranged in three longitudinal rows,
generaJly spiralling around the stem, e.g. as found in
Hymenostylium, Molendoa, Reimersia and Triquetrella.
twist: see clockwise and counterclockwise.
upper: usually refering to the portion of a structure distal to the
stem or base of a plant.
vee: the lett.e r "v"; in certain genera, basal larninal ceJls are
strongly differentiated from the upper laminal ceJls by
larger size and thinner walls, and arranged so as to reach
higher along the margins than mediaJly (along the costa),
thus forming a vee-shaped wedge of usuaJly sharp differentiation.
ventral: as used here meaning the adaxial surface of a leaf.
INDEX
Species names are included when they occur with treatments other than their own genera. Subfamilies, tribes, subgenera, sections,
and subsections, including synonyms, are all listed as single words. Recognized genera of the Pottiaceae are set in boldface.
Acaulon 253, also 201,231,273
Acaulonopsis 253
acropleurocarpous sexuality 120
Acutae 157, 218
Acutiformes 157
Aesiotortula 236, 264, 251
Agrariae 145
alar cells 185
Alaticosta 201, 253
alcohol150
A/oideae 203
Aloidel/a 203
Aloina 203, also 27, 196, 222
Aloinella 206, also 237
Amblystegioideae 113
Amphidiopsis 218
Amphidium 274
amphithecial ceJllayers 82
Anacalypta 230
analysis, phylogenetic 16
Annua 231
Anoectangiaceae 136
Anoectangium 136, also 5, 169
aestivum 215, 279
Anomalae 68
Argyrobarbula 196
Arvildia 274
Aschisma 213, also 216, 232, 250
Asteriscium 160, also 94, 157
Astomaceae 163
Astomataceae 163
Astomiopsis 275
Astomum 2, 88, 176, 279
austral taxa 46
axillary hairs 146
Barbiferus 145
Barbula 145,also86, 107,158,173
agraria 173
arenicola 116
calycina 211
indica 230, 373
pachyloma 82
Barbuleae 136, also 49
Barbu/ie/la 146
Barbuloideae 26, 136
Barnesia 2, 94
basal ceJls, transversely slit 71
Bauriella 218, 242
Beccaria 218
Beckettia 242
Begleiter 5
Bellibarbula 140
Bermuda 85
bibliography 348
Brachymenium 249
domingense 278
Brachymitrion 4
brood bodies 273
Brotherus 32 ·
Bruchia
brevifolia 216
carinata 278
Bryella 237
Bryobartramia 18, 213, 274
Bryoceuthospora 216, also 213, 218
Bryoerythrophylleae 107, also 49
374
GENERA OF THE POTTIACEAE
Bryoerythrophyllum 112, also 84, 112, 140, 150, 211, 261, 239,
255
columbianum 237
Bryum 163, 195
Bulbibarbula 145
bulge, asymmetric, of 1aminal cells 5
Buxbaumia 12
Caespitosae 100
Calymperaceae 12, 84, 193
Calymperastrum 83
Calymperella 142
Calymperes 164, 176
Calyptopogon 98, also 127, 136
calyptra 8
Camptopodiae 231
Campy/opus 86
Canescentes 227
capsule
amphithecial cell layers 82
systilious 14
Catillaria 218
Caulescentes 145
Cavanillea 176
cells, basallaminal5, 6
central strand, satellite 84
Ceuthospora 216
character state
changes 47
polarity 22
at major ancestral nodes 28
characters 3, 12
22 reduction-associated 25
autapomorphic 15
marker 15
quantitative 22
Cheilothela 275
Chen 32,50
Chenia 255, also 247
leptophylla 253
Chionoloma 76, also 88, 98
Chionolomoideae 76, also 49
Chloronotus 196
Chrysoblastella 2
Cinclidotaceae 1
Cinclidotoideae 2
Cinclidotus 110, 276
Circum-Tethyan region 31
cladograms 25, 33
Cladophascum 274
classification 31
cleistocarpous capsules, in Tortella 101
Climacocaulon 206
colliculate, ventral surface of leaf 5
Collotortula 264
color 3
Controversae 176
convergence 13, 26
Convolutae 86, 90, 135, 140, 145, 148, 209, 211
copper mosses 156, 242
Coronopapillata 132
cortex 4
costa 5, 146
Craspedophyllon 157
Crassicostata 132
Crassicostatae 227
Crassinerves 227
Crispuliformes 90, also 86, 186
Crossidium 196, also 202, 206, 222, 230, 231
aberrans 252
Crumia 240
Cuneifoliae 231
cyanobacteria 157
Cycnea 237
cylinder, central4
Dactylhymenium 157
Daltonia angustifolia 145
data set 50
DELTA 54
Dendia 232
Dermatodon 227
desiccation 5
Desmatodon 174,222,227,231
Dialytrichia 107, also 84, 107
Diastoma 151
Dicranella 275
Dicranoweisia 275
Dicranum 185
Didyctium 227
Didymodon 157, also 146
australasiae 279
johansenii 169
nigrescens 142
rigidulus 279
vinealis 279
Diphyscium 12, 68, 274, 275
Diplolepideae 17
Ditrichum 12, 14, 18, 148
Dolotortula 248, also 247
Edentella 176
Encalypta 12, 18, 37
Entosthymenium 274
environments, patchy 14
Ephemerum 216
epidermis, of costa 5
Erpodiopsis 274
Erythrobarbula 112
Erythrophyllastrum 72
Erythrophyllopsis 71
Erythrophyllopsoideae 71, also 49
Erythrophyllum 112
Euanoectangium 136
Euastomum 178
Eubarbula 145
Eucladieae 82
Eucladioideae 2, 82
Eucladium 84, also 94, 107, 276
Eudesmatodon 227
Eudidymo[ion 157
Eugymnostomum 151
Euhydrogonium 146
375
INDEX
Euhymenostomum 178
Euphascum 231
Eupottia 231
Eupottiaceae 163
Eustreptopogon 142
Eusyntrichia 258
Eutortula 227
Eutrichum 68
Euweisia 176
Euweisiaceae 163
Euwillia 233
evolution, reticulate 12
exothecial cells 16
in palisade 232
Fa/ax 145
Fa/laces 135, 157, 160
Fallaciformes 158
filaments, costal 5
formaldehyde 4, 150
Fragiles 100
Funaria macrocarpa 278
Funariaceae 14
Ganguleea 186, also 167, 176
Geheebia 157
gel, lactophenol 9
genera, non-pottiaceous 25
geographic distributions 30, 46
Gertrudia 74
Gertrudiella 74, also 17, 96
Gertrudielloideae 74, also 49
Globulina 115, 202
Globulinella 202, also 115, 174, 237
glossary 372
Gomphoneuron 157
Gondwanaland 31, 98
Graciles 158
Grimmia 38
Grimmiaceae 276
guide cells, more than one layer 96
Gymnohyophila 170
Gymnomitriella 156
Gymnostomiella 156, also 252
Gymnostomum 151, also 140, 148, 178,209
lessonii 278
Gymnoweisia 138
Gymnoweisieae 136
Gymnoweissia 123
Gyroweisia 138, also 148, 151, 209
hairs, axillary 4, 146
Hapalostomae 178
Helicopogon 145
Hennedia 242
Hennediella 242, also 219, 258, 261
Herzog 32
heterogeneity, taxonomic, measure of 54
Hilpert 32, 50
Hilpertia 270, also 195, 230
Holomitria 151
homoplasy 12
Husnotiella 2, 157
glossophylla 115
hyalodermis 4, 104
Hyalophyllum 195
hybrids 184
Hydrogonium 145, 148
hydroid strand 84, 160
encircled 251
satellite 208
Hygrophila 170
hymenium 3
Hymenoste~ium 123
Hymenostomum 2, 88, 178, 279
Hymenostyliella 166
Hymenostylium 123, also 85, 123, 135, 151, 169
Hyophila 170, also 88, 104, 167, 174,273
involuta 176
Hyophilaceae 163
Hyophiladelphus 145, 148
Hyophileae 163, also 28, 49, 148
Hyophilodonta 189
Hyophilopsis 233, also 174, 233, 249
Hypodontium 164, also 13, 79, 142
identification 10
key
to genera 54
to suprageneric taxa 53
keys 10
Kingiobryum 76
Kleioweisia 178
Kleioweisiopsis 274
KOH 3, 30, 150
lactophenol gel 9
lamellae, costal 5
lamina, basal margins serrulate 14
Laminanchium 90, also 86, 173
Lancifolia 86
leaves 4
panduriform 235
perichaetial 7
Leptobarbula 208, also 44, 107, 140, 148, 151
Leptodon 132
Leptodontieae 120, also 49
Leptodontiella 130
Leptodontiopsis 2
Leptodontium 132, also 4, 79, 84, 110, 120, 127, 130
viticulosoides 279
Leptomitrium 86
Leptophascum 255
Leptopogon 145
Limbatae 227
Limneria 157
lineages, ancient, remnants of 30
Luisierella 193, also 186
lumping 15,4
Luridae 157
lutant 9
Lydiaea 237
Macrobryum 178
Macroglossum 277
Macromitrium 275
376
magnifications 3
mamillae 5
margins, leaf 5
Melophyllum 274
membrane, of capsule mouth 181
Merceya 153
Merceyaceae 136
Merceyeae 136
Merceyoideae 105, also 43, 49, 101
Merceyopsis 153, 176
methods, phylogenetic analysis 18
Microbryum 237, also 231, 253
Microstoma 178
Microweisia 176
Mildea 231
Mildeella 231
Mironia 112, also 84, 255, 261
Mitthyridium 164, 275
Molendoa 167, also 98, 153, 164
Mollia 145
monographs 10
Morinia 167
mucilage 4
Muraliformes 227
Muticae 237
Nanotortella 176
nematode 195
Neobarbula 242
Neocardotia 110
Octoblepharum 164
Odontophylla 145
operculum 8
optical section 373
Oreoweisia 164
brasiliensis 277
Orthocarpae 112
Orthopodiae 218
Orthotrichaceae 13, 277
Orthotrichella 153
outgroup 16, 25
Oxystegus 90, also 79, 80, 86
Ozobryum 167
Pachyneuropsis 80
Pachyneurum 80, 227
Pachynoma 146
pad, costal 5
paedomorphism 157
Paenecrossidium 196
papillae 5, 6
Paraleptodontium 86, 88
perichaetia 7
perigonia 7
peristome 8
twisted 14, 68, 372
Phascaceae 1, 163
Phasconica 178, 183, 213
Phascopsis 249, also 23'/
Phascum 230
carinatum 278
phenocopy 153
GENERA OF THE POTTIACEAE
Physcomitrium mirabile 278
Physedium 230
Piliferae 218
Platyneuron 116
Plaubelia 174, also 202
Pleuridium 274
Pleurochaete 96, also 79, 104
Pleuroweisia 167
Pleuroweisieae I, 27, 163
Pleuroweisioideae 163
Pleurozygodon 136
Pohlstoffe 8 ·
polarization, character state 19
Polytrichum 17, 22, 33, 34, 39, 40
pores, resorption 7
Pottia 230, also 163,201,231,245,249
macrocarpa 278
mirabilis 278
Pottiaceae 52, also 1
look alikes 10
Pottiales 12
Pottieae 208, also 49
Pottiel/a 237
Pottioideae 163, also 26, 44, 49
preparation, microscopic 9
previous authors 32
Prionidium 157
propagula 2, 7, 14, 157, 258, 273
Pseudaloina 196
Pseudocrossidiella 146
Pseudocrossidium 116, also 76, 96, 110, 115, 116, 148, 164,
196,211,230,247,271
crinitum 267
Pseudodesmatodon 227, 231
Pseudodidymodon 158
Pseudohyophila 275
Pseudopottia 178
Pseudosymblepharis 79, also 88, 98, 101, 181, 183
Pseudotimmiella 275
Pterygoneurum 199
Ptychomitriaceae 12
Ptychomitrium 12, 17, 22, 27, 36, 42,94
incurvum 277
Purpureaeformes 146
Pusilli 176
Pycnocaulon 211
Pycnocaulus 151
Pycnophyllum 86
Quaesticula 186, also 94
Racopilum 278
Rechingerella 176
reduction 2, 13, 14, 19, 26, 30
series 32, 153, 183
Reflexae 158
refugia 31
Reimersia 123
relationships, intrafarnilial 13
relicts 30
revisions 10
Revolutae 116
INDEX
Rhabdoweisia 275
Rhachitheciaceae 277
Rhachithecium 277
Rhamphidium 2, 275
montanus 278
Rhexophyllum 110
rhizoids, insertion patterns 3
Rhystobarbu/a 145
Rigidu/ae 157
Rott/eria 170
Rubiginosae 158
Rufidulus 158
Rurales 258
Rura/iformes 258
Sagenotortula 273, 247, 249
Saito 32, 50
Saitoa 236
Saitoella 236, also 202, 232
Sarconeurum 255
Saxicolae 84
Schistidiel/a 242
Schizophascum 232, also 195,211, 216
sclerodermis 4, 104
Scopelophila 153, also 4, 242
ligulata 237
Scou/eria 276
Sebi/lea 275
section, optical 6
sectioning 8
Selagine/la 4
Seligeria 115, 208
Senophyl/aria 157
Senophy/lum 145
sequencing convention 29
series, transformation 14
Serpotorte/la 275
seta, direction of twist 372
sexuality 9, 7, 193
Simophyllum 176
sister group 25
Spartina 4
Spathulidium 138
species, definition 16
Sphaerangium 176, 253
Spha/erostomae 176
Splachnobryaceae 157, 276
Splachnobryel/a 156
Splachnobryum 276
spore 8, 14
ornamentation 11, 222
sporophytes, borne laterally 120, 189
Spruceella 277
Squamigerae 196
Squarrosa 96
states, of characters 19
Stegonia 195, also 202, 232, 273
latifolia 123 7
'\
stem 4
I
Stephanodictyon 86, 88
stereid band 5, 13, 163
377
stock, ancient 31
Stonea 252
strand
central, of stem 5
hydroid or leaf 4, 5
satellite 4
strategies, life history 14
Streblotrichum 145, 209
Streptocalypta 94, also 76
Streptopogon 142
Streptopogonel/a 142
Streptotrichum 130
subfamilies 29
Subulatae 227
Subulataeformes 227
summary, phylogenetic analysis 16
Symb/epharis 80
synonyms 53
Syntrichia 258, also 2, 219, 233, 237, 245
abruptinervis 138
caninervis 252
inermis 216
pagorum 100
papillosa l 00
Syntrichiae 258
Syrrhopodon 84, 164
Syrrhopodontaceae 12
Systegium 178
taxa, infrageneric 53
Taxonomic Section 52
technique 8
Teniola 176
Teniolophora 176
terminology 7
Tetracoscinodon 105, also 43, 166
Tetracoscinodontieae 105, also 49
Tetrapterum 209, also 88, 90, 148, 216, 232
theca 7
ribbed 277
Thermopsis 6
Timmia 12, 17, 22, 35, 41
Timmiel/Q 68, also 17, 45, 167, 275
Timmielloideae 68, also 49
Tisserantiella 173, 277
torsion 7
Tortel/Q 100, also 80, 90, 98, 193, 213
tortel/oides 255
Tortelleae 82
Tortel/oidea 100
Tortilia 178
Tortobarbula 145
Tortu/Q 218, also 27, 196, 203, 227, 239, 242, 247
acaulon 378
ammonsiana 138
[atherodes, inval. see Tortula acaulon] 199, 239
cuneifo/ia var. blissii 223
deciduidentata 242
domingensis 249, 278
maritima 237
montana 278
378
GENERA OF THE POTTIACEAE
Uleastrum 277
Uleobryum 218, also 213, 216
Uleopsis 277
Unguiculatae 145
upper !aminal cells, asymmetrically bulging 181
Vallidens 258
variation, morphologicall6
Vinea/es 158
Vinealiformes 158
Viridivellus 181
Viridulae 176
Walther 11,32
wastebasket genus 140
weighting 15, 25
Weisiodon 138
Weisiopsis 189, also 2, 138, 176
nigeriana 213
Weissia 176, also 79, 88, 98, 163
breutelii 185
macrospora 213
W eissiaceae 1, 163
Weissiodicranum 184
Willia 233
Williamsia 132
Williamsiella 132
Zygotrichia 227
Zygotrichodon 157
paulsenii 258
pygmaea 255
revolvens 237, 271
Tortulaceae 1
Tortulae 227
Tortularia 178
Tortuloideae 163, 222
Tortuosae 100
Trachycarpidium 211, a!so 216, 218
Trachyodontium 127
transformation series 15
tribes 29
Trichostomaceae 1, 82
Trichostomeae 1, 82
Trichostomoideae 82, also 27, 49
Trichostomopsis 2, 157
Trichostomum 86, also 79, 88, 101, 170, 181,211,279
brachydontium 193
caespitosum 219
crispulum 186, 261
hibernicus 101
Tridontium 276, also 107, 132, 166
trigones 5
Triquetrella 120
Tuerckheimia 94, also 90, 167, 169, 186
twist, direction of 8, 372
Ulea 277
CORRIGENDUM
Because it is superfluous under I.C.B.N. Art. 63, the name
Tortula atherodes Zand., given as a nomen novum on p. 222 and
used through the text, is not acceptable and therefore invalid
under Art. 34.1. All the new infraspecific combinations with this
binomial, none of which are "alternative names" sensu Art. 34.3,
are also invalid for the same reason. Tortula acaulon, below, is
the correct name in Tortula, and correct infraspecific combinations are also provided below.
New combinations:
Tortula acaulon (L. ex With.) Zand., comb. nov. (Phascum
acaulon L. ex With., Syst. Arr. Brit. Pl. ed. 4., 3: 768, 1901
"acaule.")
Tortula acaulon var. arcuata (Hermstadt & Heyn) Zand., comb.
nov. (Phascum cuspidatum var. arcuatum Hermstadt & Heyn,
Bryologist 94: 175, 1991).
Tortula acaulon var. affinis (Nees & Hornsch.) Zand., comb. nov.
(Phascum affine Nees & Hornsch., Bryol. Germ. 1: 74, 1823;
Phascum cuspidatum var. affine (Nees & Hornsch.) Hampe).
Tortula acaulon var. curviseta (Dicks.) Zand., comb. nov.
(Phascum curvisetum Dicks., Pl. Crypt. Brit. 4: 2, 1801; Phascum cuspidatum var. curvisetum (Dicks.) Nees & Hornsch.).
Tortula acaulon var. diaphora (Hag.) Zand., comb. nov.
(Phascum acaulon var. diaphorum Hag., K. Norsk. Vid.
Selsk. Skrift. 1928(3): 19, 1929; Phascum cuspidatum var.
diaphorum (Hag.) C. Jens.).
Tortula acaulon var. elata (Brid.) Zand., comb. nov. (Phascum
elatum Brid., J. Bot. (Schrader) 1800(1): 269, 1801).
Tortula acaulon var. intertexta (Brid.) Zand., comb. nov.
(Phascum intertextum Brid., Mant. Muse. 8, 1819.)
Tortula acaulon var. marginata (Hernnstadt & Heyn) Zand.,
comb. nov. (Phascum cuspidatum var. marginatum
Hernnstadt & Heyn, Bryologist 94: 175, 1991).
Tortula acaulon var. mitraeformis (Limpr.) Zand., comb. nov.
(Phascum cuspidatum var. mitraeforme Limpr., Laubm.
Deutsch!. 1: 187, 1885).
Tortula acaulon var. papi/losa (Lindh.) Zand., comb. nov.
(Phascum papillosum Lindh., Oefv. K. Vet. Ak. Foerh. 21:
217, 1864; Phascum cuspidatum var. papillosum (Lindh.)
Roth).
Tortula acau/on var. pilifera (Hedw.) Zand., comb. nov.
(Phascum piliferum Scherb. ex Hedw., Spec. Muse. 20,
1801; Phascum cuspidatum var. piliferum Scherb. ex
Hedw.) Hook. & Tayl.).
Tortula acaulon var. retortifolia (Guerra & Ros in Guerra,
Jimenez, Ros & Carri6n) Zand., comb. nov. (Phascum
cuspidatum var. retortifolium Guerra & Ros in Guerra,
Jimenez, Ros & Carri6n, Cryptogamie, Bryol. Lichenol. 12:
390, 1991).
Tortula acaulon var. schreberiana (Dicks.) Zand., comb. nov.
(Phascum schreberianum Dicks., Pl. Crypt. Brit. 4: 2,
1801).