Banisteriopsis, Diplopterys (Malpighiaceae)

Organization for Flora Neotropica
Banisteriopsis, Diplopterys (Malpighiaceae)

Author(s): Bronwen Gates
Source: Flora Neotropica, Vol. 30, Banisteriopsis, Diplopterys (Malpighiaceae) (Feb. 18, 1982),
pp. 1-237
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
Stable URL: http://www.jstor.org/stable/4393754 .
Accessed: 22/08/2011 08:54
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
New York Botanical Garden Press and Organization for Flora Neotropica are collaborating with JSTOR to
digitize, preserve and extend access to Flora Neotropica.
http://www.jstor.org
BANISTERIOPSIS, DIPLOPTERYS
(Malpighiaceae)
BRONWEN GATES
^.\
C
f?,
T
TROPIC
...
Of
CANCER
FLORA!
NEOTROPICA,
TROPIC
Of
CAP R ICORN
FLORA NEOTROPICA MONOGRAPH

Number 30
The New York Botanical Garden

Bronx, New York 10458
Issued 18 February 1982
Copyright ? 1982
The New York BotanicalGarden

Published by
The New York Botanical Garden

Bronx, New York 10458
International Standard Serial Number 0071-5794
This material is based upon research supported by the National Science Foundation under
Grants No. GB-37314 and No. DEB 76-01757 and is published with support of National
Science Foundation Grant No. DEB-8005535.
The Foundation provides awards for research and education in the sciences. The awardee
is wholly responsible for the conduct of such research and preparation of the results for the
publication. The Foundation, therefore, does not assume responsibility for such findings or
their interpretation.
Any opinions, findings, conclusions, or recommendations expressed in this publication are
those of the author and do not necessarily reflect the views of the National Science Foundation.
Library of Congress Cataloging in Publication Data
Gates, Bronwen, 1945Banisteriopsis, Diplopterys(Malpighiaceae)

(Flora neotropica ; monographno. 30)
Bibliography:p.
Includes index.
1. Banisteriopsis.2. Diplopterys. 3. BotanyLatin America-Classification. I. Title.
II. Series.
81-11278
QK495.M26G37 583'.214
AACR2
ISBN 0-89327-238-8
All material subject to this copyright may be photocopied for the non-commercial purpose of
scientific or educational advancement.
A MONOGRAPH OF BANISTERIOPSIS AND

DIPLOPTERYS, MALPIGHIACEAE
BRONWEN GATES'
CONTENTS
Introduction .............................................
.............................
Circumscription, distribution and subdivision of the genera .................................
Morphology ...................................................................
.......
H abit .............................................................................
H airs ... ...... ..... ............................
.... ..............
.. ...............
Stems ............................................................................
Leaves ...........................................................................
Inflorescences .....................................................................
Floral parts .......................................................................
Calyx ........................................................................
Corolla .......................................................................
Stamens ......................................................................
Gynoecium ...................................................................
Fruit ...............................................1..............
..............
Chromosome numbers .................................................................
Taxonom y ........................................
...................................
Banisteriopsis .....................................................................
.......................
.............
Diplopterys ...................................
Acknowledgments .....................................................................
Literature Cited .......................................................................
Numerical list of taxa ..................................................................
List of exsiccatae .............................................................
.........
Index to scientific and common names ...................................................
1
2
7
7
8
8
10
12
12
12
14
16
17
17
18
18
19
208
216
217
218
220
231
INTRODUCTION
The genus Banisteriopsis is one of the largest and most widespread genera in
the Malpighiaceae. The genus has presented considerable difficulty to botanists
interested in the correct identification of plants in this genus because many of the
species are very variable, have considerable synonymy, and even the name of
the genus has presented nomenclatural problems. The last monographic revision
of the genus by Franz Niedenzu (1928) recognized 74 species. Since that time,
much additional material of Banisteriopsis has accumulated, and in the present
revision I recognize 92 species of which 34 are new. Many of the widespread and
variable species still present problems, but I hope that I have been able to identify
these problem areas for further study.
The genus Diplopterys is included with this treatment of Banisteriopsis because
the two genera appear to be very closely related; in flower it is very difficult to
distinguish them.
Plants now included in the genus Banisteriopsis were first circumscribed under
the name Banisteria L. by Adrien de Jussieu (1832 in Saint Hilaire, 1840, 1843).
The name Banisteria L. as used by Humboldt, Bonpland and Kunth (1822) included also plants now segregated as the genus Stigmaphyllon. Jussieu's usage
of Banisteria L. was followed by many subsequent workers, such as Grisebach,
who treated the Malpighiaceae for Martius' Flora brasiliensis (1858), and Nie1 The
University of Michigan Herbarium, Ann Arbor, Michigan 48109, U.S.A.
Flora Neotropica
denzu (1900, 1912), who monographedthe Malpighiaceaefor Das Pflanzenreich

(1928). Unfortunately, Jussieu did not include any of the Linnaean species of
Banisteria in his usage of the name. Of the seven species included by Linnaeus
in Banisteria, one is in the Rhamnaceae,and six belong to the genera Stigmaphyllon, Heteropterysand Hiptage in the Malpighiaceae.This fact was noted by
Robinson and Small in Small's treatmentof the Malpighiaceaefor North American Flora (1910). Small (1910) advocated the use of the name Banisteria L. to
replace Heteropterys H.B.K. (since three of the Linnaean species of Banisteria
belong to the genus Heteropterys),and Robinson (in Small, 1910)introducedthe

name Banisteriopsis to replace the name Banisteria as used by Jussieu. Small's
usage of Banisteria L. did not find general acceptance, and the name Heteropterys was conserved by the InternationalBotanical Congress in Cambridgein
1930. The name Banisteria continued in general usage for plants of the genus
Banisteriopsis, and was used by floristicianssuch as Kostermans (1938), Macbride (1950), and Standley & Steyermark (1946). A few botanists, such as
O'Donell and Lourteig in their treatment of the Malpighiaceaeof Argentina
(1943), and Cuatrecasas in his Prima Flora Colombiana (1958), used the name
Banisteriopsis. However, it was not until proposals to conserve the name Banisteria for this genus (Anderson, 1967; Morton, 1967)were rejected (McVaugh,
1968)that the name Banisteriopsisgained more generalacceptance as the correct
name for this importantgenus. Unfortunately,Robinson (in Small, 1910)designated Banisteria brachiata L., which is a species of Heteropterys, as the type
of Banisteriopsis. Cuatrecasas (1958) selected B. cornifolia as lectotype, but
without proposing conservation of the name Banisteriopsis with the new type.
Such a proposalwas publishedrecently (Gates, 1977a),but it has been withdrawn
because it was deficient in several respects. That proposal will be revised and
resubmittedin the near futurein the hope that soon the name of this large group
of neotropicalplants will finally be stabilized.
CIRCUMSCRIPTION,DISTRIBUTIONAND SUBDIVISION
OF THE GENERA
The Malpighiaceaeare characterizedby opposite leaves, stipules, a five-parted

calyx which is often glandular,a corolla of five free clawed petals, usually ten
stamens, and a three-partedpistil. Niedenzu divided the family into two subfamilies, the Planitoraewith a flat torus and unwingedfruits, and the Pyramidotorae
with a pyramidaltorus and winged or bristly fruits. These subfamilynames used
by Niedenzu are invalidunderthe Rules of BotanicalNomenclature,and Morton
(1968)substitutedvalid names for the names used by Niedenzu in his infrafamilial
classification.Thus, the Pyramidotoraeto which Banisteriopsis and Diplopterys
belong was renamedthe Gaudichaudioideae.However, Niedenzu's infrafamilial
classificationis unsatisfactoryand the infrafamilialtaxonomyis now being revised
by Dr. W. R. Andersonat The University of Michigan,so that the names as well
as the groupingsare subjectto change(see Anderson, 1977).Niedenzu recognised
two tribes in the Pyramidotorae,the Hiraeeae, with the principal wing of the
samara lateral in position, and the Banisterieae with the principal wing of the
samaradorsal in position. Banisteriopsis is a member of the Banisterieaesince
the principalwing of the samarais dorsal in position. The genus is characterised
by a fruit of three samaras, each with a well-developed dorsal wing which is
thickenedalong the upper margin,minuteinterpetiolarstipules, pedicels that are
usually sessile, ten fertile stamens, and three styles with terminalstigmas. Diplopterys has been traditionallyincludedin the Hiraeeae as its fruits lack a dorsal
Banisteriopsisand Diplopterys
wing and have well-developedlateralwings. However I considerthat Diplopterys

is most closely related to Banisteriopsis on the basis of the similarityof flower
structure,and because the fruit can be interpretedas a banisteriopsoidfruit with
the dorsal wing reduced to a crest. The genus Diplopterys is characterisedby a
fruitof 3 mericarpswith well-developedlateralwinglets interconnectedby crests,
minute interpetiolarstipules, spreadinglingulate bracts and bracteoles, sessile
pedicels, petals which are externally sericeous, ten fertile stamens and three
styles with terminalstigmas.
Both Banisteriopsis and Diplopterys are entirely New World in their distribution. There are a few species of Banisteriopsis which extend their range into
the subtropicsin Mexico, Paraguayand Argentina,but most species are restricted
to the tropics, with nearly two-thirds of the species entirely Brazilian in their
distribution.Almost half of the species of Banisteriopsis are savanna species,
with more than two-thirdsof these species in the Planaltoregion of Brazil, in the
characteristicsavanna vegetation of this area known as cerrado. Possibly the
diversificationof the genus occurredduringthe Pleistocene, duringthe postulated
periods of greatlyincreased savannavegetation (cf. Haffer, 1969;Simpson, 1971;
Vanzolini, 1970).Diplopterysconsists of only four species. Two are more or less
widespread species, one in Amazonian Brazil, Venezuela, Colombia and Peru,
and the other in NE Brazil, French Guiana, Surinamand Guyana. The other
two are apparentlyendemic species, one from Veracruz, Mexico, and the other
from the Rio Cururuin AmazonianBrazil.
In his monographof the genus Banisteria in 1900, Niedenzu recognized three
subgenera, based on differences in the form of the inflorescence and fruit. The
subgenus Hemiramma was characterizedby paniculate or racemose inflorescences, rarely umbels, pedicels borne on short peduncles, glabrous styles and
petals, and the nut of the samaravery smooth or sometimes with a single lateral
wing or crest on each side, the subgenus Eubanisteriaby pink, white or rarely
dull-yellow,glabrouspetals, capitellatestigmas,andthe nut of the samarasmooth,
rugose, alulateor aculeate, and the subgenusPleiopterys by yellow petals drying
black and often externally sericeous, long hairs on the ovary, the nut of the
samara with crests or several lateral wings, and the wing of the samara more
or less contractedat the base and with the upper marginappendiculate.
I have looked at all the species in the genus with regard to these characters
used by Niedenzu to define the subgenerain order to evaluate his subgeneric
groupings, and have also looked for additionalcharactersto define subgenera.
The form of the inflorescence, the presence of a peduncle and the surface of the
nut of the samara are unsatisfactorybases for separatingthe subgenera Hemiramma and Eubanisteria, and additionalcharactershave caused me to redefine
these two subgenera. Niedenzu's section Pseudobyrsonima of his subgenus
Hemiramma does not belong in the genus Banisteriopsis; his section Monoctenia
of subgenus Hemirammaforms the basis for subgenus Banisteriopsis, and his

section Leiococca of the subgenus Hemirammatogether with his subgenus Eu-
banisteria form the amended subgenus Hemiramma. The subgenus Pleiopterys
is maintained,based on some of the charactersused by Niedenzu, such as the

appendageon the upper marginof the wing of the samaraand the long hairs on
the ovary; the group is also supportedby additionalcharactersnot used by Niedenzu (see Table I). It is possible that germinationmay also be useful for defining
the subgenera;the cotyledons of seeds of the single species of subgenusBanisteriopsis which I successfully germinatedwere epigeal, whereas those of the four
species of the subgenusHemirammaand the two species of subgenusPleiopterys
were all hypogeal.
Comparison
Table I
of the Subgenera
Subgenus Banisteriopsis
(spp. 1-8)
of Banisteriopsis
Subgenus Hemiramma
(spp. 9-66)
Vines, shrubs, shrublets, trees
On the lamina, or a single pair
petiole, rarely marginal
Habit
Shrubs or vines
Leaf glands
Marginal
Inflorescence
Paniculate, flowers borne alternately

or in pairs
Pedicels
Short-pedunculate, often thickened

and woody in fruit
Cymes or dichasia of four-flowe

umbels, or paniculate with th
flowers borne alternately or i
Sessile or rarely pedunculate, n
thickened in fruit
Calyx glands
Flower color
Calyx glandular, glands free apically

and often decurrent onto the
pedicel, borne below free part of
sepals on receptacle
Yellow
Calyx glandular, rarely eglandu

glands attached throughout th
length, not decurrent, borne
free part of the sepals
Yellow, pink or white
Stamen connectives
Eglandular
Eglandular or glandular, the gla

tissue in the form of large inf
polygonal cells
Gynoecium
Ovary short-appressed-sericeous,
styles glabrous or basally
appressed-pubescent
Ovary velutinous or sericeous,

hairs often with unequal arm
longer arm up to 1 mm; style
glabrous or basally appressed
pubescent
Carpophore
Short, up to 1 mm, broad, nonfunctional
2-4 mm long, functional
Nut
Spherical; surface smooth laterally or

with 1 winglet on each side
Laterally compressed, rarely

spherical; surface smooth to
rugose, tuberculate or alulate
Basal appendage on fruit

wing
Shallow, up to 2 mm high
None or rarely a shallow appen

1-2 mm high
Table II
Comparisonof the Sections of Subgenus Pleiopterys
Section Sciurostylis(spp. 67-73)
Leaf glands
Inflorescence
One to two pairs on the petiole or

the laminaat the base, sometimes
with a single stalkedgland on the
marginon each side of the midrib
Four-floweredumbels borne in the
leaf axils or arrangedin dichasia
Calyx glands
Absent
Petals
Glabrous,except B. heterostyla
Styles
Curved,either sigmoidor incurved

apically, and densely hairy with
spreadinghairsfor 1/3 to 2%of their
length, except B. valvata
Fruit
Carpophoreup to 2 mm long,
apparentlyfunctional;hairs on
fruit appressed,stiff and irritating
to the skin
Slender, not thickenedand woody in
fruit
Section Anisopterys(spp. 74-92)

Marginal,often more numerous
towardsthe apex, rarelyon the
laminanear the margin
Short axillaryracemes or fourfloweredumbels borne in the leaf
axils, rarelythe umbels arranged
in cymes
Present, except in two species,
borne below the free part of the
sepals
Externallysericeous, except B.
caduciflora
Fruitingpedicel
Straightand diverging,sometimes
the anteriorstyle apicallyinflexed,
basallyhairy or the hairs
extendingup to half the length of
the anteriorstyle
Carpophoreup to 1 mm long, broad,
non-functional;fruitglabrateor
with sparse, appressed,nonirritatinghairs
Often thickenedand woody in fruit
I have subdivided the subgenus Pleiopterys into two sections, section Sciurostylis and section Anisopterys, based on such charactersas the leaf glands,
form of the inflorescence, calyx glands, form of the styles and certainfruit characters (see Table II). It is to the species of section Anisopterys that the genus
Diplopterys appears to be most closely related, based on similarity of flower
structure.
Each subgenus or section of Banisteriopsiscan be divided into several groups
of species, with all members of a group sharing the same flower and/or fruit
characteristics;some groupscontainonly one species. TableIII is an enumeration
of some of the charactersused to separate these groups of species and is meant
as an aid to identificationby providinga means of allocatinga plant to one of the
species groups within the genus. It can be used as an alternativeto the key or to
corroborateit.
List of CharactersUsed in Table III
1. Position of leaf glands
A. Numeroussmallglandson the marginor on the laminanearthe margin;sometimesthe basal
pair of glandsborne on the petiole
B. One (two) pairs on the petiole, sometimeswith one gland on the marginon each side of the
midrib
C. One to six pairs of glands on the laminaabaxiallybetween the midriband margin
D. One (two) pairs beside the midribat the base
Table III
Comparison of Species Groups within Banisteriopsis. Spp. 1-8 Comprise Subgenu
Hemiramma, and Spp. 67-92 Subgenus Pleiopterys; Spp. 67-73 Comprise Sectio
terys.
1
A
B. cornifolia group, spp. 1-7
A
B. longipilifera, sp. 8
C
B. laevifolia, sp. 9
C
B. argyrophylla group, spp. 10-14
C
B. schizoptera group, spp. 15-17
D
B. malifolia group, spp. 18-23
B
B. megaphylla group, spp. 24-25
C/D
B. campestris group, spp. 26-30
C
B. calcicola, sp. 31
C
B. membranifolia group, spp. 32-34
C
B. caapi, sp. 35
C
B. schwannioides, sp. 36
C
B. prancei, sp. 37
C
B. multifoliolata, sp. 38
C
B. quadriglandula, sp. 39
B/C
B. muricata group, spp. 40-43
A
B. martiniana group, spp. 44-54
B
B. nummifera group, spp. 55-59
C/D
B. parviflora group, spp. 60-64
B
B. pseudojanusia, sp. 65
D
B. magdalenensis, sp. 66
B
B. valvata, sp. 67
B
B. heterostyla, sp. 68
B
B. lutea group, spp. 69-70
B
B. leiocarpa group, spp. 71-73
A
B. lucida group, spp. 74-83
A
B. pubipetala group, spp. 84-88
C
B. caduciflora, sp. 89
A
B. krukoffii, sp. 90
A
B. patula, sp.91
A
B. sepium, sp. 92
A
A
B
B
B
B
B
B
B
B
B
B
B/C
B
B
B
A/B
A/C
B
B
B
B
B
B
B
B/C
B/C
C
B
B
B
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
B
B
A
C
C
C
C
B
A
C
C
A
A
A
A
B
B
C
C
C
C
C
C
C
C
C
B/C
C
C
C
C
C
C
C
A
A
B
B
B
B
B
B
B
B
B
B
B
B
A
A
B
A
B
B
B
B
B
B
B
B
B
B
B
B
B
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
B
A
A
A
A
A
A
A
A
A
B
B
C
C
C
C
C
C
C
C
A/C
C
C
C
C
A/C
A/C
C
A/C
C
C
A
A
A
A
B
B
B
B
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A/B
B
A
A
B
B
A
B
B
B
A
A
A
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
B
A
B
B
B
B
B
B
--
--^~~~~~~~~~~~
Banisteriopsis and Diplopterys
2. Form of inflorescence
A. Flowers borne on ultimatebranchesin racemes of 8-45 flowers
B. Flowers borne on ultimatebranchesin 4-floweredumbels
C. Flowers borne on ultimatebranchesin short axillaryracemes of 4-8 flowers
3. Bracts and bracteoles
A. Erect or appressed
B. Spreading,divergingwidely from the axis
4. Persistenceof bractsand bracteoles
A. Deciduousin bud or flower
B. Deciduouspost-flowering
C. Persistent
5. Floriferouspeduncle
A. Present
B. Absent
6. Sepal aestivation
A. Imbricate
B. Valvate
7. Position of calyx glands
A. Absent
B. Borne below free part of sepals or receptacle
C. On free part of sepal
8. Petal pubescence
A. Glabrous
B. Externallysericeous
9. Petals
A. Posteriorpetal erect, similarto lateralpetals; lateralpetals reflexedor suberect
B. Posteriorpetal erect, differentfrom lateralpetals; lateralpetals reflexedbetween sepals
C. All petals stronglyreflexedbetween sepals
10. Petal color
A. Yellow
B. White or pale pink, becomingoff-whiteor creamin age
C. Whiteor pale pink, becomingbutteryellow in age
D. Pink, often becomingwhite in age
11. Filaments
A. Subequal
B. Filamentsopposite sepals longerthan those opposite petals
C. Filamentsopposite 3 anteriorsepals longest
D. Filamentsoppositethe 3 styles (i.e. oppositeanteriorsepal and postero-lateralpetals) longest
E. Filamentsopposite anteriorsepal, postero-lateralpetals and postero-lateralsepals longest
12. Connectives
A. Connectives of stamens opposite 3 anteriorsepals greatly enlarged,those opposite anterolateralsepals overtoppinglocules 0.6-2.0 mm
B. Connectives of stamens opposite 3 anteriorsepals not overtoppinglocules, or overtopping
the locules 0.2-0.4 mm
13. Style cross-section
A. Round
B. Flattened
14. Nut locule
A. Glabrous
B. Densely hairy
15. Wingrotation
A. Wings of samarasequidistant
B. Wingsof posteriorsamarasat 90?to wing of anteriorsamara
C. Wingsof posteriorsamarasat more or less 45?to wing of anteriorsamara
16. Fruithairs
A. Stiff, irritatinghairs present
B. Stiff, irritatinghairs absent
MORPHOLOGY
Habit
The genus Banisteriopsis includes species which are shrublets, shrubs, small
trees and vines. The shrubby habit, with many stems arising from a thick woody
Flora Neotropica
base, is the most common habit of Banisteriopsis species in savannavegetation,

but often the ends of the branches will nutate and twine around support if it is
available. Thus, many of these shrubby species are facultative vines, and will
even sometimes vine on themselves forming dense thickets. However, in the
cerrado vegetation of the Planalto region of Brazil, which is apparentlya fire
climax vegetation (Eiten, 1972;Ratteret al., 1973),many species of Banisteriopsis have the habit of a shrublet. These plants have an undergroundxylopodium
from which unbranchedstems are producedannuallyor every two to three years
dependingon the frequency of burningor other damage to the aerial parts. But
more than half of the species of Banisteriopsis are always vines, and most of
these are forest species. A few savanna species such as B. anisandra and B.
gardnerianafrom the cerradoare always vines and form dense thickets by vining
on themselves. These differenthabits are not evenly distributedamong the subgenera of Banisteriopsis. The subgenusBanisteriopsis includes shrubs with vining tendencies and vines, and the subgenus Pleiopterys consists mainly of large
forest vines with a few species in the Planaltowith the habit of shrubor shrublet,
whereas the subgenus Hemirammaincludes some vines, but also many species
which are shrubs, shrubletsand a few species which are small trees. Most of the
species in the subgenus Hemirammaoccur in the Planalto, and it is mainly the
vining species which occur outside of this area.
All of the species of the genus Diplopterys are vines, and often grow along
river margins;it is probablethat the fruits are dispersed by water.
Hairs
Both Banisteriopsisand Diplopteryshave typical "malpighiaceous"hairs, i.e.
unicellular,medifixedhairs. They are of two basic kinds, T-shapedand Y-shaped
(Fig. 1). There is much variationin the size and form of the individualhairs, in
the length and thickness of the stalk and arms, and the surface texture. In Banisteriopsis the stalk is often attached somewhat eccentrically, and one arm is
shorter than the other. In some instances, one arm of a Y-shaped hair is greatly
reducedor even absent and thus gives rise to basifixedhairs. Such hairsare found
on the stems of B. grandifolia, B. basifixa and B. parviglandula, and on the
gynoecium of species in the subgenus Pleiopterys. Most of the hairs in both

Diplopterysand Banisteriopsisbearpapillaeon theirsurface,but in a few species,
B. laevifolia and some of the subgenusBanisteriopsis,the hairsare quite smooth.
Stems
All species of Banisteriopsis are woody. The young stems are often flattened
and in many species are hairy;older stems are always terete and usuallyglabrate.
Stem pubescence is usually a good characterat the species level, though a few
species such as B. variabilis show variationin this characterwithin the species
as I have defined them. Seedlings frequently exhibit a different kind of stem
pubescence, but rapidly assume the type of pubescence typical of matureindividuals. In species in which the young stems are glabrous, the stems are green
and photosynthetic.
Older stems develop a cork cambiumand form bark, but the bark is usually
not very thick, and is smooth or somewhat corrugatewith numerouslenticels.
The lenticels are either small, round, often raisedareas, or small, elongatefissures
which sometimes coalesce to give a more extensive network of shallow fissures.
In Banisteriopsis latifolia, B. lutea, and B. heterostyla, the cork layer on the
stem often becomes very thick and forms a series of prominentridges projecting
:X
-
.f
I'
'
. .....
'
"
' ;>S
?r
.-
~~~~~~,
'
-?
?.
.
/
-.
....
X'
~~~~~~~~~~~~~~~~~~~~~~~~~~~~?
...?.~
?I?~~~~~~~~.
"'v/
rFw
..?.
?
...
'
<t.
'
....
'
..'
.':"
'
Z ';! ~'.:.
?:1..
'~~~~~~~~~~~~~~~~~~~~~~~~~
.
;}
...Z"..
.::."*';
.':'
'
.'
'
'.
i~~~~~~~~~~~~~~~~~~~~~~~~~~~~;
0-
'-
:X0A
.. 'i[
..
'
-~?1
~
_' - *-".., '
.' . -... ~:
. - -;: :: 1... .
0-
;.
}
'>itd 0 -; - '//0 8
:
:~~~~~~~~~~~~~~~~~~~~~~~~~'
"
:'?~~~~~~~~~~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~~~r
... ?
<
'
'
~~~~~~~~~~~~~~~~~~~~~..
,~~~~~~~~~~~
~ ~ ~
:,.
'
?~
i....:~
.
.:?
.~~ .
.. " }
:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..r
. '.,.:..'-?..'}
?:'~~~
? ~ ~ ~~~..
:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
:~....
.;..;...':
"'
' ~.}'.
"'
?~~p~~~~~
~ :
"
Xvs
..
;'
:~i;/t
..
?I
61:
;r|~~~~~~~~~
...
pi
"I
'
. :
;.;,.
~'-'"
.?
" ...
'~
': ...~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.....:
. .
~~~~r
~~~~-~..
.. : .. ?..?r~F...:C.?
r."... ....
. -I
.....:.
...~,'~. .:
i-'?.
.,%
,.
f:
FIG.~~~~~~~~~~~
aiseipi.?AadB?-hpd
1~:.Tpso?aisi
~~~~~~~
iE~~~~~~~~~~~~~~~~~~~~~~
i:
ar;CadD Tsae
ar;E
ovary hairs of B. hypericifolia. Pubescence composed of hairs of types A or B is described as

velutinous pubescence. Pubescence composed of hairs of type C is described as sericeous pubescence.
Pubescence composed of hairs of type D is described as tomentose pubescence.
up to 2 cm fromthe stem. This thickcork layerdevelopsonly on old stems of

B. lutea andB. heterostylawhichare oftennot collected,butis developedeven
on youngstemsof B. latifolia.Banisteriopsislatifoliais a treein the cerradosof
the Planaltoof Brazilandthe thickcorkybarkis veryadaptivein thisfireclimax
10
Flora Neotropica
vegetation; such thick corky bark is found on many cerradoplants in many different families. Some collections with old branches of B. parviflora and B. elegans exhibit a papery barkwhich peels off in successive layers like a paperbirch
tree.
Apparentlyin B. caapi the xylem parenchymaform wide bands which traverse
the secondary xylem and subdivideit into discrete lobes (see Niedenzu, 1928,p.
7, fig. 2A). These lobes are often apparenton the outside of the stem as longitudinalridges spirallinground the axis.
The stems of Diplopterys are woody. Several collections of Diplopterys show
the secondaryxylem dividedinto two lobes which spiralaroundthe axis and give
the stem a flattened appearance. This is similar to the lobing of the xylem in
Banisteriopsis caapi. The lobes are not always developed and their significance
is unknown;perhapsit is associated with the liana habit of these species.
The stipules of both genera are interpetiolarand are usually borne near the
middleof the stem but sometimesare more lateralat the base of the petiole. They
are usually small, less than one mm in length, and triangular,but in the Banisteriopsis argyrophyllagroup they are linear and up to five mm long. The bases
of the stipules are often connected by an interpetiolarline. In B. oxyclada and
a few other vining species, this line develops into a prominentridge on which the
stipules are borne and gives the appearanceof large, fused interpetiolarstipules.
Leaves
The leaves of both Banisteriopsis and Diplopterys, as in most Malpighiaceae,
are simple, opposite and decussate. In a few species of Banisteriopsisthe leaves
are sometimes ternate or even four at a node. In some of these species, such as
B. parviflora, the leaves are ternate with sufficientfrequency so as to be almost
a specific character,whereas in other species, like B. campestris, stems from the
same xylopodium, or even the same stem, may have leaves one, two, three or
four at a node. One species, B. alternifolia,has alternateleaves; its close relative
B. elegans has opposite or suboppositeleaves. Possibly this is a developmental
anomaly, althoughon the basis of other charactersit appearsto warrantspecific
recognition.The laminais entire in all species of Banisteriopsisand Diplopterys,
though one young plant of B. scutellata had its first-formedleaves somewhat
lobed.
Many species of Banisteriopsis from the cerrado of Brazil exhibit features,
such as thick cuticles, rollingof the leaf marginand stomatasunkenand restricted
to the lower epidermis, which are xeromorphiccharacters. It has been shown
(Rawitscher, 1948; Ferri, 1963)that even at the height of the dry season in the
cerrado,water is freely availablewithinthe rootingdepth of most plants, so these
features are apparentlynot adaptationsto withstand water shortage. The most
satisfactoryalternativeexplanationfor these featuresis that they help to preserve
minerals in the plants by reducingfoliar leaching (Moraes & Arens, 1971);this
would be very adaptive in the oligotrophiccerrado soils.
Leaf size, as petiole length and laminalength and width, is very variablewithin
a species and even within individualsin both Banisteriopsis and Diplopterys. In
particular,the leaves associated with the inflorescence are usually very much
smallerthan the other leaves, and often reducedto bracts. Petiole lengthis sometimes useful at the specific level for separatingspecies within groups of closely
related species. Viningspecies usually have longerpetioles than shrubbyspecies,
and their longer petioles are capable of differentialgrowth to orient the lamina
11
to receive maximallight. The range of lamina size and the length/widthratio are
sometimes useful for separatingspecies within groups of species. Some shrubs
from the Planalto of Brazil have lanceolate or linear leaves while their nearest
relatives have ovate or elliptic leaves; most Banisteriopsis species, includingall
the vining species, have ovate or elliptic leaves which are usually acuminateand
often falcate. The lamina size of the cerrado species is, on the whole, smaller
than in the rest of the genus.
The venation of both Diplopterys and Banisteriopsis is brochidodromous.In
section Sciurostylis and most members of subgenus Hemiramma of Banisteriop-
sis the secondary veins connecting the lateral veins are more prominent than
other second order veins, and give a parallel or scalariformappearance.In Di-
plopterys and subgenus Banisteriopsis, section Anisopterys, and the B. nummi-
fera group of Banisteriopsis, all second and lower order veins are more or less
equally prominentand the venation appearsreticulate.
The leaves of some species of Banisteriopsisare entirelyglabrous, or hairy at
first and soon glabrate, but many species are persistently hairy and pubescence
characters are very useful at the specific level. In a few species, such as B.
nummifera and B. variabilis, hairy and glabrous leaves are represented in the
same species, but usually leaf pubescence is constant within a species. The pubescence of the adaxial and abaxial surfaces of the lamina is different, and that
of the midribis different from the pubescence of the rest of the surface of the
lamina. Usually the midribpubescence is the same as that of the petiole, and this
is often the same as the stem pubescence. All species of Diplopterysare sparsely
appressed-sericeousabaxially;in dry materialthe leaves look glabrous without
examinationunder a microscope.
Glands are present on the leaves of both Banisteriopsis and Diplopterys,
thoughthere is sometimes variationwithinan individualin the prominenceof the
glands, and sometimes the glands are only well developed on the inflorescence
leaves. However, I have found the position, type and size of the leaf glandsuseful
for defininggroups of species, especially within the subgenus Hemiramma. In
Diplopterys, and the subgenus Banisteriopsis and section Anisopterys of Ban-
isteriopsis, the glands are usually borne on the marginof the lamina; they are
very smalland sessile and are eitherevenly distributedor morenumeroustowards
the apex. The B. martinianagroupof the subgenusHemirammaalso has marginal
glands, but here the glands are often enlarged and cup-shaped and sometimes
developed into ciliate extensions at the apex of the leaf. In the B. nummifera
group of subgenus Hemirammaand in section Sciurostylis of subgenus Pleiopterys, the glands are usually borne on the petiole near the apex; sometimes the
petiolar glands are absent, or sometimes in section Sciurostylis they are borne
on the base of the lamina. Some species with marginalglands also have glands
on the petiole. This occurs in some Diplopterys species, and some membersof
the B. martinianagroup. In subgenus Banisteriopsis the basal marginalglands
are often enlarged and are sometimes borne on the petiole and not the lamina.
Most species in the subgenus Hemirammaof Banisteriopsis have glands on the
lamina, with one to six pairs of glands borne on the lateralveins, or a single pair
beside the midribnear the base. These glands are usually more or less stalked,
they are usually yellow, and they secrete sugarsolution which apparentlyattracts
ants. Some species of Banisteriopsis (e.g. B. megaphylla) are constantly frequented by ants. The presence of the ants perhaps discouragesherbivory;often
the glands are particularlywell developed on the inflorescenceleaves and damage
to the inflorescence would be particularlyundesirableto the plant.
12
Flora Neotropica
Inflorescence
The flowers of Diplopterys and Banisteriopsis are borne on pedicels which in
some species in the subgenus Banisteriopsis and section Anisopterys of Banis-
teriopsis become woody and thickened in fruit. The pedicel is borne on the floriferous peduncle which is very reduced and nearly or quite absent in Diplopterys
and most species of Banisteriopsis. It is well developed in the subgenus Banis-
teriopsis and the B. muricata and B. nummifera groups. The apex of the peduncle
is denoted by a distinctjoint; the peduncle bears a pair of bracteoles which are

often more or less apical and appearto directly subtendthe pedicel (Fig. 2). The
peduncle itself is subtended by a single bract on the flowering axis. In species
where the peduncle is reduced the bracts and bracteoles have the appearanceof
being clustered at the base of the pedicel on the floweringaxis. The bracts and
bracteoles are deciduous in bud or flower in some species of Banisteriopsis(e.g.
B. membranifolia group), but are persistent in most species of Banisteriopsis,

and in Diplopterys. In Diploterys and two species of Banisteriopsis the bracts
and bracteoles diverge widely from the pedicel and are described as spreading;
in most species of Banisteriopsis the bracts and bracteoles are appressed to the
pedicel or diverge at a small angle from the pedicel (Fig. 2). There is considerable
variationin the size and shape of the bracts and bracteoles between species, and
also in their abaxial pubescence. The bracts and bracteoles are always glabrous
adaxially.
When the flowers are clustered together on a branchand have the appearance
of an umbel, the branch subtendingthis umbel is termed the common peduncle.
Peduncleused without qualificationrefers to the floriferouspeduncle. Sometimes
the length of the common peduncle is useful for separatingspecies; it is usually
more or less constant in length for a taxon.
The ultimate branches of the inflorescence are racemose but there is great
variationbetween species in the numberof flowers on the ultimatebranches. In
the subgenus Banisteriopsis, the B. nummifera group and in part of the B. mar-
tiniana groupof Banisteriopsisthe ultimatebranchesbear many flowers arranged

in pairs or in a spiral,and the inflorescenceis paniculate(Fig. 2). In some species
of section Anisopterys of the subgenus Pleiopterys of Banisteriopsis, the ultimate
branches bear fewer flowers and the internodes between the flowers are short;
the inflorescence is of the form of a condensed axillary raceme (Fig. 2). In Di-
plopterys and most species of subgenus Hemiramma and subgenus Pleiopterys,
the ultimate branches usually bear two pairs of flowers with the internodes between them unelongated,so that the flowers appearto be borne in four-flowered
umbels. The umbels are borne in the leaf axils as single umbels in the B. lutea
groupand some species of section Anisopterys, but in the subgenusHemiramma
and most species of section Sciurostylis of Banisteriopsis and in Diplopterys, the
umbels are arrangedin cymes or dichasiawith the terminalumbel on each inflorescence opening first (Fig. 2). The B. parviflora group bears its flowers in a
"raceme" of umbels; this arrangementof umbels can be derived from a cymose
arrangementby regularisationof the branchingpattern.
A few species of Banisteriopsis, such as B. latifolia and rarely other species
of the B. malifolia group, and the B. lutea group, will produce flowers on old
stems in the axils of leaf scars of previous seasons' leaves.
FLORAL PARTS
Calyx
The calyx is always five-parted; most flowers of Banisteriopsis and Diplopterys exhibit a markedbilateralsymmetry,with two pairs of sepals, the antero-
13
~~A;; ~
*I
,
.
~ ~~.
:::..: c..
.
.r
'
, ;
FIG. 2. Types of inflorescencein Banisteriopsis, and arrangementof bracts and bracteoles. A,

4-floweredumbel;B, enlargementof A to show bract(b ), bracteoles(b2), peduncle(p ), and pedicel
(p2); C-E, dispositionof bracts and bracteoles:C, appressed;D, erect; E, spreading;F, paniculate
inflorescence;G, cymose arrangementof umbels;H, dichasialarrangementof umbels;J, "raceme" of
umbels; K, axillary raceme. Numbers 1-6 refer to order of opening of flowers or umbels; those
numbered"1" open first, etc.
14
Flora Neotropica
lateral and postero-lateralsepals, and an unpairedsepal, the anteriorsepal (Fig.

3). The anteriorsepal is usually eglandularor rarely bears a single small gland.
Each of the four lateral sepals usually is biglandular;these glands apparently
secrete an oily secretion which is collected by certain Hymenopterawhich are
also effective as pollinators(Vogel, 1974). In B. caapi, and the B. muricata, B.
parviflora and B. martinianagroups glandularand eglandularflowers are found
in the same species, whereas in some species of section Anisopterys and all
species of section Sciurostylis the calyx is always eglandular.I do not think all
these eglandularspecies are closely related to each other, so the loss of calyx
glands has apparentlyoccurredmore than once. In all these species, some other
attractantfor pollinatorsmust be in operation.
When calyx glands are present, they are borne on the relatively large free part
of the sepal and attached throughouttheir length in the subgenus Hemiramma,
whereas in the subgenus Banisteriopsis and section Anisopterys of subgenus
Pleiopterys and in Diplopterysthey are borne below the free part of the sepal on
the receptacle (Fig. 3). In the species with the glands borne on the receptacle,
the free partof the sepal is relativelysmall, and the glandsare usuallyfree apically
and project onto the free part of the sepal. In the subgenus Banisteriopsis the
glands are often decurrentonto the pedicel.
The color of the calyx glands varies with petal color: yellow flowers have
yellow glands, white flowers have green or brownish glands and pink flowers
have pink or red glands.
Sepal aestivation is imbricatein Diplopterys and all species of Banisteriopsis
except B. valvata which is valvate. Sepal pubescence, especially of the adaxial
surface, and sepal shape are useful for separatingsome groups of species. The
sepals are always persistent in fruit, and sometimes become enlarged in fruit.
This enlargementis particularlymarkedin B. valvata.
Corolla
The corolla in Banisteriopsis and Diplopterys consists of five clawed petals
which are imbricatein bud with the unpairedpetal innermost;throughoutI shall
refer to the unpairedpetal as the posterior petal and the paired petals as the
postero-lateraland antero-lateralpetals respectively (see Fig. 3). This is the terminology used by Niedenzu in his treatmentof the Malpighiaceaefor Das Pflanzenreich. Moreover, this terminologyrecognizes the bilateralsymmetryof these
flowers with the plane of symmetrypassing throughthe anteriorsepal and posteriorpetal. The posteriorpetal is posteriorwith regardto any pollinatorvisiting
the flower, and the flowers are always borne such that the posterior petal is
closest to the axis. The antero-lateralpetals are usually a little larger and more
concave than the postero-lateralpetals.
In Diplopterys and most species of Banisteriopsis the posteriorpetal is markedly differentfrom the four lateralpetals; usually the posteriorpetal is held erect
and acts as a "flag" to attractand orient pollinatinginsects, with the four lateral
petals reflexed between the sepals. There is much variationin the form of the
posterior petal. The limb can be orbicularor obovate, and the basal part may
bear two to four pairs of gland-tippedteeth. The claw can be laminar,enlarging
distally to merge graduallyinto the limb, or it can be thick and fleshy and more
or less constricted apically. In some species of subgenus Banisteriopsis and in
B. laevifolia, the posterior petal is scarcely distinguishablefrom the four lateral
petals and all the petals are suberect. In the B. leiocarpa group all the petals are
the same and strongly reflexed between the sepals.
..
15
''.::
.i/
"":"':-
'."
''
'
'a
FIG. 3. Diagramof floralparts of Banisteriopsis,and illustrationof gland position on the sepals.
a, axis; sl-5, sepals: sl, anteriorsepal; s2, s3, antero-lateralsepals; s4, s5, postero-lateralsepals;
pl-5, petals: pl, p2, antero-lateralpetals; p3, p4, postero-lateralpetals; p5, posterior petal; 1-10,
stamens; st 1-3, styles: stl, anteriorstyle; st2, st3, posteriorstyles. A, B, glands borne on the free
partof the sepal:A, side view; B, in section; C, D, glandsbornebelow free partof sepal on receptacle:
C, side view; D, in section.
16
Flora Neotropica
The petals of Diplopterys and of subgenera Banisteriopsis and Pleiopterys of

Banisteriopsis are yellow. In subgenus Hemiramma, B. laevifolia and the B.
martiniana and B. nummiferagroups have yellow petals; the other species in

subgenus Hemiramma have white or pink petals, and usually exhibit a color
change with age. Thus pink-floweredspecies become white in age, and whiteflowered species become cream-coloredin age. A few species such as B. membranifolia and B. adenopoda have pink or white flowers at anthesis, but become
pale butter yellow in age. These species have large inflorescences, and the old
flowers persist for several days so that they contributegreatly to the showiness
of the inflorescence and perhaps attract pollinatorsfrom a distance. In several
species of Banisteriopsis, such as B. hypericifolia, B. pulcherrima in subgenus
Pleiopterys and B. stellaris, and sometimes B. campestris in subgenus Hemiram-
ma, there is a red or deep pink patch in the middle of the antero-lateralpetals.
These are the petals which are outermostin bud and the pigmentationcorresponds
to the part of the corolla exposed between the sepals in bud. Perhapsthis anthocyanin provides protection from ultraviolet radiation at high altitudes for the
anthers at meiosis.
All species of Diplopterys and most species of Banisteriopsis section Anisop-
terys have externallysericeous petals. In addition, B. heterostyla and B. valvata

in section Sciurostylishave hairypetals. The form of the petal marginalso varies
between species. All species of Diplopterys and some species of Banisteriopsis
such as the B. parvifloragroup have fimbriatemargins. Iost species of Banisteriopsis have petals with lacerate or laciniatemargins,but a few species such as
B. laevifolia and subgenusBanisteriopsishave subentireor denticulatemargins.
Stamens
There are always ten fertile stamens in Diplopterys and Banisteriopsis, although one or two stamens are sometimes absent in B. pseudo-janusia, and in
the B. parviflora group, the locules of the stamens opposite the antero-lateral
sepals are aborted. To distinguishindividual stamens where necessary, I have
referredto their position with respect to the sepals and petals.
The filaments are glabrous and basally connate in Diplopterys and Banisteriopsis; they can be subequal, but more usually those opposite the sepals are
longer than those opposite the petals. One of the trends recognizablein the form
of the androeciumis an exaggerationof this differencein filamentlength, usually
associated with increasing differentiationof the anther connectives. In other
species the longest filamentsare those of the stamens associated with the styles
(i.e. the stamens opposite the anteriorsepal and the postero-lateralpetals), and
in these species the filaments of the three posterior stamens are sigmoid and
usually inflexed between the posterior styles. In B. schwannioides and some
specimens of B. gardneriana, only the anteriorstyle develops and the five posterior stamens are greatly reduced in size.
The anthers are basifixed and appendicular.The pollen sacs are longitudinal
and adaxial on the connectives near the lateral margins. Dehiscence is always
introrse and occurs at or slightly before anthesis. The medianarea of the pollen
sac where dehiscence occurs is membranousand forms a white membranousfrill
along the length of the marginof the pollen sac on each side. The pollen sacs can
be glabrous or more or less hairy. The connectives can be subequal and not
glandularas in the subgenus Banisteriopsis and some species in the subgenera
Hemiramma and Pleiopterys of Banisteriopsis or can be glandular and often
enlarged. The glandulartissue in Diplopterys and in the subgenus Pleiopterys
17
consists of papillae which are usually more numerous towards the apex of the
connective. In the subgenus Hemiramma,the glandulartissue consists of greatly
enlargedpolygonal cells which usually cover the apex of the connective in the
glandularspecies. The glandularcells appear to be thin walled and filled with
watery fluid in life, and in dry specimens the glandulartissue has a wrinkled
appearance;non-glandulartissue, which is composed of small thick walled cells,
maintainsits shape and a smooth surface on drying. When stamens are fixed in
FAA the glandulartissue of the connective becomes white, whereas non-glandular tissue on the connectives becomes dark brown. In the B. muricata group
and B. martiniana, the connectives of the stamens opposite the anterolateral
sepals are greatly enlarged,but only part of this tissue is glandular.The function
of this glandulartissue is unknown. Field examinationof these structuresdid not
reveal any exudation from them. The connectives are situated such that they
would come into contact with the ventral surface of a pollinatinginsect, and
perhaps the connectives make a secretion which is effective as a "glue" for the
adhesion of pollen to the body of the insect.
The pollen of Banisteriopsis is polyporate with residual furrows. Its form is
quite peculiar and is under study by S. Lowrie and Dr. W. R. Anderson at The
University of Michigan. It does not appear to be useful at the species level
in this genus.
Gynoecium
The gynoeciumin Diplopterysand Banisteriopsisconsists of three free carpels
adnate to a pyramidaltorus. The ovary is hairy, and these hairs persist on the
nut of the fruit. In the subgenus Pleiopterys, the apex of the ovary also bears
long hairs with unequalarms which are sometimes also present on the style. The
future dorsal wing or dorsal crest of the fruit is evident on the ovary as a shallow
crest. The single ovule in each locule is adaxial, pendent and anatropous,and the
nucellarbeak usually protrudesbeyond the micropyleinto the locule.
There are usually three styles in both Banisteriopsis and Diplopterys, distinguished as the anteriorand posteriorstyles (see Fig. 3). In B. schwannioidesand
sometimes B. gardnerianathe posterior styles are much reduced;in some other
species, the posterior styles are slightlylonger than the anteriorstyle. The styles
arise from the inner (adaxial)face of the ovary and are subterminal;usually they
are round in cross section, but in the B. argyrophyllagroup, B. laevifolia and
some individualsin the B. schizoptera group they are laterallycompressed. The
styles can be straightand parallel, or more or less curved and diverging.In the
subgenera Banisteriopsis and Hemiramma, the styles are usually glabrous or
sometimes appressed-pubescentat the base, but in the subgenusPleiopterys and
in Diplopterys, one or more of the styles are densely and loosely pubescent. In
the B. parviflora group, the two posterior styles are parallel throughouttheir
length and sometimes fused together apically.
The stigmasare terminal,and in many species stronglycapitate. The styles and
stigmas are persistent in fruit.
Fruit
The fruit in most species of Banisteriopsis consists of three samaras which
separate at maturity. Each samara has a well-developed dorsal wing which is
thickenedalongits uppermargin;in three species, B. cipoensis, B. paraguariensis
and B. sepium, the wing is reduced to a crest. In the subgenus Hemirammaand
in section Sciurostylis, there is typically a carpophoreassociated with each sa-
18
Flora Neotropica
marawhich separatesfrom the lower marginof the nut and supportsthe samara
after its separationfrom the torus. Presumablythis requires a minimumwind
speed to breakthe carpophorebefore dispersalcan occur. In the subgeneraBanisteriopsis and Pleiopterys the carpophoreis smalland apparentlynon-functional.
The seed completelyfills the locule of the nut when fully mature.In the subgenus
Hemirammathe areole or area of attachmentof the samarato the torus is usually
large and concave; in the subgeneraBanisteriopsis and Pleiopterys, the areole
is small.
In most species of subgenusPleiopterys, there is a prominentappendageat the
base of the wing. In the subgenus Banisteriopsis, the appendage is small and
shallow whereas in most species of subgenus Hemirammait is absent.
The fruits of many species of Banisteriopsis such as the B. malifolia and B.
martinianagroups and section Sciurostylisbear stiff hairs which are irritatingto
the skin. The type of pubescence on the fruit and the form of the fruit nut vary
greatly between species and are useful for defining some groups of species. In
section Anisopterys, the nut is usuallylaterallyalulate, with the crests or winglets
parallelto the areole in the B. pubipetala group and radiatingfrom the areole in
the B. lucida group.
In the Banisteriopsis malifolia group and the B. membranifolia group, the
wings of the two posterior samarasare not equidistantfrom each other and from
the anteriorwing, but are orientedmore nearly parallelto the anteriorwing (Fig.
15). The fruits of the B. malifolia group are furtherunusualin that the locule of
the nut is densely hairy inside; this characteris also found in B. caapi, B. multifoliolata and the B. megaphylla group.
The fruits of Diplopterys have a dorsal crest instead of a dorsal wing, welldeveloped lateralwinglets on the nut which are more or less interconnectedwith
ridges (Fig. 45), and no carpophore.The nut is usually very large and apparently
the embryo does not fill the locule. Perhapsthe air-filledlocule and lateralprojections serve as buoyancy devices for water dispersalof the fruits.
CHROMOSOMENUMBERS
There are only four chromosome counts for Banisteriopsis in the literature.I
have made counts for seven additional species, with tentative counts for two
more, using pollen mother cells from materialfixed in the field in Carnoy'sfluid
or Newcomer's solution. To obtain the counts, the pollen mother cells were
squashed in 45%aceto-carmineor 45%propioniccarmine. Some of the material
fixed in the field showed poor preservation with dark granularcytoplasm and
poorly defined chromosomes so that counts were not obtained for an additional
six species which were attempted. Those counts which were successfully made
were achieved relatively easily with good stainingand differentiation.
I have not seen vouchers of the materialstudiedby Pal (1964)and Fouit (1966),
so the taxonomic identity of these species is uncertain.The counts availableare
listed in TableIV. It appearsthat the base chromosomenumberfor Banisteriopsis
is x = 10.
There are no recorded chromosomecounts for Diplopterys.

TAXONOMY
Separation of the Genera Banisteriopsis and Diplopterys
1. Bracts and bracteoles appressed or slightly diverging (except B. patula and B. krukoffii);
fruit with large, well-developed dorsal wing (except B. paraguariensis, B. cipoensis and B.
I. Banisteriopsis.
sepium, and these with no or only one lateral winglet on the nut).
1. Bracts and bracteoles lingulate and spreading; fruit with shallow dorsal crest and well deII. Diplopterys.
veloped lateral winglets or crests on each side of the nut.
19
Table IV
ChromosomeNumbers in Banisteriopsis
1. Numbersfrom the literature
Species
B. muricata (as Banisteria
argentea)
"B. laevifolia"
B. caapi
Source of the count
Count
Fouit, 1966, mitotic root

cell by sectioning
Pal, 1964
Baldwin, 1946, mitotic
2n = 20
n = 20
2n = 20
leaf cell count
2. Numbersobtainedin the currentstudy

(All counts were madeof pollen mothercells; A meansa collectionby W. R. Anderson,andG means
a collection by B. Gates.)
Count
Voucher
Species
B. acapulcensis var.
llanensis
B. acerosa
B. argyrophylla
B.
B.
B.
B.
B.
B.
hypericifolia
laevifolia
muricata
oxyclada
pulchra
vernoniifolia
G
A
A
G
A
A
A
A
A
A
307
11177
11142
399
11548
11143
11148
11144
11789
11490
n
n
n
n
n
n
n
n
n
n
=
=
=
=
=
=
=
=
=
=
10
10 (many figures)
10 (many figures)
10 (many figures)
10 (many figures)
10 (many figures)
ca. 20
10 (many figures)
10 (several figures)
10 (many figures)
Because it is difficultto distinguishDiplopterys from some species of Banisteriopsis in flower, I have also included Diplopterysin the key to floweringspecimens of Banisteriopsis.
I. BANISTERIOPSIS
Robinson in Small, North AmericanFlora 25: 131. 1910.
Banisteria sensu Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 36. 1832[1833];Ann. Sci. Nat.
Bot., Ser. 2, 13: 280. 1840;Arch. Mus. Hist. Nat. 3: 388. 1843.
Banisteria sensu Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem. 1900, et p. hiem. 1901; Verz.
Vorles. Ak. Braunsberg:13. 1912;et in A. Engler, Das PflanzenreichIV, 141: 386. 1928.
Jubistylis Rusby, Mem. New York Bot. Gard.7: 273. 1927.Type species. Banisteriopsislutea,
as J. mollis Rusby.
Lianas, shrubs, vining or not, treelets, or shrublets with woody xylopodium.

Stipules interpetiolar,small to minute, Leaves simple, opposite or ternate, rarely
alternate or 4 at a node, the petiole short and eglandularor longer and often
bearing 1-2 pairs of glands distally, the laminaentire, bearing 1-several pairs of
glands along the marginor abaxially on the lamina or on the costa at the base,
adaxiallyglabrous, velutinous or appressed-pubescentto glabrate,abaxiallyglabrous, velutinous, tomentose or sericeous, with the venation camptodromous,
the secondary veins often scalariform,prominentto impressed adaxially, prominent abaxially. Inflorescencepaniculateor cymose, the flowers borne in pairs or
alternatelywith up to 45 flowers on each branch or reduced to 2 pairs with the
internodes between the 2 pairs reduced to give the appearanceof a 4-flowered
umbel. Bracts and bracteoles eglandular,persistent or deciduous, pubescent or
glabrateabaxially, glabrousadaxially. Pedicels usually sessile, more rarely with
20
Flora Neotropica
a short peduncle 0.5-4.0(-7.0) mm long. Flowers hermaphroditic.Calyx of 5

sepals, in a few species eglandular,but usually the 4 lateral sepals each bearing
2 abaxial glands, the anteriorsepal usually eglandular,persistent in fruit. Petals
5, white, pink, lemon- or yolk-yellow, clawed, the limb glabrous or abaxially
sericeous, cochleate to convex, with the margin fimbriate or laciniate, rarely
crenulateor subentire, sometimes glandular,the posteriorpetal usually different
from the 4 lateral petals with a longer claw and smaller limb, deciduous after
flowering. Stamens 10, all fertile, the filaments subequal or variously differentiated, the locules glabrousor pilose, the marginof the pollen sacs along the line
of dehiscence membranaceous,the connectives buff or yellow, eglandularor
glandular, the glandulartissue composed of large polygonal cells or papillae.
Gynoecium of 3 carpels, each uniovulate, free, adnate to a pyramidaltorus, the
ovule adaxial, pendent, anatropous. Styles 3, equal or sometimes the anterior
style different from the 2 posterior styles, rarely only the anterior style developing, glabrousor hairy, persistentin fruit. Stigmasterminal,often capitate.Fruit
of 3 samarasor mericarpseach usually with a functionalcarpophore.Nut of the
samarawith a hardwoody or bony pericarp,smooth to rugose or bearingvarious
lateral appendages, sometimes with a tooth at the base of the distal margin,with
a well-developeddorsal wing with the upper marginthickened, rarelyreduced to
a short crest, sometimes with an outgrowthat the base of the upper marginof
the wing, in some species the wings of the 2 posterior samaraslying subparallel
to the wing of the anteriorsamara. Locule of the samaramore or less spherical
or kidney-shaped, sometimes pubescent within, the seed filling the locule, the
embryo with large cotyledons, equal or subequal,fleshy and pear-shapedor narrowly oblong and folded distally. Germinationepigeal or hypogeal.
Type species. Banisteriopsis cornifolia (Humboldt, Bonpland & Kunth) Robinson in Small (Heteropteryscornifolia Humboldt, Bonpland& Kunth).
The name Banisteriopsis is derived from the name Banisteria L. which was
named for John Baptiste Banister, an English missionaryand author of the first
catalog of plants of Virginia,with the suffix "-opsis," "with the appearanceof."
Distribution. Found throughoutthe New World tropics, with a few species
extending north into Mexico and south to Argentina. The greatest number of
species are found in Brazil, and more than one-thirdin the Planalto region of
central Brazil.
Explanationof terms in the keys (see also Morphology)
Leaf length measurementsare taken from the apex of the petiole to the apex
of the leaf; leaf width measurementsare taken at the widest point. All leaf measurementsrefer to matureleaves, not leaves in the inflorescence.
Primaryveins are the main lateralveins which arise from the costa; secondary
veins are the veins connecting the primaryveins; other lesser-order veins are
described as the reticulation.
The secondary veins are described as scalariformwhen they are more or less
paralleland relatively more prominentthan the reticulation(Figs. 32, 36).
Leaf glands are not always present on every leaf of an individual;sometimes
it is necessary to examine several leaves to ascertaintheir numberand distribution. They are usually well-developedon the inflorescenceleaves and sometimes
their position and number on these leaves can be used as an indicator of their
distributionon the vegetative leaves.
Hairs are malpighiaceousand principallyof two types, T-shapedand Y-shaped.
If the hairs are T-shaped and more or less sessile with a straighttrabecula,and
21
the trabeculae more or less parallel, the pubescence is described as sericeous

(Fig. 1); if the hairs are T-shapedand more or less stalked, often with a flexuous
trabecula, and the trabeculae not parallel to each other, the pubescence is described as tomentose (Fig. 1). The pubescence is also described as tomentose
when the hairs are Y-shaped with the arms very widespread,i.e. divergingmore
than 90? (Fig. 1). If the hairs are Y-shaped with the arms formingan angle less
than 90? between them, the pubescence is described as velutinous. Sometimes
the vesture includes more than one hair type, when compounddescriptionssuch
as tomento-velutinous,tomento-sericeousare used. Some species bear Y-shaped
hairs in which one arm is very reduced or absent and the hairs then appear
basifixed(Fig. 1).
If the density of the pubescence is such that the hairs do not form a continuous
covering, but instead the surface is exposed between the hairs, the pubescence
is described as sparse, and very sparse if the hairs are scattered. Leaves which
are sparselyappressed-sericeousoften requireexaminationunderthe microscope
for the hairs to be apparent.If the hairs are contiguous and form a continuous
covering, the pubescence is described as dense, and if the hairs form a covering
more than one hair thick, the pubescence is described as very dense.
The peduncle without qualificationrefers to the floriferous peduncle which
subtends the pedicel; it is usually absent in Banisteriopsis, but in some species
it is 0.5-4.0(-7.0) mm long. The pedicel is the stalk which subtends a single
flower. The common peduncle is the portion of an inflorescence branch which
directly subtends the peduncles (if present) and pedicels.
Bracts and bracteoles without qualificationrefer to the floriferousbracts and
bracteoles which subtend the individualflowers (Fig. 2); they are considered to
be deciduous if there is abscission of any bracts and bracteoles independentof
abscission of the pedicel. Most species have erect or appressedbracts and bracteoles; the bracts and bracteoles are described as spreadingonly if they diverge
greatly from the pedicel, and are more or less reflexed (Fig. 2).
Measurementsof the diameter of flower buds are made on mature buds just
prior to anthesis; they are sometimes useful as a measure of overall flower size.
All flower measurementsand measurementsof pedicel diameterare made on
soaked material;pedicel diameteris measuredin the middle of the pedicel.
The calyx glands are describedas being borne below the free part of the sepal
on the receptacleif the attachmentof the glandis below the free part of the sepal;
often the apices of the glands are enlargedand free, and extend over part of the
base of the sepal. They are describedas being borne on the free part of the sepal
when the attachmentlies on the sepal (Fig. 3). It is necessary to look at open
flowers to see this characteras some species with glands on the sepal develop
the glands late in flower development,just before anthesis, and it is difficultto
determinewhetherthe glandprimordiumis on or below the free part of the sepal.
All flowers of Banisteriopsis except the B. leiocarpa group exhibit marked
bilateral symmetry, with the plane of symmetry passing through the posterior
petal and the anterior sepal. The anterior sepal is the unpaired sepal which is
usually eglandularin species with calyx glands, and the paired sepals are distinguished as the antero-lateralsepals (nearer the anterior sepal) and the posterolateral sepals. The posterior petal is the unpairedpetal, which is usually borne
erect, and the paired petals are usually reflexed between the sepals and distinguished as the antero-lateraland postero-lateralpetals (see Fig. 3).
Occasionalflowers in species with hairy antherlocules will have antherlocules
which are glabrate;it is often necessary to examine several flowers to establish
whether or not the antherlocules are hairy. Sometimes the hairs are very sparse,
22
FloraNeotropica
' -'
~"r
*'
'
.;'
~ ~
~
~"" ~
'"
~ '' ~ ~\' ~ ~' ~'"" ' 'd
' ' ' '
..
' L
:- '';: - '
..
*
/
'.
? -- - \
i.:i..
: /-?~
....
?' . -.-;
::A i" '-"
.
', ; '..........................."" :
/.
., "!'"-,''.....
.; ? .'":......
19' '
-.
':'".'Z'.
'
'..
' * -
:",:> ..
'"
*r .?
..
. i
.?,.
:-
.-
-";.'}
? . ..}j
,
?.
'..:
. <.
,,!1..
,'
'
..'-
-.' 'i.'
? . ? ?-., .~? ',.
..
....
... .,.
d?
. .
sffi
; T ,ffi. ?
~~~~~~~~~~~~~~~~~
:.;;-
. i
.. .,
~
'
. .
... ?
....
...
....
(\
Banisterio;-is.
?,
.:to illustrate usag' of terms
'?
nut
at
wies
ventral..
par;
A, sm
.:~,~i
,
mat'c;
,1a,ld_agra.
I0 0
:I
\#
C,
. ??;i ?
Dig,am'of :. sam_,_
winglet;
of win
s, style.
winle.
winglet|~~~~~~;
sr,~~~~~
/ s ,l.
viw
?;
.-
r?
;: ; -.
?-
mwit
?.-
??
'
id
.:,
*
?~~~~~~~~~
B,nt
'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'
.
,
>?
.
.
.
... ..- ..?
::
FIG
i?~~~~~~~~~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
:.
..
:.
~~~~~~~~~~~~~~~~~~~~~~~
C
?~'
-.
.~
- ,
vie.
n-
(a,
laea
nut
b,
wingetS;
aroe
.Z
c.
legt
cappoe
of laea
W-;
0 ^ti>iX
;,;
e,
apedae
s,syle.
wige;
.~
.?.......
q- hegtoaea
_f*
wing..................;
23
or are only present as a tuft of hairs at the base or apex of the locules. The hairs
appearto be attachedto the locules, and are usually not readily visible in soaked
specimens. Occasionally, loose hairs from other parts of the plant will adhere to
the anthers and give the illusion of locule hairs; this artifact can usually be detected by careful observation since such hairs will also be found adheringto the
stigmas or other parts of the stamen. If the anther locules are described as glabrous they are always quite glabrous;if any hairs are found on the antherlocules
they must be considered hairy.
Straightparallel styles sometimes become more or less divergingin dried material; upon soaking they usually become straightand parallelagain.
The areole is the point of attachmentof the samarato the receptacle;the lateral
winglets on the nut of the samara are described as radiatingfrom the areole if
their bases are derived from the marginof the areole, and they are described as
parallelto the areole if the bases of the winglets are parallelto the marginof the
areole (Fig. 4). The height of the fruit nut is the dimensionparallelto the surface
of the areole, and the length of the nut is that perpendicularto the surface of the
areole (Fig. 4). The terms "high" and "long" used with reference to the lateral
winglets mean respectively the distance the wing projects from the nut and the
length of the attachmentof the lateral winglets to the nut (Fig. 4).
Keys to the Species of Banisteriopsis
Key to Flowering Specimens
1. Corollayellow.
2. Petals hairyexternally.
3. Calyx eglandular.
67. B. valvata.
4. Sepals valvate, 6.5-8.0 mm long.
4. Sepals imbricate,up to 3.0 mm long.
92. B. sepium.
5. Petiole 4-9 mm long, laminavery coriaceous,glabrous.
5. Petiole 5-20(-36) mm long, laminachartaceous,rarely subcoriaceous,tomentose or appressed-sericeousabaxially,sometimes sparselyso.
6. Bracts and bracteoles erect or appressedto pedicel; anteriorstyle longer
68. B. heterostyla.
than posteriorstyles, hairyin the middle.
91. B. patula.
6. Bracts and bracteoles spreading;styles subequal,glabrous.
3. Calyx glandular.
7. Bracts and bracteolesspreading,lingulate.
8. Petals densely sericeousexternally,denticulateto short-fimbriate. 90. B. krukoffii.
8. Petals sparsely sericeous externally,long-fimbriate.
Diplopterys.
7. Bracts and bracteoleserect or appressedto pedicel, triangular.
9. Laminanearlyor quite glabrousabaxially.
10. Leaves sessile or the petiole up to 3 mm long, laminatruncateto amplexicaul at the base.
11. Laminaelliptic;anteriorstyle longerthan posteriorstyles and densely
84. B. pubipetala.
hairy in the basal half.
11. Laminaovate to orbicular;styles equal, glabrousor basally strigose.
12. Laminabroadlyovate to orbicular,5.5-10.6 cm long and 5.2-9.2
82. B. amplectens.
cm wide, claspingthe stem.
12. Laminaovate to orbicular,2.5-7.5 cm long and 1.2-6.4 cm wide,
cordateat the base.
13. Liana; inflorescence branches and pedicels sparsely ap81. B. virgultosa.
pressed-sericeous.
13. Shrub or subshrub;inflorescencebranches and pedicels glabrous.
80. B. hypericifolia.
10. Leaves with the petiole 3-17 mm long, laminacuneateto subcordateat the
base.
14. Petals sparselysericeousexternally;laminawith 2-4 pairs of glandular
areas /2 to 2%of the distancefrom the midribto the margin.
89. B. caduciflora.
Flora Neotropica
24
14. Petals densely sericeous externally; lamina with marginalglands or

numerousminuteglands on the laminanear the margin.
15. Petiole (6-)10-27 mm long; lamina (8.0-)12.0-25.5 cm long,
(3.5-)4.5-13.0 cm wide.
16. Numerous glandularareas up to 2 mm wide on the lamina
margin;flowers borne in 4-floweredumbels or with a pair of
flowers subtendinga 4-floweredumbel.
88. B. schunkei.
16. Minuteglandsup to 0.8 mm diameteron marginor on lamina
near margin;flowers borne in condensed axillaryracemes of
3-8 pairsof flowers,the terminal2-3 pairssometimesforming
an umbel.
86. B. longialata.
15. Petiole 2-8(-12) mm long; lamina 5.1-13.0(-16.0) cm long, 2.17.0(-8.3) cm wide.

17. Anteriorstyle longer and thicker than posteriorstyles, hairy
for up to half its length.
18. Bracts and bracteoles up to 1 mm long; pedicels appressed-brown-sericeous;calyx glands 1.0-1.8 mm long,
74. B. lucida.
0.8-1.2 mm wide.
18. Bractsand bracteoles0.8-1.6(-2.0) mm long; pedicels appressed-white-sericeousto glabrate;calyx glands 1.5-3.0
mm long, 0.8-2.0 mm wide.
84. B. pubipetala.
17. Styles equal or subequal,glabrousor hairy.
19. Styles hairyfor up to half their length.
85. B. platyptera.
19. Styles glabrousor basally strigose.
75. B. nutans.
9. Laminahairyabaxially
20. Abaxialleaf pubescencetomentoseor velutinous.
21. Leaves subsessile, petiole up to 2 mm.
83. B. rondoniensis.
21. Leaves petiolate, petiole 6-10 mm.
22. Branches tomento-sericeous;bracts and bracteoles 1.5-2.2 mm
79. B. cachimbensis.
long.
22. Branchesappressed-sericeous,old branchesglabrate;bracts and

77. B. woytkowskii.
bracteoles0.8-1.2 mm long.
20. Abaxialleaf pubescence sparselyappressed-sericeous.
23. Leaves with 8-18 pairs of lateralveins; petiole 10-27 mm long, lamina
87. B. erianthera.
8.2-23.2 cm long and 4.3-12.2 cm wide.
23. Leaves with 4-8 pairsof lateralveins; petioleup to 10 mmlong, lamina
5.1-13.0(-16.0) cm long and 2.1-7.0(-8.3) cm wide.
24. Bractsand bracteolesup to 1.0 mm long; calyx glands 1.0-1.8 mm
74. B. lucida.
long, 0.8-1.2 mm wide.
24. Bracts and bracteoles 1.0-2.0 mm long; calyx glands 1.5-3.8 mm
long, 1.0-2.0 mm wide.
25. Anteriorstyle longer than posteriorstyle, up to 3.2 mm long
and incurvedapically;posteriorstyles 2.4-2.6 mm long.
76. B. peruviana.
25. Styles more or less equal, 2.2-2.8 mm long, straightand parallel or divergingapically.
26. Pedicels sparselygolden-sericeous;petals sericeousin the
middle externally;4 lateral sepals 1.5-2.0 mm long and
wide.
78. B. nigrescens.
26. Pedicels densely brown-sericeous;petals densely sericeous externally, except at the margin;4 lateral sepals
75. B. nutans.
2.0-2.8 mm long and wide.
2. Petals glabrousexternally.
27. Flowers borne when plant is leafless.
B. lutea group.
28. Bractsand bracteoles 1.0-2.0 mm long, lanceolateto linear,sparselytomentose
to glabrateabaxially;sepals tomento-sericeousabaxially.
69. B. lutea.
28. Bractsand bracteoles1.0-1.5 mmlong, triangular,sparselyappressed-sericeous
70. B. cristata.
abaxially;sepals appressed-sericeousabaxially;Venezuela.
27. Flowers borne on leafy plant.
29. Leaves with sessile marginalglands, sometimes only well developed on inflorescence leaves and sometimesdevelopedinto ciliate extensions apically, often
with a well-developedpairof glandson the laminabeside the midribat the base.
B. martiniana group.
25
30. Flowers borne in 4-(6-)floweredumbeis.

31. Connectivesof stamens opposite antero-lateralsepals overtoppinglocules 0.6-1.0 mm.
32. Flowers large, the limb of the 4 lateralpetals 8.0-11.0 mm long and
wide; connectives of stamens opposite antero-lateralsepals globose, 1.4-1.8 mm wide.
54. B. pulcherrima.
32. Flowers small, the limb of the 4 lateralpetals 3.0-6.5 mm long and
wide; connectives of stamens opposite antero-lateralsepals pearshaped, 0.9-1.3 mm wide.
33. Laminaglabrousor very sparsely sericeous to glabrateabaxially.
44. B. martiniana.
33. Laminavelutinous,sometimesglabrateadaxially.
45. B. velutinissima.
31. Connectivesof stamensopposite antero-lateralsepals overtoppinglocules 0-0.5 mm.
34. Inflorescence tomentose or tomento-velutinous;lamina abaxially

sparselytomentose or loosely sericeous.
35. Pedicels sparsely tomento-sericeous, 10-14 mm long; sepals
sericeousabaxially,glandsup to 1.2 mmlong and0.8 mmwide.
50. B. pubescens.
35. Pedicels glabrous, 12-18 mm long; sepals glabrousabaxially,

49. B. wilburii.
glands 1.5-2.6 mm long and 0.8-1.4 mm wide.
34. Inflorescenceglabrousor appressed-sericeous;laminaabaxiallyglabrous or sparselyappressed-sericeous.
36. Lamina abaxially appressed-sericeous,rarely glabrate;inflorescence densely appressed-golden-sericeous;pedicels ap53. B. maguirei.
pressed-sericeous;GuayanaHighland.
36. Laminaabaxiallyglabrousor rarely very sparsely appressedsericeous;inflorescenceglabrousor sparselysilvery-appressed46. B. elegans.
sericeous;pedicels glabrous.
30. Flowers borne alternatelyor in pairs on the ultimatebranches,not in umbels.
52. B. alternifolia.
37. Flowers and leaves borne alternately.
37. Flowers and leaves borne in pairs or subopposite.
38. Leaves with 3-5 pairs of main lateralveins; stem with sparse basifixed hairs up to 5 mm long.
51. B. grandifolia.
38. Leaves with 5-8 pairs of main lateralveins; stems glabrousor appressed-sericeous.
39. Petiole 5-14 mm long, laminavery coriaceous, cordate at the
base, the margin revolute; inflorescence densely golden-ap53. B. maguirei.
pressed-sericeous;GuayanaHighland.
39. Petiole 3-8 mmlong, laminachartaceousto subcoriaceous,cuneate to truncateat the base, the marginplane; inflorescence
tomento-sericeousor silvery-appressed-sericeous.
40. Flowersborneas new leaves expand;laminawith 3-5 pairs
of prominentcupulateglands on the margin;pedicels to48. B. padifolia.
mento-sericeous.
40. Flowers borne with mature leaves; leaves eglandularor
with numerousminuteglands on or near the margin;ped47. B. polygama.
icels appressed-sericeous.
29. Leaves with 2-6 pairs of glands on the laminaabaxially,and/orwith 1-2 pairs
of glands at the apex of the petiole, and sometimeswith a stalkedgland on the
marginon each side of the midribat the base.
41. Calyx eglandular.
67. B. valvata.
42. Sepals valvate, 6.5-8.0 mm long.
42. Sepals imbricate,up to 3.0 mm long.
43. All 5 petals stronglyreflexedbetween the sepals; styles similarand
B. leiocarpa group.
apicallyinflexed.
44. Lamina 1.4-4.5 cm long and 0.9-2.5 cm wide.
72. B. parvifolia.
44. Lamina3.9-8.4 cm long and 2.0-7.4 cm wide.

45. Laminaabaxiallyloosely tomentose;petiole 9-24 mmlong;
4-7 pairs of lateralveins.
71. B. leiocarpa.
45. Laminaabaxiallyshining-sericeous;petiole 7-14 mm long;
73. B. populifolia.
7-9 pairs of lateralveins.
Flora Neotropica
26
43. Posteriorpetal erect or suberect, 4 lateralpetals reflexedbetween

the sepals; anteriorstyle differentfrom posterior styles, erect or
sigmoidand directedaway from the axis.
46. Anteriorstyle hairy for up to two-thirdsof its length, slender
and sigmoid.
47. Flowers borne in 4-flowered umbels in the axils of the
leaves; leaves elliptic to broadly elliptic; sepals glabrous
adaxially.
68. B. heterostyla.
47. Flowers usually borne in 4-(6)-floweredumbels in the axils

of previous seasons' leaf scars; leaves ovate to orbicular;
B. lutea group.
sepals tomento-sericeousadaxially.
48. Bracts and bracteoles 1.0-2.0 mm long, lanceolate to
linear,sparselytomentoseto glabrateabaxially;sepals
69. B. lutea.
tomentose abaxially.
48. Bracts and bracteoles 1.0-1.5 mm long, triangular,
sparsely appressed-sericeous abaxially; sepals ap70. B. cristata.
pressed-sericeousabaxially;Venezuela.
46. Anteriorstyle glabrousor basally strigose, stout and erect.
49. Laminanearlyto quite glabrateat maturity. 60. B. parviflora.
49. Laminaabaxiallyhairythroughout.
50. Brancheshispid, with interspersedbasifixedhairs;petiole and lamina on both sides hispid to hispido-sericeous.
63. B. basifixa.
50. Branchestomento-sericeous,hairsall malpighiaceous;

petiole and lamina tomento-sericeousabaxially, glabrous adaxially.
61. B. salicifolia.
41. Calyx glandular.

51. Flowers borne in 4-(6-)floweredumbels, sometimesthe umbel subtended by a pair of flowers.
52. Bracts and bracteolesdeciduousin bud.
32. B. membranifolia.
53. Laminaabaxiallyappressed-sericeous.
53. Laminaabaxiallytomento-velutinous.
34. B. adenopoda.
52. Bracts and bracteolespersistent.
54. Leaves nearlyor quite glabrousabaxially.
55. Laminawith 2-4 pairs of glandularareas 12-2 of the dis89. B. caduciflora.
tance between the midriband margin.
55. Laminawith a pair of sessile or stipitate glands near the
midribat the base.
56. Leaves sessile; pedicel 19-22 mm long.
66. B. magdalenensis.
56. Leaves with the petiole 8-18 mm long; pedicel 5-10

mm long.
60. B. parviflora.
54. Laminahairyabaxially.
57. Lamina densely white- or golden-sericeous or whitetomento-sericeousabaxially.
58. Leaves sessile or subsessile, petiole 0-3(-8) mm long;
styles equal or subequal,erect or diverging.
59. Pedicels glabrous, 19-22 mm long.
59. Pedicels white-sericeous,9-13 mm long.
9. B. laevifolia.
58. Leaves petiolate, petiole 6-16(-20) mm long; posterior styles longeror shorterthan anteriorstyle.
60. Posterior styles longer than anterior style, ly59. B. lyrata.
rate; petiole apicallybiglandular.
60. Posteriorstyle shorterthan anteriorstyle, straight
and parallelthroughouttheir length, coherentapically.
61. B. salicifolia.
57. Laminasparselysericeous, tomento-velutinousor hispidotomentoseabaxially.

61. Petiole 10-20(-35) mm long; laminaelliptic or ovate,
tomento-velutinousor hispido-tomentoseabaxially.
62. Pedicels glabrous;stem with basifixedhairs.
63. B. basifixa.
27
62. Pedicels sparselyvelutinous;stem velutinous.
62. B. scutellata.
61. Petiole 3-12 mm long; lamina lanceolate, abaxially

sparselyappressed-sericeous.
63. Umbelsbornesinglyin leaf axils;calyx glands 1.22.6 mm long.
65. B. pseudojanusia.
63. Umbels borne in racemes on axillary branches;

64. B. parviglandula.
calyx glands 0.4-0.6 mm long.
51. Flowers borne alternatelyor in pairs on the ultimatebranches.
64. Calyx glandsborne on free part of sepals, not on receptacle.
B. nummifera group.
65. All or some leaves quite glabrousabaxially.

66. Pedicels 5-7 mm long; calyx glands0.6-1.2 mm long, 0.50.7 mm wide; styles slender, apicallyinflexed.
56. B. sellowiana.
66. Pedicels7-13 mmlong; calyx glands 1.0-2.0 mmlong, 0.61.4 mm wide; styles stout, diverging.
55. B. nummifera.
65. All leaves abaxiallyhairy.
67. Posterior styles slender, less than 0.2 mm in diameter,
sometimesabsent, longeror shorterthanthe anteriorstyle.
68. Posteriorstyles longer than anteriorstyle, lyrate.
59. B. lyrata.
68. Posteriorstyles shorterthan anteriorstyle or absent.

58. B. gardneriana.
67. Styles equal or subequal, stout, 0.2-0.4 mm in diameter,

divergingfrom the base.
57. B. anisandra.
69. Laminaabaxiallytomentose.
69. Laminaabaxiallyappressed-shining-sericeous.
55. B. nummifera.
64. Calyx glands borne below free part of sepal, on receptacle.

70. Laminawith 2-4 pairsof glandularareason the abaxialsurface
2-2/3 of the distancebetween the midriband margin.
89. B. caduciflora.
70. Laminawith numerousminuteglands on the margin.
Subgenus Banisteriopsis.
71. Styles 2.0-2.6 mm long, divergingat the base and inflexed

towards the axis apically;filamentsof the 3 anteriorstamens almostfree to the base, those of the stamensopposite
the antero-lateralsepals flexuousand directedtowardsthe
posteriorpetal.
72. Laminaabaxiallysparselyappressed-sericeous.
6. B. wurdackii.
72. Lamina abaxially tomentose or tomento-sericeous.

7. B. cinerascens.
71. Styles 1.0-2.0 mm long, straightand parallelor diverging

throughouttheirlength;stamenssubequal,erect, all somewhat connate at the base.
73. Laminaabaxiallyglabrousor tomentose.
74. Inflorescencesaxillary, up to 6 cm long, the ultimate branchesbearing 16-40 flowers; laminaglabrous; Brazil.
3. B. ferruginea.
74. Inflorescencesaxillaryup to 20 cm long, the ultimate branchesbearing 10-25 flowers; laminaglabrous or sparselyto densely tomentose.
2. B. acapulcensis.
73. Laminaabaxiallysparselyappressed-sericeous.
75. Ultimate branches of inflorescencebearingup to
45 flowers.
76. Pedicel 2-5 mm long; petiole 7-15 mm long.
5. B. brevipedicellata.
76. Pedicel 7-10 mm long; petiole 6-10 mm long.
3. B. ferruginea.
75. Ultimate branches of inflorescencebearingup to

20 flowers.
Flora Neotropica
28
77. Petals entire; styles 1.6-1.8 mm long.

78. Petiole (7-)10-20 mm long; ultimate
branches of inflorescence with up to 20
flowers; Brazil.
4. B. megaptera.
78. Petiole 3-10(-15) mm long; ultimate

branchesof inflorescencewith usually up
to 12 (occasionally 20) flowers; Central
America and NW South America.
1. B. cornifolia.
77. Petals denticulate;styles 1.0-1.2 mm long.

8. B. longipilifera.
1. Petals pink or white, sometimesbecomingcreamor pale butter-yellowin age.

79. Antherlocules quite glabrous.
80. Posteriorstyles reduced;anteriorstyle stout, up to 0.6-0.9 mm in diameter.
36. B. schwannioides.
80. Posteriorstyles as long as or longerthanthe anteriorstyle; anteriorstyle up to 0.4

mm in diameter.
81. Connectivesof stamensopposite the antero-lateralsepals greatlyenlargedand
projecting0.8-1.7 mm beyond the locules.
B. muricatagroup.
82. Pedicel borne on a peduncle 1-4 mm long.
83. Non-viningshrub;laminalanceolate, abaxiallydensely shining-whitesericeous; styles straightand slightlydiverging;Goias, Brazil.
43. B. goiana.
83. Vines; laminaovate or ellipticto orbicular,abaxiallysparselyto densely sericeous or tomentose;styles diverging,the posteriorstyles lyrate
at the base.
84. Lamina 1.3-6.7 cm long, elliptic to ovate, the marginrevolute,
abaxiallysparselysericeous, with the petiole 5-8 mm long; Cuba.
42. B. pauciflora.
84. Lamina (3.5-)5.3-18.0 cm long, ovate to orbicular,the margin

plane, abaxiallysparsely to densely sericeous or tomentose, the
petiole 5-25 mm long.
85. Petiole 14-25 mmlong; inflorescencebranchesdensely whitetomentose; laminaadaxiallyvelutinousto glabrate,abaxially
densely white-tomentoseor sericeous;pedicel 1.2-1.4 mm in
diameter,0.8 mmwithoutthe hairs;flowerslarge,the 4 lateral
41. B. oxyclada.
petals with the limb 9-13 mm long.
85. Petiole 5-16 mm long; inflorescencebranches golden or silvery-velutinousor appressed-pubescent;laminaadaxiallyappressed-pubescentto glabrate,abaxially sparsely to densely
silver or golden appressed-sericeous;pedicel 0.6-0.8 mm in
diameter,0.3-0.5 mm withoutthe hairs;flowers small, the 4
40. B. muricata.
lateralpetals with the limb 4.0-8.5 mm long.
82. Pedicels sessile.
31. B. calcicola.
86. Petioles 2-8 mm long; laminavelutinousabaxially.
86. Petioles 5-16 mm long; laminaappressed-pubescentabaxially.
40. B. muricata.
81. Connectivesof stamensoppositethe antero-lateralsepals not greatlyenlarged,

projecting0.1-0.4(-0.7) mm beyond the locules.
B. megaphyllagroup.
87. Claw of the posteriorpetal widest at its apex.
88. Leaves elliptic or broadly elliptic; inflorescencelax; flower bud diameter4.0-6.0 mm;petals pale pink, the 4 lateralpetals with the limb
8-10 mm long.
25. B. irwinii.
88. Leaves broadly elliptic to orbicular;inflorescencecongested; flower

bud diameter3.0-4.5 mm; petals dark pink or white with dark pink
margins,the 4 lateralpetals with the limb 5-7 mm long.
24. B. megaphylla.
87. Claw of the posteriorpetal constrictedat the apex.

B. campestrisgroup.
89. Petiole 1-8(-10) mm long; non-vining.
90. Laminaabaxiallywith a pair of peltate glands on the basal lateral
veins one-thirdto two-thirdsof the distancefromthe midribto the
margin.
30. B. arborea.
91. Slendertree to 4 m, much branched.
29
91. Shrubor shrubletto 1.5(-2.5) m tall, unbranchedbelow the

inflorescence.
26. B. campestris.
90. Laminaabaxiallywith 1(-2) pairsof peltateglandsbeside the midrib at or near the base.
92. Laminaglabrousabaxially,except the midriband mainlateral
veins.
93. Slender treelet or shrub; lamina held erect, more than
three times as long as wide, with only the primaryand
29. B. andersonii.
secondaryveins impressed.
93. Shrub;laminadivergingfrom the stem, less than 3 times
as long as wide, rugose.
26. B. campestris.
94. Lamina abaxially very densely tomento-sericeous, not
visible throughthe pubescence, more than 2.5 times as
long as wide, with only the primaryand secondaryveins
28. B. angustifolia.
prominentabaxially.
94. Laminaabaxiallyvery densely to sparselytomentose,visible throughthe pubescence, less than 2.5 times as long
as wide, with the reticulationof veins very prominent
abaxially.
95. Shrublet to 4 dm tall, lamina hispido-tomentose,
mostly less than 5.5 cm long; Serra do Cip6, Minas
Gerais, Brazil.
27. B. cipoensis.
95. Shrub or shrublet 1.5(-2.5) m tall; leaves softly tomentose, mostly more than 5.5 cm long.
26. B. campestris.
89. Petiole (4-)8-19 mm long; vines.

96. Leaves abaxiallyvelutinous.
37. B. prancei.
96. Leaves abaxiallysericeous.
39. B. quadriglandula.
79. Antherlocules hairy, sometimeswith only basal or apical tufts of hairs.
97. Bracts and bracteolespersistent.
98. Claw of the posteriorpetal constrictedat the apex.
99. Pedicel more than 11 mm long; sepals 4.5-4.4 mm long.
33. B. pulchra.
99. Pedicel 7-11 mm long; sepals up to 3.5 mm long.
35. B. caapi.
98. Claw of the posteriorpetal widest at the apex.
100. Stamens subequal, the connectives all more or less the same size, not
glandularand enlarged,not projectingbeyond the locules (Fig. 9).
B. argyrophylla group.
101. Leaves narrowlyelliptic to linear (more than 3 times as long as

wide).
102. Shrubletor shrub;leaves chartaceousto subcoriaceouswith
only the primaryveins impressed;Goias, Brazil.
14. B. vernoniifolia.
102. Shrubor vine; leaves chartaceousor very coriaceous, rugose

or bullate.
103. Leaves very coriaceous, bullate;Bahia.
13. B. harleyi.
103. Leaves chartaceous,with the reticulationimpressedand
the areoles between the veins shallowlyinflated.
10. B. argyrophylla.
101. Leaves elliptic to ovate (less than 3 times as long as wide).

104. Shrubletwith erect stems to 6 cm; leaves with the primary
veins stronglyimpressed;Paraguayand adjacentMatoGrosso,
Brazil.
12. B. paraguariensis.
104. Shrubor vine to 3 m tall;leaves more or less rugose or bullate,

with the reticulationimpressed.
13. B. harleyi.
105. Leaves stronglybullate, coriaceous.
105. Leaves more or less rugose, chartaceous.
106. Liana or vining shrub; lamina adaxially soon glabrate and with the areoles between the veins shallowly inflated;stem pubescence velutinous to sub10. B. argyrophylla.
tomentose;petiole 3-10 mm long.
106. Shrub, not vining; lamina adaxially cobwebby-pubescent, eventually glabrate, rugose, with the ar-
30
Flora Neotropica
eoles between the veins not inflated;stem pubescence cobwebby;petiole 2-5 mm long. 11. B. byssacea.
100. Stamens not equal, with the filaments opposite the sepals longer than
those opposite the petals; connectives, at least those opposite the 3 anterior sepals, glandularand enlarged,projecting0.1-0.7 mm beyond the
locules.
107. Petioles more than 2 mm in diameterand more than 5 mm long.
B. megaphylla group.
108. Leaves ellipticor broadlyelliptic;inflorescencelax, flowerbud

diameter4.0-6.0 mm;petals pink, the 4 lateralpetals with the
25. B. irwinii.
limb 8-10 mm long.
108. Leaves broadly elliptic to orbicular;inflorescencecongested;
flowerbud diameter3.0-4.5 mm;petals darkpinkor white with
darkpink margin,the 4 lateralpetals with the limb up to 7 mm
24. B. megaphylla.
long.
107. Petioles less than 2 mm in diameter, or, if more, less than 5 mm

long.
109. Laminaabaxiallywith a single pair of glands 1 mm or more in
diameterbeside the midribat the base.
110. Tree with thick corky bark; flowers borne on old wood
(cauliflorous),sometimes also with short terminalflowering shoots bearing reduced leaves; lamina glabrous.
20. B. latifolia.
110. Shrubor shrublet,the bark not thick and corky; flowers

in terminal or axillary inflorescences, not cauliflorous;
laminadensely to sparselyappressed-sericeous.
18. B. malifolia.
109. Laminaabaxiallywith 1-5 pairs of glands less than 1 mm in

diameterabove the base.
111. Pedicels white-sericeo-tomentose;sepals white-sericeous
throughoutabaxially,oblong, roundedand often revolute
at the apex.
38. B. multifoliolata.
not revolute.
B. schizoptera group.
111. Pedicels white- or golden-velutinousor sparsely pubescent; sepals golden-sericeous abaxially, often glabrate
apically, broadly ovate, with the apex obtuse to acute,
112. Slenderwillow-liketree or shrubwith pendentnonviningbranches;laminaless than 7 mm wide, acicular, glabrate,inflorescenceof a pair of flowers ter17. B. acerosa.
minatingbranches.
112. Shrubletor shrubwith viningbranches;lamina1.06.5 cm wide, ovate or elliptic to lanceolate, pubescent or glabrous;inflorescenceof 4-floweredumbels
borne singly or in condensed axillary or terminal
cymes.
113. Laminaglabrousabaxiallyor with sparse appressed, sessile hairs, the trabecula 0.2-0.6
mm long.
16. B. stellaris.
113. Laminapubescent abaxially, the hairs sessile

or stipitate, the trabecula 1.0-2.4 mm long.
15. B. schizoptera.
97. Bracts and bracteolesdeciduousin bud, or duringor immediatelyafterflowering.

114. Stamenswith the filamentsopposite the postero-lateralpetals almost as long
as and parallelto the divergingposteriorstyles and longerthan the filaments
of the stamensopposite the antero-lateralsepals.
B. membranifoliagroup.
115. Laminavelutinouson both sides, bearingstalked glands abaxially,the
34. B. adenopoda.
stalk 0.5-1.5 mm long.
115. Laminaappressed-pubescentor glabrateon both sides, bearingsessile
or subsessile glandsabaxially.
116. Pedicels 7-10 mm long.
35. B. caapi.
116. Pedicels more than 11 mm long.
117. Sepals 4.5-5.5 mmlong; hairson the abaxialleaf surfacewith
the trabecula0.2-0.6 mm long.
33. B. pulchra.
31
117. Sepals up to 3.5 mm long; hairs on the abaxial leaf surface

with trabecula0.1-0.25 mm long.
114. Stamenswith the filamentsoppositethe antero-lateralsepalslongerthanthose
opposite the postero-lateralpetals, those opposite the postero-lateralpetals
shorter and not parallelto the posterior styles, the posterior styles straight
and parallelor slightlydivergingor apicallyrecurved.
118. Lamina with a pair of glands abaxiallybeside the midribat the base;
sepals abaxiallywhite-sericeousthroughout,oblong, roundedand often
revolute at the apex, loosely investingthe corollain bud. B. malifoliagroup.
119. Laminaquite glabrouson both sides.
120. Tree with thick corky bark;young branchessmooth, reddishbrown, not lenticellate;flowers borne on old wood (cauliflorous), sometimes also with short terminalflowering shoots
20. B. latifolia.
bearingreducedleaves.
120. Shrubor shrublet;the old branchesnot with thick corky bark,
punctate with numerouslenticels; flowers borne in terminal
and axillaryinflorescences,not cauliflorous.
121. Shrubletwith reclinate branchesto 1.5 m long; leaves
falcate; sepals abaxiallysparselysericeous, apicallyglabrate;Chapadados Veadeiros,Goias, Brazil.
23. B. hatschbachii.
121. Shrubor shrubletwith erect branchesto 2.5 m; leaves

plane; sepals abaxiallydensely white-sericeousthroughout.
21. B. variabilis.
119. Laminahairy, at least abaxially.

122. Laminasessile or with the petiole up to 3 mm long and more
than 2 mm thick.
123. Laminahispid-pubescentadaxially,the hairs Y-shaped,
andhispido-sericeousabaxially;pedicels golden-velutinous, more than 1.2 mm in diameter,0.6-1.0 mm without
hairs; Chapada dos Veadeiros, Goias, Brazil.
22. B. hirsuta.
123. Lamina appressed-pubescentto velutinous adaxially,

and tomento-sericeous abaxially; pedicels whitetomento-sericeous,0.8-1.2 mm in diameter,0.6-0.8 mm
without hairs.
21. B. variabilis.
122. Laminawith the petiole 2-15 mm long, less than 2 mm thick.

124. Lamina appressed-pubescent to glabrate adaxially,
sparselyto densely appressed-pubescentabaxially;pedicels appressed-sericeousor subtomentose. 18. B. malifolia.
124. Laminavelutinousto glabrateadaxially,loosely tomento-sericeousabaxially;pedicels velutinousto subtomentose.
125. Petiole 6-15 mm long; lamina (3.5-)5.9-12.0 cm
long, plane; inflorescence usually in the axils of
full-sizedleaves or cauliflorous.
19. B. confusa.
125. Petiole 2-7 mm long; lamina 2.8-7.4(-10.2) cm
long, rugose;inflorescenceusuallyon long terminal
shoots with the umbels borne in the axils of reduced leaves.
18. B. malifolia.
118. Laminawith 1-2(-5) pairs of glands abaxiallyon the lateralveins near
the base; sepals abaxially golden-sericeous,broadly ovate, obtuse to
acute and not revolute at the apex, firmlypressed againstthe corollain
bud.
126. Slenderwillow-liketree or shrubwith pendentbranches, not vining; laminaless than 7 mm wide, acicular,glabrate;inflorescence
of a pair of flowers terminatingbranches.
17. B. acerosa.
126. Shrubletsor shrubswith viningbranches;lamina 1.0-6.5 cm wide,
ovate to elliptic or lanceolate,pubescentor glabrous;inflorescence
of 4-floweredumbels borne singly or in condensed axillaryor terminalcymes.
127. Laminaglabrousabaxiallyor with sparse appressed, sessile
16. B. stellaris.
hairs, the trabecula0.2-0.6 mm long.
32
Flora Neotropica
127. Laminahairy abaxially, the hairs T-shaped, stipitate or ses15. B. schizoptera.
sile, the trabecula1.0-2.4 mm long.
Key to FruitingSpecimens
1. Calyx withoutglands.
2. Fruit nut smooth laterally,withoutlateralwings or crests.
3. Samaradensely appressed-sericeousthroughout,the hairs soft and not irritating,the
trabeculaup to 0.5 mm long.
40. B. muricata.
3. Samarasparselyappressed-sericeousto glabrate,the hairs stiff and irritating,the trabecula 0.6-3.0 mm long.
4. Laminaabaxiallydensely hairy.
5. Styles incurvedtowardsthe axis apically, densely hairyin the basal third,the
hairs spreading,flexuous.
B. leiocarpa group.
6. Lamina 1.4-4.5 cm long and 0.4-2.5 cm wide.
72. B. parvifolia.
6. Lamina3.0-8.4 cm long and 2.0-7.4 cm wide.
7. Laminaabaxiallyloosely tomentose, with 4-7 pairs of lateralveins; petiole 9-24 mm long.
71. B. leiocarpa.
7. Laminaabaxiallyshining-sericeous,with 7-9 pairs of lateralveins; petiole 7-14 mm long.
73. B. populifolia.
5. Styles erect or divergingand directed away from the axis, glabrous or appressed-sericeousat the base.
8. Secondaryveins interconnectingprimarylateralveins parallel;Peru.
45. B. velutinissima.
8. Secondaryveins betweenprimarylateralveins formingprominentreticulum.

9. Brancheshispid with interspersedbasifixedhairs;petiole and laminaon
both sides hispid to hispido-sericeous.
63. B. basifixa.
9. Branchestomento-sericeous,hairsall malpighiaceous;petiole andlamina
61. B. salicifolia.
abaxiallytomento-sericeous,laminaglabrousadaxially.
4. Laminaabaxiallyglabrousor very sparselysericeous or tomentose.
10. Flowers mostly borne in 4-floweredumbels, rarely in 6-floweredumbels or
racemes.
11. Pedicels sparselytomentose;sepals sericeousabaxially;leaf base strongly
50. B. pubescens.
cordate, subsessile;petiole 2-7 mm long.
11. Pedicels glabrous;sepals glabrousor very sparsely sericeous abaxially;
leaf base truncateto cordate;petiole 5-14 mm long.
12. Commonpeduncle0.4-1.0(-1.5) cm long; inflorescenceminutelyvelutinousor tomentose.
44. B. martiniana.
12. Common peduncle up to 2.5 cm long; inflorescence glabrous or
46. B. elegans.
sparsely appressed-sericeous.
10. Flowersbornein pairsor alternatelyon the side branchesof the inflorescence,
never in 4-floweredumbels.
13. Leaf marginwith 3-5 pairs of prominentcup-shapedglands; pedicel tomento-sericeous.
48. B. padifolia.
13. Leaf margineglandularor with numerousminuteglands on or withinthe
margin;pedicels sparselyappressed-sericeousto glabrate. 47. B. polygama.
2. Fruit nut bearingcrests or wingletsor muricate.
14. Fruitnut muricateor cristatelaterally,densely appressed-sericeousthroughout,the
hairs soft and not irritating,the trabeculaless than 0.5 mm long.
40. B. muricata.
14. Fruitnut with wingletslaterally,appressed-sericeousto glabrous,the hairs stiff and
irritatingto the skin, the trabecula0.6-3.0 mm long.
15. Lateralwinglets radiatingfrom the areole.
B. lutea group.
16. Sepals abaxiallysparselytomentose;fruit wing 19-26 mm long, appendage
2-5 mm tall.
69. B. lutea.
16. Sepals abaxiallyappressed-sericeous;fruitwing 25-34 mmlong, appendage
6-7 mm tall.
70. B. cristata.
15. Lateralwinglet(s)parallelto the areole.

17. Nut with a single winglet on each side; laminaglabrousor with a few hairs
on the midrib.
33
18. Nut greatly enlarged, more than 1 cm tall and long, the dorsal wing
crest-like.
92. B. sepium.
18. Nut up to 9 mm tall and 6 mm long; dorsal wing well developed.
17. Nut with 2-3 winglets on each side; laminadensely hairy abaxially,rarely
sparsely so.
19. Calyx valvate, sepals more than 7 mm long.
67. B. valvata.
19. Calyx imbricate,sepals less than 3 mm long.
20. Bracts and bracteoles appressed;laminaabaxiallytomentose; Colombia.
68. B. heterostvla.
20. Bracts and bracteolesspreading;laminaabaxiallysericeous;Rio de

Janeiroand EspiritoSanto, Brazil.
91. B. patula.
1. Calyx glandular.
21. Calyx glandsborne on free part of sepals.
22. Flowersmostlybornein 4-floweredumbels,rarelyin 6-floweredumbelsor racemes.
23. Umbels borne in "racemes" of umbels with 5-7 pairs of umbels borne on an
axillary branch up to 7 cm long, the common peduncles less than 1 cm long.
B. parviflora group.
24. Laminaquite glabrous,or with a few hairs.

60. B. parviflora.
24. Laminavery hairy, at least abaxially.
25. Pedicels glabrous,rarelywith 1 or 2 hairs.
26. Petiole 3-12 mm long; lamina lanceolate, appressed-sericeous
abaxially.
64. B. parviglandula.
abaxially.
63. B. basifixa.
26. Petiole 10-20(-35) mm long; lamina ovate, hispido-tomentose
25. Pedicels sparselyhairy.

27. Laminaadaxiallyglabrous.
61. B. salicifolia.
27. Laminaadaxiallyvelutinous.
62. B. scutellata.
23. Umbels borne singly, or in dichasiaor cymes, or if with the appearanceof a
raceme of umbels, the commonpedunclemore than 1.5 cm long.
28. Laminawith marginalglands, sometimesonly well developedon the inflorescence leaves, and sometimes developed into ciliate extensions at the
B. martinianagroup.
apex of the leaf.
29. Inflorescencebranchestomentousor velutinous, sometimes minutely
so.
30. Sepals tomento-sericeousabaxially;base of lamina stronglycordate, almost claspingthe stem, petiole 2-4(-7) mm long.
50. B. pubescens.
30. Sepals glabrous or very sparsely appressed-sericeousabaxially;

base of laminatruncateto cordate, petiole (3-)5-20 mm long.
31. Pedicels 12-18 mm long; sepals 2.8-3.2 mm long and 2.0-2.5
mm wide; nut up to 6 mm tall and 3-5 mm long; hairs on
49. B. wilburii.
samarawith trabecula1.5-2.5 mm long.
31. Pedicels 4-14 mm long; sepals 1.4-2.5 mm long and wide; nut
up to 4 mm tall and 2-3 mm long; hairs on samarawith trabecula 0.6-1.5 mm long.
32. Branchesand leaves densely velutinous. 45. B. velutinissima.
32. Branches and leaves glabrous or sparsely appressed-sericeous.
44. B. martiniana.
29. Inflorescencebranchesglabrousor appressed-sericeous.

33. Base of leaf strongly cordate, almost clasping stem; petiole 2-6
mm long; Guayana Highland.
54. B. pulcherrima.
33. Base of laminatruncateto cordate;petiole 5-14 mm long.

34. Lamina abaxially sparsely appressed-sericeous,rarely glabrate; inflorescencegolden-sericeous;pedicels appressed-se53. B. maguirei.
riceous; GuayanaHighland.
34. Lamina abaxially glabrous, rarely sparsely sericeous; inflorescence glabrous or sparsely silvery-appressed-sericeous;
46. B. elegans.
pedicels glabrous.
28. Leaves with 1-4 pairs of glands on the lamina,or a single pair at the base
beside the midrib.
35. Bractsand bracteolesbroadlytriangular,papery,glabrous,persistent.
65. B. pseudojanusia.
36. Petiole 7-11 mm long; laminalanceolate.
34
Flora Neotropica
36. Petiole up to 3 mm long; laminaovate to rotund.
35. Bracts and bracteoles triangularto lanceolate, not papery, abaxially

hairy, deciduousor persistent.
37. Locule of nut densely hairy within.
38. Lamina abaxially sparsely appressed-sericeousto glabrate
with 1-5 pairs of sessile or short-stalkedglands on the lamina
betweenthe midriband the marginabove the leaf base, sometimes with an additionalpair beside the midribat the base or
with 1-2 pairs of cup-shapedglands on the petiole.
39. Pedicels 7-11 mm long; nut of samarasmooth to rugose.
40. Lamina sparsely appressed-sericeous abaxially.
35. B. caapi.
40. Laminadensely and loosely sericeous abaxially.
38. B. multifoliolata.
39. Pedicels 11-17 mm long; nut of samaraprominentlyalu-
late.
38. Laminaabaxiallytomentose,appressed-sericeousor glabrous,

with a single largepairof glandsbeside the midribat the base
of the leaf or at the apex of the petiole.
41. Wings of posterior samaras more or less rotated to lie
parallelto the wing of the anteriorsamara;petioles less
than 8 mm long, or if more, less than 2 mm thick.
B. malifolia group.
42. Petiole of largerleaves more than 2 mm thick, up to

3 mm long.
43. Laminaglabrouson both sides.
44. Tree with thick corky bark; fruits borne on
old wood (cauliflorous),sometimes also on
short terminal branches bearing reduced
leaves; young branches smooth, red-brown,
20. B. latifolia.
not lenticellate.
44. Shrubor shrublet,barklenticellate,not thick
andcorky;fruitsborneon axillaryor terminal
branches;young brancheslenticellate.
45. Shrublet with reclinate branches to
1 m; lamina falcate; inflorescence
branches sparsely sericeous to glabrate,
reducedleaves in inflorescenceglabrate;
Chapadados Veadeiros, Goias, Brazil.
23. B. hatschbachii.
45. Shrubor shrubletwith erect branchesto

2.5 m; lamina plane; inflorescence
branches densely sericeous or velutinous, reducedleaves in inflorescenceappressed-sericeousabaxially. 21. B. variabilis.
43. Lamina loosely tomentose or hispid-pubescent
abaxially.
46. Lamina densely hispid-pubescenton both
sides; pedicel 1.3-2.2 mm in diameter;Chapada dos Veadeiros,Goias, Brazil. 22. B. hirsuta.
46. Lamina appressed-pubescent or glabrate,
rarely velutinous, adaxially, loosely sericeous abaxially;pedicel 1.0-1.3(-1.7) mm in
diameter.
21. B. variabilis.
42. Petiole of largerleaves up to 2 mm thick, more than

3 mm long.
47. Pedicels and inflorescencevelutinous to tomentose.
48. Petiole 3-7 mm long; laminarugose adaxially, 2.8-7.4 cm long; fruits borne on long terminalbranchesin the axils of reducedleaves.
18. B. malifolia.
35
48. Petiole 6-15 mmlong;laminaplaneadaxially,

(3.5-)5.9-12.0 cm long; fruits borne on short
branches in the axils of full-sized leaves or
cauliflorous.
19. B. confusa.
47. Pedicels and inflorescenceappressed-pubescent.
18. B. malifolia.
41. Wings of posterior samarasequidistantfrom each other

and from the wing of the anteriorsamara;petioles more
than 8 mm long and more than 2 mm thick.
B. megaphylla group.
49. Laminaellipticor broadlyelliptic;pedicel 1.5-2.0 mm

in diameter; wing of samara 18-27 mm long.
25. B. irwinii.
49. Laminabroadly elliptic to orbicular;pedicel 1.0-1.6

mm in diameter;wing of samara 13-21 mm long.
24. B. megaphylla.
37. Locule of nut glabrouswithin.

50. Bracts and bracteolesdeciduous.
51. Petiole 1-7 mm long.
52. Fruitnut prominentlyalulatewith one or more lateral
wingletson each side more than 4 mm high and with
31. B. calcicola.
an attachmentmore than 4 mm long.
52. Fruit nut smooth or rugose, rarely with one small
winglet up to 4 mm long on one or both sides, with
an attachment1-2 mm long.
53. Laminaabaxiallyvelutinous;anteriorstyle stout,
up to 0.6 mm thick in diameter,posterior styles
36. B. schwannioides.
reduced.
53. Laminaabaxiallyglabrousor loosely tomentose;

styles equal, 0.2-0.4 mm in diameter.
54. Slender treelet or shrub, not vining; leaves

pendent, acicular, 1.5-7.0 mm wide, glabrate, inflorescenceof single pairs of flowers
17. B. acerosa.
terminatingbranches.
54. Shrubor shrublet,sometimes vining; leaves
divergingor reflexed,lanceolateto orbicular,
1.0-6.5 cm wide, glabrateor pubescent; inflorescenceof 4-floweredumbelsbornesingly
or in cymes on axillaryor terminalbranches.
55. Lamina glabrous to sparsely pubescent
abaxially,the hairs sessile, the trabecula
16. B. stellaris.
0.2-0.6 mm long.
55. Lamina pubescent abaxially, the hairs
stipitate,T-shaped,the trabecula1.0-2.4
15. B. schizoptera.
mm long.
51. Petiole of largerleaves (5-)7-27 mm long.

56. Fruit nut smooth to slightly rugose; styles subequal,
straight,parallelto slightlydiverging,not lyrateat the
37. B. prancei.
base.
56. Fruit nut prominentlyalulate; posterior styles longer than anterior style, divergingand lyrate at the
base.
B. membranifolia group
57. Laminavelutinouson both sides, sometimesglabrate adaxially, bearing 1-2(-4) pairs of stalked
glands, the stalk 0.5-1.5 mm long.
34. B. adenopoda.
57. Laminaappressed-pubescentor glabrateon both

sides, bearing 1-2(-4) pairs of sessile or subsessile glands.
58. Sepals 4.5-5.5 mm long; laminadensely appressed-white-sericeous,the hairssubsessile,
33. B. pulchra.
the trabecula0.2-0.6 mm long.
58. Sepals up to 3.5 mm long; lamina sparsely
36
Flora Neotropica
appressed-sericeous to glabrate, the hairs
sessile, the trabecula0.1-0.25 mm long.
50. Bracts and bracteolespersistentin fruit.

59. Pedicels borne on peduncles0.5-4.0 mm long.
B. muricata group.
60. Non-viningshrub;laminalancolate, densely shiningwhite-sericeousabaxially;wing of samara 11-18 mm

43. B. goiana.
long; Goias, Brazil.
60. Vines; lamina ovate or elliptic, sparsely to densely
sericeous abaxially;wing of samara16-40 mm long.
61. Lamina 1.3-6.7 cm long, elliptic, the marginrevolute, abaxiallysparselysericeous, with the petiole 5-8 mmlong; wing of samara16-23 mmlong;
Cuba.
61. Lamina (3.5-)5.3-13.0(-18.0)
42. B. pauciflora.
cm long, ovate to
orbicular,the marginplane, abaxiallysparselyto

densely sericeous or tomentose, with the petiole
5-25 mm long; wing of samara20-40 mm long.
62. Petiole 14-25 mm long; pedicels 1.0-1.4 mm
in diameter, 0.8 mm without hairs.
41. B. oxyclada.
62. Petiole 5-16 mm long; pedicels 0.6-0.8 mm

in diameter,0.3-0.5 mm withouthairs.
63. Styles slender,up to 0.2 mmin diameter,
posterior styles longer than anterior
40. B. muricata.
style, lyrate.
63. Styles stout, 0.3-0.4 mm in diameter,
equal, parallel or diverging apically.
39. B. quadriglandula.
59. Pedicels sessile.

64. Styles flattenedlaterally,divergingand recurvedapically, 0.4-0.8 mm in diameterat the apex.
65. Laminawith the reticulationprominulousadaxi9. B. laevifolia.
ally; pedicels 6-10 mm long.
65. Laminawith at least the lateralveins impressed
adaxially;pedicels 8-19 mm long.
B. argyrophylla group.
66. Laminanarrowlyellipticto linear(morethan

3 times as long as wide).
67. Shrubletor shrub, never vining; lamina
chartaceouswith only the primaryveins
impressed, not rugose or bullate; Goias,
Brazil.
14. B. vernoniifolia.
67. Shrub or vine; lamina chartaceous or

very coriaceous, rugose or bullate.
68. Lamina very coriaceous, bullate.
13. B. harleyi.
68. Lamina chartaceous, with the reticulation impressed and the areoles
betweenthe veins shallowlyinflated.
66. Laminaelliptic to ovate (less than 3 times as

long as wide).
69. Fruit of 3 mericarps,the dorsal wing reduced to a shallow crest 1-2 mm high.
12. B. paraguariensis.
69. Fruit of 3 samaras,the dorsal wing well

developed, 12-40 mm long.
70. Laminacoriaceous, stronglybullate.
13. B. harleyi.
37

70. Lamina chartaceous, more or less
rugose.
71. Non-vining shrub or shrublet;
wing of samara 12-17 mm long,
its lower edge meetingthe nut in
the middle; stems and lamina
adaxially cobwebby-pubescent.
11. B. bvssacea.
71. Liana or vining shrub; wing of

samara(20-)25-36 mm long, its
lower edge embracingthe base
of the nut; stems velutinous to
subtomentose;lamina adaxially
glabrate.
64. Styles not flattenedlaterally, straightand parallelor

divergingat the base but not recurvedapically, 0.10.3 mm in diameterat the apex.
72. Petiole 5-16 mm long, laminasmooth adaxially,
abaxially sparsely to densely appressed-seri40. B. muricata.
ceous, broadlyovate.
72. Petiole 1-8 mm long, laminawith the reticulation
impressed adaxially, abaxially velutinous, loosely tomentose, glabrousor densely tomento-sericeous,if tomento-sericeousthe lamina
narrowlyelliptic to linear.
73. Nut of samaraprominentlyalulate with one
or morelateralwings on each side morethan
31. B. calcicola.
4 mm high.
73. Nut of samararugose rarely with one small

wingletup to 3 mmhighon one or both sides.
B. campestris group.
74. Fruit wing reduced to 1.5-6.0(-9.0) mm

long; shrublet to 4 dm; Serra do Cipo,
Minas Gerais, Brazil.
27. B. cipoensis.
74. Fruit wing well-developed, 13-32 mm

long; shrublet,shrubor slendertree.
75. Laminaabaxiallywith a pair of peltate glandson the basal lateralveins
one-third to two-thirds of the distance from the midribto the margin.
76. Slender tree to 4 m tall, much
branched;nut of samara often
with a shortlateralwing 2-3 mm
high; wing 18-32 mm long.
30. B. arborea.
76. Shrubto 1.5 m tall, unbranched

below the inflorescence;nut of
samararugose; wing (11-)15-27
mm long.
26. B. campestris.
75. Laminaabaxiallywith 1(-2) pair(s)of

peltateglandsbeside the midribat or
near the base of the leaf.
77. Lamina glabrous abaxially except the midriband main lateral
veins.
78. Slender treelet or shrub;
leaves held erect, morethan
3 times as long as wide with
only the lateral veins impressed; wing of samara612 mm wide.
29. B. andersonii.
38
Flora Neotropica
78. Shrub;leaves divergingfrom
the stem, less than 3 times
as long as wide, rugose;
wing of samara 9-19 mm
wide.
26. B. campestris.
79. Lamina abaxially very
densely tomento-sericeous,
not visible throughthe pubescence, more than 2.5
times as long as wide, the
margin revolute, with only
the primaryand secondary
veins impressed;wing of samara 13-22 mm long.
28. B. angustifolia.
79. Laminaabaxiallyloosely tomentose, or if appressedpubescent less than 2.5

times as long as wide and
the marginnot revolute, rugose; wing of samara(11-)
15-27 mm long.
26. B. campestris.
22. Flowers borne alternatelyor in pairson the ultimatebranches,never in 4-flowered

umbels.
80. Leaves with marginalglands, sometimesonly well-developedon inflorescence
leaves, and sometimesdevelopedinto ciliate extensionsat the apex of the leaf.
81. Flowers and leaves all borne alternately.
52. B. alternifolia.
81. Flowers and leaves opposite or subopposite,rarelysome alternate.
82. Laminawith 3-5 mainlateralveins all or mostlyarisingin the proximal
half of the leaf; stem with sparse basifixedhairs, often persistingonly
at the nodes; inflorescenceminutelyvelutinous;inflorescenceleaves
51. B. grandifolia.
long-ciliate.
82. Laminawith 5-9 main lateral veins, arisingthroughoutthe length of

the midrib;stems glabrous or appressed-sericeous,hairs malpighiaceous; inflorescence appressed- or tomento-sericeous;inflorescence
leaves with marginalglands.
83. Leaves very coriaceous;petiole up to 14 mm long; leaf base truncate to cordate, marginrevolute;inflorescencegolden-appressed53. B. maguirei.
sericeous; GuayanaHighland.
83. Leaves chartaceousto subcoriaceous;petiole up to 8 mm long;
leaf base cuneateto truncate,marginplane;inflorescencetomentosericeous or silvery-appressed-sericeous.
84. Leaf marginwith 3-5 pairs of prominentcupulateglands;in48. B. padifolia.
florescencetomento-sericeous.
84. Leaf margineglandularor with numerousminuteglandson or
withinthe margin;inflorescenceappressed-sericeous.
47. B. polygama.
80. Leaves bearing2-6 pairs of glands on the lamina abaxiallyand/or with 1-2
pairs at the apex of the petiole.
85. Leaves with 2-6 pairs of glands on the laminaabaxially, sometimes also
with 1-2 pairs on the petiole.
86. Stipules minute,0.5-1.0 mm long; laminaabaxiallyvelutinous.
37. B. prancei.
86. Stipules linear, 1-5 mm long; laminaabaxiallydensely white-shiningsericeous
85. Leaves with a single pair of glands at the apex of the petiole.
B. nummifera group.
87. All or some leaves quite glabrousabaxially.

88. Pedicels 5-7 mm long; fruit nut up to 5 mm tall and 3 mm long,
39
bearingone small winglet on each side; wing 20-24 mm long and

9-11 mm wide.
56. B. sellowiana.
88. Pedicels 7-13 mm long; fruit nut 6-8 mm tall, 3-5 mm long, reticulate laterally; wing 22-32 mm long and 11-15 mm wide.
55. B. nummifera.
87. All leaves hairyabaxially.

89. Styles moreor less equal, stout, 0.2-0.4 mmin diameter,diverging
from the base.
90. Laminaabaxiallyappressed-tomentose.
57. B. anisandra.
90. Laminaabaxiallyappressed-shining-sericeous. 55. B. nummifera.
89. Posterior styles longer or shorter than the anteriorstyle, sometimes absent, slender,less than 0.2 mm diameter.
91. Posteriorstyles longerthan anteriorstyle.
59. B. lyrata.
91. Posteriorstyles shorterthan anteriorstyle or absent.
58. B. gardneriana.
21. Calyx glands borne below free part of sepal on receptacle.

92. Bracts and bracteoleslingulate,spreading.
93. Fruitwith dorsal wing representedby shallow crest, nut with 2-several lateral
winglets or crests.
Diplopterys.
93. Fruitwith dorsal wing, nut with 1 winglet on each side.
90. B. krukoffii.
92. Bracts and bracteolestriangular,erect or appressedto pedicel.
94. Lamina abaxially with 2-4 pairs of glandularareas, V? to 2/3 of the distance
between the midriband margin.
89. B. caduciflora.
94. Laminawith numerousminuteglands on the laminanearthe margin.
95. Flowers borne in panicles, the ultimatebranchesbearing8-45 flowers alternatelyor in pairs;fruit nut smooth or with a single lateralwing parallel
to the areole.
SubgenusBanisteriopsis.
96. Hairs on samara1.5-2.0 mmlong; lateralwingletsconfluentbelow the
8. B. longipilifera.
nut.
96. Hairs on samara0.4-0.8 mm long; lateral winglets absent or with a

wingleton each side of the nut, not confluentbelow.
97. Ultimatebranchesof inflorescencewith up to 45 flowersarranged
in a spiral.
98. Laminalanceolate,up to 11.0 cm long and 4.5 cm wide; ped3. B. ferruginea.
icels 7-10 mm long.
98. Laminaelliptic, up to 15.4 cm long and 8.6 cm wide; pedicels
2-5 mm long.
5. B. brevipedicellata.
97. Ultimatebranchesof inflorescencewith up to 25 flowersarranged

in a spiralor in pairs.
99. Fruit nut 9-11 mm tall, 7-9 mm long; wing (25-)30-53 mm
4. B. megaptera.
long and (11-)14-23 mm wide.
99. Fruitnut 4-9 mm tall, 4-8 mm long; wing 16-38 mm long and
7-16 mm wide.
100. Styles 2.0-2.6 mm long, divergingat the base and more
or less inflexed towards the axis apically; filamentsof
stamens opposite the antero-lateralsepals flexuous and
directedtowardsthe posteriorpetal.
101. Lamina abaxially sparsely appressed-sericeous.
6. B. wurdackii.
7. B. cinerascens.
100. Styles 1.0-2.0 mmlong, straightandparallelor diverging
apically;stamens subequal,erect.
102. Lamina abaxially sparsely appressed-sericeous.
102. Laminaabaxiallyglabrateor tomentose.
1. B. cornifolia.
2. B. acapulcensis.
95. Flowers borne in 4-floweredumbels or short axillaryracemes of up to 12

flowers; fruit nut reticulate, cristate or with 2-3 winglets parallel to or
radiatingfrom the areole.
103. Fruit nut with 2-3 winglets on each side parallelto the areole.
B. pubipetala group.
Flora Neotropica
40
104. Leaves with 8-18 pairs of lateral veins, abaxiallysparsely ap87. B. erianthera.
pressed-sericeous.
104. Leaves with 4-8 pairs of lateralveins, abaxiallyglabrous.
105. Petiole 2-8(-12) mm long; lamina5.1-13.0(-15.5) cm long
and 2.1-6.0(-8.3) cm wide.
106. Flowers borne in 4-flowered, rarely 6-floweredumbels; pedicel thickened in fruit up to 1.5 mm in diameter;nut 11-15 mm tall and 9-11 mm long; styles
stout, equal and diverging.
85. B. platyptera.
terior styles.
84. B. pubipetala.
106. Flowers borne in condensed axillaryracemes of 3-5

pairs of flowers, the terminal4-6 flowers often forming an umbel; pedicels not thickenedin fruit, up to
1.0 mmin diameter;nut 4-6(-11) mmtall and 3-5(-9)
mm long; anteriorstyle longerand thickerthan pos-
105. Petiole (6-)10-27 mm long; lamina (8.0-)12.0-25.5 cm

long, 4.5-12.0 cm wide.
107. Leaves with numerousglandularareas on the lamina
near the margin;flowers borne in 4- to 6-flowered
umbels or condensed racemes of 3 pairs of flowers;
fruitnut 13-17 mmtall and 11-13 mmlong;fruitwing
(56-)60-73 mm long and 20-28 mm wide. 88. B. schunkei.
107. Leaves with minute glands on or near the margin;
flowersbornein axillaryracemesof 4-8 pairsof flowers; fruit nut 7-13 mm tall and 7-10 mm long; fruit
wing (38-)44-55 mm long and 15-20 mm wide.
86. B. longialata.
103. Fruit nut reticulateor cristate or with 2-3 winglets on each side raB. lucida group.
diatingfrom the areole.
108. Leaves sessile or subsessile, petiole up to 3 mm long.
83. B. rondoniensis.
109. Laminaglabrous.
110. Laminabroadlyovate to orbicular,5.5-10.6 cm long
and 5.2-9.2 cm wide, claspingthe stem; fruit nut up
to 9 mm tall and 8 mm long. Mato Grosso, Brazil.
82. B. amplectens.
110. Lamina ovate or elliptic to orbicular,2.0-7.5(-9.0)

cm long and 1.0-6.4(-7.5) cm wide, cordate at the
base; fruit nut up to 7 mm tall and 6 mm long.
111. Liana; stems and pedicels appressed-sericeous;
nut laterallysmooth to reticulate,densely golden-sericeous.
81. B. virgultosa.
111. Shrubor subshrub;stems andpedicelsglabrous;

nut laterally ridged or cristate, sparsely sericeous to glabrate.
80. B. hypericifolia.
108. Leaves petiolate,petiole 6-9 mm long.

112. Laminaabaxiallytomentose or velutinous.
113. Lamina4.4-6.5 cm long, abaxiallytomentose; Serra
79. B. cachimbensis.
do Cachimbo,Para.
113. Lamina4.5-14.6 cm long, abaxiallytomento-velutinous; Peru.
77. B. woytkowskii.
112. Laminaabaxiallysparselyappressed-sericeousto glabrate.

114. Nut up to 20 mm tall and 15 mm long. 76. B. peruviana.
114. Nut up to 10 mm tall and 10 mm long.
78. B. nigrescens.
115. Fruit wing 20-25 mm long.
115. Fruitwing 30-63 mm long.
116. Calyxglands 1.0-1.8 mmlong, 0.8-1.2 mm
wide;anteriorstyle longerand stouterthan
posterior styles, apically inflexed; fruit
with appendageon upper marginof wing
up to 10 mm tall; wing usually narrowat
74. B. lucida.
the base, wideningdistally.
116. Calyxglands2.0-3.8 mmlong, 1.2-2.0 mm
41
wide; styles equal, stout, diverging;fruit
with appendageon upper marginof wing
up to 5.5 mm tall; wing usually parallelsided and embracingthe nut at the base.
75. B. nutans.
Banisteriopsis subgenus 1. Banisteriopsis, species 1-8.

Banisteria section MonocteniaNiedenzu, Ind. Lect. Lyc. Brunsb.:4. 1900.Lectotype species.
Banisteriopsis cornifolia (H.B.K.) Robinson in Small as Banisteria maracaybensis.
Leaf glands marginal, sessile. Flowers borne in panicles. Bracts and bracteoles
persistent, pedicels short pedunculate, thicker in fruit. Calyx 8-glandular, the
glands borne on the receptacle below the free part of the sepal. Petals yellow,
entire to denticulate. Stamens with eglandular connectives. Styles equal, glabrous
or basally strigose. Ovary short-white-sericeous. Fruit nut ca. orbicular, smooth
or with a single winglet on each side parallel to the areole, bearing a shallow
appendage on its upper edge, nut locule glabrous within.
Type species. Banisteriopsis cornifolia (H.B.K.) Robinson in Small.
1. Banisteriopsis cornifolia (H.B.K.) Robinson in Small, North Amer. Flora 25:
132. 1910.
Heteropterys cornifolia H.B.K., Nov. Gen. et Sp. PI. 5: 165. 1821 [1822].
Banisteriacornifolia(H.B.K.) Sprengel,Syst. Veg. 2: 388. 1825.
Banisteria maracaybensis Adr. Jussieu var. cornifolia (H.B.K.) Niedenzu, Ind. Lect. Lyc.
Brunsb.:5. 1900.
Shrub or vine, branches terete, at first appressed-sericeous, soon glabrate, bark

greyish- or reddish-brown with pale prominent lenticels. Stipules triangular up to
1 mm long, often joined by an interpetiolar line. Leaves with the petiole 3-15 mm
long, appressed-sericeous or glabrous, eglandular, the lamina 4.3-11.5(-17.9) cm
long, 2.2-6.7(-8.8) cm wide, lanceolate to elliptic, cuneate, rarely truncate, at the
base, obtuse to acute to apiculate at the apex, with the margin slightly revolute
and bearing minute glands, the gland at the base of the leaf on each side greatly
enlarged, glabrous adaxially, minutely appressed-sericeous abaxially, the hairs
with the trabecula 0.2-0.4 mm long, with the reticulation prominent adaxially and
the lateral veins prominent abaxially. Inflorescence paniculate, axillary, the
branches up to 30 cm long, appressed-sericeous, the side branches bearing 8-20
flowers arranged in a spiral; bracts and bracteoles 0.6-1.0 mm long, triangular,
appressed-sericeous abaxially, involucrate, persistent; pedicels sessile or borne
on a peduncle up to 1.5 mm long, 4-10 mm long, 0.3-0.6 mm in diameter, appressed-sericeous, thicker and glabrate in fruit. Sepals 1.0-1.8 mm long, 0.6-1.2
mm wide, oblong, rounded at the apex, appressed-sericeous abaxially, glabrous
adaxially, the 4 lateral sepals biglandular, projecting 0.6-1.4 mm beyond the
glands, the glands 1.5-2.8 mm long, 0.8-1.5 mm wide. Petals yellow, glabrous,
the 4 lateral petals reflexed between the sepals, the claw 0.5-1.0 mm long, the
limb 3.5-5.0 mm long and 2.5-3.5 mm wide, broadly obovate, entire, a little
concave, the posterior petal with the claw suberect, 1.0-1.2 mm long, only a little
longer and thicker than those of the other petals, the limb 3.5-5.0 mm long, 2.03.0 mm wide, obovate, basally denticulate, plane. Stamens with the filaments
erect, those opposite the sepals subequal, 1.4-2.0 mm long, those opposite the
petals subequal, 1.0-1.4 mm long, glabrous; anthers reflexed, the locules 0.9-1.4
mm long, glabrous or sparsely hairy, the connectives 0.8-1.2 mm long, 0.3-0.7
mm wide, 0.2-0.4 mm deep, those opposite the sepals a little wider and deeper
42
Flora Neotropica
than those opposite the petals. Ovary 1.0-1.2 mm tall, hairy, the styles stout and
equal, 1.6-1.8 mm tall, diverging apically. Fruit of 3 samaras, the carpophore
short and stout, the nut 5-8 mm tall and 4-5 mm long, appressed-sericeousto
glabrate, smooth or bearing on 1 or both sides a ridge or short winglet up to 6
mm along its base and projectingup to 2 mm from the nut, the wing 16-34 mm
long, 7-15 mm wide, appressed-sericeousto glabrate,the hairs 0.3-0.5 mm long,
with an appendagealong its upper marginat the base 3-7 mm long and 1-2 mm
tall.
Key to the Varieties of Banisteriopsis cornifolia
1. Petiole (9-)10-15 mm long; fruit wing 16-27 mm long and 7-11(-13) mm wide.
a. var. cornifolia.
1. Petiole 4-7(-9) mm long; fruit wing 22-34 mm long and 10-15 mm wide.
2. Side branches of inflorescence up to 2 cm long, bearing 10-20 flowers; fruit nut with
lateral crest or wing.
b. var. elliptica.
2. Side branches of inflorescence less than 1 cm long, bearing up to 8 flowers; fruit nut
c. var. standleyi.
quite smooth.
la. Banisteriopsiscornifoliavar. cornifolia

Branches dark- or grey-brown with pale lenticels. Leaves with the petiole
(9-)10-15 mm long, the lamina4.3-12.8 cm long, 2.2-6.2 cm wide, lanceolate to
elliptic, with the reticulumof veinlets prominentadaxially, the smallest areoles
0.5-1.0 mm2.Inflorescenceof narrowelongate axillarypanicles, the branches830 cm long, the side branches, often 2 or 3 in each axil, up to 2.5 cm long and
bearing 8-12 flowers. Stamens with glabrous anther locules. Fruit with the nut
5-6 mm tall and 3.5-4.5 mm long, smooth or usually bearinga ridge or winglet
on one or both sides, the wing 16-27 mm long, 7-11(-13) mm wide, erect or
diverging.
Type. Humboldt&Bonplands.n., Colombia,Tolima,nearIbague,fl (holotype,

P; isotype, P).
Distribution. (Fig. 5). In the Magdalenavalley in Colombia at elevations of
250-1700 m, extending N to Venezuela.
Collected in flower in February,July, November and December and in fruit in
July and December.
COLOMBIA. Cundinamarca: Andre 1874 fl (F, NY); Garcia-Barriga 11834 bud (US); 12332 fr
(US); Holton 800 fl (G, NY); Killip et al. 38193 bud (F, US); Lehmann B. T. 1093 fl (G, GH, NY);
Lehmann 7558 fl (F, G, NY, US); Pennell 2838 fl fr (GH, NY, US); 2929 fl fr (GH, NY, US); Perez
Arbeldez 2106 fl fr (US); Triana 3394 fl (BR, G, NY); Uribe-Uribe 2878 fl (U). Tolima: Haught 6258
fl (US); Humboldt & Bonpland s.n. fl (P); K0ie 4980 fr (US). Huila: Mason 13838 fl (GH, US); 13881
fr (US); Rusby & Pennell 319 fl fr (NY, US); 354 fl (NY); S. G. Smith 1135 fl (GH, IAN, MO, US).
No other locality: Mutis 2420 fl (US). VENEZUELA. Saer 755 fl (F).
Common names. Colombia: "almizclillo

"patillo."
Banisteriopsis cornifolia var. cornifolia is restricted to the Magdalena valley
of Colombia except for one collection, Saer 755, from Venezuela. It is distinguishedfrom var. elliptica by its longerpetioles, smallernarrowerleaves, narrow
elongate axillarypanicles, and smallerfruits. The presence or absence of a lateral
winglet on the fruit is a variablecharacter;some collections have a smooth nut,
some a shallow ridge or a well-developed winglet.
lb. Banisteriopsiscornifoliavar. elliptica(Niedenzu) B. Gates, comb. nov.
Banisteria maracaybensis Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 285. 1840. Type. Ple
s.n., Venezuela, Zulia, Maracaibo, fl fr (holotype, P; isotypes, BR, F, P, P-JU).
43

*4pf" *:.
,+
'
..-'*.
...'
, f,.^,7 ,d^4
tf-?
'.
V.
+,,
,s
;-?
..
--
+"
-.'
:.:
4-
FIG. 5. Distribution of Banisteriopsis cornifolia and B. acapulcensis. Closed circle, B. cornifolia

var. cornifolia; open circle, B. cornifolia var. elliptica; triangle, B. cornifolia var. standleyi; closed
square, B. acapulcensis var. acapulcensis; open square, B. acapulcensis var. llanensis.
Banisteria maracaybensis var. elliptica Niedenzu, Ind. Lect. Lyc. Brunsb.: 5. 1900.
Banisteria cornifolia var. elliptica (Niedenzu) Niedenzu in A. Engler, Das Pflanzenreich IV, 141:
405. 1928.
Branches pale with prominent pale lenticels. Leaves with the petiole 3-5(-9)
mm long, the lamina 4.7-17.9 cm long, 3.0-8.8 cm wide, elliptic to ovate, with
the reticulum of veinlets prominent adaxially, the smallest areoles 0.3-0.6 mm2.
Inflorescence of condensed axillary panicles, the branches 2.0-4.5 cm long, the
side branches up to 2 cm long and bearing 10-20 flowers. Stamens with sparsely
hairy anther locules, rarely glabrous. Fruit with the nut 6-7 mm tall, 4-5 mm
long, with a winglet on each side, the wing (22-)25-34 mm long and (10-)12-15
mm wide, diverging.
Types. Ple'e s.n., Venezuela, Zulia, Maracaibo, fl fr (lectotype, BR; isolectotypes, F, P, P-JU). Oersted 7 (Pl. Centroamericanae 5362) Costa Rica, in monte
Aguacate, fl (syntypes, C, GOET).
Distribution. (Fig. 5). In lowland savannas of northern Venezuela, extending
into Central America.
Collected in flower in February, March, and June to December, and in fruit in
September to February.
GUATEMALA. Steyermark 38077 bud (F). NICARAGUA. L. 0. Williams & A. Molina R. 42559
fr (F, MICH). COSTA RICA. Oersted 7 (PI. Centroam. 5362) fl (C, GOET); Skutch 4737 fl fr (GH,
NY). VENEZUELA. Zulia: de Bruijn 1265 fl fr (MICH); Karsten s.n. fl fr (W); Plee s.n. fl fr (BR,
F, P). Trujillo: Pittier 10731 fr (G, GH, NY, P, US); 10779 fl fr (G, GH, NY, P, US). Carabobo:
Alston 5941 fr (NY). Aragua: Archer 3005 fr (US); LI. Williams 10304 fl (F, US). Distrito Federal:
Bredemeyer 92 fl (W); Pittier 13036 fl fr (G, GH, M, NY, US); 13452 fl (G, US); Steyermark et al.
102333 fl fr (W). Guarico: Aristeguieta 6146 fl (NY); 6414 fr (US); Curran 170 fl (NY); Davidse 3738
fr (MICH); Pittier 12234 (G, GH, M, NY, US). Anzoategui: Pittier 15119 fl (G, US); F. D. Smith 265
fl (US). No other locality: Eggers 13547 fl (C).
44
Flora Neotropica
This variety is distinguishedby its short petioles, large ovate to elliptic leaves,
and large fruits with a well-developed lateral winglet on the nut. The minute
reticulationof the veins on the adaxial leaf surface is also quite distinctive, and
exhibited by all the collections from Venezuela. It is interestingthat collections
of Banisteriopsis acapulcensis from northernColombia, which have been segregatedin the past as B. dugandii, show this same reticulationwhich is not shown
by the rest of the species. I have includedmost collections from CentralAmerica
in var. elliptica since they resemble this variety in all charactersexcept the reticulation.
Ic. Banisteriopsiscornifoliavar. standleyiB. Gates, var. nov.
Rami rufo-brunnei,lenticellis eburneis, prominentibus.Folia petiolo 5-7 mm
longo, lamina 5.0-12.0 cm longa, 1.8-3.8 cm lata, lanceolata ellipticave, reticulo
venulorum adaxialiterprominenti, areolis minimis 0.5-1.0 mm2. Inflorescentia
floribus in paniculis axillaribus brevibus, usque 3 cm longis, dispositis, ramis
lateralibususque 1 cm longis et usque 8 flores gerentibus. Stamina loculis antherarumsparsim pubescentibus. Fructus nuce 7-8 mm alta, 4-5 mm lata, laevissima, ala 26-29 mm longa, 10-13 mm lata, divergenti.
Type. P. C. Standley 88835, Guatemala,Retalhuleu,vicinity of Retalhuleu,fl
fr (holotype, F).
Distribution.(Fig. 5). Known only from 2 collections from Retalhuleu, Guatemala.
Collected in flower and fruit in February.
GUATEMALA. Standley 87795 fl (F); 88835 fl fr (F).
This variety is distinguishedfrom the other two varieties of Banisteriopsis

cornifolia by its inflorescences with short, few-flowered side branches, and its
smooth fruit nut.
It is named for Paul C. Standley, the most productivefloristicianof the plants
of CentralAmerica and Mexico.
2. Banisteriopsis acapulcensis (Rose) Small, North Amer. Flora 25: 132.
1910.
Fig. 6.
Heteropterysacapulcensis Rose, Contrib.US Nat. Herb. 5: 139. 1897.
Banisteria maracaybensis Adr. Jussieu var. angustifolia Grisebach, Vidensk. Meddel. Dansk.
Naturhist. Foren. Kj6benhavn: 46. 1854. Type. Oersted 31 (PI. Centroamericanae 5363),
Colombia,Antioquia,fl fr (holotype, C).
Banisteria guatemalensis Niedenzu, Ind. Lect. Lyc. Brunsb.: 6. 1900. Type. Seler 2428, Gua-
temala, Salama,Llano grande,fr (holotype, B destroyed, NY fragmentex B).
Banisteria maracaybensis var. leiocarpa Niedenzu, Ind. Lect. Lyc. Brunsb.: 5. 1900. Types.
Palmer 219, Mexico, Guerrero,near Acapulco, fl fr (lectotype, NY fragmentex B; isolec-
totypes, MO, NY). J. D. Smith 4121 (leg Heyde & Lux), Guatemala, Santa Rosa, Naranjo,
fl (syntypes, F, MO, NY).

Banisteriopsisguatemalensis(Niedenzu)Robinsonin Small, North Amer. Flora 25: 132. 1910.
Banisteria cornifolia var. leiocarpa (Niedenzu) Niedenzu in A. Engler, Das PflanzenreichIV,
141: 405. 1928.
Banisteriopsis cornifolia var. leiocarpa (Niedenzu) Cuatrecasas, Webbia 13: 496. 1958.
Banisteriopsis dugandii Cuatrecasas, Webbia 13: 489. 1958. Type. A. Dugand & R. Jaramillo
3335, Colombia, Bolivar, Cartagena, alrededores de Turbaco y Arjona, fl (holotype, US).
Shrub, sometimes vining and formingthickets, branches pale- to dark-brown

with pale lenticels, appressed-sericeousto glabrate. Stipules up to 1 mm long,
triangular,joined by an interpetiolarline. Leaves with the petiole 2-10 mm long,
appressed-sericeousto glabrous, eglandular,the lamina 4.3-13.7 cm long, 1.5-
45
8.0 cm wide, lanceolate to elliptic, cuneate, rarely truncate, at the base, obtuse
to acuminate at the apex, with the marginrevolute and bearing minute glands,
the gland on each side of the midribnearest the base enlarged,adaxiallysparsely
tomentose, soon glabrate, abaxiallyat first densely and loosely tomentose, soon
glabrateor persistently sparsely tomentose, the hairs flexuous with the trabecula
0.5-1.0 mm long, with the reticulationimpressedor prominentadaxially,and the
main lateral veins prominent abaxially. Inflorescence paniculate, axillary, the
branches up to 20 cm long, appressed-sericeousor tomento-sericeous, the side
branches bearing 10-25 flowers; bracts and bracteoles 0.8-1.5 mm long, triangular, appressed-sericeousabaxially, involucrate, persistent; pedicels sessile or
borne on a peduncle up to 1 mm long, 5-10 mm long, 0.4-1.0 mm in diameter,
appressed- or tomento-sericeous. Sepals 1.2-2.0 mm long, 0.6-1.3 mm wide,
oblong, roundedat the apex, sericeous abaxially,glabrousadaxially,the 4 lateral
sepals biglandularand projecting0.8-1.2 mm beyond the glands, the glands 1.82.8 mm long, 1.0-1.4 mm wide. Petals yellow, glabrous, the 4 lateral petals reflexed between the petals, the claw 0.6-1.0 mm long, the limb 4.0-7.0 mm long,
2.0-5.0 mm wide, obovate, entire, slightly concave, the posteriorpetal with the
claw suberect, 1.0-1.8 mm long, only a little longer and thickerthan those of the
lateralpetals, the limb 3.5-6.5 mm long, 2.0-3.5 mm wide, obovate to pandurate,
entire with 2-3 glandularteeth at the base on each side directed anteriorly.Stamens with the filamentserect, glabrous,connate basally, 1.6-2.5 mm long, those
opposite the sepals subequal, slightly longer than those opposite the petals; anthers reflexed, the locules 1.0-1.4 mm long, sparsely hairy or glabrous, the connectives 0.8-1.2 mm long, 0.4-0.7 mm wide, 0.2-0.6 mm deep, oblong, those
opposite the sepals a little wider and deeperthan those opposite the petals. Ovary
1.0-1.5 mm tall, densely sericeous, the styles stout and equal, 1.4-1.8 mm long,
straight and parallel or divergingapically. Fruit of 3 samaras with short broad
carpophoreup to 1 mm long and 1 mm wide, the nut 6-8 mm tall and 4-5 mm
long, appressed-sericeousto glabrate,smooth or bearinglaterallya ridgeor winglet up to 8 mm along its base and projectingup to 2 mm from the nut, the wing
19-36 mm long, 9-16 mm wide, appressed-sericeousto glabrate,with an appendage along its upper marginat the base 3-4 mm long and 1-2 mm tall.
Key to the Varieties of Banisteriopsis acapulcensis
1. Petiole (3-)4-10 mm long; fruitnut smoothor with a ridgeor wingleton each side, the wing
19-30(-36) mm long and 9-14(-17) mm wide, oblong or narrowerat the base.
a. var. acapulcensis.
1. Petiole 2-4(-8) mm long; fruitnut with a well-developedwingleton each side, the wing 28b. var. Ilanensis.
36 mm long and 13-17 mm wide, the lower marginconvex.
2a. Banisteriopsisacapulcensisvar. acapulcensis

Brancheslight- or dark-brownwith pale lenticels. Leaves with the petiole slender, (3-)4-10 mm long, the laminalanceolate to elliptic, cuneate to obtuse at the
base, abaxially very soon glabrate or sparsely and persistently tomentose, the
adaxial leaf surface plane with the reticulationof veinlets prominent. Inflorescence appressed-sericeous.Fruitnut 6-8 mm tall and 4-5 mm long, laterallyquite
smooth or bearingon one or both sides a ridge or winglet up to 6 mm along the
base and projecting 1-2 mm from the nut, the wing 19-30(-36) mm long,
9-14(-17) mm wide, parallel-sidedthroughoutor narrowerat the base.
Type. E. Palmer 219, Mexico, Guerrero,near Acapulco, fl fr (holotype, US
n.v.; isotypes, MO, NY).
46
Flora Neotropica
Distribution.(Fig. 5). From southernMexico southwardthroughCentralAmerica to northernColombia.

Collected in flower and in fruit in July to March.
Representativecollections. MEXICO.Guerrero:Palmer 219 fl fr (MO, NY). Oaxaca:Alexander
187 fl (MICH, NY); Conzatti 3725 fr (US). GUATEMALA. Kellerman 7849 fr (F); Ruana 1341 fl
(F); Seler 2428 fr (NY); J. D. Smith 4121 (leg Heyde & Lux) fl (F, MO, NY); Standley 60492 fl (F);
75778 fl yng fr (US); 77122 yng fr (F); 87532 fl (F, US); 89073 fl fr (F, US); Steyermark 31646 fl (F);
31993 fr (F). HONDURAS. A. Molina R. 694 fr (F, GH); 3909 fr (F, GH); 13217 fl fr (F, NY);
Niederlein 165 fr (NY); Rodriguez 893 fl (F); 3249 fl (F); Standley 13888 fl (F); 28725 fr (F); L. O.
Williams & A. Molina R. 10878 fl (F, GH, MICH); 11901 fr (F, GH, MICH). EL SALVADOR.
Calderon 224 fl (NY, US); Cox & Geezmdn 1147 fl yng fr (ECON); J. D. Smith 8981 (leg Velasco) fl
(F, US); Standley 19447 fl (MO, NY, US); 19691 fr (MO, NY, US); 19908 fl (MO, NY, US); 20949
fr (NY, US); 21339 fr (NY, US). COSTARICA. Almeda3983 fr (MICH);4207 fl fr (MICH);Pittier

(leg Tonduz) 13880 fl fr (F, G, GH). COLOMBIA. Atlantico: Dugand 74 fl (F, US); 5331 fr (US, W);
5784 fl fr (NY, US); Dugand & Jaramillo 2773 fr (US); 3443 fl (US); Magdalena: C. Allen 125 fl
(MO); 787 fl fr (MO); Haught 2247 fl (F, GH, NY, US). Bolivar: Dugand & Jaramillo 3335 fl (US);
Killip & Smith 14506 fr (NY, S, US). Antioquia: Oersted 31 (PI. Centroam. 5363) fl fr (C).
Common names. Guatemala: "gutute." Honduras: "ala de cucaracha." Colombia: "carasucio."
This is a very variable species in its leaf size and shape, abaxial leaf pubescence, presence of a lateral winglet on the nut and size of the fruit wing. The
type of Banisteriopsisacapulcensis is from the northernend of the species range,
and appears very differentfrom the specimens from Colombiawhich have been
segregatedin the past as B. dugandii. The type of B. acapulcensis has lanceolate,
glabrous leaves, with a smooth fruit nut, whereas B. dugandii has elliptic, persistently tomentose leaves, and a fruit nut with lateral winglets. However, these
characters vary throughoutthe range of the species. Many specimens from the
northernpart of the range have elliptic leaves which are glabrateor persistently
tomentose, and although most specimens from the northernpart of the range
have a smooth nut, Seler 2428 from Guatemalaand other specimens from El
Salvador and Costa Rica have a lateral ridge on the nut. Thus, although the
Colombianindividualsexhibit a more or less well developed lateralridgeor winglet on the nut, I am not maintainingthem as a separate taxon as there is no
discontinuityin the variation.
It is interestingthat specimens of Banisteriopsisacapulcensis from Colombia
show minute, prominentreticulationon the adaxialleaf surface, and lateralwinglets on the nut, and that specimens of B. cornifolia var. elliptica from adjacent
Venezuela show these same characters. In these characters, these plants approach each other and are different from other individuals of their respective
species in other parts of their geographicranges.
2b. Banisteriopsisacapulcensisvar. IlanensisB. Gates, var. nov.
Fig. 6.
Rami brunnei, lenticellis fuscis. Folia petiolo 2-4(-8) mm longo, lamina elliptica, basi truncata, abaxialiterdense et pertinacitertomentosa, adaxialiterirregulariterrugata,reticulovenulorumparumimpresso. Inflorescentiatomento-sericea. Fructus nuce 7-9 mm alta, 5-6 mm longa, lateraliterutrinquealula usque 8
mm secus basim et ab nuce 1-2 mm projecta, ala 28-36 mm longa, 13-17 mm
lata, margineinferioriconvexo.
Type. A. Gentry 10294, Venezuela, Guarico, 1 km E of Calabozo on road to
Represa, fl fr (holotype, MICH;isotype, MO).
Distribution.(Fig. 5). In the llanos of Guaricoand Cojedes, Venezuela.
Collected in flower and in fruit in Septemberand November to March.
47
..'
--1-
~.~
FIG. 6 Baniteriosis
mgaptea,
B.acapuensisvar.
~E
1anenis, Blong p
ilifra anB. bevipe
FIG. 6. Banisteriopsis megaptera, B. acapulensis var. llanensis, B. longipilifera and B. brevipe-
dicellata. A-E, B. megaptera. A, inflorescencebranch with buds x 2; B, flower x 4; C, androecium x 7.5 (the stamen at the left is opposite the anteriorsepal); D, gynoecium x 7.5; E, fruit x 1.
F, B. acapulcensis var. Ilanensis, fruit x 1. G, B. longipilifera, fruit x 1. H, J, B. brevipedicellata.
H, bud x 5; J, old flower to show styles x 5. (A-E, Anderson 11734; F, Gates 303; G, Krapovickas
32897; H, Krukoff8758.)
Krukoff 8758.)
& Schinini 32897'
Flora Neotropica
48
VENEZUELA. Cojedes:Delascio et al. 3400 fl (VEN); 4688 fl (VEN); 5087 fl fr (VEN). Guarico:
Aristeguieta 4097 fl (NY, U); 4894 fl (US); Davidse 4276 fl yng fr (MICH); Gates 303 fl fr (MICH);
307 fl fr (MICH); Gentry 10294 fl fr (MICH); Pittier 11470 fl (G, GH, NY, US); 12528 fl fr (F, G,
GH, M, MO, NY, US); 14741fl (US); 14943fl fr (US).
This variety is distinguishedby its short petioles, dense, tomentose abaxialleaf

pubescence, irregularly-wrinkledleaves and large fruits. It occurs only in the
llanos of Guaricoand Cojedes, Venezuela.
3. Banisteriopsisferruginea(Cavanilles)B. Gates, comb. nov.
Banisteriaferruginea Cavanilles,Nona Diss. Bot. 3: 424. 1790.
Triopterys brasiliensis Poiret, Encycl. Bot. Dict. 8: 106. 1808. Type. Commerson s.n., Brazil, fl
fr (holotype, P-LAM;isotypes, C, P, R).

Banisteriaunialata Vellozo, Flora Fluminensis:190. 1825[1829].Tab 157. Type unknown.
Tetrapterys acutifolia var. ferruginea Grisebach, Linnaea 13: 236. 1839. Type. Sellow s.n., Bra-
zil, Rio de Janeiro,Serrad'Estrella,fl (holotype, B, destroyed, NY, fragmentof holotype).

Tetrapteryscrebifloravar. lundianaGrisebach,Vidensk.Meddel.Dansk. Naturhist.Foren. Kjobenhavn: 142. 1875.Type. Lunds.n., Brazil, fl (holotype, C).
Liana, young branches appressed-sericeous, old branches glabrate, brown,

with numberousprominentlenticels. Stipulesminute,up to 1 mm long, triangular,
connected by a prominentinterpetiolarline. Leaves with the petiole 6-10 mm
long, sparsely appressed-sericeous, eglandular, the lamina 4.0-8.5(-11.0) cm
long, 1.6-4.5 cm wide, smallerin the inflorescence,lanceolate to narrowlyovate,
obtuse to truncate at the base, acute to acuminateat the apex, with the margin
plane and bearing numerous small glands, the basal gland on each side of the
midribenlarged,adaxiallyat first appressed-sericeous,very soon glabrate,shiny,
abaxiallysparselyappressed-sericeousto glabrate,the trabecula0.5-0.8 mm long,
with the reticulationprominentadaxially and the lateral veins prominentabaxially. Inflorescencepaniculate,axillaryor terminal,the branchesup to 6 cm long,
loosely brown-sericeous,the ultimatebranchesbearing16-45 flowers;bracts and
bracteoles 1.2-1.8 mm long, triangular,carinate, acute at the apex, appressed,
persistent; pedicels sessile or short-pedunculate,7-10 mm long, 0.4-0.5 mm in
diameter,thickerin fruit, sparsely appressed-sericeous.Sepals 1.4-2.0 mm long,
0.6-1.0 mm wide, oblong, roundedat the apex, sparsely sericeous on both sides
and with a fringe of hairs along the margin,the 4 lateral sepals biglandular,projecting 1.0-1.4 mm beyond the glands, the glands 1.6-3.0 mm long, 0.8-1.2 mm
wide, decurrent on the pedicel posteriorly. Petals yellow, entire or irregularly
denticulate, glabrous, the 4 lateral petals reflexed between the sepals, the claw
0.5-1.0 mm long, the limb 4.0-5.0 mm long, 2.5-3.0 mm wide, elliptic, plane or
slightly concave, shallowly keeled abaxially, the posterior petal with the claw
erect, 1.4-1.8 mm long, a little longer and thicker than the other claws, the limb
4.0-4.5 mm long, 2.0-2.5 mm wide, obovate to pandurate, reflexed, with 3-4
pairs of glandularteeth at the base directedanteriorly.Stamenswith the filaments
erect, glabrous, 1.4-2.2 mm long, those opposite the sepals a little longer than
those opposite the petals; anthers reflexed, locules 1.0-1.2 mm long, sparsely
hairy to glabrate, the connectives 0.8-1.0 mm long, 0.4-0.6 mm wide, 0.2-0.5
mm deep, oblong, those opposite the sepals a little wider and deeper than those
opposite the petals. Ovary 1.0-1.2 mm tall, hairy, the styles stout and equal, 1.21.6 mm long, divergingapically. Fruit of 3 samaraswith the carpophoreup to 1
mm long and wide, the nut 5-6 mm tall and 4-6 mm long, smooth, appressedsericeous, bearing on each side a winglet 5-7 mm along its base and projecting
2-5 mm from the nut, the wing 19-36 mm long, 11-14 mm wide, embracingthe
49
nut at its base, oblong or narrowerdistally, at first densely appressed-sericeous,

soon glabrate, the hairs with the trabecula0.2-0.4 mm long.
Type. Commersons.n., Brazil, Rio de Janeiro, near Sebastianopolis,fl fr (holotype, MA n.v.; isotypes C, P, P-JU, P-LAM, R).
Distribution.(Fig. 7). In the Atlanticcoastal forest of Rio de Janeiro,Sao Paulo
and Parana, Brazil, apparentlyextending northwardto Bahia.
Collected in flower and in fruit in May to October and December.
BRAZIL. Bahia: Belem & Pinheiro 2678 fl (IAN, NY, RB, UB). Rio de Janeiro: Beyrich s.n. fl
(P); Casaretto 1408 fl (G); Commerson s.n. fl (C, P, R); Glaziou 1325 fl (BR, P); 6489 fl fr (C, G,
GOET, P, R); 8588 fl fr (C, G, NY, P, R); Jesus 1992 fr (RB); M. Lima s.n. fr (R); Netto 90 fr (R);
Pabst 4507 fl fr (MICH); Riedel 1036 fl (C, P, US, W); Schwacke 5594 fr (NY, RB); Sellow I it. s.n.
(NY); Sucre 9599 fl (MICH, RB); Vauthier 4 fl (P); 8 fl (G). Sao Paulo: Bowie & Cunningham 522
fr (BM); Burchell 3105 fr (K). Parana: Miiller 14 fl (P); Schwacke II, 14 fl fr (P). No other locality:
Lund s.n. fl (C); Meyer s.n. fl (SP); Nadeaud s.n. fl (P).
This species is distinguishedfromBanisteriopsiscornifoliaby its smallerleaves

which are obtuse to truncateat the base and abaxiallyglabrate,its inflorescence
side brancheswith many (16-45) flowers, and its fruitwith a large, well-developed
winglet on each side of the nut and the wing narrowerdistally. I am including
Belem & Pinheiro2678 in this species althoughit differsin several respects from
typical B. ferruginea: the leaves are elliptic to ovate, cuneate to obtuse at the
base with persistent abaxial pubescence; the inflorescence side branches bear
fewer flowers, up to 12(-16); the styles are longer (up to 2.5 mm) and more
slender. In other respects, this specimen is like B. ferruginea and I prefer to
maintainit as part of the variationof this species until more and fruitingmaterial
becomes available, when the decision can be re-evaluated.
4. BanisteriopsismegapteraB. Gates, sp. nov.
Fig. 6.
Banisteria ovata Niedenzu, non Cavanilles, Ind. Lect. Lyc. Brunsb.: 7. 1900. Type. Glaziou
12492, Brazil, Minas Gerais, between Caraga and Araguai, fl fr (holotype, B destroyed, NY
fragment ex B; isotypes, BR, C, G, NY, P, S).
Banisteria ovata var. subleiocarpa Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 407. 1928.
Type. Ule 7071, Brazil, Bahia, caatinga near Tamburi, fl fr (holotype, B destroyed?; isotype, G).
Frutex usque 3 m altus, rami saepe volubiles ad apicem, brunnei, lenticellis

pallidis. Stipulae minutae, triangulares,jugo interpetiolarijunctae. Folia petiolo
(7-)10-20 mm longo, sparsimsericeo vel glabrato,ad apicembiglanduloso,lamina
4.5-11.7 cm longa, 3.1-6.9 cm lata, ovata orbicularive,basi obtusa truncatave,
apice obtusa apiculatave, marginerevoluta et glandibus minutis instructa, pari
prope costam ad basim aucto, adaxialiterglabrata,abaxialitersparsim apressosericea, pilis trabecularecta 0.2-0.5 mm longa, reticulo prominentiadaxialiter,
venis prominentibusabaxialiter.Inflorescentiaaxillarisvel terminans,paniculata,
usque 20 cm longa, tomento-sericea,demumglabrata,ramis lateralibususque 20
flores gerentibus; bracteis bracteolisque0.5-0.8 mm longis, triangularibus,glabris, involucratis,persistentibus;pedicello sessili vel pedunculousque 2 mm longo, 7-12 mm longo, 0.5-0.8 mm diametro,in fructu crassiore, brunneo-sericeo.
Sepala 5, 1.5-2.5 mm longa, 0.9-1.0 mm lata, oblonga, apice rotundata,abaxialiter sparsim appresso-sericea, adaxialiterglabra, 4 lateralia 2 glandes 1.5-2.5
mm longa et 0.8-1.4 mm lata gerentiaet 1.0-1.5 mm superantia.Petala5, vitellina,
glabra,margineintegro, 4 lateraliainter sepalapatentia,ungue 0.8-1.2 mm longo,
limbo 5.0-6.0 mm longo, 3.0-4.5 mm lato, elliptico, parum concavo, abaxialiter parumcarinato,petalumposticumungue erecto usque 1.4 mm longo, quam
50
Flora Neotropica
ceteris parumcrassioreet longiore,limbo4.5-5.5 mm longo 3.0 mm lato, obovato,

basi utrinque 1-2 dentibusglandulatismunito. Staminafilamentissubaequalibus,
illis sepalis oppositis quam petalis oppositis parumlongioribus, 1.4-2.2 mm longis, basi connatis; antheraereflexae, loculis 1.0-1.2 mm longis, pilosis, connectivis 0.8-1.0 mm longis, 0.5-0.7 mm latis, 0.3-0.4 mm profundis, oblongis, illis
sepalis oppositis quamillis petalis oppositis parummajoribus,eglandulosis.Ovarium 1.0-1.4 mm altum, candido-sericeum,styli aequales, 1.6-1.8 mm longi, crassi, paralleli vel ad apicem divergentes, stigmata truncata. Fructus ex 3 samaris
(vel abortu 1-2) constans; samara carpophorousque 1 mm longo, 2 mm lato,
nuce ovoidea, 9-11 mm alta, 7-9 mm longa, laevi vel rugulosa, dense appressosericea, pilis trabeculaflexuosa usque 0.5 mm longa, ala (25-)30-53 mm longa,
(11-)14-23 mm lata, dense appresso-sericea,basi marginissuperiorisappendicula
6-9 mm longa et 1-2 mm alta munita.
Type. W.R. Anderson11734, Brazil, MinasGerais,43 km by road S of Itaobim
on BR-116, fl fr (holotype, MBM; isotype, MICH).
Distribution.(Fig. 7). Apparentlyin salt-richsoils in eastern Brazil.
Collected in flower in January, March, May, October and December, and in
fruit in Marchand May.
BRAZIL. Ceara: Allemdo s.n. fr (P, R). Bahia: T. S. dos Santos 1318 fl (MICH); Ule 7071 fl fr
(G). Minas Gerais: Anderson 11734 fl fr (MBM, MICH); 11735 fl fr (MBM, MICH); Glaziou 12492
fl fr (BR, C, G, NY, P, S); 13607 fl (BR, C, G, IAN, NY, P); 13615 fl (BR, C, G, NY, P); Magalhdes
15304 fl fr (RB, US); 15434 fl (HB, IAN). Rio de Janeiro: Fromm 1325/E. Santos 1347/Trinta 251 fl
(MICH).
This species is distinguishedfrom Banisteriopsiscornifoliaby its long petioles,

inflorescence with many-floweredside branches, and large fruit; it can also be
distinguishedfrom B. ferruginea by its long petioles and large fruit, and in addition, by its leaf shape and persistent leaf pubescence. Two fruits of Glaziou
12492, one on the Brussels sheet and one visible on the photo of the Berlin
specimen, have a lateral winglet on the fruit up to eight mm long and two mm
high. No other specimens exhibit lateral winglets, and since the presence or
absence of lateral winglets seems to be a variable characterthroughoutthe B.
cornifolia group, I am not accordingit taxonomic significance.
The name refers to the size of the fruit wing, which is large for the subgenus
Banisteriopsis.
5. BanisteriopsisbrevipedicellataB. Gates, sp. nov.

Fig. 6.
Liana, ramijuniores appresso-sericei,vetustioresglabrati.Stipulaeminutaevel
nullae. Folia petiolo 7-15 mm longo, appresso-aureo-sericeo,eglanduloso,lamina
5.6-15.4 cm longa, 2.2-8.6 cm lata, elliptica, basi obtusa truncatave, revoluta
glandulisminimis instructa, pari prope costam ad basim majore, adaxialiterglabra, abaxialitersparsimappresso-sericea,pilis rectis trabecula0.2-0.4 mm longa,
venis densius aureo-sericeis, venis venulisque adaxialiterprominentibus,abaxialiter venis prominentibus.Inflorescentiaaxillaris, rami usque 5 cm longi, appresso-sericei, interdum2-3 in axilla, 25-45 flores gerentes, floribusin spira dispositis; bracteis bracteolisque 0.5-1.5 mm longis, triangularibus,abaxialiter
appresso-sericeis,involucratis,persistentibus,pedicellis sessilibus vel pedunculo
0.5 mm, 2-5 mm longis, 0.6 mm diametro,appresso-sericeis.Sepala 5, usque 1.0
mm longa, 0.7-0.8 mm lata, oblonga, apice rotundata,abaxialitersparsimsericea,
adaxialiterglabra, 4 lateralia 2 glandes 1.5-1.8 mm longas et 0.8-1.0 mm latas
gerentia, 0.5-0.8 mm superantia.Petala 5, lutea, glabra, denticulatavel integra,
inter sepala patentia, petalum posticum non multo dissimile, ungue 0.5-1.0 mm
51
TROPICAL
AMEfiVCA____
?
M?kr
..
.
..-'
:,,?-.?
--
br
...............
FIG. 7. Distribution of part of subgenus Banisteriopsis, species 3-8. Open square, B. ferruginea;
closed triangle, B. megaptera; closed square, B. brevipedicellata; open circle, B. wurdackii; closed
circle, B. cinerascens; open triangle, B. longipilifera.
longo, limbo 4.0-5.0 mm longo, 3.0-3.5 mm lato, elliptico obovatove, abaxialiter

parum carinato, eglanduloso. Stamina 10, filamentis subaequalibus, 1.5-1.8 mm
longis, basi connatis, crassis; antherae reflexae, loculis 1.5-1.6 mm longis, pilosis,
connectivis 1.3-1.4 mm longis, 0.6-0.7 mm latis, 0.3-0.4 mm profundis, subaequalibus, oblongis. Ovarium 0.8 mm altum, carpellis 3, uniovulatis, liberis, toro
pyramidali adnatis, candido-sericeum; styli 3, aequales, 1.6-2.0 mm longi, crassi,
divergentes ab basi, basi candido-sericei, stigmata truncata, parum complanata.
Fructus non suppetunt.

Type. B. A. Krukoff8758, Brazil, Amazonas, Mun. Sao Paulo de Olivenga,
basin of creek Belem (Rio Solimoes), fl (holotype, MO; isotypes, A, MICH, NY).
Distribution. (Fig. 7). Known only from tropical rain forest on terra firme on
the Rio Solimoes in Amazonian Brazil.
Collected in flower in October to December.

BRAZIL. Amazonas: Krukoff8758 fl (A, MICH,
MMO,
fl (GH, MICH, MO, NY).
fO(GH,
NY); 8779
MICH, MO, NY); 8873
Flora Neotropica
52
This species is unique in the Banisteriopsiscornifolia group for its many-flowered inflorescences, the short-pedicellate,small flowers with the posterior petal
scarcely different from the lateral petals, the subequal stamens, and widely diverging styles which are sericeous at the base. It is a large vine in tropical rain
forest and in this respect resembles B. wurdackii;most other species in the B.
cornifolia group are shrubs with a tendency to vine in suitable habitats.
6. BanisteriopsiswurdackiiB. Gates, Brittonia31: 109. 1979.
Fig. 8.
Banisteriacinerascens (Bentham)Grisebachvar. glabrescens Niedenzuin A. Engler,Das Pflanzenreich,IV, 141:406. 1928.Type. Ule 9477, Brazil,Acre, Rio Acr6, SeringalSao Francisco,
fl (holotype, B, destroyed;isotypes, G, MG).
Liana, branches appressed-sericeous,soon glabrate, pale to dark brown with

pale prominentlenticels. Stipules triangular,up to 1.5 mm long, sparsely sericeous to glabrate,joined by a prominentinterpetiolarline. Leaves usually falcate
with the petiole 7-19 mm long, sparsely appressed-sericeous,canaliculate,with
a pair of glands adaxially at the apex, the lamina (5.4-)8.8-21.6 cm long and
(2.6-)4.5-12.7 cm wide, lanceolate to broadly elliptic, acute to obtuse, rarely
cordate, at the base, short- to long-acuminateat the apex, with the marginsomewhat revolute and bearingminuteglands, glabrousadaxially, sparsely appressedsericeous abaxially,the hairs sessile with the trabecula0.2-0.4 mm long, with the
reticulationprominentadaxiallyand the lateral veins prominentabaxially. Inflorescence axillary,paniculate,the side branchesbearing10-20 flowers, appressedsericeous; bracts and bracteoles0.6-1.0 mm long, triangular,appressed-sericeous
abaxially, persistent, involucrate;pedicels subsessile, the peduncle 0.5-2.0 mm
long, 6-12 mm long, 0.3-0.6 mmin diameter,thickerin fruit, appressed-sericeous.
Sepals 1.2-1.6 mm long, 0.6-1.0 mm wide, narrowlyoblong to oblong, rounded
at the apex, sparsely sericeous abaxially, glabrousadaxially, the 4 lateral sepals
biglandularand projecting0.7-1.2 mm beyond the glands, the glands 1.4-2.0 mm
long, 0.6-1.0 mm wide, free and wider at the apex. Petals yellow, glabrous, the
4 lateral petals reflexed between the sepals, with the claw 0.5-0.8 mm long, the
limb 4.5-5.5 mm long and 3.0-4.0 mm wide, elliptic, entire or denticulateat the
base, with the antero-lateralpetals more concave than the postero-lateralpetals,
the posterior petal with the claw erect, 1.5-2.0 mm long, the limb 4.5-5.5 mm
long, 3.0-4.5 mm wide, obovate, short-fimbriate,the basal fimbriaeusually enlarged and gland-tipped.Stamens with the filaments 0.8-2.6 mm long, that opposite the anterior sepal longer, and that opposite the posterior petal shorter,
than the others, the other 8 subequalor those opposite the postero-lateralpetals
slightlylonger, those of the 3 anteriorstamens erect and connate only at the base,
the other 7 connate for up to half their length, those opposite the anterolateral sepals flexuous and directed towards the posteriorpetal; antherswith the
locules glabrous,0.8-1.4 mm long, the connectives of the stamens opposite the 3
anterior sepals 0.8-1.2 mm long, 0.5-0.8 mm wide, 0.3-0.8 mm deep, obovate,
papillateapically,the other7 connectives, 0.4-0.9 mmlong, 0.2-0.6 mmwide, 0.10.4 mm deep, oblong, smooth or papillateapically. Ovary0.8-1.0 mmtall, densely
hairy, the styles equal, 2.0-2.6 mm long, divergingfrom the base, more or less
inflexedtowardsthe axis apically, stigmastruncate.Fruitof 3 samaras,with short
broad carpophoreup to 1 mm long and 1.4 mm wide, the nut 6-9 mm tall, 5-8
mm long, densely appressed-sericeous,the hairs with the trabecula0.4-0.8 mm
long, bearing on each side a single winglet parallel to the areole 4-10 mm long
projecting 2-4 mm from the nut, the wing (20-)24-38 mm long, (8-)10-15 mm
53
~?.~F
_,
r..9
>'N
.
,~~~~~~~~~%..
'~..
F~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~?
"
::,
-:.
e~~~~~~~~~~~~~~~~'
FIG. 8.Banisteriopsis
pe aiax1
wurdacki
i
.
a foeigsotx05(nareeto
~~~
~,
lower
x 5);
b, flower
bud x 3.5;c,flower
x 3.5;
x 3.5;
d posterior (flag) petal
e, part ofandroecium
x 10; f, anther,
stamen
to leftopposite
anterior sepal, stamen
to
right
opposite posteriorpetal,
adaxia
view,
x 10; g, gynoecium,
anterir
left~~~~~~~~I
wit syeiideigtwt
olf,x1;h
neirsyl
frut ith2 amaasdevloedx
075
(Daw
byKain
ouhi;aWrdak
Moacino4076
b-g~~~,
,Wudc&Moaho415.
Pn 44
FIG. 8. Banisteriopsis wurdackii.a, flowering shoot x 0.5 (enlargementof upper lamina x 1,

lower x 5); b, flower bud x 3.5; c, flower x 3.5; d, posterior (flag) petal x 3.5; e,partofandroecium,
stamento left oppositeanteriorsepal, stamento rightoppositeposteriorpetal, x 10;f, anther,adaxial

view, x 10;g, gynoecium,left with anteriorstyle in middle, rightwith anteriorstyle to left, x 10;h,
fruit with 2 samarasdeveloped x 0.75. (Drawnby KarinDouthit;a, Wurdack& Monachino40976;
b-g, Pena 474; h, Wurdack & Monachino 41145.)
54
Flora Neotropica
wide, narrowlyoblong or arcuatedistally, with an appendageon its uppermargin

at the base up to 2.5 mm tall and 5-8 mm along the base.
Type. J. J. Wurdack & J. V. Monachino 41145, Venezuela, Bolivar, Rio Vil-
lacoa (Rio Auyacoa), Raudalito ca. 25 km from river mouth, fl fr (holotype,

MICH;isotypes, NY).
Distribution. (Fig. 7). In tropical rain forest at low elevations from Central
America and northernSouth America, extending throughAmazonia to Bolivia.
Collected in flower in January,June to October, and December, and in fruit in
January,March, April, July, August, November and December.
COSTA RICA. A. Jimenez M. 3788 fr (F, MO); A. Molina R. et al. 18168 fr (F, MO). PANAMA.
P. H. Allen 1129 fl (GH); 2091 fl (F, MO, NY); Foster 1478 fr (DUKE); Gentry & Dwyer 3348 fr (F);
3356 fl (NY); Hayes 770 fl (NY); Shattuck 645 fl (F); Steyermark & Allen 16776 fl (BR, G, MICH,
P, S, U); Wetmore & Abbo 63 fl (F, GH). COLOMBIA. Santander: Gentry et al. 20031 fl (MICH).
Meta: Fosberg 19458 fl (NY, US); Schwabe s.n. fl (COL). VENEZUELA. Bolivar: Wurdack &
Monachino 40976 fl (MICH, NY); 41145 fl fr (MICH, NY). FRENCH GUIANA. Mori & Bolten
8722 fr (MICH). PERU. Loreto: Klug 706 fl (F, NY, US); Plowman et al. 6488 st (F). San Martin:
Woytkowski 7265 fr (MICH). BRAZIL. Acre: Prance et al. 6355 fl (MICH, NY); Ule 9477 fl (G,
MG). Amaz6nas: Fr6es 21228 fl (IAN, US); Pena 474 fl (IAN). Para: Fr6es 30024 fl (IAN, UB);
30061 fl (IAN). Maranhao: Pires & Black 2576 fl (IAN, US). Rond6nia: Cordeiro 556 fr (MICH);
Pires 9963 fl (IAN, NY, UB); Prance et al. 5883 fl yng fr (MG, MICH, NY). Mato Grosso: J. G.
Kuhlmann 441 fr (RB). BOLIVIA. Pando: Prance et al. 5707 fl (MG, MICH, NY); 6273 fr (MG,
MICH, NY); 6483 fl yng fr (MG, MICH, NY).
Commonnames. Brazil: "Lucu-tu-6"(Caripiname).

This species is distinguishedfrom Banisteriopsis cornifolia by its habit as a
large liana, its largerleaves and fruits, and its longer, more slender styles which
are divergingfrom the base. There is considerable variation within the species
with regardto leaf size and shape, and in the time of flowering. Specimens have
been collected in flower in Costa Rica in December and January,in Brazil from
June to October, and in Colombia in December and September. Perhaps additional collections will make this differencein floweringtime disappear,or perhaps
B. wurdackiiis reflectinga differentpeak floweringtime for the rain forests of
Costa Rica and Brazil. There is also some variationin the fruits: Prance 6273
has relatively small fruits, and the specimens from Para, Brazil, in young fruit
apparentlylack a lateralwinglet on the nut. The specimens from Paraare further
unusual in that the reticulationis not prominentadaxially and the leaves may
have been somewhat fleshy when alive, but in all other respects they resemble
other collections of this species.
This species is named for John J. Wurdack,world authorityon the Melastomataceae, whose collections from Bolivar, Venezuela, providedsome of the best
materialof this species.
7. Banisteriopsiscinerascens(Bentham)B. Gates, Brittonia31: 108. 1979.
Heteropterys cinerascens Bentham, Lond. J. Bot. 7: 131. 1848.
Banisteria cinerascens (Bentham) Grisebach, Linnaea 22: 18. 1849.
Banisteriopsis sublucida Rusby, Bull. New York Bot. Gard. 8: 95. 1912. Type. R. S. Williams
102, Bolivia, La Paz, Apolo, fl fr (holotype, NY; isotype, US).
Banisteria cinerascens var. platyphylla Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 406.
1928. Types. J. Steinbach 3260, Bolivia, Santa Cruz, between Piray and Surutii, fl (lectotype,
NY, fragment ex B; isolectotypes, F, G, GH, MO, S). Steinbach 1075, Bolivia, Santa Cruz,
Buenavista, fl (syntypes, GH). Steinbach 7384, Bolivia, Santa Cruz, Buenavista, fl (syntypes,
F, G, GH, GOET, MO, NY, S, U).
Vine or spreadingshrub,branchesdensely appressed-sericeous,eventuallyglabrate, light to dark brown, with prominentpale lenticels. Stipules up to 1.5 mm
55
long, triangular,sericeous abaxially, joined by a prominentinterpetiolarridge.

Leaves with the petiole 4-10(-14) mm long, densely tomento-sericeous, eglandular or apically biglandular,the lamina 3.3-13.9(-17.5) cm long, 1.6-8.0 cm
wide, narrowlyto broadlyelliptic, cuneate to truncateat the base, acute to shortacuminate at the apex, with the marginplane to slightly revolute and bearing
minute glands, the gland nearest the midribon each side at the base enlarged,
adaxially glabrous, abaxially densely to sparsely tomentose, rarely glabrate, the
hairs sessile, the trabecula flexuous up to 1.0 mm long, with the reticulation
prominentadaxially and the lateral veins prominentabaxially. Inflorescenceaxillary, paniculate, the side branches bearing 8-20 flowers, sparsely to densely
tomento-sericeous;bracts and bracteoles 0.6-1.0 mm long, triangular,appressed,
persistent; pedicels sessile or borne on a peduncle up to 3 mm long, 6-11 mm
long, 0.4-0.5 mm in diameter, sparsely tomento-sericeous. Sepals 1.3-1.8 mm
long, 0.8-1.2 mm wide, oblong, roundedat the apex, abaxiallysparsely sericeous,
adaxiallyglabrous,the 4 lateralsepals biglandular,projecting0.8-1.2 mm beyond
the glands, the glands 1.2-1.8 mm long, 0.8-1.0 mm wide. Petals yellow, glabrous,
the 4 lateral petals reflexed between the sepals, the claw 0.5-0.8 mm long, the
limb 5.0-5.5 mm long, 3.0-4.5 mm wide, elliptic, concave, denticulate, the posterior petal with the claw erect, 1.5-2.0 mm long, the limb 4.5-6.0 mm long and
3.0-4.0 mm wide, broadly obovate, short-fimbriate,the basal fimbriae glandtipped. Stamens with the filaments subequal, 1.8-2.4 mm long, those opposite
the antero-lateralpetals and posteriorpetal a little shorterthan the others, those
of the 3 anteriorstamens erect and almost free to the base, the other 7 connate
basally, those opposite the antero-lateralsepals flexuous and directed towards
the posterior petal; anthers with the locules glabrous, 0.9-1.4 mm long, the connectives of the stamens opposite the sepals 0.8-1.1 mm long, 0.5-0.8 mm wide,
0.3-0.6 mm deep, oblong, those opposite the petals 0.6-1.0 mm long, 0.3-0.6 mm
wide, 0.1-0.3 mm deep, narrowlyoblong. Ovary 1.0-1.4 mm tall, densely hairy,
the styles equal, 2.0-2.4 mm long, divergingfrom the base, more or less inflexed
towards the axis apically, stigmas truncate. Fruit of 3 samaraswith short broad
carpophoreup to 1.5 mm long and 1.0 mm wide, the nut 4-5 mm tall and long,
pyriform,densely appressed-sericeous,bearingon each side a single wingletparallel to the areole 5-6 mm along its attachmentand projecting0.5-2.0 mm from
the nut, the wing 20-28 mm long, 8-11 mm wide, narrowly oblong, sparsely
appressed-sericeouswith a shallow appendageon its uppermarginat the base up
to 1 mm tall and 3-4 mm along the base.
Type. Rob. Schomburgk 2nd. Col. 488, Brazil, Roraima, Cotinga/Suruma
rivers, fl (holotype, K, isotypes, F, fragment, G, P, W) = Rich Schomburgk

773 (isotypes, GOET, K, NY, fragmentex B).
Distribution.(Fig. 7). In savannas of Roraima, Brazil, and of Bolivia in the
foothills of the Andes.
Collected in flower in February,April, August, Septemberand December, and
in fruit in February,August and October.
BRAZIL. Roraima: Coradin & Cordeiro 547 fr (MICH); Luetzelburg 20660 fl (M, R); Rob. Schomburgk 488/Rich. Schomburgk 773 fl (F, G, GOET, K, NY, P, W). BOLIVIA. La Paz: Boeke 1443 fr
(MICH); R. S. Williams 102 fl fr (NY, US). Santa Cruz: Cardenas 5891 fr (US); Krapovickas &
Schinini 31763 fl (MICH); Steinbach 1075 fl (GH); 3260 fl (F, G, GH, MO, NY, S); 7384 fl (F, G,
GH, GOET, MO, NY, S, U). No locality: Troll 957 fl (NY fragment ex B).
Commonname. Bolivia: "Abejucado."

The relatively long, slender styles, divergingfrom the base, of this species are
found also only in Banisteriopsis wurdackii of the B. cornifolia group; B. cine-
Flora Neotropica
56
rascens can be distinguishedreadily from B. wurdackiiby its tomentose abaxial

leaf pubescence. The Luetzelburgspecimen has much larger leaves than other
collections of this species, but its leaves nearer the inflorescence are similar in
size to those found in other collections; perhaps this collection samples more of
the variationin leaf size on an individualplant.
8. BanisteriopsislongipiliferaB. Gates, sp. nov.
Fig. 6.
Frutex usque 3 m altus, rami volubiles, appresso-sericei. Stipulae minutae,
triangulares.Folia falcata, petiolo 6-12 mm longo, appresso-sericeo,eglanduloso
vel ad apicem biglanduloso,lamina5.0-13.5 cm longa, 3.8-7.9 cm lata, elliptica,
basi truncatacordatave, apice acuminataapiculatave, margineplana vel parum
revoluta et glandibusminutisinstructa,pari prope costam ad basim aucto, adaxialiterglabra,abaxialitersparsimappresso-sericeavel glabra,pilis rectis trabecula
usque 1.0 mm longa, venis densius aureo-appresso-sericea,venis venulisque
prominulisadaxialiter,venis prominentibusabaxialiter.Inflorescentiapaniculata,
axillaris, ramis appresso-sericeisusque 10 cm longis, lateralibus6-16 flores gerentibus,in spiradispositos;bracteisbracteolisque0.4-0.8 mm longis, triangularibus, involucratis,persistentibus;pedicellis sessilibus vel pedunculo 1.0 mm longo, 5-6 mm longis, 0.4-0.5 mm diametro, sparsim appresso-sericeis. Sepala 5,
1.4-1.6 mm longa, 0.8-1.2 mm lata, ovata, apice rotundata,abaxialitersparsim
sericea, adaxialiterglabra,4 lateralia2 glandes 1.6-2.2 mm longas et 0.6-1.0 mm
latas gerentia, 0.8-1.0 mm superantia.Petala 5, lutea, glabra, denticulata,eglandulosa, suberecta, non inter sepala reflexa, petalumposticum quam 4 lateralibus
non multo dissimile, parumminorius,ungue 0.5-1.0 mm longo, limbo 2.5-4.0 mm
longo, 2.0-3.0 mm lato, elliptico, parumconcavo. Stamina 10, filamentis0.8-1.6
mm longis, basi connatis, crassis, illis sepalis oppositis quamillis petalis oppositis
parumlongioribus;antheraesuberectae, loculis 1.0-1.2 mm longis, glabris, connectivis 0.8-1.0 mm longis, 0.4-0.7 mm latis, 0.2-0.4 mm profundis, oblongis,
illis sepalis oppositis quam illis petalis oppositis parummajoribus.Ovarium0.81.0 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidaliadnatis, candidosericeum; styli 3, aequales, 1.0-1.2 mm longi, crassi, recti parallelique,stigmata
truncata, parum complanata. Fructus ex 3 samaris (vel abortu 1-2) constans;
samaracarpophorousque 1 mmlongo et 2 mmlato, nuce 6-9 mmalta, 5-7 mmlata,
utrinque alula 6-11 mm longa et 1-6 mm alta instructa, sub nuce confluenti,
sparsimappresso-sericea,pilis aureis, trabecularecta 1.5-2.0 mm longa, ala 1730 mm longa, 9-18 mm lata, sparsimappresso-sericeanuce simili, margineinferiore convexo et nuce amplectenti, marginesuperioread basim appendicula4-6
mm longa et 1-2 mm alta munito.
Type. A. Krapovickas & A. Schinini 32897, Brazil, Mato Grosso, 23 km E of
Aquidauana,fl fr (holotype, MICH).

Distribution. (Fig. 7). Known only from cerrado near Aquidauanain Mato
Grosso, Brazil.
Collected in flower and fruit in April and July.
BRAZIL. Mato Grosso: Hatschbach 38613 fl fr (MICH); Krapovickas & Schinini 32895 fl fr
(MICH);32897 fl fr (MICH).
This species is distinguishedby its glabrateleaves, fruit with sparser, longer

hairs and the well-developedlateralwingletson the nut which are confluentbelow
the nut. The name refers to the hairs on the samarawhich are much longer than
those of other species in the subgenus Banisteriopsis.
57
Banisteriopsissubgenus2. Hemiramma(Grisebach)B. Gates, comb. nov., species

9-66.
Banisteria section Hemiramma Grisebach, Linnaea 13: 190. 1839.
Banisteria section Leianthera Grisebach, Linnaea 13: 191. 1839. Lectotype species. Banisteriopsis campestris (Adr. Jussieu) Little as Banisteria campestris.
Banisteria section Trichanthera Grisebach, Linnaea 13: 197. 1839. Lectotype species. Banisteriopsis membranifolia (Adr. Jussieu) Gates as Banisteria membranifolia.
Banisteria section Eubanisteria Grisebach in Martius, Fl. bras. 12: 45. 1858. Lectotype species.
Banisteriopsis campestris (Adr. Jussieu) Little as Banisteria campestris.
Banisteria subgenus Hemiramma (Grisebach) Niedenzu, Ind. Lect. Lyc. Brunsb.: 4. 1900.
Banisteria section Leiococca Niedenzu, Ind. Lect. Lyc. Brunsb.: 7. 1900. Lectotype species.
Banisteriopsis anisandra (Adr. Jussieu) Gates as Banisteria clausseniana.
Banisteria subgenus Eubanisteria (Grisebach) Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1900.
Lectotype species. Banisteriopsis campestris (Adr. Jussieu) Little as Banisteria campestris.
Banisteria section Orthostylis Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1900. Lectotype species.
Banisteriopsis campestris (Adr. Jussieu) Little as Banisteria campestris.
Banisteria subsection Microzeugma Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1900. Lectotype
species. Banisteriopsis argyrophylla (Adr. Jussieu) Gates as Banisteria argyrophylla.
Banisteria subsection Pachyzeugma Niedenzu, Ind. Lect. Lyc. Brunsb.: 14. 1900. Lectotype
species. Banisteriopsis campestris (Adr. Jussieu) Little as Banisteria campestris.
Banisteria section Camptostylis Niedenzu, Ind. Lect. Lyc. Brunsb.: 19. 1900. Lectotype species.
Banisteriopsis membranifolia (Adr. Jussieu) Gates as Banisteria membranifolia.
Banisteria subsection Narcothamnus Niedenzu, Ind. Lect. Lyc. Brunsb.: 20. 1900. Lectotype
species. Banisteriopsis schizoptera (Adr. Jussieu) Gates as Banisteria schizoptera.
Banisteria subsection Cosmiothamnus Niedenzu, Ind. Lect. Lyc. Brunsb.: 22. 1900. Lectotype
species. Banisteriopsis membranifolia (Adr. Jussieu) Gates as Banisteria membranifolia.
Banisteria series Trichothece Niedenzu, Ind. Lect. Lyc. Brunsb.: 22. 1900. Lectotype species.
Banisteriopsis membranifolia (Adr. Jussieu) Gates as Banisteria membranifolia.
Banisteria series Psilothece Niedenzu, Ind. Lect. Lyc. Brunsb.: 45. 1900. Lectotype species.
Banisteriopsis muricata (Cavanilles) Cuatrecasas as Banisteria metallicolor.
Leaf glands marginalor on the lamina,or a single pairat the apex of the petiole,
sessile or stalked. Flowers borne in panicles or cymes of 4-(6-)floweredumbels.
Bracts and bracteoles persistent or caducous, pedicels sessile or short pedunculate, not thicker in fruit. Calyx 8-glandularor eglandular,rarely 4-glandular,
the glands borne on the free part of the sepal. Petals white, pink or yellow, often
changing color with age, denticulate, lacerate or fimbriate.Stamens rarely with
all connectives eglandular,usually at least some of the connectives glandularand
often greatly enlarged, the glandulartissue composed of large polygonal cells.
Styles equal, or the posterior styles differentfrom the anteriorstyle, glabrousor
basally strigose. Ovary short- to long-white-sericeous.Fruit nut laterally compressed or orbicular,smooth, rugose, muricateor rarelywith 1 or more winglets
on each side parallelto the areole, without an appendageor more rarely with a
shallow appendage on its upper edge, nut locule glabrous or densely sericeous
within.
Type species. Banisteriopsis anisandra (Adr. Jussieu) Gates as Banisteria anisandra.
9. Banisteriopsislaevifolia(Adr. Jussieu) B. Gates, comb. nov.
Fig. 9.
Banisteria laevifolia Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 38. 1832 [1833].
Banisteria grata Grisebach in Martius, Fl. bras. 12(1): 49. 1858. Types. Gardner 3061, Brazil,
Goias, Natividade, fl (lectotype, K). Gardner 3625, Brazil, Goias, Serra de Santa Brida, fl
(isosyntypes, K, W). Pohl s.n., Brazil, fl (paratype, M).
Banisteria laevifolia var. grata (Grisebach) Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem. 13. 1900.
Heteropterys micans Skottsberg, Kongl. Svenska Vetenskaps-akad. Handl. 35: 26. 1901. Type.
Lindman 2851, Brazil, Mato Grosso, Santa Cruz da Barra, fl (holotype, S; isotypes, A, BM,
F, G, K, MO, NY, US).
58
Flora Neotropica
Subshrubor shrub, usually with vining tips, or liana to 4 m. Young branches

terete or sometimes slightly flattened, densely white or golden-sericeous, older
branchesglabrate,darkbrown to rufous with numerousprominentlenticels. Stipules minute or absent, triangular,sericeous on both surfaces or apically glabrate
abaxially. Leaves opposite or ternate, the petiole 0-3(-8) mm long, densely sericeous like the stem, eglandular,the lamina (5.0-)6.5-10.5(-13.5) cm long, 2.55.5(-7.5) cm wide, smaller in the inflorescence, rotund to narrowlyelliptic, cordate to truncate at the base, acute, acuminate or apiculate at the apex, flat or
slightly revolute at the margin,bearing 1-4 pairs of peltate glands abaxially on
the lateralveins or sometimes beside the costa basally, the glands especially well
developed on the inflorescence leaves, adaxially sparsely sericeous to glabrate,
abaxiallymore or less densely white-sericeous,with the veins prominulousadaxially and prominent abaxially. Inflorescence terminal, dichasial, composed of
4-(6-)floweredumbels, up to 5 times branched, lax to very dense, white- or
golden-sericeous,the bractsand bracteoles 1.2-2.2 mm long, triangular,adaxially
glabrous, abaxially sericeous, sometimes glabrate apically, persistent; pedicels
sessile, (4-)6-10(-13) mm long, 1.0-1.2 mm wide, 0.8 mm wide without hairs,
white- or golden-sericeous. Sepals golden-sericeous abaxially, minutely velutinous adaxially, caudate, apicallyreflexed, the 4 lateralsepals 3.0-3.5 mm long and
2.2-3.0 mm wide, rotund, the glands green to yellow, 1.6-3.2 mm long, 0.9-1.4
mm wide, the anteriorsepal 2.4-3.2 mm long, 1.0-1.4 mm wide, narrowlyovate,
keeled. Petals yolk-yellow, the claws erect, 0.9-2.0 mm long, 0.5-0.9 mm wide,
the limb (3.0-)4.0-6.5(-8.0) mm long, slightlykeeled externally, crenulateto denticulate, the teeth often gland-tipped,especially proximally, the antero-lateral
petals 5-7 mm wide, orbicular, concave, the postero-lateralpetals 3.0-4.5 mm
wide, elliptic, plane, the posteriorpetal 2.5-4.0 mm wide, elliptic, plane. Stamens
with filaments 1.4-2.0 mm long, those opposite the sepals longer than those opposite the petals, connate for up to one third of their length; anthers reflexed,
with the locules 1.2-1.5 mm long, sparsely to densely sericeous, the connectives
of stamens opposite the petals 0.6-1.0 mm long, 0.4-0.6 mm wide and 0.2-0.5
mm deep, those of stamens opposite the sepals 1.0-1.4 mm long, 0.7-0.8 mm
wide and 0.3-0.5 mm deep, smooth, yellow to buff. Ovary 1.0-1.2 mm high,
white-sericeous, styles flatteneddistally, diverging, 1.8-2.5 mm long, 0.3-0.5 mm
wide at the apex. Fruit with carpophore 1.5-2.0 mm long, 0.5 mm wide, nut 68 mm high and 4.0-5.5 mm long, smooth to tuberculate,olive brown, tomentose,
wing 17-29 mm long, 8-16 mm wide, more or less embracingthe nut, appressedpubescent.
Type. A. Saint Hilaire, Catal. Cl, 170, Brazil, Minas Gerais, Rio das Mortes,
fl (holotype, P; isotype, P).

Distribution.(Fig. 10). Cerradoon deep red soil and marginsof galleryforests,
throughoutthe Planaltoregion of Brazil.
Collected in flower November to May.
Representative collections. BRAZIL. Mato Grosso: Anderson 11235 fl fr (MICH); Hatschbach
34022 fl (MICH); Irwin et al. 16669 fr (MICH, NY); Philcox & Fereira 4446 fl (NY); Ratter et al.
R1065 fr (NY). Goias: Anderson 6504 fl fr (MICH, NY); Barroso et al. s.n. fl (RB); Hatschbach &
Kummrow 38234 fl (MICH); Irwin et al. 21666 fl (MICH, NY); Philcox 3978 fl (NY, RB). Distrito
Federal: Gates 397 fl fr (MBM, MICH); Irwin et al. 11398 fl (MICH, NY, UB); Pires et al. 9014 fl
(UB). Minas Gerais: Assis 204 fl (R, RB, SP); Claussen 65A fl (BR, W); Claussen 543 fl (C, F, NY,
P, S); Glaziou 18163 fl (C, K, P, R); Irwin et al. 26736 fl (NY, UB); Macedo 4163 fl (IAN, MO, RB,
SP); Mexia 5546 fl fr (BM, F, G, GH, K, MICH, MO, NY, S, U, US); Regnell 11.30 pp fl fr (BR, C,
F, K, M, P, R, S, U, US, W); Warming s.n. fl fr (C, GOET, MO, W). Sao Paulo: Chaves 20 fl (GUA,
NY); Eiten 2532 fl (GH, NY, SP); D. A. Lima & Torgo 25 fl (HB); Riedel s.n. fl (M, P, S, U, W);
N. Santos s.n. fl (R). Parana: Dusen 16518 fl (S); Hatschbach 29232 fl (NY, P).
59
I..V
?:
FIG. 9. Banisteriopsis laevifolia and B. argyrophylla. A-F, B. laevifolia. A, flower bud x 3.5;
B, flower x 3.5; C, stamens 1-6, semi-diagrammatic, x 6.3 (stamen 1 is opposite anterior sepal;
stamen 6 is opposite posterior petal); D, lateral view of stamens 1 and 2 x 6.3; E, gynoecium x 6.3;
F, samara x 1.8. G-M, B. argyrophylla. G, flower bud x 3.5; H, flower x 4.2; J, stamens 1-6, semidiagrammatic, x 7 (stamen 1 is opposite anterior sepal; stamen 6 is opposite posterior petal); K,
lateral view of stamens 1 and 2 x 7; L, gynoecium x 8.4; M, samara x 1.8. (A-E, Gates 388; F,
Irwin et al. 16669; G-L, Gates 399; M, Irwin et al. 17806.)
Flora Neotropica
60
, up
WI
+"'
=__
u,
S +_+
__p
__,_
up
'
Sr
-
-,
sr-
'-
'
?'.
FIG. 10. Distribution of Banisteriopsis laevifolia.
Local names. Brazil: "Barboleta do campo," "Cipo prata."

This is a widespread species throughout the Planalto showing considerable
variation in habit, leaf shape and size, petiole length and leaf pubescence. Niedenzu recognized a variety grata, which he distinguished by its scandent habit,
up to 8 pairs of leaf glands, longer pedicels and sparser leaf pubescence. Examination of all material available shows that number of leaf glands and pedicel
length vary throughout the range of the species, and are not correlated with
sparser leaf pubescence. Specimens with sparse leaf pubescence are found in
Goias, Distrito Federal, Mato Grosso and northern Minas Gerais in Brazil; these
plants are usually vines, but throughout its range B. laevifolia is a facultative
vine, apparently vining wherever support is available. Since density of pubescence is a very variable character throughout the range of the species, I do not
feel that recognition of this variety is justified unless study of living populations
provides additional data,
data, such as ecological
separation, to support
support segregation.
provides
ecological separation,
segregation.
61
The affinities of B. laevifolia appear to lie with the B. argyrophyllagroup,

based on similaritiesbetween them in the following characters:position and type
of leaf glands; flowers in four-floweredumbels, the umbels borne in cymes or
dichasia; stamens with short filaments,with the anthersreflexed, subequal, with
pilose locules and eglandularconnectives; short styles which are diverging,and
recurvedand laterallycompressed apically;fruit with a smooth nut and the wing
embracingthe nut. Many of these charactersare also shared with the B. schizoptera group, but in this group, some of the connectives are glandularand the
styles are less divergingand scarcely compressed laterally. However, B. laevifolia differs from both of these groups in its yolk-yellow petals with crenulateto
denticulate margins, which are held suberect with the posterior petal scarcely
differentiatedfrom the four lateralpetals; the subgenusBanisteriopsishas petals
which are yolk yellow, stamens with pilose or glabrous locules and eglandular,
subequal connectives, and the fruit with a smooth nut and the wing embracing
the nut. But the inflorescences are paniculate, with the flowers in pairs and not
in umbels in the subgenus Banisteriopsis, and the leaf glands are marginal.
Banisteriopsis laevifolia thus appears to provide a link between these groups of
species.
The Banisteriopsis argyrophylla group (species 10-14)
This groupof five species consists of one widespreadspecies, B. argyrophylla,

common in the cerrados of the Planaltoregion of Brazil, and four species which
are restrictedin their distribution(Fig. 11). These four species, with the possible
exception of B. harleyi, are shrublets,and grow in rocky campos. Banisteriopsis
harleyi may sometimes be a vine; it shows many xeric features and may replace
B. argyrophyllain the more xeric vegetation of Bahia where it occurs.
The characterssharedby membersof this group are (see Fig. 9 for illustration
of flowerand fruittype): one to several pairs of leaf glands abaxiallyon the lateral
veins; leaves adaxially densely white-tomento-sericeous;sepals rounded at the
apex and appressed in bud; calyx glands green or brown; petals long-fimbriate,
the claw of the fifth petal not constricted;filaments short (up to 2.6 mm), thick,
subequal;antherssubequal,locules pilose, the connectives oblong, not glandular;
styles diverging,flattenedlaterallyand flaringdistally, the anteriorstyle thicker,
0.8-1.0(-1.4) mm in diameter;fruit nut velutinous.
10. Banisteriopsisargyrophylla(Adr. Jussieu) B. Gates, comb. nov.
Figs. 9, 12.
BanisteriaargyrophyllaAdr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 45. 1832[1833].

Banisterianitens Grisebach,Linnaea 13: 198. 1839. Type. Sellow s.n., Brazil, fl (holotype, B,
destroyed;isotype, K).
Shrubto 3 m with vining branches, or vigorous liana. Young branches terete,

densely golden- or silvery-velutinousto subtomentose,older brancheseventually
glabrate, dark brown to rufous, with numeroussmall pale brown punctate lenticels. Stipules 2.0-3.5(-5.0) mm long, linear, velutinous throughoutor sometimes
glabrate apically, appressedon young branches, projecting,with the tip inflexed
and the base thickened, on old branches. Leaves opposite, the petiole 3-10 mm
long, densely golden- or silvery-velutinouslike the stem, eglandular,the lamina
(4.5-)6.0-13.0(-16.0) cm long, (2.5-)3.5-8.0(-10.5) cm wide, smallerin the inflorescence, chartaceous, elliptic or rarely narrowlyelliptic, obtuse to subcordate
at the base, short-acuminateto apiculate at the apex, revolute at the margin,
bearing 2-6 pairs of peltate glands abaxially on the lateral veins, especially on
the inflorescenceleaves, adaxiallyglabrate,abaxiallydensely silvery- or golden-
62
Flora Neotropica
FIG. 11. Distribution of Banisteriopsis argyrophylla group, species 10-14. Closed circle, B. argyrophylla; open circle, B. byssacea; closed square, B. paraguariensis; triangle, B. harleyi; open
square, B. vernoniifolia.
sericeous, the primaryand secondary veins very impressed adaxially and prominent abaxially, the areoles between the veins shallowly inflatedadaxially. Inflovelutinous
and
at the apex,
revolutepairs
at the
adaxially,
appressed
margin,
rescence
dichasial
of rounded
4-flowered
or racemes
of 2-4(-10)
of flowers,
umbels,
terminatingnumerous axillary branches, very lax, densely golden- or silveryvelutinous; bracts and bracteoles 1.0-2.4 mm long, triangular,velutinous abaxially, sometimes glabrateapically, glabrousadaxially, persistent; pedicels sessile,
(6-)9-19 mm long, 1.4-1.6 mm wide, 0.8-1.0 mm wide without the hairs, goldenvelutinous. Sepals golden-sericeous abaxially, sparsely so at the apex, minutely
4 the
lateral sepals
and wide, the glands green, 2.43.4-3.6 mm long
3.2-3.5 mm
3.2 mm long and 1.2-1.8 mm wide, the anterior sepal 2.5-2.8 mm long, 1.6-2.2
mm wide, elliptic, keeled. Petals white, becoming cream in age, not keeled, den-
63
tate, the 4 lateral petals reflexed between the sepals, the claw 2.2-3.3 mm long
and 0.6-0.8 mm wide, the limb 7.5-10.0 mm long, 8.0-9.5 mm wide, orbicular,
eglandular,the anteriorpetals more concave thanthe posteriorones, the posterior
petal with the claw erect, 4.3-5.5 mm long, distally widened and flattened, up to
2.8 mm wide at the apex, the limb 4.5-6.0 mm long and 4.0-5.0 mm wide, broadly
obovate, reflexed, convex, with the basal teeth enlargedand glandular.Stamens
with filaments2.0-2.6 mm long, thick, white, connate for up to two-thirdsof their
length;anthersreflexed,pilose, with the locules 1.5-2.6 mm long, the connectives
0.8-1.8 mm long, 0.6-0.9 mm wide, 0.3-0.6 mm deep, smooth, not glandular,
yellow, oblong. Ovarywhite-sericeo-velutinous,1.2-1.4 mmtall, styles diverging,
flatteneddistally, 2.0-2.2 mm long, the anteriorone flaringdistally to 0.8-1.0 mm
wide, the posterior ones 0.4-0.6 mm wide, only slightlyflared. Fruit with carpophore 2.0-2.2 mm long and 0.5 mm wide, the nut 10-13 mm tall and 6-8 mm long,
smooth to slightly rugose, short golden-brown-velutinous,the wing (20-)25-36
(-40) mm long, (9-)11-18(-23) mm wide, embracingthe nut, sericeous, glabrate
apically, the hairs T-shaped, short-stalked,with flexuous arms.
Chromosome number. n = 10. Voucher: Gates 399.
Type. A. Saint Hilaire, Catal. Bl, 860, Brazil, Minas Gerais, Nossa Senhora
da Conceitao, fl (holotype, P; isotypes, P-2 sheets).

Distribution.(Fig. 11). Cerradoand gallery marginsthroughoutthe Planaltoof
Brazil, extending southwardto Paraguay.
Collected in flower Januaryto May, and in fruit April to June.
Representativecollections. BRAZIL. Para: N. T. Silva 729 fr (IAN). Goias: Anderson 6350 fl
(MICH, NY); Burchell 6946 fl (GOET, K, P); Hatschbach 36672 fl (MICH); Irwin et al. 17806 fr (F,
IAN, MICH,NY, SP, US); Pohl 1514.1528dfl (F, K, NY, W). DistritoFederal:Gates 399 fl (MICH);
Pires et al. 9023 fl (IAN, NY); Irwin et al. 15811 fl fr (MICH, NY). Mato Grosso: Anderson 9680 fl
fr (MICH, NY); Hatschbach 23705 fl (MICH, P); Irwin et al. 16673 fr (MICH, NY); Malme 3328 fr
(G, R, S); R. R. de Santos et al. R1243 fl (MO, NY, RB). Minas Gerais: Barreto 7562 fl (F, US);
Glaziou 18162 fl (C, G, NY, P, R); Hatschbach & Ahumada 31319 fl (MICH, NY); Irwin et al. 26081
fl (MICH, NY, UB); Macedo 1045 fl (S, US); Martius 722 fl (M); Schwacke 7194 (R, RB). Sao Paulo:
Anderson 11142 fl (MICH, NY); Brade 7312 fl (SP); J. C. Gomes 240 fl (RB); Lofgren 569 fr (C,
GH); Riedel s.n. fl (C, K, P, US, W). PARAGUAY. Hassler 10403 fl fr (G, GH, NY); Hassler 10832
fr (G, GH).
Local names. Brazil: Cipo prata.

Banisteriopsisargyrophyllais variableas to leaf size and shape, in the colour
and form of its pubescence and in general habit. In open cerrado, the species is
a shrub with vining tips; in more closed cerrado or at gallery margins, it is a
vigorous vine or liana. There is also variationin the form of the inflorescence;
usually the flowers are borne in terminalor axillary racemes consisting of up to
ten pairs of flowers.
11. BanisteriopsisbyssaceaB. Gates, sp. nov.
Fig. 12.
Frutex usque quidem 7.5 dm altus, non volubilis, ramisjunioribus teretibus,
laxe byssaceo-tomentosis, ramis vetustioribusglabratis,laevibus, teretibus, purpureo-fuscis, lenticellis minutis punctatispallidis dispersis. Stipulae 1.0-2.2 mm
longae, triangulares,penitus tomentosae. Folia opposita vel subopposita;lamina
(5.0-)6.4-8.0(-10.5) cm longa, (2.5-)3.5-5.0 cm lata, vel in inflorescentiaminori,
anguste elliptica vel elliptica, basi truncata, apice brevi-acuminata,apiculata
acutave, margineplana integraque,4-6 glandes peltatas in nervis lateralibussubtus gerenti, suprabyssaceo-tomentosavel glabrata,subtuspertinaciterdense candido-sericea, rugulosa, venis primariis secundariisque supra impressis, subtus
prominentibus;petiolo 3-5 mm longo, laxe cinereo-tomentoso,eglanduloso. In-
64
Flora Neotropica
florescentia floribus in umbellis 4-floriferisvel racemis usque 12-floriferisfasciculatis, umbellis in cymis dispositis, semel vel bis composita, laxa densave, cinereovel aureo-byssacea;bracteis bracteolisque 1.7-3.0 mm longis, lanceolatis
vel triangularibus,abaxialitercinereo-sericeis, adaxialiterglabris, persistentibus;
pedicellis sessilibus 7-16 mm longis, 0.9-1.2 mm diametro,0.5-0.7 mm sine indumento, dense albido-tomentosis.Sepala 5, abaxialitercinereo-sericea, adaxialiter basi minute velutina, apice obtusa introflexaque,4 lateralia 2.8-3.0 mm
longa, 2.4-3.0 mm lata, glandulis 1.4-1.8 mm longis, 0.6-1.0 mm latis, sepalo
antico 2.8-3.0 mm longo, 1.7-2.0 mm lato, elliptico, medio carinato. Petala 5,
alba, longe fimbriata,orbicularia,glabra, 4 lateraliainter sepala patentia, ungue
2.0-2.2 mm longo, 0.6-0.7 mm diametro,limbo 6.0-8.0 mm longo, 6.0-7.0 mm
lato, eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribus et parum
majoribus;petalo postico ungue erecto 3.0 mm longo, distalitercomplanatodilatatoqueusque 0.8 mm diametroversus apicem, limbo 3.5 mm longo, 3.0 mm lato,
recurvato, convexo vel plano, fimbriisbasalibus auctis glandulosisque.Stamina
10, filamenta 1.8-2.2 mm longa, crassa, proximaliterconnata, glabra; antherae
recurvatae, subaequales, oblongae, loculis 1.8-2.0 mm longis, sparsim pilosis,
connectivis 1.2-1.6 mm longis 0.6-1.0 mm latis, 0.3-0.6 mm profundis,laevibus,
eglandulosis, antherarumsepalis oppositarumquam petalis oppositarumparum
majoribus.Ovarium1.8-2.0 mm altum, carpellis 3, uniovulatis,liberis, toro pyramidali adnatis, candido-sericeis; styli 3, aequales, 1.8-2.0 mm longi, divergentes, basaliter sericei, apice complanatidilatatique,stylo antico magis, usque
0.9 mm lato apice, posticis usque 0.4-0.5 mm latis. Fructus ex 3 samaris (vel
abortu 1-2) constans; samara carpophoro2.0-3.0 mm longo, 0.4-0.6 mm lato,
nuce ovoidea vel orbiculari,8-9 mm alta, 6-7 mm longa, laevi vel parumrugosa,
dense aureo-velutina,ala 12-17 mm longa, 8-11 mm lata, appresso-pubescenti,
ab dimidio superiorimarginisabaxialisnucis excrescenti.
Type. J. Vidal 1-778, Brazil, Minas Gerais, Mun. de Thermopolis, Chapadao
de Jacuhy, fl fr (holotype, R 43406).
Distribution.(Fig. 11). This species is known only from southernMinas Gerais
near the border with Sao Paulo. The specimens show no indications of vining,
and the species seems to be a shrublet growing in a rocky campo habitat on
shallow soil.
Collected in flower and fruit in Februaryand April.
BRAZIL. Minas Gerais:W of CamposAltos, Davidse & Ramamoorthy10840fl imm fr (MICH);
Sao Sebastiao do Paraiso, Vidal 1-392 pp fl (R 75755); Pohl 489 fl (W).
This species is distinguishedfrom B. argyrophyllaby its non-vininghabit, its

cobwebby stem pubescence, which also gives the species its name, its shorter
petioles, smallerrugose leaves, and its shorterfruit wing which does not encircle
the nut. Banisteriopsis byssacea is distinguished from B. paraguariensis by its
larger rugose leaves, and the fruit with the nut not rugose and with a well developed wing.
12. BanisteriopsisparaguariensisB. Gates, sp. nov.
Fig. 12.
Fruticulus erectus usque 6 dm, renovationibus annuis ab xylopodio. Ramis
teretibus, cineraceo-sericeis, basi glabratis. Stipulae 2-6 mm longae, lineares,
utrinquesericeae vel abaxialiterapice glabratae.Folia opposita vel ternata;lamina (1.7-)4.5-7.5(-9.3) cm longa, (0.8-)3.0-4.0 cm lata (vel in inflorescentiaminori), elliptica vel late elliptica, basi obtusa subcordatave,apice acuminataapiculatave, margine plana revolutave integraque, 2-6 glandes peltatas in nervis
lateralibussubtusgerenti, supratomentosavel glabrata,subtuspertinaciterdense
65
I ~;..
FIG. 12. Banisteriopsis paraguariensis, B. harleyi, B. byssacea and B. argyrophylla. A-B, B.
paraguariensis. A, habit x 0.25; B, mericarpx 2.5. C-D, B. harleyi. C, portion of branch x 0.75;
D, immaturesamara x 2.3. E-G, B. byssacea. E, portionof shoot x 0.75; F, samara x 2; G, stem
hairs x 30. H, B. argyrophylla, stem hairs x 30. (A, Hassler 9856, 9832a; B, Hassler 9832a; CD, Harley 13875A; E-G, Vidal I-778; H, Anderson 6350.)
66
Flora Neotropica
albido-sericea, venis primariis supra impressis, subtus prominentibus;petiolo

crasso 2-7 mm longo, cineraceo-sericeo, eglanduloso. Inflorescentiafloribusin
umbellis 4-floriferisfasciculatis, umbellis in cymis dispositis, semel vel bis composita, laxa densave, cinereo-sericea;bracteis bracteolisque 1.0-2.0 mm longis,
late triangularibusvel ovatis, abaxialitercineraceo-sericeis, adaxialiterglabris,
persistentibus;pedicellis sessilibus, 10-14 mm longis, 1.0-1.2 mm diametro,0.60.8 mm sine indumento,dense sericeis. Sepala 5, abaxialiteraureo-sericea,sparsim versus apicem, adaxialiterbasi minute velutina, apice obtusa vel rotundata,
introflexa, 4 lateralia 3.4-4.2 mm longa, 2.4-3.2 mm lata, ovata, glandulis 1.62.2 mm longis, 1.2-1.4 mm latis, sepalo antico 3.0-4.0 mm longo, 2.2-3.2 mm
lato, elliptico, medio carinato. Petala 5, alba, longissime fimbriata,glabra, 4 lateraliainter sepalapatentia,ungue 1.4-2.4 mm longo, 0.4-0.7 mm diametro,limbo
7.0-10.0 mm longo latoque, orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribuset parum majoribus,petalo postico ungue erecto, 4.05.0 mm longo, distalitercomplanatodilatatoque,usque 1.4 mm lato apice, limbo
5.0-6.4 mm longo et 4.0-5.0 mm lato, late obovato, recurvato,convexo vel plano,
fimbriisbasalibusauctis et glanduliferis.Stamina 10, filamenta1.2-2.2 mm longa,
crassa, basaliter vel omnino connata, glabra;antheraerecurvatae, subaequales,
oblongae, loculis 1.8-2.2 mm longis, sparsim pilosis vel glabratis, connectivis
1.2-1.6 mm longis, 0.6-0.9 mm latis, 0.4-0.6 mm profundis, laevibus, eglandulosis. Ovarium1.1-1.4 mm altum, carpellis3, uniovulatis,liberis, toro pyramidali
adnatis, candido-sericeis;styli 3, aequales, 2.0-2.6 mm longi, divergentes, basaliter sericei, apice complanatidilatatique,stylo antico magis usque 1.4 mm lato
apice, posticis usque 0.6-0.8 mm latis. Fructus ex 3 mericarpiis(vel abortu 1-2)
constans; mericarpiumsine carpophoro,nuce ovoidea 11-14 mm alta, 7-8 mm
longa, rugosa, velutina, ala dorsali ad cristambrevem 1-2 mm altam redacta.
Type. Hassler 9832a (leg. Rojas), Paraguay,Sierrade Amambay,fr (holotype,
NY; isotypes, A, G, K, P, W).
Distribution.(Fig. 11). Open grassy campos of the Sierra de Amambay,Paraguay, and adjacentBrazil.
Collected in flower in December and in fruit in December and May.
BRAZIL. Mato Grosso: Hatschbach 38669 fr (MICH). PARAGUAY. Hassler 9832 (leg. Rojas)
fl (C, G, GH, K, MICH, MO, NY, P, RB, S, US, W); 9856 fl (G); Rojas 4159 fl (M).
This species is distinguishedfrom B. argyrophyllaby its suffruticosehabit, its

appressedstem pubescence, shorterpetioles, smallerleaves with only the primary
veins impressed, and reduced fruit wing. It is distinguishedfrom B. byssacea
also by its reducedfruitwing and its leaves with only the primaryveins impressed.
13. Banisteriopsisharleyi B. Gates, sp. nov.
Fig. 12.
Frutex erectus usque 1.5 m, interdumscandens, ramisjunioribusteretiusculis,
laxe cineraceis-arachnoideo-tomentosis,ramis vetustioribus demum glabratis,
teretibus, fuscis, lenticellis numerosis prominentibusfuscis. Stipulae 1.0-2.0
(-5.0) mm longae, lineares, omnino cineraceo-tomentosaevel abaxialiterapice
glabratae.Folia opposita; lamina (1.0-)1.7-6.0 cm longa, 1.0-3.0 cm lata (vel in
inflorescentiaminori), late elliptica vel rotunda, basi truncata cordatave, apice
brevi-acuminataapiculatave, marginerevoluta integraque, 2-4 glandes peltatas
in nervis lateralibussubtus gerenti, supra pubescenti mox glabrata, subtus pertinaciter dense candido-sericea, bullata, nitida, venis primariis secundariisque
supra impressis, subtus prominentibus;petiolo crasso 0.5-2.0 mm longo, cineraceo-tomentoso, eglanduloso. Inflorescentiafloribus in umbellis racemisve 4-flo-
67
riferis fasciculatis, umbellis in cymis dispositis semel vel bis composita, laxa
densave, cineracea-arachnoideo-tomentosa;bracteis bracteolisque 1.0-1.8 mm
longis, late triangularibus,abaxialitercineraceo-tomentosis, adaxialiterglabris,
persistentibus;pedicellis sessilibus 8-13 mm longis, 1.0-1.2 mm late, 0.6-0.8 mm
sine indumento, laxe candido-sericeis.Sepala 5, abaxialiteralbo- vel aureo-sericea sparsimversus apicem, adaxialiterbasi minute velutina, apice rotundataintroflexaque,4 lateralia2.8-3.6 mm longa, 3.0-3.5 mm lata, glandulisviridis 2.02.4 mm longis, 1.2-1.4 mm lata, sepalo antico 2.8-3.0 mm longo, 2.2-2.5 mm
lato, late elliptico, medio carinato. Petala 5, alba demum erubescentia, postica
interdumrosea, longissime fimbriata,orbicularia,glabra, 4 lateraliainter sepala
patentia,ungue 2.4-3.0 mm longo, 0.6-0.7 mm diametro,limbo 5.0-8.0 mm longo
latoque, eglanduloso vel interdumfimbriisglanduliferis,2 anterioribusquam 2
posterioribusconcavioribusparum majoribus,petalo postico ungue erecto 4.05.0 mm longo, distalitercomplanatodilatatoque,usque 1.2 mm lato apice, limbo
3.5-5.0 mm longo, 4.0-6.0 mm lato, recurvato,convexo vel plano, fimbriisbasalibus auctis glanduliferisque.Stamina 10, filamenta 0.8-1.8 mm longa, crassa,
proximaliterconnata, glabra;antheraerecurvatae,subaequales,oblongae, loculis
1.4-1.8 mm longis, sparsim pilosis, connectivis 0.8-1.2 mm longis, 0.7-0.9 mm
latis, 0.3-0.6 mm profundis,laevibus, luteis, eglandulosis. Ovarium1.0-1.4 mm
altum, candido-sericeum;styli 3, aequales, 2.0-2.2 mm longi, divergentes, basaliter sericei, apice aliquantumcomplanati,stylo antico magis, usque 0.9 mm lato
apice, posticis usque 0.4-0.5 mm latis. Fructus ex 3 samaris (vel abortu 1-2)
constans; samara immaturasine carpophoro, nuce ovoidea, 9.0-12.0 mm alta,
5.0-7.0 mm longa, laevi, velutina, ala usque 22 mm longa, 15 mm lata, nucem
amplectenti, appresso-pubescenti.
Type. Harley 15875A, Brazil, Bahia, Serrado Sincora, 2 km N of Cascavel on
road to Mucuge, fl fr (holotype, CEPEC;isotype, MICH).
Distribution.(Fig. 11). Open grasslandwith occasional shrubs overlying white
sand and crystalline quartz, alt. 1000-1200m, central Bahia, Brazil.
Collected in flower in Januaryand February,and in young fruit in February.
BRAZIL. Bahia: Serra do Sincora, 6 km N of Barra da Estiva, Harley et al. 15640 fl bud (CEPEC,
MICH); Serra da Agua de Rega, 23 km N of Seabra; Irwin et al. 30879 fl (MICH, NY, UB); 30909A
fl young fr (MICH, NY, UB).
This species is distinguishedfrom B. argyrophyllaby its smaller, sessile, coriaceous, strongly bullate leaves. The fruit wing of B. harleyi is shorter than in
B. argyrophylla,but more, and mature,fruits of B. harleyi are needed for confirmationof this.
Harley 15640 is unusual in many respects. It is a vine, with petioles up to 5
mm long, the leaves are narrowly elliptic up to 9 cm long, and are not very
bullate, sometimes with only the primaryveins impressed adaxially. Since little
materialof this species has been collected, little is known of the variationwithin
the species, and other collections may be made which are intermediatein leaf
size, shape and rugosity.The vininghabitmay indicatethatthis plantwas growing
in a habitatwhere supportwas available.The differencesshown by this collection
may reflect a less xeric habitat, or less recent or less frequentburning.
This species is namedfor RaymondHarley, whose moderncollections in Bahia
include many hithertounknowntaxa.
14. Banisteriopsis vernoniifolia (Adr. Jussieu) B. Gates, comb. nov.
Banisteria vernoniifolia Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 144. 1843.
68
Flora Neotropica
Subshrubwith many unbranchedstems from a common rootstock, or shrubto

2 m tall. Young branchesterete or somewhatflattened, sericeous, older branches
glabrescent, smooth, terete, dark purplish brown, with numerous small, dark
brown, punctatelenticels. Stipules minute,up to 1 mm long, triangular,sericeous
to glabrate. Leaves opposite, sometimes alternateor 3 or 4 at a node, the petiole
1-4(-7) mm long, loosely grey-sericeous like the stem, eglandular,often canaliculate, the lamina(1.5-)3.5-11.0(-16.0) cm long, 0.5-2.0(-3.0) cm wide, smaller
in the inflorescence, linear to narrowlyelliptic or narrowly oblong, truncate to
obtuse at the base, acute to short-acuminateor apiculateat the apex, revolute at
the margin, bearing (0-)1-2(-4) pairs of peltate glands abaxially on the lateral
veins, especially on the inflorescenceleaves, adaxiallyglabrateand shiny, abaxially densely white-sericeous, with the primary veins impressed adaxially and
prominentabaxially. Inflorescencea terminaldichasiumof 4-floweredumbels, 23 times compound,lax to dense, loosely grey-sericeous,the bracts and bracteoles
1.4-2.0 mm long, lanceolate to triangular, sparsely grey-sericeous adaxially,
sometimes glabrate apically, glabrous abaxially, persistent; pedicels sessile,
(6-)8-13 mmlong and0.9-1.2 mmin diameter,0.5-0.7 mmwithoutthe hairs.Sepals
grey-sericeousabaxially,apicallyglabrous,minutelyvelutinousadaxially,rounded and appressed at the apex, the 4 lateral sepals 3.0-3.5 mm long, 2.5-3.5 mm
wide, the glands green, 2.0-3.0 mm long and 1.0-2.4 mm wide, the anteriorsepal
2.5-3.2 mm long and 2.0-2.5 mm wide, elliptic, keeled. Petals white, becoming
creamin age, fimbriate,orbicular,eglandular,the 4 lateralpetals reflexedbetween
the sepals, the claw 2.0-2.8 mm long and 7-9 mm wide, the anteriorpetals more
concave than the posterior ones, the posteriorpetal with the claw erect, 4.2-5.0
mm long, distally flattened and widened to 2 mm at the apex, the limb 4.0-4.6
mm long, 4.0-6.0 mm wide, reflexed, flat to convex. Stamens with filaments 1.62.2 mm long, thick, connate basally, the anthers reflexed, with the locules 1.62.0 mm long, sparselypilose, the connectives 1.0-1.5 mm long, 0.8-1.0 mm wide
and 0.3-0.6 mm deep, smooth, not glandular,oblong, yellow. Ovary 1.0-1.3 mm
tall, white-sericeous, styles flattened,diverging,the anteriorone 1.9-2.5 mm long
and flareddistally to 0.8-1.0 mm wide, the posterior ones 1.8-2.0 mm long, 0.40.6 mm wide, only slightlyflared. Fruitwith carpophore2.0-2.5 mm long and 0.5
mm wide, the nut smooth to slightly rugose, short-velutinous,the hairs goldenbrown, the wing 14-22(-26) mm long, 9-14 mm wide, golden sericeous, the hairs
appressed with flexuous arms, glabratedistally, embracingthe nut, red.
Type. Martiuss.n., Brazil, fl (lectotype, M). There are two sheets at Munich
annotatedas Banisteria vernoniifoliaby Jussieu, one in flower, the other in fruit.
The fruiting sheet is Banisteriopsis angustifolia (Adr. Jussieu) Gates. I have
selected the floweringsheet as lectotype of Banisteria vernoniifolia,since it corresponds more closely to Jussieu's originaldescription. All collections attributable to B. vernoniifoliaare from Goias, and it is probablethat the type is also
fromGoias, since it is the fruitingsheet whichbears the locality "in summaMonte
Itambe, Prov. Minas Geraes" cited in Jussieu's original description.
Distribution. (Fig. 11). Rocky campos at 1000-1700 m in the Chapada dos

Veadeiros, Serra Geral do Paranaand Serra Dourada, Goias, Brazil, on shallow
sandy soil overlying sandstone and quartzite.
Collected in flower December to May, and in fruit, Marchto May.
Representative collections. BRAZIL. Goias: Anderson 7694 fl fr (MICH, NY); Barroso et al. 773
fl (NY, UB); Burchell 7026 fl (GOET, K, P); Hatschbach 36882 fl (MICH); Heringer 8928 fl (HB);
Irwin et al. 24109 fl (MICH, NY, UB); Macedo 4419 fl (HB, IAN, RB); Pohl 1328 fl fr (F, NY, W);
Rizzo 3976 fl fr (RB); Ule 423 fl (P, R).
69
There is considerable variation in leaf size and shape in this species. Most
specimens collected from the Chapada dos Veadeiros have relatively short
(length/widthratio = 3/1) narrowlyelliptic to oblong leaves, whereas some specimens from the SerraDouradahave long (length/widthratio = at least 10/1)linear
leaves. Few recent collections have these long leaves, so that perhaps short leaf
length is indicative of more frequentburningor other habitatdisturbance.
The Banisteriopsis schizoptera group (species 15-17)
This groupincludestwo widespreadspecies of vines fromthe Planaltoof Brazil:

B. schizoptera is predominantlynorthernin its distribution,and B. stellaris is
mainly western and southern (Fig. 14). The third species, B. acerosa, is nonvining, and is restricted to the Sierra de Amambay in Paraguay,and southern
Mato Grosso and southwesternMinas Gerais in Brazil.
The species in this group share the following characters(see Fig. 13 for illustration of flower type): one or two pairs of glands abaxiallyon the lateral veins;
sepals obtuse, calyx glands green to brown;petals white or pink becomingcream
in age, the claw of the posterior petal not constricted; stamens with filaments
relatively short and thick, longer opposite the sepals, the anther locules pilose,
the connectives glandularon all or at least the five anteriorstamens;styles straight
to slightly diverging, slightly flaringand flatteneddistally.
This group of species is close to the B. argyrophyllagroup and B. laevifolia,
differingchieflyin the more differentiatedandroecium;in its androecialcharacters
this group resembles the B. malifolia group.
15. Banisteriopsisschizoptera(Adr. Jussieu) B. Gates comb. nov.
Fig. 13.
BanisteriaschizopteraAdr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 43. 1832[1833].

Banisteria adamantiumMartiusex Adr. Jussieu, Ann. Sci. Nat. Bot., S6r. 2, 13: 283. 1840.
Type. Martiuss.n., Brazil, MinasGerais, Tejuco, distr. Adamantium,fl fr (holotype, M).
BanisteriaintermediaAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840. Types. Martius
s.n., Brazil, Bahia, Monte Santo, fl fr (holotype, M). Blanchet 2860, Brazil, Bahia, Serra
Agurua,fl (paratype,G; isoparatypes,BM, BR, C, G, F, K, NY, P, W).
Banisteria stellaris Grisebachvar. intermedia(Adr. Jussieu) Grisebachin Martius, Fl. bras.
12(1):48. 1858.
Shrubwith viningtips, becominga vine if supportis available.Young branches

terete, velutinous, sometimes with an overlay of sparse golden T-shaped hairs,
older branchesglabrate, darkbrown, the bark splittinginto fissures with numerous pale prominentpunctate lenticels. Stipules minute or absent. Leaves opposite, rarely ternate on vining branches, the petiole 1-3(-7) mm long, golden-velutinous to glabrate, eglandular,the lamina 1.7-7.0(-8.5) cm long, 1.1-4.0 cm
wide, smaller in the inflorescence, elliptic or rarely narrowlyelliptic, truncateto
cordate at the base, acute to apiculateat the apex, with the marginflat to slightly
revolute, with 1(-5) pair(s)of stipitateglands abaxiallyon the basal lateralveins,
sparsely appressed-sericeousto velutinousadaxially,eventuallyglabrate,loosely
white- or golden-sericeousabaxially,the hairs T-shapedwith the trabecula(1.0-)
1.4-2.4 mm long and the stalk 0-0.3 mm long, with the veins prominulousto
prominentadaxiallyand prominentabaxially.Inflorescenceof 4-floweredumbels,
borne in cymes, terminalor on short axillarybranchesbearing 1-5 pairs of more
or less reducedleaves, densely velutinous to glabrate;bracts and bracteoles 1.01.8(-2.2) mm long, broadly triangular, sparsely sericeous abaxially, glabrous
adaxially, deciduous in bud or during flower, rarely persistent; pedicels sessile
(6-)11-18 mm long, 0.6-1.0 mm wide, 0.6-0.8 mm without the hairs, densely to
70
Flora Neotropica
sparsely velutinous, sometimes with an overlay of golden T-shapedhairs. Sepals

golden-sericeous, often glabrousaroundthe marginabaxially, adaxiallyminutely
velutinous distally, the anterior sepal (2.5-)3.0-4.0 mm long, 1.2-1.8 mm wide,
narrowlyelliptic, keeled, the 4 lateral sepals (3.2-)3.5-4.5 mm long, (1.8-)2.83.5 mm wide, broadly elliptic, obtuse at the apex, with the glands 2.5-3.4 mm
long, 1.0-1.6 mm wide, green to brown, the sepals projecting1.5-1.8 mm beyond
the glands. Petals white, sometimes with pink median zone on the petals outermost in bud, the 4 lateral petals reflexed between the sepals, the claw 1.5-2.5
mm long, 0.5-0.6 mm wide, the limbs (6.5-)7.5-10.0 mm long, (5.5-)6.5-9.0 mm
wide, dentate, orbicular,eglandularor with the teeth glandularespecially basally,

the antero-lateralpetals slightly larger and cucullate, the postero-lateralpetals
flat to slightlyconcave, the posteriorpetal with the claw erect (3.5-)4.0-4.5(-5.0)
mm long, up to 1.8 mm wide at the apex, channelleddistally, not constricted,the
limb (5.0-)5.5-6.0 mm long, 4.5-6.0 mm wide, rotund, reflexed, flat to slightly
convex, with at least the basal fimbriaeglandular.Stamens with the filaments
2.0-3.0 mm long, connate basally, the locules densely pilose, 1.2-1.8 mm long,
the connectives creamto buff, all more or less enlargedand glandularat the apex,
those of the stamens opposite the petals 0.8-1.4 mm long, 0.6-0.8 mm wide, 0.30.5 mm deep, those opposite the sepals 1.1-1.6 mm long, 0.8-1.0 mm wide, 0.60.9 mm deep. Ovary 1.0-1.5 mm tall, white-sericeous, the styles equal, straight
and parallelto slightly diverging,basally strigose, (2.0-)2.5-3.2 mm long, up to
0.6 mm wide at the apex, flaringand slightly flattenedapically, stigmas capitate.
Carpophoreup to 2 mm long, 0.6-0.8 mm wide, the nut 7-9 mm tall, 4-6 mm
long, golden-brown,appressed-sericeous,tuberculateto alulate, the wing 17-28
mm long, 9-14 mm broad with appressed stiff golden hairs, becoming glabrate,
scarlet, the upper and lower marginsmore or less parallel.
Type. Saint Hilaire, Catal. BI, 1365, Brazil, Minas Gerais, "immense Chapada
entre Alto du bois et Villa de Fanado," fl (lectotype, P; isotypes, P-2 sheets).
Saint Hilaire, Catal. B], 1132, Brazil, MinasGerais, Morrod'Andaia,fl (syntype,

P; isosyntype, P). I have chosen as lectotype the sheets annotated by Jussieu,
with his cross after the name.
Distribution.(Fig. 14). This species occurs on crystallineoutcrops in the Planalto region of Brazil, such as in the Serra do Espinhago in Minas Gerais, the
Serra dos Pirineus in Goias, and in the Distrito Federal. It extends northwards
into Bahia, Piaui and Maranhao.
Collected in flower November to July, and in fruit in December to July.
Representative collections. BRAZIL. Maranhao: Eiten & Eiten 10512 fl (NY, US); 10600 fl (NY,
US); 10663 fl (NY, US); 10666 fl (NY); 10804 fl (NY). Piaui: Castellanos 25309 fr (GUA, NY);
Liitzelburg 446 fl (M). Goias: Anderson 10226 fl fr (MICH, NY, UB); Irwin et al. 10901 fl (NY, UB);
34044 fl (NY, UB); Magalhaes 9800 fl (IAN). Distrito Federal: Anderson 11429 fl fr (MICH); Clayton
4889 fl (NY); Gates 392 fl fr (MBM, MICH); Heringer 8863/1067 fl (UB); Irwin et al. 11634 fl (MICH,
NY, UB). Bahia: Anderson 11750 fr (MICH); 36421 fl (MICH, NY, UB); 36584 fl (UB); Harley 15227
fl (MICH); Ule 7394 fl (G, K). Minas Gerais: Barreto 9321 fl (F, UB); Glaziou 12485 fl (BR, C, G,
K, MG, P, R); Hatschbach & Ahumada 31633 fl (MICH, NY); Markgraf & Brade 3240 fl (F, HB,
RB); Mexia 5711 fl (BM, F, G, GH, K, MICH, MO, NY, R, U, US).
This is a very variable species, varying in leaf shape, size, and pubescence.
Plants exhibit an elliptic leaf shape in Goias and western Bahia, becoming ovate
to lanceolate farthersouth in Minas Gerais. The stem pubescence shows considerablevariationin its density and hairsize throughoutthe range.Plantssegregated
in the past under the name B. intermedia show the most dense leaf and stem
pubescence and the largest flower size, and approachB. malifolia in these characters. There is a continuous gradient northwardin Goias, eastward through
71
Bahia to Piaui, and southwardto Minas Gerais, in which the leaf pubescence
becomes sparser and more appressed and golden, and the stem pubescence less
dense, less persistent and of hairs with shorter arms. This species appearsto be
a predominantlynorthernspecies in its distributionand I found it growing on
crystalline quartz outcrops or in quartz sand. Banisteriopsisstellaris appears to
be more southern in its distribution,occurringon deep red cerrado soils. Thus
in the Serra do Espinhagonear Diamantina,the glabrousB. stellaris is rare and
I found it only in the deep soil of river valleys at relatively low elevations. It is
in Maranhaowhere the ecology is very complex that the distinctionbetween B.
stellaris and B. schizoptera becomes problematic.Throughoutthe rest of their
range, in addition to the leaf pubescence character, B. schizoptera may be distinguishedfrom B. stellaris by its larger flower size, thicker pedicels and all its
stamens with glandularconnectives. However, in Maranhao,individuals with
leaves exhibitingthe pubescence characterof B. schizoptera have flowers within
the size range of B. stellaris, and with only the anteriorstamens with well developed glandularconnectives. For the purpose of naming these plants, I am
using the hair pubescence character to distinguish between these two species
since it is an easily and accuratelyobservable character,recognisingthat here it
represents a somewhat arbitrarydistinction. I hope that future work in this ecologically diverse area will resolve the problem.
16. Banisteriopsisstellaris(Grisebach)B. Gates, comb. nov.
Fig. 13.
Banisteria stellaris Grisebach, Linnaea 13: 192. 1839.

Banisteria comans Martius ex Adr. Jussieu, Ann. Sci. Nat. Bot., Sir. 2, 13: 283. 1840. Type.
Martiuss.n., Brazil, Piaui, Sao Goncalod'Amarante,fl (holotype,M; isotypes, M, P-herb.
Jussieu, fragment).
Banisteria minarum Martius ex Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840. Type.
Martiuss.n., Brazil, MinasGerais, Serro Frio, fl (holotype, M).
Banisteria comans var. stellaris Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 407. 1843. Type. Claussen
s.n., Brazil, Minas Gerais, fl (holotype, P).
Banisteria comans var. latifolia Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 407. 1843. Type. Blanchet
2845, Brazil, Bahia, Serra Agurua, fl (holotype, G; isotypes, BM, BR, C, F, G, K, MG, MO,
P, W).
Banisteria comans var. pubifolia Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 407. 1843. Type. Martius
s.n., Brazil, Bahia, Monte Santo, fl (holotype, M).
Banisteria schizoptera var. minarum (Martius ex Adr. Jussieu) Grisebach in Martius, Fl. bras.
12(1):48. 1858.
Shrubletor shrub,becominga vine where supportis available.Young branches

terete, smooth, darkbrown, sparsely and minutelypubescent, especially on vining branches, or glabrate,old branchesdarkbrown, glabrouswith numerouspale
lenticels. Stipules minute or absent. Leaves opposite, rarely 3 or 4 at a node on
vining branches or regeneratingshoots, the petiole 1.0-4.0 mm long, slender,
sparsely appressed-pubescentor glabrous, eglandular,the lamina 1.5-12.2 cm
long, 1.0-6.5 cm wide, smallerin the inflorescence,lanceolate to ovate to rotund,
truncateto cordate at the base, acute at the apex, with the marginflat to slightly
revolute, with 1-2(-4) pairs of stipitateglandson the basal lateralveins abaxially,
glabrous adaxially or rarely sparsely appressed-sericeous,abaxially glabrous or
very sparsely appressed-sericeous, the hairs T-shaped, golden, trabecula appressed 0.4-1.0 mm long, sessile, with the veins prominenton both sides. Inflorescence of 4-flowered umbels, borne in cymes, terminal or on short axillary
branches bearing 1-3 pairs of more or less reduced leaves, glabrousor sparsely
minutely velutinous; bracts and bracteoles 0.8-2.0 mm long, broadly triangular
to lanceolate, sparselygolden-sericeousor glabrousabaxially,glabrousadaxially,
Flora Neotropica
72
'Is:.
A
:
FIG. 13. Banisteriopsis schizoptera group. A-E, B. stellaris. A, flower bud x 3.2; B, flower x 5;
C, stamens 1-6, semi-diagrammatic,x 9 (stamen 1 is opposite anteriorsepal; stamen 6 is opposite

posterior petal); D, lateral view of stamens 1 and 2 x 20; E, gynoecium x 8.4. F, B. schizoptera,
samara x 2.1. G-H, B. acerosa. G, gynoecium x 8.4; H, samara x 2. (A-E, Gates 394; F, Anderson
10226; G-H, Anderson 11177.)
73
caducous before or duringflowering,rarely persistent;pedicels sessile 8-14 mm

long, 0.4-0.6 mm wide, 0.4-0.5 mm without the hairs, sparsely short-velutinous
to glabrate, sometimes with a sparse overlay of golden T-shaped hairs. Sepals
golden-sericeous abaxially, minutely velutinous distally adaxially, the anterior
sepal 2.5-3.5 mm long, 1.0-1.5 mm wide, narrowlyelliptic, keeled, the 4 lateral
sepals 3.0-4.0 mm long, 2.0-3.0 mm wide, broadly elliptic, with the apex obtuse
to caudate, with the glands 1.8-3.0 mm long, 0.6-1.5 mm wide, green to brown,
with the sepals projecting1.5-1.9 mmbeyond the glands. Petals white, sometimes
with a pink median zone on the petals outermost in bud, fimbriate,the 4 lateral
petals reflexed between the sepals, the claw 1.6-2.0(-2.5) mm long, 0.4-0.6 mm
wide, the limb 5.0-7.0(-8.5) mm long, 5.0-7.0(-8.0) mm wide, orbicular,at least
the basal fimbriaegland-tipped,the antero-lateralpetals cucullate and slightly
larger than the postero-lateralpetals, the postero-lateralpetals flat to slightly
concave, the posterior petal with the claw erect, 3.0-4.0(-5.5) mm long, 1.2-1.5
mm wide distally, not constricted, the limb 3.5-5.0(-6.0) mm long, 3.0-5.0 mm
wide, broadly oblong to orbicular,reflexed, with the fimbriaegland-tipped.Stamens with the filaments 1.5-2.5(-3.5) mm long, basally connate, the locules
densely pilose, 0.8-1.8 mm long, the connectives cream to buff, oblong, those of
the stamens opposite the petals 0.4-1.2 mm long, 0.3-0.8 mm wide, 0.1-0.4 mm
deep, only those opposite the antero-lateralpetals somewhat glandular,those
opposite the sepals 0.8-1.6 mm long, 0.5-0.9(-1.2) mm wide, 0.3-1.0 mm deep,
with those opposite the 3 anteriorsepals glandularand enlarged. Ovary 1.0-1.2
mm tall, white-sericeous, the styles equal, straightand parallelto diverging,basally strigose, 1.4-2.0(-2.5) mm long, flatteneddistally, up to 0.5 mm wide at the
apex, stigmascapitate. Fruitwith carpophore2.0-2.5 mm long, 0.3-0.5 mm wide,
the nut 7-8 mmtall, 5-6 mm long, appressed-golden-sericeous,rugoseto muricate
to tuberculate, the wing 15-21 mm long, 8-11 mm wide, with appressed, stiff,
golden hairs, becoming glabrate, scarlet, the upper margin straight to slightly
convex, the lower marginarcuate to semi-circular,embracingthe nut.
Type. Sello s.n., Brazil, Minas Gerais, Sao Joao das Guidas, fl (holotype, B,
destroyed; NY, fragment,probablyof the holotype).
Distribution.(Fig. 14). Cerradoon deep red soil throughoutthe Planaltoregion
of Brazil, extending westward to Rondoniaand northwest to "cerrado" outliers
in Para such as the Serra dos Carajas.
Collected in flower December to October, and in fruit Januaryto October.
Representative collections. BRAZIL. Para: Cavalcante 2168 fl fr (MG); 2637 fl (MG); 2691 fl (MG);
Fr6es 29698 fl (IAN, UB). Maranhao: Eiten & Eiten 4220 fl (NY, US); 4619 fl (US); Pires & Black
2297 fl (IAN); 2338 fl (IAN, US). Piaui: Gardner 2074 fl (BM, G, GH, K, P, W); Jobert & Schwacke
1079 fl (P, R); Liitzelburg 394 fl (M); Sucre 9231 fl (RB); Ule 7192 fl (G, K). Ceara: Castellanos &
Duarte 520 fl (HB); Duarte 1428 fl (MICH, RB); Guedes 444 fl fr (IAN); Lifgren 601 fr (R, S);
Liitzelburg 26000 fl (M). Rond6nia: Ribeiro 1077 fl (MICH). Mato Grosso: Anderson 11389 fl (MICH,
NY); Carauta 773 fl fr (GUA, NY, RB); Hatschbach 31894 fl (NY); Macedo 37 fl fr (S, SP); Occhioni
5767 fl (MICH). Goias: Anderson 9394 fl (MICH, NY, UB); Burchell 8729-2 fl (GOET, K, P); Hunt
5442 fr (NY); Irwin et al. 12453 fl (MICH, NY, UB); Pohl 2524.3273 fl fr (F, NY, W). Distrito
Federal: Anderson 11427 fl (MICH); Gates 406 fl fr (MICH); Irwin et al. 5772 fl fr (MICH, NY);
7843 fl (F, G, IAN, MO, NY, S, SP, US); Sastre 1105 fl (P). Bahia: Anderson 36833 fr (MICH, NY,
UB); Blanchet 2845 fl (BM, BR, C, F, G, K, MG, MO, NY, P, W); Frvxell 1961 fl (NY); Harley
16108 fl fr (MICH); Irwin et al. 30762 fl (MICH, NY, UB). Minas Gerais: Brade & Altamiro 17811
fl (R, RB); Felippe 149 fl (HB, RB, SP); Irwin et al. 23918 fl (MICH, NY, UB); Magalhdes 240 fl
(IAN, UB); Mexia 5605 pp fl (BM, G, GH, MO, NY, S, U, US). Sao Paulo: Eiten & Eiten 2573 fl
(NY, UB, US); Gottsberger 219 fl (NY); Handro 4 fl (RB, SP); Mosen 3841 fl (P, S); Riedel 1898 fl
(C, K, NY, US, W).
Local name. Brazil: "Marmelinha da flor branca."
74
Flora Neotropica
sigecleto
frre peristn
o m Rn6i
l
et
FIG. 14. Distributionof Banisteriopsisschizoptera group, species 15-17. Open circle, B. schizoptera; closed circle, B. stellaris; square, B. acerosa.
This is one of the most widespreadspecies of Banisteriopsis in the Planaltoof

Brazil, and is certainly the most variable. Not only is there great variationin leaf
size and shape, but also in flower and fruit form. Flower size increases and some
of the flowers have all the connectives glandularin the southernpart of its range
in Sao Paulo, and there is a change in the styles from diverging and flattened
apically in Sao Paulo and Minas Gerais to styles which are straightand parallel
and not flattenedin the Distrito Federal and Goias.
Most adult leaves of B. stellaris are quite glabrous; hairs are usually present
on the young leaves, and these hairs are sessile, with short appressed trabecula
0.4-1.0 mm long. Occasional plants are found throughoutthe range which are
persistently though sparsely pubescent; the few collections from Para and the
single collection from Rondoniaare persistently pubescent.
17. Banisteriopsisacerosa (Niedenzu) B..
Fig. 13.
Banisteria acerosa Niedenzu in Engler, Das Pflanzenreich IV, 141: 426. 1928.
S. 4:
., Wash. Acad.
8.
1955. Type. Maceo
Banisteriopsis macedoana L
3226,
Brazil, Minas Gerais, Mun. Cruz Verde, Km 210 along highwayfrom Sao Paulo to Cuiaba,
fl fr (holotype, US; isotypes, MO, RB, SP).
A much branchedshrubor slender tree to 2 m tall, not vining. Young branches

pendent, terete, slender, smooth, dark brown, glabrous, older branches smooth,
75
terete, darkpurplish-brown,glabrous,with pale-brownlenticels. Stipules absent.

Leaves pendent with the petiole 4-7 mm long, slender, glabrous, eglandular,the
lamina 2-5 cm long, 1.5-7.0 mm wide, smallerin the inflorescence, asymmetric,
narrowlylanceolate, obtuse to truncateat base, acute at the apex, with the margin
revolute, with a single pair of stipitateglands abaxiallyon the basal lateralveins,
glabrousadaxially,with sparse appressed-goldenhairs or glabrateabaxially,with
the veins prominulousadaxially, prominentabaxially. Inflorescenceof 2-3-flowered umbels, terminatingnumerous branchlets, sparsely pubescent to glabrate;
bracts and bracteoles 0.8-1.2 mm long, triangular,sparsely sericeous to glabrate
abaxially, glabrousadaxially, caducous before or duringflowering.Pedicels sessile, 8-15 mm long, 0.3 mm wide, minutelyvelutinous. Sepals sericeous abaxially,
minutelyvelutinous near the base adaxially, the anteriorsepal 2.4-3.0 mm long,
1.0-1.4 mm wide, narrowlyelliptic, keeled, the 4 lateralsepals 2.4-3.6 mm long,
2.0-2.4 mm wide, broadlyelliptic, narrowedat the apex, with the glands 1.2-2.0
mm long, 1.0-1.4 mm wide, dark red brown. Petals white, those outermost in
bud often with a red median zone, dentate, the 4 lateralpetals reflexed between
the sepals, the claw 2.0-4.0 mm long, 0.4-0.5 mm wide, the limb 6.0-8.5 mm
long, 6.0-8.0 mm wide, orbicular,eglandular,the antero-lateralpetals cucullate
and the postero-lateralpetals flat to slightly concave, the posteriorpetal with the
claw erect, 4.5-5.5 mm long, up to 1.4 mm wide at the apex, not constricted, the
limb 4.5-6.0 mm long, 4.0-6.0 mm wide, orbicular,reflexed, with the basal teeth
gland-tipped.Stamens with the filaments 1.0-2.4 mm long, connate basally, the
locules densely pilose, 0.8-1.5 mm long, the connectives white, those of the
stamens opposite the petals 0.5-1.0 mm long, 0.4-0.7 mm wide, 0.1-0.4 mm deep,
not glandular,those opposite the sepals 0.8-1.4 mm long, 0.5-1.0 mm wide, 0.41.0 mm deep, with the apex enlargedand glandular.Ovary 1.2-1.4 mmtall, whitesericeous, styles equal, diverging,2.0-2.5 mm long, flatteneddistally to 0.6 mm
wide, stigmas capitate. Fruit with carpophoreup to 5 mm long and 0.5 mm wide,
nut 6-7 mm tall, 4-5 mm long, golden-brown,rugose to tuberculate,with golden
appressed pubescence, the wing 15-21 mm long, 7-10 mm wide, with appressed
stiff golden hairs or glabrate,darkred, the upper and lower marginssubparallel,
the lower marginmeeting the nut ca. halfway along its distal edge.
Chromosomenumber. n = 10. Voucher, Anderson 11177.
Type. Hassler 10481 (leg. Rojas), Paraguay, Cerro Cora, fl fr (holotype, G;
isotypes, A, G).
Distribution.(Fig. 14). Open grasslandin SE Mato Grosso and the Triangulo
Mineiro of Minas Gerais in Brazil, extending northwardto the Distrito Federal
and Rio Pardo in Minas Gerais and southward to the Sierra de Amambay in
Paraguay.
Collected in flower Februaryto July, and in fruit Marchto July.
BRAZIL. Mato Grosso: Anderson 11177 fl fr (MICH, NY); 11387 fl (MICH); Hatschbach 23706
fl (MICH, P); 31940 fl fr (MICH, NY); 34723 fl fr (MICH). Distrito Federal: Ratter et al. 3444 fl fr
(E, MICH). Minas Gerais: Riedel 489 fl fr (LE).
This species is very distinctive with its pendent, narrow leaves and pendent
branches, giving the whole plant a willowy aspect, and the flowers are borne in
pairs and not in umbels. Banisteriopsisacerosa is most closely relatedto B. stellaris, from which it can easily be distinguishedby the charactersdescribedabove.
The Banisteriopsis malifolia group (species 18-23)
This group consists of three widespread species and three localised endemic
species; the group can be divided into 2 subgroups, the B. malifolia group and
76
Flora Neotropica
the B. variabilis group. The widespreadspecies show great variationover their

range, and also often exhibit characters from other species in the group with
which they are sympatricat a given place, suggesting that gene flow within the
group does occur. As a group, the species cohere very well, with the same flower
and fruit type (see Fig. 15). The fruits are unusual in the rotation of the two
posterior wings to lie parallel to the anteriorwing, and the locule of the nut is
densely hairy inside.
The B. variabilis group (B. variabilis, B. latifolia, B. hirsuta, B. hatschbachii)
and the B. malifolia group (B. malifolia, B. confusa) can be distinguished by the
following characters:
Petiole length
Petiole width
Leaf width
Pedicel width:
With hairs
Without hairs
B. malifolia group
(2-)3-8(-13) mm
up to 2 mm
2.5-7.5(-12.0) cm
B. variabilis group
0.3(-4) mm
more than 2 mm
5.6-14.1 cm
0.6-1.0 mm
0.4-0.6 mm
(0.8-)1.0-2.2 mm
0.6-1.0 mm
18. Banisteriopsismalifolia(Nees et Martius)B. Gates, comb. nov.
Fig. 15.
Malpighia malifolia Nees et Martius, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 12: 21. 1824.
Banisteria crotonifolia Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 36. 1832 [1833]. Types.
Saint Hilaire, Catal. B], 1104, Brazil, Minas Gerais, summit of Morro de Mondo Velho, fl
(lectotype, P; isolectotype, P). Saint Hilaire s.n., Brazil, Minas Gerais, near Barbados (syntype, P).
Banisteria crotonifolia var. angustata Grisebach in Martius, Fl. bras. 12(1): 52. 1858. Type.
Martius s.n., Brazil, Minas Gerais, Itacolumi, near Marianna, fr (holotype, M).
Subshrub,shrub or bushy tree to 3 m, the branchends weakly vining. Young

branches terete, densely golden-brown-velutinousto appressed-sericeous,older
branches with pale punctate lenticels, eventually glabrate,the bark dark-to reddish-brown.Stipules minute or up to 1 mm long, triangular,appressed-sericeous
to glabrate.Leaves with the petiole 2-7 mm long, up to 2 mm in diameter,goldenvelutinous or appressed-sericeous,eglandular,with the lamina2.8-7.4(-10.2) cm
long, 1.7-5.7 cm wide, smallerin the inflorescence, narrowlyelliptic to broadly
elliptic or ovate, plane to rugose, obtuse to cordateat the base, acute to apiculate
at the apex, with the marginplane to revolute, bearing a single pair of peltate
glands 0.8-1.0 mm in diameter beside the costa at the base abaxially, shortvelutinous or appressed-pubescent becoming glabrate adaxially, sparsely to
densely tomentose or sericeous abaxially, the hairs T-shapedwith the trabecula
1.2-2.8 mm long, straightto flexuous, and the stalk 0.04-0.20 mm long, flat to
rugose, with the veins very impressed or not adaxially, prominent abaxially.
Inflorescence of 4-floweredumbels on leafy shoots, the umbels borne in cymes
on terminal shoots, densely golden-velutinous or white-appressed-sericeous;
bracts and bracteoles 1.8-2.8 mm long, lanceolate to broadly oblong, sericeous
FIG. 15. Banisteriopsis malifolia. A, flower bud x 4.2; B, flower x 4.2; C, stamens 1-6, semidiagrammatic, abaxial and lateral views, x 11 (stamen 1 is opposite anterior sepal; stamen 6 is opposite posterior petal); D, gynoecium x 8.4; E, fruit to show rotation of wings of posterior samaras x
2.5; F, longitudinal section of locule of samara x 3.5. (A-F, Gates 395.)
77
.S. .B
.....
if
..
Ob
~~~~~~~~~~~~~7;.
*-."
i.i34!
?
.
.;,..'
. ..',
"
'^ ? .
; .
.D
78
Flora Neotropica
abaxially, glabrous adaxially, caducous in bud; pedicels sessile, 9-16(-24) mm

long, 0.6-1.0 mm wide, 0.4-0.6 mm without hairs, velutinous or appressed-sericeous. Sepals white-sericeousabaxially, sparsely minutelyvelutinousthroughout
adaxially, 3.5-5.3 mm long, 2.0-2.5 mm wide, elliptic to oblong, rounded and
sometimes revolute at the apex, with the glands 2.3-3.0 mm long, 0.9-1.6 mm
wide, the sepals projecting 1.6-2.3 mm beyond the glands. Petals white or pale
pink, sometimes with pink veining, becoming pale yellow in age, dentate to laciniate, the 4 lateral petals reflexed between the sepals, the claw 1.5-2.5 mm
long, 0.4-0.8 mm wide, the limb 8.0-11.0 mm long, 7.5-10.0 mm wide, the anterolateral petals cochleate, the postero-lateralpetals plane, orbicular, eglandular,
the posterior petal with the claw erect, 3.0-4.0 mm long, up to 1.3 mm wide at
the apex, not constricted, the limb 6.5-9.5 mm long, 6.0-8.5 mm wide, plane to
convex, with the basal teeth not glandularor gland-tipped. Stamens with the
filaments2.0-4.5 mm long, connatebasally, the locules 1.0-1.6 mm long, sparsely
pilose, the connectives yellow, glandular,those opposite the petals 0.6-1.2 mm
long, 0.4-0.9 mm wide, 0.2-0.7 mm deep, those opposite the sepals 0.9-1.4 mm
long, 0.7-1.2 mm wide, 0.4-0.9 mm deep, projectinga little beyond the locules.
Ovary 1.0-1.4 mm long, white-sericeous;styles equal, straightand parallel, 2.53.5 mm long, taperingto 0.3-0.5 mm wide, stigmas capitate. Fruit with carpophore 0.6-1.2 mm long, 0.3-0.6 mm wide, the nut 5-7 mm tall, 4-5 mm long,
appressed-pubescent,the wing (10-)15-25 mm long, 7-14 mm wide, with appressed T-shaped hairs, the trabeculaup to 1.8 mm long, glabrescent, with the
wings of the 2 posterior samarasrotated to lie parallel with that of the anterior
samara,the locule densely hairy inside.
Key to the Varieties of Banisteriopsis malifolia
1. Branches and petioles velutinous; leaves rugose, abaxially tomentose with the hairs
T-shaped, with the trabecula 1.2-2.8 mm long and the stalk 0.08-0.20 mm long. a. var.
malifolia.
1. Branches and petioles appressed-sericeous; leaves plane, abaxially appressed-tomentose to
sericeous, the hairs T-shaped with the trabecula 0.8-1.6 mm long, the stalk 0.04-0.10 mm
b. var. appressa.
long.
18a. Banisteriopsismalifoliavar. malifolia

Branches densely grey- or golden-velutinous.Petioles velutinous. Leaves rugose, adaxially velutinous, rarely appressed-pubescent,but then velutinous on
the veins, sometimes glabrate, adaxially loosely grey-tomentose, the hairs
T-shaped, the stalk 0.08-0.2 mm long, the trabecula1.2-2.8 mm long, straightor
flexuous. Pedicels velutinous to tomentose. Bracteoles caducous early in bud,
lanceolate.
Type. Prinz von Neuwied s.n., Brazil, Minas Gerais, between Tamboriland
Valos (holotype, BR; isotype, BR).
Distribution.(Fig. 16). In red soil cerradosthroughoutthe northernpart of the
Planaltoregion of Brazil.
Collected in flower October to July, and in fruit Februaryto September.
Representative collections. BRAZIL. Goias: Anderson 8105 fl (MICH, NY); Glaziou 20752 fl (BR,
C, G, K, P, R, RB, S); Irwin et al. 10303 fl (G, IAN, MO, NY, S, SP, US); 13440 fl fr (MICH, NY);
Pohl 2858 pp fl fr (F, NY, W). Distrito Federal: Belem & Barroso 4013 fl (NY, UB); Ferreira 28 fl
(UB); Heringer 10102 fl (IAN, NY, UB); Irwin et al. 7961 fl fr (MO, NY, S, SP, US); Pires et al.
9093 fl (NY, P, UB, US). Bahia: Belem 3655 fl (IAN, MICH, NY); Castellanos 25037 fl (NY, GUA);
Duarte & Castellanos 313 fl (MICH); R. Silva Santos s.n. fl (MICH); Sobrinho 261 fl (US). Minas
Gerais: Barreto 7619 fl (HB); Brade 13783 fr (RB); Glaziou 12486 fl (BM, BR, C, G, K, NY, P, R);
79

70,
FIG. 16. Distribution of Banisteriopsis malifolia and B. confusa. Closed square, B. malifolia var.
malifolia; open square, B. malifolia var. appressa; triangle, B. confusa.
Magalhades19127fl (MICH, NY, U); Mexia 5895 fl (BM, F, G, GH, K, MICH, MO, NY, R, S, U,
US).
Local names. Brazil:flor do dia.

This variety is very widespread throughout the cerrados of the Planalto of
Brazil, and appears to have considerable success in disturbedhabitats, such as
the roadsides of northern Minas Gerais between Salinas and Diamantina, and
some areas cleared for building in the cerrado of the Universidade de Brasilia.
There is also considerable variation in leaf size and shape in this variety; individuals with largerless rugose leaves are found in the Distrito Federal and southern Minas Gerais near Belo Horizonte, where B. malifolia is sympatricwith B.
variabilis, which has large non-rugoseleaves. It is possible that some gene flow
between these two species is occurringin these areas (e.g., Gates 414, 415). In
northern Minas Gerais and Bahia, var. malifolia exhibits characters typical of
var. appressa, and has leaves which are not rugose and with appressed pubescence on the upper epidemis.
18b. Banisteriopsismalifoliavar. appressaB. Gates, var. nov.
Rami sparsim vel dense aurei- vel albidi-appresso-sericei.Folia plana, lamina
adaxialitersparsimvel dense appresso-pubescenti,interdumglabrata,abaxialiter
sparsim vel dense sericea vel tomentosa, pilis T-formibus, pede 0.04-0.10 mm
80
Flora Neotropica
longo, trabecula0.8-1.6 mm longa, recta vel parumflexuosa;petiolo laxe sericeo.

Inflorescentiapedicellis sericeis; bracteisbracteolisqueper anthesincaducis, late
obovatis.
Type. Irwin et al. 14459, Brazil, Goias, Serra Geral de Goias, Rio da Prata,
ca. 6 km S of Posse, fl fr (holotype, UB; isotypes, MICH, NY).
Distribution.(Fig. 16). ThroughoutnortheasternBrazil in Bahia, Pernambuco,
Ceara, Piaui, Maranhao,Goias and extending to the Serra dos Carajasin Para,
at altitudes of 200-1200 m.
Collected in flower Februaryto May and July to September, and in fruit in
April, May and August.
BRAZIL. Para: Cavalcante 2108 fr (MG 36688); 2110 fl (MG 36690); 2137 fl (MG 36717); 2638 fl
(MG 37863); Pires & Passos 13191 fl fr (IAN, MICH); N. T. Silva & Ribeiro 3656 fr (IAN). Maranhao:
Lisboa 2490 fl (MG, RB). Ceara: Cutler 8405 fl (F, MO, R, US); Duarte & Ivone 1293 fl (RB);
Gardner 1484 fl fr (BM, K) (syntype B. schwannioides Grisebach); Guedes 377 fl (IAN, UB); Lutzelburg 12493 fl (M, NY). Pernambuco: Heringer et al. 74 fl (R, UB); 349 fl (R, RB, UB). Goias:
Anderson 6686 fl (MICH, NY, UB); 7954 fl (NY, UB); Hatschbach & Kummrow 38316 fl (MICH);
Irwin et al. 14341 fl fr (MICH, NY, UB); 21530 fr (MICH, NY, UB). Bahia: Anderson 11743 fl
(MICH); Anderson et al. 36590 fl (NY, UB); 36905 fl (NY, UB); Harley 16855 fl (K); Ule 7534 fl (G,
K). Without locality: Burchell 8682 fl (GOET); 8746-6 fl (GH, GOET, K, P).
Local name. Brazil: "Rama de moco".
This variety occupies the northernpart of the rangeof B. malifolia, and is very
variable in the density of leaf pubescence, leaf size and shape, and habit. Specimens from the SerraGeralde Goias and the Espigao Mestrein Bahia show dense
sericeous leaf and stem pubescence, whereas in Ceara and eastern Bahia, the
pubescence is more sparse, and some individualsfrom northernGoias and Maranhaoare glabrate.This variationmay be correlatedwith altitude, since all the
glabrateor sparsely sericeous individualsare from relatively low altitudes (200500 m). The collections from Pernambucohave subvelutinousstem pubescence,
but in their leaf pubescence they lie within var. appressa. Anderson 11743 from
southern Bahia is somewhat intermediatein its pubescence between var. malifolia and var. appressa.
Leaf shape in var. appressa varies from ovate to elliptic or orbicular.Individuals which are described as subshrubshave larger more ovate leaves, and individuals described as shrubs or vines have smaller elliptic to orbicularleaves.
Collections from the Serra dos Carajasin Para have very revolute leaves, but in
other respects are like var. appressa.
One specimen, Anderson 6686 from the Chapadados Veadeiros in Goias, has
very short petioles and approachesB. variabilis in this respect; however, it has
the slenderpedicels and dense pubescence of B. malifolia var. appressa. Gardner
1484, a syntype of B. schwannioides, belongs to this variety. It is a specimen in
youngfruitwhich has been galled so that the developingfruitsare greatlyenlarged
and distorted. This phenomenonis quite common in B. malifolia.
19. BanisteriopsisconfusaB. Gates, sp. nov.

Banisteria velutina Adr. Jussieu (non Martius ex Colla), Ann. Sci. Nat. Bot., Ser. 2, 13: 282.
1840. Type. A. R. Ferreira s.n., Brazil, fl (holotype, P-JU, Catal. 11590; isotype, P).
Frutex ramosususque 3 m altus, interdumvolubilis vel dumetaformans. Rami

juniores teretes, dense aureo-brunneivelutini, rami vetustiores demum glabrati,
lenticellis numerosisin rimis suberosispraesentibus.Stipulaeminutaevel nullae.
Folia opposita;lamina(3.5-)5.9-12.0 cm longa, 2.1-6.1 cm lata, in inflorescentia
minori, ovata vel elliptica vel anguste ovata, basi obtusa cordatave, apice acuta
81
apiculatave, margineplana revolutave, subtus basi prope costam 2 glandes peltatas 1.0-1.2 mm diametrogerenti, supra aureo-velutina,demum glabrata,pilis
Y-formibusstipitatis, brachio0.1-0.3 mm longo, subtus laxe albido-sericea,pilis
T-formibus, pede 0.1-0.2 mm longo, trabecula recta vel flexuosa 0.8-1.4 mm
longa, venis venulisque supra prominulisvel parumimpressis, subtus prominentibus; petiolo 6-13 mm longo, gracili, aureo-velutino,eglanduloso. Inflorescentia
floribus in umbellis 4-floriferisfasciculatis foliis adultis vel redactis subtentis,
umbellis in cymis dispositis, aureo-brunnea-velutina;
bracteis bracteolisque2.03.0 mmlongis, lanceolatis,abaxialitersericeis vel glabratis,adaxialiterglabrisante
vel per anthesincaducis;pedicellissessilibus 10-20 mmlongis, 0.9-1.1 mmdiametro, 0.5-0.7 mm sine indumento,velutinis. Sepala5, abaxialitersericea, minutevelutina adaxialiter,3.8-5.0 mm longa, 2.0-3.2 mm lata, elliptica oblongave, apice
rotundata et saepe revoluta, 4 lateralia glandes virides vel badias 2.0-3.4 mm
longas et 1.0-1.5 mm latas gerentia, glandes 2.2-2.8 mm superantia.Petala 5,
alba, aetate luteola, fimbriata,4 lateraliainter sepala patentia, ungue 1.5-2.5 mm
longo, 0.5-0.8 mm lato, limbo 7.5-10.0(-11.5) mm longo, 7.0-9.0(-10.5) mm lato,
orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribus, petalumposticum ungue erecto 3.5-4.0 mm longo, 0.6-1.0 mm lato versus apicem,
non contracto, limbo 7.0-9.0(-10.5) mm longo, 6.0-8.5(-9.5) mm lato, orbiculari,
retroflexo, piano convexove, fimbriisbasalibusglandulosis. Stamina 10, filamentis 2.0-4.0 mm longis, proximaliterconnatis, glabris;antheraeretroflexae,loculis
0.8-1.4 mm longis, pilosis pilis apice et basi caespitosis, connectivis canarinis,
petalis oppositis 0.6-1.0 mm longis, 0.4-0.7 mm latis, 0.2-0.6 mm profundis,
parumglandulosisvel solum 2 petalis anterioribusoppositis glandulosis, illis sepalis oppositis (0.9-)1.2-1.6 mm longis, 0.6-1.0 mm latis, 0.5-1.0 mm profundis,
glandulosis dilatatisque, illis 3 sepalis anticis oppositis loculos 0.4-0.6 mm superantibus. Ovarium 1.0-1.4 mm altum, carpellis 3, uniovulatis, liberis, toro
pyramidaliadnatis, candido-sericeis;styli 3, aequales 2.5-3.0 mm longi, recti parallelique, decrescentes versus apicem usque 0.4 mm diametro, stigmatibus
capitatis. Fructusex samaris(vel abortu 1-2) constans; samaracarpophorousque
3.0 mm longo, 0.6 mm lato, nuce ovoidea 5-9 mm alta, 3.5-5.0 mm longa, parum
rugosa vel alulata, alulis usque 8 mm longis, sparsim appresso-pubescenti,pilis
rigidis, basi distaliterprominentedentata, ala 14-24(-30) mm longa, 10-13(-15)
mm lata, appresso-sericeavel glabrata,pilis sessilibus trabecularecta 1.4-1.6 mm
longa; alis 2 posterioribusad alam anticam subparallelibus;loculo nucis interius
dense piloso.
Type. Anderson 11277, Brazil, Mato Grosso, Serra da Pimenteira,25-35 km
SW of town of Rio Verde, fl fr (holotype, MBM; isotypes, MICH, NY) .
Distribution. (Fig. 16). Collected in Rondonia and Mato Grosso and also in
Minas Gerais and Sao Paulo in Brazil, and in SE Paraguayand Bolivia.
Collected in flower Januaryto March, May to July, September, October and
December, and in fruit in February,March, July and December.
Representative collections. BRAZIL. Rond6nia: Black & Cordeiro 52-14593 fl (IAN, UB); 52-15189
fl (IAN); Cordeiro 917 fl (MICH). Mato Grosso: Hatschbach 23759 fl (MICH); 24295 fl (MICH);
33946 fl (MICH); Commissao Rond6n (leg. Hoehne) 1629 fl (R); 4896 fl (R); Riedel s.n. fl (W). Minas
Gerais: Regnell 111.316 fl fr (P, S, US); Hipolito s.n. fl fr (MICH). Sao Paulo: Hemmendorff 86 fl
(S). BOLIVIA: Buchtien 912 fl (NY); 1804 fl (NY, fragment, US); Kuntze s.n. fl (F , NY). PARAGUAY: Balansa 4581 fl (G, P); Fiebrig 689 fl (BM, F, G, GH, GOET, K, M, MO, NY); Hassler
6726 fl (BM, R, G, GH, K, MO, NY, P, S, W); Krapovickas et al. 12476 fl (CTES); Schinini 4417 fl
fr (CTES).
Banisteriopsis confusa may be distinguished from B. malifolia by its larger,
plane leaves with longerpetioles and shorterhairs abaxially,its more vininghabit
82
Flora Neotropica
and its inflorescence. In some of these characters, B. confusa approaches B.

adenopoda and is somewhat intermediate between B. malifolia and B. adeno-
poda; B. confusa can be distinguishedfromB. adenopoda by its basalleaf glands,

less enlarged stamen connectives and straight styles, and its fruit without the
prominentlateral wings of B. adenopoda. It is possible that some hybridisation
between these two species occurs in southernMinas Geraisand Sao Paulo where
they occur sympatrically,but biosystematic studies are needed to resolve this.
The few collections of B. confusa from this area are disjunctfrom the rest of the
taxon; this disjunctionmay be only apparent,however, due to the destructionof
the naturalvegetation of Sao Paulo.
There is considerable variation in leaf shape and size in this species; in the
case of collections from vining individuals,this may be because the largerleaves
were not collected. One of the collections from Rondonia,Black & Cordeiro525189, is unusual in its narrowly elliptic, rugulose, glabrate leaves; with so few
collections from Rondonia, the significanceof this variance is hard to evaluate.
20. Banisteriopsislatifolia(Adr. Jussieu) B. Gates, comb. nov.
Banisterialatifolia Adr. Jussieuin Saint Hilaire,Fl. bras. mer 3: 40. 1832[1833].
Gnarledtree to 4 m high, trunkto 15 cm in diameter. Young branches round,

at first densely tomentose, soon glabrous, the bark smooth, red-brown, developing corky splits. Old branchesgrey-brownwith thick corky barkformingprominent corky ridgesand furrows.Stipulesminuteor absent. Leaves with the petiole
2-3 mm long, glabrousor sparsely tomentose, eglandular,the lamina (3.5-)6-14
cm long, (3.0-)4.5-10.0 cm wide, muchreducedor bract-likein the inflorescence,
elliptic or ovate, acute, apiculateor retuse at the apex, truncateto cordateat the
base, with the marginflat, bearing abaxially on the costa at the base a pair of
sessile glands 1.5-2.0 mm in diameter, loosely sericeous adaxially when first
expanded, soon glabrous, abaxially densely and loosely white-sericeous when
first expanded, soon glabrous, except in the inflorescence, the hairs T-shaped
with the trabecula1.0-2.0 mm long and the stalk 0.1-0.2 mm long, with the veins
prominulousadaxially and prominentabaxially. Inflorescenceof 4-floweredumbels, cauliflorous and borne at the nodes, sometimes also on short terminal
branchesbearingreducedleaves, the umbels borne in condensed cymes, densely
sericeous; bracts and bracteoles 2.2-3.0 mm long, lanceolate, glabrousadaxially,
sericeous abaxially, caducous just before or during flowering; pedicels sessile
(14-)19-27 mm long, 0.8-1.2 mm in diameter,0.6-0.8 mm without hairs, loosely
white-sericeous. Sepals white-sericeousabaxially, adaxiallyminutelyvelutinous,
3.8-6.0 mm long, 2.2-3.2 mm wide, roundedand often revolute at the apex, with
the glands 2.4-3.5 mm long, 1.0-2.2 mm wide, the sepals projecting2.8-3.0 mm
beyond the glands. Petals white, becomingpale yellow in age, dentateto laciniate,
the 4 lateral petals reflexed between the sepals, the claw 2.0-2.5 mm long, 0.81.0 mm wide, the limb 7.0-11.0 mm long, 7.0-10.0 mm wide, orbicular,eglandular, the antero-lateralpair of petals more concave and largerthan the posterolateral pair, the posterior petal with the claw erect, 4.0-5.0 mm long, up to 1.2
mm wide, narrowedat the apex, the limb 6.0 mm long and wide, orbicular,plane
to convex, with the basal teeth eglandularor sometimes gland-tipped.Stamens
with the filaments1.8-4.0 mm long, connatebasally, the locules 1.0-1.6 mm long,
pilose, the connectives yellow, those opposite the petals 0.8-1.2 mm long, 0.70.8 mm wide, 0.4-0.6 mm deep, these opposite the antero-lateralpetals glandular,
those opposite the sepals 1.0-1.4 mm long, 0.7-1.0 mm wide, 0.6-0.8 mm deep,
glandularand enlargedat the apex. Ovary 1.5-1.8 mm high, white-sericeous,the
83
styles equal, straightand parallel, 3.0-3.2 mm long, taperingto 0.4 mm wide at

the apex, the stigmascapitate. Fruitwith carpophorefiliform,up to 3 mmlong, 0.5
mm wide, the nut 5-8 mmtall, 4-5 mmlong, smoothto slightlyrugose with a basal
tooth, appressed-sericeous,the wing 15-25 mm long, 9-15 mm wide, appressedsericeous to glabrate, the hairs T-shaped, with the trabecula 1.0-1.2 mm long,
the wings of the 2 posteriorsamarasrotatedto lie parallelwith that of the anterior
samara,the locule densely hairy within.
Type. Saint Hilaire, Catal. Cl, 642, Brazil, Goias, near Riberao Frio, fl (holotype, P; isotypes, P-2 sheets).
Distribution.(Fig. 17). Found only in the cerradosof the Distrito Federal, and
the Serra dos Pirineusand Serra Geral do Paranain adjacentGoias in Brazil.
Collected in flower in Marchto May, and in fruit in March.
BRAZIL. Goias: Anderson 7578 fl fr (MICH, NY); 10294 fl (MICH, NY); Glaziou 20750 fl (BR,
G, P); 20751 pp fl (K, NY, P, R); Irwin et al. 15446 fl (MICH, NY); 34107 st (MICH, NY); Macedo
3673 st (RB); Onishi et al. 068 fl (NY); Pires et al. 9660 fl (US); Pohl 1677 fl (F, NY, W); Weddell
2073 fl (P). Distrito Federal: Heringer 9096/1290 yng fr (MICH, UB); 11206 fl (NY); 11511 fl (NY);
Irwin et al. 14099 bud (MICH, NY); Lima & Barroso 255 fl (NY, UB); Philcox & Onishi 4825 fl (NY,
UB); Pires et al. 9051 fl (UB); 9428 fl (GH, UB, US).
This is a very distinctive species with its thick corky bark, large glabrous
leaves, caulifloroushabitand long pedicels. The branchesduringtheirfirst season
of growth are quite smooth, red-brownin colour and without lenticels; by their
second season of growth the branchesexhibit prominentcorky splits in the bark,
and by the next season the branches are already corky. Banisteriopsis latifolia
is also unusual in its habit as a small tree, with a single erect unbranchedtrunk
up to 15 cm in diameter;most cerrado species of Banisteriopsis are shrubswith
several woody stems from the base. I did, however, observe one individualof
B. latifolia in the cerrado of the Universidadede Brasiliawith several stems; in
all other respects the plant was typical B. latifolia, and this growth form may
representregenerationafter damageto the main trunk.
The foliage leaves in this species are densely and loosely tomentose on both
sides when they first expand, but become quite glabrous very soon-probably
withina few days. One specimen, Irwinet al. 34762, has leaves which are densely
hairy abaxially, associated with thick corky bark and a tree habit; many of the
leaves are from the previous growingseason, and the young stems show lenticels
as well as corky splits; this individualmay representa hybridbetween B. latifolia
and B. variabilis. Since the materialis sterile, it was not possible to look for
other indicationsof hybrid origin, such as low pollen fertility.
21. BanisteriopsisvariabilisB. Gates, sp. nov.
Banisteria latifolia var. paraguariensis Niedenzu, Verz. Vorles. Ak. Braunsberg: 18. 1912.
Types. Hassler 10252 (leg. Rojas), Paraguay, Sierra de Amambay, fl (lectotype, NY; isolectotypes, A, G, K). Hassler 10252a (leg. Rojas), Paraguay, Sierra de Amambay, fr (syntype, A).
Banisteria velutina sensu Niedenzu, non Jussieu.
Fruticulus virgatus vel frutex ramosus usque 3 m altus. Rami teretes, dense
aureo- vel albido-velutini vel sericei, interdum glabrati, lenticellis numerosis
punctati, interdumsubter indumentovix manifestis. Stipulae minutaevel usque
1.0 mm longae, triangulares,sericeae vel glabratae.Folia opposita; lamina(4.5-)
7.1-14.1 cm longa, (3.0-)5.0-9.2(-12.6) cm lata, orbiculariet minori in inflorescentia, ovata vel rotunda, basi truncatacordatave, apice acuta vel apiculatavel
acuminata,margineplanavel parumrevoluta, subtusbasi prope costam 2 glandes
84
Flora Neotropica
peltatas 1.0-1.6 mm diametrogerenti, supra velutina vel sparsimsericea et mox

glabrata,subtus sparsimet laxe sericea, interdummox glabrata,pilis T-formibus,
pede 0-0.2 mm longo, trabecularecta vel flexuosa 0.4-2.4 mm longa, venis venulisque supraplanis vel prominulis,subtus prominentibus;petiolo 1-6 mm longo, crasso, velutino vel sericeo vel glabrato, eglanduloso. Inflorescentiafloribus
in umbellis4-floriferisfasciculatis, umbellisin cymis dispositis, in axillis foliorum
redactorumin ramis terminalibusgerentis, velutina vel sericea; bracteis bracteolisque 1.8-3.0 mm longis, lanceolatis ellipticisve, albido-sericeis adaxialiter,
glabris abaxialiter, ante vel per anthesin caducis; pedicellis sessilibus, (7-)1016(-20) mm longis, 1.0-1.7 mm diametro, 0.6-1.0 mm sine indumento, laxe albido-seiceis. Sepala 5, abaxialiteralbido-sericea,adaxialiterminute pubescentia,
basi glabra,4.0-5.5 mm longa, 2.0-3.8 mm lata, ellipticaovatave, apice rotundata
revolutave, 4 lateralia glandes virides vel badias 2.0-3.6 mm longas et 0.8-1.8
mm latas gerentia, glandes 1.8-2.0 mm superantia.Petala 5, alba vel subrosea,
aetate luteola vel salmonea, dentata, 4 lateraliainter sepala patentia, ungue 1.02.4 mm longo, 0.4-0.8 mm diametro,limbo 8.0-11.5 mm longo, 7.5-9.0 mm lato,
orbiculari,eglanduloso,2 anterioribusquam2 posterioribusconcavioribuset parum majoribus,petalum posticum ungue erecto 3.2-4.5 mm longo, 1.0-1.7 mm
lato, canaliculato,apice non contracto,limbo 7.0-8.0 mm longo, 6.5-8.0 mm lato,
orbicularivel obovato, retroflexo,plano convexove, eglandulosovel 1-2 dentibus
basalibus utrinqueglandulosis. Stamina 10, filamentis2.0-4.0 mm longis, proximaliter connatis, glabris; antherae subaequales, retroflexae, loculis 1.0-1.8 mm
longis, pilosis pilis apice et basi caespitosis, connectivis canarinis,petalis oppositis (0.6-)0.9-1.4 mm longis, 0.6-1.3 mm latis, 0.2-0.6 mm profundis,glandulosis
vel interdumillis 3 posterioribusoppositis eglandulosis,illis sepalis oppositis 1.01.8 mm longis, 0.8-1.2 mm latis, 0.6-1.0 mm profundus, glandulosis. Ovarium
1.0-1.8 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidaliadnatis, candido-sericeis;styli 3, aequales, 2.6-3.6 mm longi, recti parallelique,decrescentes
versus apicem usque 0.4-0.6 mm diametro, stigmatibuscapitatis. Fructus ex 3
samaris(vel abortu 1-2) constans; samaracarpophorousque 1.5-2.0 mm longo,
0.5-0.6 mm lato, nuce ovoidea 7-10 mm alta, 3.5-5.0 mm longa, laevi vel rugosa,
appresso-pubescentivel glabrata,ala 12-30 mm longa, 11-14 mm lata, appressosericea vel glabrata,pilis stipitatis trabecularecta 0.8-1.2 mm longa, alis 2 posterioribusad alam anticam subparallelibus;loculo nucis interius piloso.
Type. Anderson 10000, Brazil, Goias, SerraDourada,ca. 15 km (straightline)
S of Goias Velho, fl fr (holotype, UB; isotypes, MICH, NY).
Distribution.(Fig. 17). Cerradosof Mato Grosso, Goias, southern Minas Gerais, Rio de Janeiro, Sao Paulo, and Paranain Brazil, extending southwardsinto
Paraguay.
Collected in flower Septemberto July, and in fruit Februaryto July.
Representative collections. BRAZIL. Mato Grosso: Anderson 11252 st (MICH); Hatschbach 31867
fl fr (MICH, NY); 31945 fl (MICH, NY); D. Smith 202 fl fr (K). Goias: Anderson 9382 fl fr (MICH,
NY, UB); Gardner 4100 fl (K); Hatschbach & Kummrow 34958 fl (MICH); Irwin & Soderstrom 7414
fl (NY); Irwin et al. 13555 fl fr (MICH, NY, UB). Distrito Federal: Belem & Barroso 4022 fl (NY,
UB); Gates 413 fl (MBM, MICH); 418 fl yng fr (MBM, MICH); Heringer 8009 fr (SP); Irwin et al.
18091 fr (MICH, NY). Minas Gerais: Barreto 7612 fl (F, R, US); Claussen 66A fl (BR, F, W); Duarte
2442 fl (RB); Hatschbach 36999 fl (MICH); L. B. Smith 6955 fl (NY, P, R, US). Rio de Janeiro:
Occhioni 5699 fl (MICH). Sao Paulo: Avalone et al. 5 fl (IAN); Eiten & Eiten 2784 fl (NY, SP, US);
Gottsberger 49R-4573 fr (MICH); Riedel 2210 fl (K, US); Valio 265 fl (RB, SP, US). Parana: Anderson
11160 fl (MICH, NY); Hatschbach 8842 fl fr (MICH, US); 14247 fl fr (MICH, P, US); 21586 fl fr
(MICH, NY); Lindeman & Haas 973 fl fr (MBM, NY, U). PARAGUAY: Hassler (leg. Rojas) 10831
fl (GH).
85
I
FIG. 17. Distribution of Banisteriopsis variabilis subgroup of B. malifolia group, species 20-23.
Circle, B. latifolia; closed square, B. variabilis; triangle, B. hirsuta; open square, B. hatschbachii.
This is an exceedingly variable taxon, and I recognise that the treatment of this
variation as one species is not entirely satisfactory. However, I am unable to
subdivide the variation in any coherent way on either a geographical or morphological basis. I shall give a rather full discussion of the variation, and hope that
in the future a better understanding of this variation can be achieved.
1. Habit. This species may be a shrublet with many erect stems to 1 m from
an underground base, or a shrub, often spreading, with many stems to 2.5 m or
more.
It is possible
that the individuals
with the habit of a subshrub
may represent
young or regenerating individuals; they do produce flowers and fruit. In the cerrados of Brasonelia,this species is very abundant, both as a shrublet and a shrub;
the individuals with the habit of a subshrub usually have larger leaves which are
pubescent to glabrate, whereas the shrubs usually have smaller leaves which are
86
Flora Neotropica
nearly to quite glabrous. This suggests an age-based habit correlation to me,

especially since the cerradounder study was almost undisturbed;in other areas
where disturbanceby man has been great, the subshrubs may be regeneration
forms.
2. Leaf pubescence. Throughoutmost of its range, this species has adultleaves
which are densely pubescent abaxially. However, individuals from Paraguay,
Mato Grosso, Sao Paulo and the DistritoFederalhave leaves which are glabrous.
Since in these same areas and throughoutthe range of the species, individuals
with glabrateleaves are found, it would seem unwise to segregatethese glabrous
individualsas they appearto representone extreme of a characterwhich shows
considerable variability. In the Distrito Federal the glabrous-leavedindividuals
are usually shrubs;if habit of a shrubis age-related,this would mean that older
individualsproduce glabrousleaves. However, many of the glabrous-leavedindividuals from Paraguayand Mato Grosso, Brazil, are described as subshrubs,
and many plants described as shrubshave persistentlypubescent leaves. I could
find no consistent correlationbetween glabrousleaves and other charactersshowing variation. The glabrous-leavedindividualsfrom Paraguayand Mato Grosso
and Sao Paulo in Brazil have large broadly elliptic to rotund leaves, but in Sao
Paulo individualswith hairy leaves have this leaf shape. In the Distrito Federal
in Brazil, the glabrous individualsusually had smaller leaves, but I also found
individualswith small hairy leaves. There appears to be some gene flow in this
area between this species and B. malifolia, which has small hairy leaves (e.g.,
Gates 415), and possibly also with B. latifolia, which is glabrous(e.g., Prance
and Silva 58167).
Thereis also variationin the pubescence of the upperepidermisin B. variabilis,

from appressed-pubescentto glabrateto velutinous. The velutinous pubescence
occurs mainly in individualsfrom Minas Gerais near Belo Horizonte. Plants of
B. variabilisfrom this area also have smallerleaves, and both of these characters
are found in B. malifolia with which it is here sympatric.It is possible that gene
flow is responsiblefor the convergence of these two species in this area. A single
collection from the Serrado Caiapoin Goias, Irwin & Soderstrom7414, also has
velutinous pubescence on its upper epidermis.
3. Stem pubescence. Throughoutmost of the southern part of its range in
Parana,Sao Paulo, Minas Geraisand parts of Goias, plants of B. variabilis show
velutinous stem pubescence; in Goias and the Distrito Federal, the pubescence
becomes appressed,and in certainindividualsthroughoutthe rangeof the species
the stems are glabrate.
4. Leaf shape and size. The leaves exhibit great variationin shape from ovate
to elliptic or orbicular,with the base truncateto very cordateand the apex acute
to acuminateor apiculate, but I was unable to find any geographicalcorrelation
for this diversity. There is a possible correlationbetween large ovate leaves and
young or regeneratingindividuals, but this statement requires more field observations before it can be offered with any confidence. Individuals from Minas
Gerais near Belo Horizonte, as already stated, have somewhat smallerleaves.
22. Banisteriopsishirsuta B. Gates, sp. nov.
Fruticuluserectus usque 1 m altus. Rami teretes, dense aurei hispido-sericei
vel velutini, lenticellis numerosis punctati subter indumento vix manifestis.
Stipulae minutae vel nullae. Folia opposita; lamina 9.4-15.2 cm longa, 4.8-7.5
cm lata, in inflorescentiaminori, ovata, parumfalcata, basi truncata,apice acuta
vel retusa, margineplana, subtus basi prope costam 2 glandes peltatas 0.9-1.2
87
mm diametro gerenti, supra aureo-hispida,pilis Y-formibus, stipitatis brachio

0.4-0.5 mm longo, subtus hispido-sericea, pilis T-formibus, pede 0.1-0.2 mm
longo, trabecula 2.0-2.6 mm longa, supra plana, subtus venis prominentibus;
petiolo 1-3 mm longo, aureo, hispido-sericeo,eglanduloso.Inflorescentiafloribus
in umbellis4-floriferisfasciculatis, umbellisin dichasiis dispositis, laxa, duplo vel
triploramosa,aureo-hispida;bracteisbracteolisque2.2-3.0 mm longis, adaxialiter
glabris, abaxialiteraureo-sericeis, ante vel per anthesin caducis; pedicellis sessilibus 9-18 mm longis, 1.3-2.2 mm diametro,0.6-1.0 mm sine indumento,aureovelutinis. Sepala 5, abaxialiteraureo-sericea,adaxialitersparsimvelutina distaliter, 4.6-5.0 mm longa, 2.0-3.0 mm lata, elliptica, apice rotundata, 4 lateralia
glandes 2.0-2.8 mm longas et 1.0-1.5 mm latas gerentia, glandes 2.0-3.0 mm
superantia.Petala 5, alba vel subrosea, aetate luteola vel salmonea, dentata, 4
lateraliainter sepala patentia, ungue 1.5-2.0 mm longo, 0.5-0.8 mm lato, limbo
9.5-11.0 mm longo, 8.0-10.0 mm lato, orbiculari, eglanduloso, 2 anterioribus
quam 2 posterioribusconcavioribus;petalumposticum ungue erecto 3.0-4.0 mm
longo, usque 1.4 mm lato versus apicem, margineinvoluto, canaliculato, apice
non constricto, limbo 8.0-9.0 mm longo, 7.0-8.0 mm lato, orbicularivel obovato,
eglanduloso. Stamina 10, filamentis 2.7-4.2 mm longis, proximaliterconnatis,
glabris;antheraeretroflexae,loculis 1.0-1.8 mm longis, glabratisvel apice et basi
pilis caespitosis, connectivis canarinis,petalis oppositis 1.0-1.2 mm longis, 0.60.8 mm latis, 0.1-0.5 mm profundis, parum glandulosis vel solum 2 petalis anterioris oppositis glandulosis, illis sepalis oppositis 1.3-1.9 mm longis, 0.5-0.8
mm latis, 0.4-1.0 mm profundis, glandulosis dilatatisque, praesertim 3 sepalis
anterioribusoppositis. Ovarium 1.0-1.6 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidaliadnatis, candido-sericeis;styli 3, aequales 3.6-3.8 mm longi,
recti parallelique,decrescentes versus apicem usque 0.3-0.4 mm diametro,stigmatibus valde capitatis. Fructus ex 3 samaris(vel abortu 1-2) constans; samara
carpophorousque 2.2 mm longo, 0.4 mm lato, nuce 6-8 mm alta, 4-5 mm longa,
laevi vel parumrugosa, laxe sericea vel glabrata,ala 11-15 mm longa, 8-9 mm
lata, appresso-sericeavel glabrata,pilis T-formibus,trabecularigida0.8-1.2 mm
longa, alis 2 posterioribusad alam anticam subparallelibus;loculo nucis interius
piloso.
Type. Anderson6658, Brazil, Goias, Chapadados Veadeiros, fl (holotype, UB;
isotypes, MICH, NY).
Distribution.(Fig. 17). Known only from a few collections on rocky hillsides
of the Chapadados Veadeiros, Goias, Brazil, 1250-1700m.
Collected in flower in Januaryto March, and in fruit in February.
BRAZIL. Goias: Anderson 11480 fl (MBM); Gates & Estabrook37 fl (MICH, UB); 224 fl fr (MICH,
UB).
This species is distinguishedby its large hispid leaves, thick pedicels and large
flowers.
23. BanisteriopsishatschbachiiB. Gates, sp. nov.
Fruticuluscaulibus non ramosis cernuis ab xylopodio usque 2 m altus. Rami
teretibus, appresso-sericei mox glabrati, lenticellis prominentibuspunctati, demum rimis et fissuris suberosis evolutis. Stipulae minutae vel nullae. Folia opposita vel ternata; lamina 5.6-9.8 cm longa, 2.4-5.4 cm lata, in inflorescentia
minori, ovata, basi truncata subcordatave, apice acuta, margine plana, subtus
basi prope costam 2 glandes sessiles usque 1.0 mm diametrogerenti, suprainitio
appresso-sericea mox glabra, subtus initio laxe sericea, mox glabra, venis pri-
88
Flora Neotropica
mariis supra prominulis, subtus prominentibus,reti venularumutrinqueparum

impresso. Inflorescentiafloribus in umbellis 4-floriferisfasciculatis umbellis in
cymis dispositis in axillis foliorum redactorumin ramis terminalibusgerentis,
sparsim appresso-sericeavel glabrata;bracteis bracteolisque 1.2-2.2 mm longis,
adaxialiter glabris, abaxialiter appresso-sericea, ante, per, vel post anthesin
caducis; pedicellis sessilibus (8-)11-23 mm longis, 0.8-1.0 mm diametro,0.6-0.8
mm sine indumento, appresso-sericeis. Sepala 5, abaxialitersparsim appressosericea, saepe apice glabra,adaxialiterglabravel minute sericea, apice rotundata
et saepe revoluta, sepalum anticum 2.8-3.5 mm longum, 1.7-2.4 mm latum, 4
lateralia3.5-5.0 mm longa, 2.3-3.0 mm lata, glandes 1.3-2.8 mm longas et 0.61.4 mm latas gerentia,glandes 1.5-2.5 mm superantia.Petala5, alba vel subrosea,
aetate luteola vel salmonea, dentatalaciniatave, 4 lateraliainter sepala patentia,
ungue 1.5-2.0 mm longo, 0.4-0.6 mm lato, limbo 8.0-11.5 mm longo, 6.0-8.0 mm
lato, orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribus;
petalum posticum ungue erecto 4.5-5.0 mm longo, usque 1.3 mm lato versus
apicem, apice non contracto,limbo 5.0-7.5 mm longo, 4.0-6.0 mm lato, obovato,
retroflexo, plano vel convexo, eglanduloso. Stamina 10, filamentis 2.5-4.2 mm
longis, proximaliterconnatis, glabris; antherae retroflexae loculis 0.8-1.6 mm
longis, sparsimpilosis pilis basi caespitosis, connectivis canarinis,petalis oppositis 0.6-1.0 mm longis, 0.4-0.6 mm latis, 0.2-0.4 mm profundis, apice parum
glandulosis, illis sepalis oppositis 0.6-1.2 mm longis, 0.5-1.0 mm latis, 0.5-0.9
mm profundis, glandulosis dilatatisque,praesertim3 sepalibus anterioribusoppositis. Ovarium 1.0-1.4 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidali adnatis, candido-sericeis;styli 3, aequales, 3.0-3.2 mm longi, recti parallelique, decrescentes versus apicem usque 0.3-0.4 mm diametro, stigmatibus
capitatis. Fructus ex 3 samaris (vel abortu 1-2) constans; samara carpophoro
usque 2 mm longo, 0.4 mm lato, nuce ovoidea 4-5 mm alta, 2.5-3.5 mm longa,
laevi vel parum rugosa, appresso-pubescentivel glabrata,ala 13-16 mm longa,
8-11 mm lata, appresso-sericeavel glabrata,pilis T-formibus,trabecularecta 0.81.2 mm longa, alis 2 posterioribusad alam anticam subparallelibus;loculo nucis
interiuspiloso.
Type. Hatschbach 36745, Brazil, Goias, mun. Alto Paraiso, fl fr (holotype,
MBM; isotype, MICH).
Distribution.(Fig. 17). Found only on rocky campos of the Chapadados Veadeiros, Goias, Brazil, at 1250-1500m.
Collected in flower in December, Februaryto Marchand May, and in fruit in
May.
BRAZIL. Goias: Barroso et al. 608 fl (NY, RB, UB); Anderson 11493 fl (MICH); Irwin et al. 12419
fl (MICH, NY, UB); 24752 fl (MICH, NY, UB); 33114 fl (UB).
This species is distinguishedfrom B. variabilis by its glabrousfalcate leaves,

its sparsely sericeous sepals and its habit as a decumbent shrublet. It is named
for Dr. Gert Hatschbach,collector of the type, specialist on the floraof southern
Brazil and enthusiasticcollector of all Brazilianplants.
The Banisteriopsis megaphylla group (species 24-25)
The B. megaphyllagroup consists of two species, B. megaphylla, widespread

in the cerradosthroughoutthe Planaltoof Brazil, and B. irwinii, restrictedto the
Chapada dos Veadeiros and Serra Geral do Parana in Goias. They share the
following characters (see Fig. 18 for illustration of flower and fruit type):
large hispido-tomentoseleaves with a large basal pair of glands on the costa and
89
a long thick petiole; sepals broadlyovate, appressedin bud; petals long-fimbriate,

the posterior petal with the limb not constricted; anther locules more or less
pilose; stamen connectives all somewhat glandular, those opposite the sepals
quite enlarged; styles equal, straight and parallel; fruit not rugose, the wing
greyish-red,the locule of the nut densely sericeous within.
Some of these charactersare also typical of the B. argyrophyllagroup (sepal
and petal characters, and pilose antherlocules), and of the B. campestris group
(style and fruit characters,glandularconnectives). In addition, this group shares
the characterof the locule of the fruit nut densely sericeous within with the B.
malifolia group.
24. Banisteriopsismegaphylla(Adr. Jussieu) B. Gates, comb. nov.
Fig. 18.
BanisteriamegaphyllaAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 282. 1840.
Shrubto 2 m tall. Branches terete, densely tomentose, the vestiture complex,

of short- and long-stalkedT-shapedhairs. Stipules minute. Leaves with the petiole (6-)8-12(-18) mm long, thick, tomentose, eglandular,or bearing a pair of
large sessile glands at its apex, the lamina 9.5-19.5(-26.1) cm long, 8.5-16.0
(-22.6) cm wide, smallerin the inflorescence,broadlyovate to ellipticor orbicular,
truncateto cordate at the base, acute to apiculateor emarginateat the apex, the
marginflat, sometimes bearinga pair of large sessile glands on the costa basally,
hispido-velutinousadaxially,the hairs T-shaped,the stalk 0.20-0.35 mm long, the
arms slightly flexuous, 0.2-0.3 mm long, densely tomentose abaxially, the hairs
T-shaped, stalked, the stalk 0.15-1.40 mm long, the trabecula 1.3-2.6 mm long,
flexuous, with the reticulation impressed adaxially, prominentabaxially. Inflorescence of 4-floweredumbels, terminal,the umbels borne in dichasia, very condensed and much branched, tomentose, the trabecula 0.4-0.6 mm long; bracts
and bracteoles 1.0-1.6 mmlong, broadlytriangular,tomentose abaxially,glabrous
adaxially, persistent; pedicels sessile, (5-)7-12 mm long, 1.2-1.4 mm wide, 0.8
mm wide without the hairs, loosely sericeous. Sepals sericeous abaxially, minutely pubescent distally adaxially, appressed in bud, broadly ovate, rounded
and inbent at the apex, 2.0-2.4 mm long, 1.4-2.4 mm wide, the glands red, 2.62.8 mm long, 0.8-1.2 mm wide, the 4 lateralsepals projecting1.2-1.8 mm beyond
the glands. Petals dark pink or white with pink margin,fimbriate,the 4 lateral
petals reflexed between the sepals, the claw 1.5-2.0 mm long, 0.4-0.5 mm wide,
the limb 5.5-6.4 mm long, 5.0-6.0 mm wide, orbicular,eglandular,the anterolateralpetals very cochleate, the postero-lateralpetals slightly concave, the posteriorpetal with the claw erect, 2.5-3.0 mm long, up to 1.4 mm wide at the apex,
not constricted, the limb 3.5-4.0 mm long, 3.0-3.5 mm wide, broadly elliptic,
reflexed, plane to convex, the basal fimbriae enlarged and glandular.Stamens
with the filaments1.5-2.2 mm long, connatebasally, the locules 1.0-1.2 mm long,
basally and apically pilose or glabrate, the connectives yellow, all more or less
glandularapically, those opposite the petals 0.6-0.8 mm long, 0.7-0.8 mm wide,
0.4-0.6 mm deep, those opposite the sepals 0.8-1.0 mm long, 0.9-1.2 mm wide,
0.6-0.8 mm deep. Ovary 0.9-1.2 mm tall, white-sericeous, the styles equal,
straight and parallel, 2.2-2.4 mm long, 0.2-0.3 mm wide, tapering slightly apically, stigmas capitate. Fruit with carpophoreup to 2 mm long, 0.8 mm wide, the
nut 7-9 mm tall, 5-6 mm long, rugose or with one short lateralwing on each side,
appressed-pubescent,grey to green, the wing 13-21 mm long, 11-17 mm wide,
the upper margin straight to slightly arcuate, the lower margin semi-orbicular,
appressed-sericeous,greyish-crimson,the locule hairy within.
Flora Neotropica
90
r9?
i
i'
,:
'
''
?ij
"
..
,..
*:
.-
..
FIG. 18. Banisteriopsis megaphylla group. A-F, B. megaphylla. A, flower bud x 4.2; B, flow-
er x 4.2; C, stamens 1-6, semi-diagrammatic,x 8.4 (stamen 1 is opposite anteriorsepal; stamen 6

is opposite posterior petal); D, lateral view of stamens 1 and 2 x 8.4; E, gynoecium x 8.4; F, samara x 1.8. G, B. irwinii, samara x 1.8. (A-F, Gates 387; G, Anderson 11471.)
?..- ..
91
. .
..
. . .
. ...
*1
phylla; circle, B. irwin i.
Type. Claussen s.n., Brazil, Minas Gerais, Curvello, fl fr (holotype, P; isotypes, P).
Distribution. (Fig. 19). In cerrado on deep red soil throughout the Planalto of
Brazil.
Collected in flower in December to May, and in fruit January to August.
Representative collections. BRAZIL. Mato Grosso: Anderson 11249 fl fr (MICH); Hatschbach
33981 fl (MICH); Philcox & Fereira 3862 fl (K, NY); Ratter et al. R1338 fl fr (IAN, K, MO, NY,
RB); R896 fl (NY). Goias: Anderson 6835 fl fr (MICH, NY); Gardner 3068 fl (BM, K, W); Gates 412
fl (MBM, MICH); Hatschbach & Kummrow 38243 fl (MICH); Irwin et al. 14318 fl fr (NY). Distrito
Federal: Anderson 11426 fl (MICH); Ferreira 34 fl fr (NY); Gates 391 fl fr (MBM, MICH); Irwin et
al. 1184 fl (MICH, NY); Pires et al. 9526 fl (US). Minas Gerais: Atala 59 fl fr (R); Barreto 7606 fl
(F, US); Claussen 542 fl (C, NY, P, S); Duarte 2441 fl fr (RB);_ Glaziou 18938 fl fr (C, K, P);
Magalhaes 1441 fl (UB, US); Mexia 5679 fr (BM, G, GH, MO, NY, US); Paula 58 fl (SP); Warming
s.n. fr (C, GOET).
92
Flora Neotropica
This species is very distinctive with its large greyish hispid-pubescentleaves,

its very congested and much branchedinflorescences, and its small, deep pink
petals with very fimbriatemargins.
The inflorescence of every individualof B. megaphylla which I examined was
frequentedby large numbersof ants which seemed to visit all parts of the inflorescence, but were sometimes seen to feed at the enlargedglands on the reduced
leaves of the inflorescence.
25. Banisteriopsisirwinii B. Gates, sp. nov.
Fig. 18.
Frutex minime ramosus usque 2 m altus vel volubilis usque 5 m; ramis teretibus, pertinaciterdense cinereo-sericeis.Stipulaeusque 1 mmlongae, triangulares,
sericeae, apice glabratae. Folia opposita; lamina 8.3-18.5(-24.0) cm long, 3.38.7(-15.0) cm lata, vel in inflorescentiaminori, elliptica vel late elliptica, basi obtusa truncatave,apice obtusa acutave apiculatave,margineplana, subtus basi in
costa 2 glandes sessiles gerenti, supra sparsim hispida, subtus laxe tomentosa,
venis venulisque supraimpressis, subtusprominentibus.Inflorescentiaramosterminansex dichasiis composita, floribusin umbellis4-floriferisfasciculatis, usque
ter ramificans,laxissima, laxe sericea, trabecula0.6-1.2 mm longa;bracteisbracteolisque 1.5-1.8 mm longis, late triangularibus,abaxialitersericeis, adaxialiter
glabris,persistentibus;pedicellis sessilibus8-12 mmlongis, 1.5-2.0 mm diametro,
0.5-0.6 mm sine indumento,laxe sericeis. Sepala5, abaxialitersericea, adaxialiter
omnino minute pubescentia, late elliptica, apice rotundata, 3.8-4.0 mm longa,
3.0-3.6 mm lata, 4 lateraliaglandes roseas 2.5-3.0 mm longas et 0.8-1.8 mm latas
gerentia, glandes 1.0-1.5 mm superantia.Petala 5, alba vel pallido-rosea,longissime fimbriata,4 lateraliainter sepala patentia, ungue 1.0-1.5 mm longo, 0.6-0.9
mm diametro, limbo 8.5-10.0 mm longo, 5.0-9.0 mm lato, orbicularivel late
obovato, eglanduloso, 2 anterioribuscucullatis, 2 posterioribusplanis vel parum
concavis; petalum posticum ungue erecto 3.2-3.5 mm longo, usque 1.2 mm diametro, apice parum constricto, limbo 6.0-7.0 mm longo, 4.5-6.0 mm lato, late
obovato, retroflexo, plano convexove, eglanduloso. Stamina 10, filamentis2.53.8 mm longis, proximaliterconnatis, glabris; antherae recurvatae, loculis 1.21.6 mm longis, basi et apice pilosis, connectivis canarinis,petalis oppositis 0.81.0 mm longis, 0.8-1.0 mm latis, 0.2-0.3 mm profundis, vix dilatatis et parum
glandulosis, illis sepalis oppositis 1.0-1.4 mm longis, 0.9-1.4 mm latis, 0.8-1.2
mm profundis,dilatatisglandulosisque,globosis, praesertim3 sepalis anterioribus
oppositis. Ovarium0.8-1.0 mm altum, carpellis 3, uniovulatis,liberis, toro pyramidali adnatis, candido-sericeis;styli 3, aequales, 2.8-3.0 mm longi, recti parallelique, glabri, versus apicem decrescentes usque 0.3 mm diametro,stigmatibus
usque 2 mm longo, 0.5 mm lato, nuce ovoidea 8.0-11.0 mm alta, 4.0-6.0 mm
longa, rugosa, appresso-pubescenti,cinerea, ala 18-27 mm longa, 12-17 mm lata,
appresso-sericea, oblonga, marginibussuperioribusinferioribusquesubparallelibus, cinereo-carmesina;loculo interiuspiloso.
Type. Irwin et al. 24705, Brazil, Goias, Chapadados Veadeiros, ca. 20 km S
of Alto Paraiso, fl yng fr (holotype, UB; isotypes, MICH, NY).
Distribution.(Fig. 19). Only on steep rocky slopes of the SerraGeraldo Parana
and Chapadados Veadeiros, Goias.
Collected in flower in Februaryto May, and in fruit in April and May.
BRAZIL. Goias: Anderson 11471 fl (MICH, NY); Gates 409 fl fr (MBM, MICH); 411 fl (MBM,
MICH); Hatschbach 36793 fl (MICH).
93
This species is distinguishablefrom B. megaphyllaby its largerpale pink flowers, its looser less branchedinflorescencewith longer hairs, narrowerleaves, and
its habitaton rocky hillsides. It is named for Dr. Howard S. Irwin, whose extensive collections in the Planaltoregion of Brazil have done much to documentthe
flora of this area.
Banisteriopsis irwinii also differs from B. megaphylla in its vining potential. In
undisturbedhabitats it will vine over other shrubs to form thickets, whereas B.

megaphylla never vines.
The Banisteriopsis campestris group (species 26-30)
The B. campestris group consists of one widespread species, B. campestris,

which occurs in the cerrados throughoutthe Planalto of Brazil, and four other
species with more or less restricted distributionswithin the Serra do Espinhafo
(Figs. 21, 23). All of these species are very similar with regardto flower form,
with only minor differences in size and proportionsbetween them. The species
sharethe followingcharacters(see Fig. 20 for illustrationof flowerandfruittype):
plants never vining;bracteolespersistent;sepals broadlyoblong, white-sericeous,
calyx glands pink to red; petals laciniate, the posterior petal with the claw constricted at the apex; stamen filamentsslender, those opposite the three anterior
sepals longest, those opposite the three posteriorpetals shortest; connectives on
stamens opposite sepals enlarged and glandular, especially opposite the three
anterior sepals; anther locules glabrous; styles straight and parallel; fruit with
rugose nut, fruit wing greyish-red,persistently appressed-pubescent.
26. Banisteriopsiscampestris (Adr. Jussieu) E. L. Little, Phytologia 6: 506.
1959.
Fig. 20.
BanisteriacampestrisAdr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 36. 1832[1833].
Banisteria campestris Adr. Jussieu var. rotundata Grisebach, Linnaea 13: 195. 1839. Type.
Sellow s.n., Brazil, near Aral das Elbas, fr (holotype, B? destroyed).
Banisteria campestris var. glaucescens Grisebach, Linnaea 13: 196. 1839. Type. Sellow s.n.,
Brazil, fr (holotype, B? destroyed;probableisotypes, BR, G, K, W).
Banisteria campestris var. ovata Grisebach, Linnaea 13: 196. 1839. Type. Sellow s.n., Brazil,
fr (holotype, B? destroyed;probableisotypes, BR, K).

Banisteria wettsteinii Kralik, Denkschr. kaiserl. Akad. Wiss., Math.-Naturwiss.Kl. 79: 279.
1908. Types. Wachsmunds.n., Brazil, Sao Paulo, near Sao Bernardo,fl (lectotype, W).
Wacket s.n., Brazil, Sao Paulo, fl (syntype, W). Jose de Campos Novaes s.n., Brazil, Sao
Paulo, Camposde Bocaina, Campinas(n.v.).
Virgate shrublet or shrub to 1.5 m with woody xylopodium. Branches terete,

loosely sericeous, rarely glabrate, the bark dark brown. Stipules minute, triangular, sericeous on both sides. Leaves sometimes ternate or 4 at a node, the
petiole 1-4(-7) mm long, loosely sericeous, eglandular,the lamina3.1-7.5(-11.5)
cm long, 1.5-4.1(-7.8) cm wide, narrowly elliptic to orbicularor ovate, rarely
obovate, smaller in the inflorescence, cuneate to cordate at the base, acute to
apiculateor rarelyemarginateat the apex, with the marginflatto slightlyrevolute,
abaxially with 1-2 pairs of peltate glands beside the costa basally, adaxiallyhispido- to velutino-pubescent,abaxially sparsely to densely tomentose, the hairs
stalked, T-shaped, the trabeculaflexuous to straight,rugose, with the veins impressed adaxially, prominent abaxially. Inflorescence terminal, dischasial, of
4-floweredumbels up to 3(-5) times branched,lax to dense, sericeous, bracts and
bracteoles 1.2-2.2 mmlong, lanceolateto triangular,sericeous abaxially,glabrous
adaxially, persistent; pedicels sessile, 8-17 mm long, 0.8-1.2 mm wide, 0.5-0.8
mm without the hairs, sericeous. Sepals sericeous abaxially, minutely sericeous
Flora Neotropica
94
i''
4?
..:.,.'....-,'
.', '
.-.';d'.'!''..-'/J,.
.'.
;'.
FIG. 20. Banisteriopsis campestris group. A-E, B. campestris. A, flower bud x 4; B, flower x
3.3; C, stamens 1-6, semi-diagrammatic, abaxial and lateral views, x 6.7 (stamen 1 is opposite anterior sepal, stamen 6 is opposite posterior petal); D, gynoecium x 6.7; E, samara x 2.3. F, B.
36
ats
s>amarax 2.3.
.3......................
cioni,smr
(A-D, Gates 357; E. An.derson 678;F,
6738; F, Gates
386.)
cipoensis,
95
adaxially near the base, glabrous apically, ovate, rounded at the apex, 2.6-3.6
mm long, 2.0-2.6 mm wide, the glands red, 1.2-2.2 mm long, 0.6-1.0 mm wide,
the 4 lateralsepals projecting1.5-2.5 mm beyond the glands. Petals pink, becoming paler in age, lacerate, the 4 lateral petals reflexed between the sepals, the
claw 1.0-1.8 mm long, 0.5-0.8 mm wide, the limb 7.0-10.5 mm long and 6.0-10.0
mm wide, orbicular,eglandular,the antero-lateralpetals more concave than the
postero-lateralones, the posteriorpetal with the claw erect, 3.0-4.5 mm long, up
to 1.0 mm wide at the apex, the limb 6.0-9.0 mm long, 5.0-8.5 mm wide, broadly
obovate, reflexed, plane to convex, with the basal fimbriaeenlargedand glandtipped. Stamens with the filaments 1.6-4.0 mm long, connate basally, the locules
glabrous,0.9-1.6 mm long, the connectives yellow, those of the stamensopposite
the petals 0.3-0.9 mmlong, 0.3-0.7 mm wide and 0.1-0.4 mm deep, not glandular,
those of the stamens opposite the sepals 0.6-1.4 mm long, 0.4-1.2 wide, 0.2-1.2
mm deep, with the middle glandularand enlarged, especially in the stamens
opposite the 3 anteriorsepals. Ovary0.8-1.0 mm tall, white-sericeous,the styles
equal, straightand parallel, 3.0-3.8 mm long, taperingapically to 0.3-0.4 mm in
diameter, stigmas capitate. Fruit with carpophoreup to 3.5 mm long, 0.4 mm
wide, the nut 6.0-8.0 mm tall, 4.0-5.0 mm long, rugose to tuberculate,appressedpubescent, grey to green, the wing (11-)15-27 mm long, 9-19 mm wide, appressed-sericeous, the upper marginstraightto arcuate, the lower marginsemicircular.
Type. A. Saint Hilaire s.n., Brazil, Minas Geraisfl (holotype, P; isotypes, P2 sheets).
Distribution.(Fig. 21). Commonin the cerradoon deep red soil throughoutthe
Planaltoregion of Brazil.
Collected in flower in every month of the year, and in fruit in Januaryto May
and August to November.
Representativecollections. BRAZIL. Maranhao:Fr6es 34033 fl (IAN). Bahia: Anderson et al.
36424 fl (NY, UB); 36906 fl fr (MICH, NY); Irwin et al. 14628fr (MICH, NY, UB). Mato Grosso:
Anderson 11204 fl fr (MICH); 11255 fl (MICH); 11396 fl (MICH); Hoehne 5801 fl (R); Philcox &
Fereira 3716 fl (IAN, MO, NY, RB). Goias: Anderson 7905 fl fr (MICH, NY); Barroso et al. s.n. fl
(RB); Heringer 11475 fl (UB); Irwin et al. 10952 fl (NY, US); Ule 421 fl (P, R). Distrito Federal:
Gates 396 fl fr (MBM, MICH); Heringer 13260 fl (NY, RB); Irwin et al. 10719 fl (F, IAN, MO, NY,
S, SP, UB, US); Pires et al. 9025 fl (IAN); 9072 fl (UB). Minas Gerais:Assis 166 fl (R, RB, SP);
Barreto 10298 fl (HB, R); Claussen s.n. fl (BM, BR, F, G, GH, K, P, R, RB); Magalhdes 17173 fl
(IAN, UB); Regnell 1.29pp fl (C, F, NY, P, R, S, U, US); Riedel 837 fl (BM, K, P, W, US). Sao
Paulo: Burchell4602 fl (GOET, K); Felippe 6 fl (NY, RB, SP, US); Glaziou 13602fl (BR, C, G, P,
US); L6fgren 90 fr (R); Martius 1165 fl (BR, G, M, NY, P). Parana: Dusen 11010 fl (BM, GH, MO,
NY, S, US); 14991 fr (F, G, S); Hatschbach 3055 fl (MICH, RB); 6851 fl (MICH); van der Veen 21
fl (U).
This is a very common, widespread and distinctive species, although within

the species there is considerablevariationwith regardto leaf shape and size, type
and density of pubescence on the leaves, degree of branchingof the inflorescence,
and size and shape of the fruit wing. Their appear to be certain syndromes of
characters within this variation, but there are no discontinuities that make a
satisfactorysubdivisionof the species possible. In addition, althoughthese combinations of charactersare usually typical of a particulargeographicalarea, individualswith these charactersmay be found throughoutthe rangeof the species.
A further consideration is that some pubescence variation appears to be age
related;from my own observationsin the field, in the herbariumand on seedlings
I have grown, it is apparentthat the pubescence of the firstformedleaves is very
differentfrom that of later formed leaves. This factor furtherconfuses the situation. Thus it seems that here is a group that needs detailed field observations
Flora Neotropica
96
l{~
{m,
lip
{l~
w
0*
a+
ik,
'
:
+&
l-i
+, +~
,,-1"S.~~
+
W I'
m
ob
FIG. 21. Distribution of part of Banisteriopsis campestris group and B. calcicola, species 26, 28,
31. Circle, B. campestris; square, B. angustifolia; triangle, B. calcicola.
and biosystematic study to establish the nature of its variation. In the absence of
such data, I shall simply record the variation pattern as I have observed it.
1. In rocky campos of the Serra do Espinhago, near Diamantina, decumbent
individuals are common (e.g., Anderson 11558, 35204). In addition, many of these
plants have leaves which are not rugose, only the main lateral veins being impressed above, and have a smaller fruit wing (10-18 mm long, cf. 15-27 mm long
for "typical" campestris), and many have dense sericeous pubescence. However,
in travelling from Diamantina to Sao Joao da Chapada, a continuous transition
was observed between these decumbent, sericeous-leaved individuals to erect
individuals with sericeous pubescence and non-rugose leaves, to plants with rugose leaves, sparse pubescence, and plants with larger fruits. Southwards from
Diamantina, beyond Datas, decumbent individuals were found with small fruit
wings but sparse pubescence and rugose leaves. Since the edaphic conditions of
97
the rocky campos are so unusual,transplantand breedingexperimentsare needed

to determineto what extent the variationis genetic in origin. Any decision about
the taxonomic status of these individualsmust await such data.
2. Near Lagoa Santa and Belo Horizonte in southeasternMinas Gerais, many
specimens have obovate emarginateleaves, longer petioles, hispid leaf pubescence, and a more branchedhabit.
3. Specimens from the Chapadados Veadeiros, Goias, have leaves which are
glabrousto glabrateabaxially,althoughhairs are sometimespresent on the veins.
However, southwardstowards the Distrito Federal, the density of hairs on the
veins increases, and hairs occur on the laminabetween the veins.
4. Some collections from near Corinto, Minas Gerais, and eastwards towards
Diamantina,show muchbranchedinflorescences(4-7 times branched),formlarge
shrubs, and their leaves and branchesare glabrate.
In addition, there are a few specimens of anomalousindividualswhich should
be mentioned. Martius 1165, Riedel 1839 and Claussen s.n. pp have narrowly
lanceolate leaves. They are readilydistinguishedfrom B. angustifolia, which has

narrowlylanceolate leaves, by the sparsepubescence adaxially,the rugose leaves
and the leaf marginwhich is flat or only slightly revolute.
27. BanisteriopsiscipoensisB. Gates, sp. nov.

Fig. 20.
Fruticuluserectus usque 4 dm ab xylopodio ligneo crasso. Ramiteretes, aureosericei. Stipulaeminutae,triangulares,sericeae. Folia oppositavel ternata;lamina
(1.9-)3.2-5.5(-8.0) cm longa, (1.2-)1.8-2.7(-3.5) cm lata, elliptica, basi obtusa
truncatave,apice apiculata,margineplana vel parumrevoluta, subtus basi prope
costam 2 glandes peltatae gerenti, suprapilis stipitatisrigidisT-formibussparsim
hispido-sericea, vetustiora glabrata, subtus pilis T-formibus stipitatis flexuosis
laxe hispido-tomentosa,venis venulisque supra impressis, rugulosa, subtus prominentibus;petiolo 2-5 mm longo, sericeo, eglanduloso. Inflorescentiacaules
terminansex dichasiiscomposita, floribusin umbellis4-floriferisfasciculatis, laxe
sericea; bracteis bracteolisque 1.5-2.2 mm longis, lanceolatis, apice rotundatis,
abaxialitersericeis, adaxialiterglabris, persistentibus;pedunculis0-1.5 mm longis, pedicellis (6-)9-14 mm longis, 0.8-0.9 mm diametro,0.6-0.7 mm sine indumento, laxe sericeis. Sepala 5, abaxialitersericea, adaxialiterbasi minutepubescentia, ovata vel anguste ovata, apice rotundatasaepe revolutaque, 3.0-4.0 mm
longa, 1.6-2.2 mm lata, 4 lateraliaglandes roseas 1.6-2.0 mm longas et 0.8-1.0
mm latas gerentia, 1.6-3.0 mm glandes superantia.Petala 5, rosea, aetate interdum albicantia,fimbriatalaciniatave,4 lateraliainter sepala patentia, ungue 1.82.5 mm longo, 0.5-0.6 mm diametro, limbo 9.0-12.0 mm longo et 8.5-11.0 mm
lato, orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribus;
petalumposticum ungue erecto 4.4-5.5 mm longo, usque 1.2 mm diametro,apice
constricto, limbo 7.0-10.0 mm longo, 6.5-10.0 mm lato, late obovato, retroflexo,
plano convexove, fimbriis basalibus dilatatis glandulosisque. Stamina 10, filamentis 2.0-4.5 mm longis, gracilibus, proximaliterconnatis, glabris. Antherae
loculis 0.9-1.4 mm longis, glabris, connectivis canarinis,petalis oppositis 0.4-1.0
mm longis, 0.3-0.6 mm latis, 0.1-0.4 mm profundis, illis 2 petalis anterioribus
oppositis parumglandulosis,illis sepalis oppositis 0.5-1.4 mm longis, 0.4-1.2 mm
latis, 0.2-1.0 mm profundis, dilatatisglandulosisque,praesertim3 sepalis anterioribus oppositis. Ovarium0.9-1.2 mm altum, carpellis3, uniovulatis,liberis, toro
pyramidaliadnatis, candido-sericeis; styli 3, aequales, 3.2-4.2 mm longi, recti
parallelique,glabri,versus apicem decrescentes usque 0.3-0.4 mm diametro,stigmatibuscapitatis.Fructusex 3 mericarpiis(vel abortu 1-2) constans; mericarpium
Flora Neotropica
98
carpophoro0-0.6 mm longo, non fungenti, nuce ovoidea, 6-7 mm alta, 4-5 mm

lata, laevi vel parum rugosa, appresso-sericea,cinerea vel viridi, ala ad cristam
1.5-6.0(-9.0) mm longam redacta, appresso-sericea,cinereo-carmesina,ab dimidio superiorimarginisabaxialis nucis excrescenti.
Type. Gates 386, Brazil, Minas Gerais, Serra do Cip6, ca. Km 122 on road to
Belo Horizonte from Conceigaodo Mato Dentro, fl fr (holotype, MBM; isotype,
MICH).
Distribution. (Fig. 23). Known only from the Serra do Cipo, Minas Gerais,
Brazil, growing on rocky campo in quartz sand at 1000-1300m.
Collected in flower October to March, and in fruit in March.
BRAZIL. Minas Gerais: Anderson et al. 36159 fl (MICH, NY, UB); Barreto 7609 fl (F, R); Dravassos s.n. fl (RB 169904); Duarte 1958 fl (MICH); Eiten & Eiten 10987 fl fr (MICH); 11086 fl fr
(MICH); Hatschbach & Ahumada 31540 fl (MICH, NY); Irwin et al. 20002 fl (MICH, NY, UB); Joly
1035 fl (SP); Tryon & Tryon 6791 fl (GH, US); Vidal 11.6268 pp fl (R).
This species is distinguishedfrom B. campestrisby its habitas a small shrublet

to 4 dm tall, its small elliptic, hispido-pubescentleaves, and its reduced fruit
wing.
The scarcity of fruitingcollections perhaps reflects the inconspicuousness of

the fruits; since the plants are small, and the fruits are also small and more or
less hidden by the foliage, they are easily overlooked.
28. Banisteriopsisangustifolia(Adr. Jussieu) B. Gates, comb. nov.
BanisteriaangustifoliaAdr. Jussieuin Saint Hilaire,Fl. bras. mer. 3: 38. 1832[1833].
Banisteriamontana Adr. Jussieu in Saint Hilaire,Fl. bras. mer. 3: 37. 1832[1833].Type. Saint
Hilaire, Catal. Bl, 1117, Brazil, Minas Gerais, Morro do Mondo Velho, fl (holotype, P;
isotypes, P-2 sheets).
Banisteria campestrisvar. montana Adr. Jussieu ex Grisebachin Martius,Fl. bras. 12(1):51.
1858.
Banisterialanceolata Niedenzu, Ind. Lect. Lye. Brunsb.p. hiem.: 16. 1900.Types. Pohl 5719,
Brazil, Minas Gerais, Oreas, Rio Jequitinhonha,fl (lectotype, W). Glaziou 13606, Brazil,
MinasGerais,Serrade Caraca,fl (syntype, B? destroyed;isosyntypes, C, K, NY, fragment,
P). Schwacke8367pp fl (syntype, B, destroyed).
Banisteria campestris var. lancifolia Niedenzu, Verz. Vorles. Konig. Ak. Braunsb.: 17. 1912.
Type. Ule 2460, Brazil, MinasGerais, Caraga,fl (lectotype, R).
Subshrubor slender shrub to 1.5 m tall. Young branches terete, appressedsericeous, older branches glabrate, dark purple-brownwith numerouspunctate
lenticels. Stipules up to 0.4 mm long, triangular,sericeous to glabrate. Leaves
divaricate, the petiole 2.0-5.0(-8.0) mm long, appressed-sericeous,eglandular,
lamina (3.0-)8.0-12.5(-15.5)
cm long, 0.2-1.5(-3.1) cm wide, smaller in the inflo-
rescence, linear to narrowlyelliptic (length:width= 3-30:1), obtuse to truncate

at the base, acute at the apex, with the marginflat to strongly revolute, bearing
a single pair of peltate glands basally on the lamina beside the costa, rarely on
the basal lateralveins, adaxially sparsely appressed-sericeous,abaxiallydensely
white-sericeo-tomentose,the vestiture of two kinds of hairs, the outermosthairs
T-shaped, the trabecula straight, 1.8-2.2 mm long, the stalk 0.1-0.2 mm long,
overlying a layer of T-shapedhairs, the trabeculaflexuous, 0.6-1.0 mm long, the
stalk 0.04-0.08 mm long, rugulose, with the primary and secondary veins impressed adaxially, prominent abaxially. Inflorescence terminal, dischasial, of
4-floweredumbels, up to twice branched,loose, appressed-sericeous;bracts and
bracteoles 1.0-2.2 mm long, narrowlyelliptic, the apex rounded, appressed-sericeous abaxially, glabrous adaxially, persistent; pedicels sessile, (9-)13-20 mm
long, 0.5-0.7 mm wide, 0.4-0.5 mm wide without the hairs, appressed-sericeous.
99
Sepals appressed-sericeousabaxially, sometimes glabrateapically, adaxiallyminutely pubescent near the base, rounded and somewhat appressed at the apex,
the 4 lateral sepals 2.5-4.0 mm long, 1.8-2.5 mm wide, projecting2.5-3.0 mm
beyond the glands, the glands red, 1.8-2.5 mm long, 0.8-1.3 mm wide, the anterior sepal 2.5-3.5 mm long, 1.8-2.2 mm wide. Petals pink, becomingpaler with
age, not keeled, laciniate, the 4 lateral petals reflexed between the sepals, the
claw 1.0-1.5 mm long, 0.4-0.7 mm wide, the limb (6.0-)8.5-10.0 mm long,
(6.5-)8.0-10.0 mm wide, orbicular,eglandular,the antero-lateralpetals more concave than the postero-lateralones, the posteriorpetal with the claw erect, 3.0-4.0
mm long, apically constricted, up to 1.2 mm wide, thick and fleshy, the limb
(5.0-)7.5-8.0 mm long, (5.0-)7.5-9.0 mm wide, broadly obovate, reflexed, convex, with the basal laciniaeenlargedand gland-tipped.Stamenswith the filaments
1.8-4.0 mm long, basally connate, the locules 0.8-1.1 mm long, glabrous, the
connectives yellow, those of the stamens opposite the petals 0.5-0.8 mm long,
0.3-0.6 mm wide, 0.1-0.6 mm deep, sometimes slightly glandularin the middle,
those of the stamensopposite the sepals 0.8-1.6 mm long, 0.7-1.4 mm wide, 0.51.4 mm deep, with the middle enlargedand glandular,especially in the stamens
opposite the 3 anteriorsepals, those opposite the antero-lateralsepals overtopping
the locules by 0.2-0.7 mm. Ovary white-sericeous, 0.8-1.3 mm tall, the styles
equal, straight and parallel, 3.0-4.0 mm long, taperingapically, 0.3-0.4 mm in
diameter, stigmas capitate. Fruit with carpophoreup to 3.5 mm long, 0.5 mm
wide, the nut 8.5-10.0 mm tall, 5.5-6.5 mm long, rugose to tuberculate, appressed-sericeous, the wing 13-22 mm long, 12-17 mm wide, appressed-sericeous, the upper marginerect to widely diverging,the lower marginmeeting the
nut at the base or up to halfway along its distal edge.
Type. Saint Hilaire, Catal. B], 428, Brazil, Minas Gerais, Santa Quiteria, fl
(lectotype, P). Saint Hilaire Catal. Bl, 1142, Brazil, Minas Gerais, Morro
d'Andaia, fl (syntype, P).

Distribution.(Fig. 21). Commonthroughoutthe Serra do Espinhagoon rocky
slopes from Bahia southwardto near Belo Horizonte in Minas Gerais, alt 10001600m.
Collected in flower December to April, and in fruit, Februaryto June.
Representativecollections. BRAZIL.Bahia:Fr6es 20238fl fr (IAN, NY); Harley 15856fl (MICH);
16789fl fr (MICH);Irwin et al. 30909 fl fr (MICH, NY); 31081 fl (MICH, NY, UB). MinasGerais:
Anderson 8445 fl fr (MICH, NY); Belem & Mendes 375 fl (IAN, RB, UB, US); Gates 335 fl fr
(MICH); 351 fl (MICH); Glaziou 14567 fl (BR, C, G, NY, fragment, P); Hatschbach & Ahumada
31546 fl (MICH, NY); Irwin et al. 29562 fl (NY, UB); Joly 1122 fl (SP); Machado 47 fl (HB);
Magalhaes 1219 fl (IAN, UB); Martius 1127 fl (M); Pohl 3056 fr (F, W); Ule 2458 fl (R).
This species is very variable with regardto leaf size and shape, varyingfrom
narrowlyelliptic (L:W = 3:1), to linear (L:W = 12-30:1). Individualsrecognised
as B. montana in the past representone extreme of this variation(the relatively
short-leavedindividuals),but there appearsto be a continuumof variationfrom
individuals with very long, narrow leaves to individuals with narrowly elliptic
leaves. I was unable to find any geographicalbasis for this variation. I found
individuals with the whole range of leaf size and shape growing together on a
single hillside, and I could find no correlationbetween leaf variabilityand other
variable characters, such as flower size or fruit size and shape. Also, I found
individualsshowing variationin leaf size and shape on the same plant; Gates 335
was a shrub with very long linear leaves (L:W = 30:1), but some new shoots
from the base bore much broaderlanceolate leaves (L:W = 6-10:1).
Gates 351 is unusual in its very small flower size, but this specimen was collected in northeasternMinasGeraisin 1976duringa severe drought,so that water
Flora Neotropica
100
4.1~~
__
Irwin
et
al.
20904a
; D,
Irwin
et
al.
~i
?~~~~~-s
20827~~~~~~~~~~~~~~~~~~~~~~~~~~.)
FIG. 22. Banisteriopsis andersonii and B. arborea. A-C, B. andersonii. A, habit x 0.5; B,
flower x 0.75; C, samara x 2.5. D, B. arborea, branch with fruits x 0.5. (A-B, Anderson 35203; C,
Irwin et al. 20904a; D, Irwin et al. 20827.)
101
stress may be responsiblefor the size variation.Two specimens from near Ouro
Preto in southernMinas Gerais, Irwin29562 and Magalhaes 1219, are interesting
in that their leaves abaxiallyshow only the short-stalked,flexuous hairs found as
the inner layer in typical B. angustifolia. This gives the leaves a loose silver-grey
tomentose pubescence.
29. BanisteriopsisandersoniiB. Gates, sp. nov.
Fig. 22.
Suffrutexvel arbusculagracilisusque 2 m alta, erecta salignaque.Ramijuniores
teretes, appresso-sericei,rami vetustiores glabrati,cortice rubiginosovel fusco,
lenticellis numerosispunctato. Stipulae minutaetriangulares,utrinqueappressosericeae. Folia erecta, opposita vel ternata,lamina2.0-9.0(-13.0) cm longa, 0.21.0 cm lata, vel in inflorescentiaminori,anguste ellipticavel lineari, basi truncata
cordatave, apice acuta, margine revoluta, subtus basi prope costam 2 glandes
peltatas gerenti, supra pubescenti mox glabrata, subtus glabra praeter costam
(appresso-sericeaut in petiolo), venis primariissupra parum impressis, subtus
prominulisvel prominentibus;petiolo 1.5-5.0 mm longo, appresso-sericeo,eglanduloso. Inflorescentiaramiterminansex dichasiis composita, floribusin umbellis
4-floriferisfasciculatis, semel vel bis composita, appresso-sericea;bracteis bracteolisque 1.7-2.0 mm longis, triangularibus,abaxialiterappresso-sericeis,adaxialiter glabris, persistentibus;pedicellis sessilibus, (9-)13-20(-25) mm longis, 0.60.8 mm diametro,0.5-0.6 mm sine indumento,appresso-sericeis.Sepala 5, abaxialiter sericea, adaxialiterbasi minutepubescentiaerecta, apice obtusa revolutave,
3.0-3.8 mm longa, 2.2-2.4 mm lata, sepala 4 lateraliaglandes rosas 2.5-3.0 mm
longaset 1.0-1.2 mmlatasgerentia,glandes 1.2-1.5 mmsuperantia.Petala5, rosea,
aetate interdumalbicantia, limbo orbiculari, marginelaciniato, 4 lateralia inter
sepala patentia, ungue 1.0-1.5 mm longo, 9.0-10.5 mm lato, eglanduloso, 2 anterioribus quam 2 posterioribusconcavioribus;petalum posticum ungue erecto
3.8-4.0 mm longo, usque 1.0 mm diametro, apice constricto, limbo 9.0-9.6 mm
longo, 9.3-10.0 mm lato, retroflexo,plano convexove, laciniis basalibusdilatatis
glandulosisque.Stamina 10, filamentis2.5-4.0 mm longis, basaliterconnatis, glabris; antherae loculis 0.9-1.4 mm longis, glabris, connectivis canarinis, petalis
oppositis 0.3-0.6 mm longis, 0.4-0.6 mm latis, 0.1-0.4 mm profundis, solum
connectivis petalis anterioribusoppositis leviter glandulosis,illis sepalis oppositis
0.7-1.4 mm longis, 0.5-1.0 mm latis, 0.4-1.3 mm profundis,dilatatisglandulosisque. Ovarium0.9-1.0 mm altum, carpellis3, uniovulatis,liberis, toro pyramidali
adnatis, candido-sericeis.Styli 3, aequales, 3.0-3.4 mm longi, recti parallelique,
versus apicem decrescentes, glabri, stigmatibuscapitatis. Fructus ex 3 samaris
(vel abortu 1-2) constans; samaracarpophorousque 3.5 mm longo, 0.4 mm lato,
nuce ovoidea 7.5-8.0 mm alta, 4.0-5.0 mm longa, rugosa vel rarius muricata,
appresso-sericea,ala (10-)14-24 mm longa, (6-)10-12 mm lata, pertinaciterappresso-sericea, cinereo-carmesina.
Type. Anderson 8383, Brazil, Minas Gerais, Serra do Espinhago, 10 km by

road NE of Diamantinaon road to Rio Jequiti, fl fr (holotype, UB; isotypes,
MICH, NY).
Distribution.(Fig. 23). Mainly collected in rocky hillsides at 1100-1400 m in
the Serra do Espinhago, from Diamantinasouthwardto the Serra do Cipo. One
collection, Glaziou 18164, is supposed to be from Sabara, near Belo Horizonte,
Minas Gerais, and Lutzelburg7165 is supposedly from the Rio Doce in Espirito
Santo.
Collected in flower Januaryto July, and in fruit Februaryto July.
BRAZIL. Minas Gerais: Anderson 8500 fr (MICH, NY, UB); 11594 fl (MICH); Anderson et al.
35203 fl (MICH, NY, UB); 36124a fl (MICH, NY, UB); Brade 13772 fl fr (RB); Duarte 7879 fl (MICH,
102
Flora Neotropica
t
-44
-0Q
'
A.
..
40" .
' ' '

*
**
** **/ . ;7.
i0
23.
FIG.
Distribution
cipoesis;closd
FIG
of
of part
cirle,
Banisteriopsis
anerson\
group,
campestris
cicle,
opn
arore
27,
species
29,
30.
B.~~~~~~~~
Triangle,
Distribution of part of Banisteriopsis campestris group, species 27, 29, 30.
angle, B.
fl fr (MBM,MICH);
NY, RB);Eglers.n.fl (RB);Gardner4472fl fr (BM,K); Gates351Glauou~~~~~~~~~~~~~~'~i:
18164
Irwin
(BM,
fl (R);
Hatschbach
pp
et
al. 20904a
GH,
NY,
Thispecie
fr (NY);
fl
US); Pereira
&
Aumada
27976
1720
fl
is railditgisefrm
..ve
.hc
glabrous~~~~~~.
.r
31631
fl (MICH,
fr
(RB).
...... ....t
fl
(MICH,
UB);
Espirito
2267^
Santo:
NY);
Heriger
fl (MICH,
&
NY,
Lutzelburg
7165
. nutfiabissh
.;dno,..:-cae,its
Castellanos
UB);
fl
f"'(R'22163
Mexia
...
f:
5732...p.
(M).~.
tr,
fr
alos
Z.'wes
; ' it
SCALE
ciponsis;
closed circle, B. andersonii; open circle, B. arborea.
NY, RB); Egler s.n. fl (RB); Gardner 4472 fl fr (BM, K); Gates 351 fl fr (MBM, MICH); Glaziou
18164 pp fl (R); Hatschbach & Ahumada 31631 fl (MICH, NY); Heringer & Castellanos 22163 fl (R);
Irwin et al. 20904a fl fr (NY); 21916 fl (MICH, UB); 22678 fl (MICH, NY, UB); Mexia 5732 pp fl fr
(BM, GH, NY, US); Pereira 1720 fl fr (RB). Espirito Santo: Liitzelburg 7165 fl (M).
This species is readily distinguishedfrom B. angustifolia by its shorter, almost

glabrous leaves which are held erect and not divaricate, its larger flowers with
103
erect sepals, and its habit as a slender erect treelet. AlthoughB. angustifolia will
become quite large and woody in protected situations, its appearanceis quite
different, being spreadingand bushy, unlike the willowy, wand-like habit of B.
andersonii. I found a single plant, Anderson 11595, on a burnt and cut-over
hillside which appearsto be a hybridbetween B. angustifolia and B. andersonii;
it has leaves which are relatively short, sparsely pubescent abaxially, and which
are held suberect. The plant had few flowers and was not setting fruit. I have
seen no other individualswhich could be described as hybrids.
The Liitzelburgspecimen is unusualin its very long leaves (up to 13 cm), and
its supposed location in Espirito Santo is unexpected. It is possible that the
collection is from the Rio Doce fartherinland in Minas Gerais.
This species is named for Dr. WilliamR. Anderson, foremost student of the
Malpighiaceaetoday, whose help and encouragementmade this treatmentpossible.
30. BanisteriopsisarboreaB. Gates, sp. nov.
Fig. 22.
Arbusculagracilisusque 4 m alta, non volubilis. Ramijuniores teretes, cinereovel aureo-sericei,ramivetustioresglabrati,cortice rubiginosovel fusco, lenticellis
numerosispunctato. Stipulaeusque 0.5 mm longae, triangulares,penitus sparsim
sericeae vel glabratae.Folia opposita;lamina 1.7-6.1(-9.0) cm longa, 0.8-3.3 cm
lata, vel in inflorescentiaminori,ellipticavel lanceolata, obovata in renovationes,
basi obtusa truncatave,apice acuta apiculatave, margineplana revolutave, 2(-4)
glandes peltatas basi in nervis lateralibussubtus gerenti, suprasparsimappressosericea, subtus laxe tomentosa(rariussparsim),pilis stipitatis,trabeculaflexuosa,
venis primariissupraimpressis, subtusprominentibus;petiolo 3-5(-8) mm longo,
cinereo-sericeo, eglanduloso.Inflorescentiaramiterminansex dichasiis composita, floribus in umbellis 4-floriferisfasciculatis, laxa densave, appresso-sericea;
bracteis bracteolisque (1.2-)1.6-2.5 mm longis, lanceolatis, apice rotundatis,
abaxialiter sericeis, adaxialiterglabris, persistentibus vel in fructu cadentibus;
pedunculis 0-1.5 mm longis, pedicellis 12-21 mm longis, 0.7-1.2 mm diametro,
0.5-0.6 mm sine indumento, appresso-sericeis. Sepala 5, abaxialiter sericea,
adaxialiterbasi minute pubescentia ovata, apice rotundata, 2.8-3.2 mm longa,
2.0-2.7 mm lata, 4 lateraliaglandes roseas 1.6-2.0 mm longas et 0.8-1.0 mm latas
gerentia, glandes 1.2-1.5 mm superantia. Petala 5, rosea, aetate interdum albicantia, laciniata,4 lateraliainter sepala patentia, ungue 1.0-1.4 mm longo, 0.30.4 mm diametro,limbo 7.5-10.0 mm longo, 8.0-10.0 mm lato, orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribus concavioribus; petalum posticum
ungue erecto 3.5-4.0 mm longo, usque 1.0 mm diametro,apice constricto, limbo
6.0-7.0 mm longo latove, late obovato, retroflexo,plano convexove, laciniis basalibus dilatatis glandulosisque. Stamina 10, filamentis 1.2-3.8 mm longis, gracilibus, proximaliterconnatis, glabris;antheraeloculis 0.9-1.4 mm longis, glabris,
connectivis canarinis,petalis oppositis 0.6-0.8 mm longis, 0.4-0.6 mm latis, 0.20.5 mm profundis,eglandulosis,illis sepalis oppositis 0.7-1.3 mm longis, 0.4-1.0
mm latis, 0.4-1.0 mm profundis, dilatatis glandulosisque, praesertim 3 sepalis
anterioribusoppositis. Ovarium0.9-1.2 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidaliadnatiscandido-sericeis;styli 3 aequales, 3.0-3.2 mm longi,
recti parallelique,glabri, versus apicem decrescentes usque 0.25 mm diametro,
stigmatibuscapitatis. Fructus ex 3 samaris (vel abortu 1-2) constans; samara
carpophorousque 2.5 mm longo, 0.6 mm lato, nuce ovoidea, 6.0-8.0 mm alta,
4.0-5.0 mm longa, rugosa tuberculatave,plerumqueutrinquealulamlateralem23 mm altamad areolamparallelamgerenti, appresso-sericea,ala 18-32 mm longa,
104
Flora Neotropica
13-19 mm lata, margine dorsali recta curvatave, margine ventrali arcuata semiorbiculatave,pertinaciterappresso-sericea,cinereo-carmesina.
Type. Anderson 11564, Brazil, Minas Gerais, Serra do Espinhago, ca. 10 km
SW of Diamantinaon BR-259, fl fr (holotype, MBM; isotype, MICH).
Distribution.(Fig. 23). On rocky hillsides at 1000-1300m in the Serra do Espinhago in Minas Gerais, from Diamantinasouthwardto the Serra do Cipo.
Collected in flower Januaryto March, and in fruit Januaryto April.
BRAZIL. Minas Gerais: Anderson 8971 fr (MICH, NY, UB); 11585 fl (MICH, NY); 11601 fl
(MICH); Anderson et al. 35191 fl (MICH, NY, UB); 35506 fl (MICH, NY, UB); Egler s.n. fl (RB);
Gates 359 fl fr (MBM, MICH); 361 fl fr (MBM, MICH); 378 fr (MBM, MICH); Irwin et al. 20827 fl
fr (MICH,NY, UB); 22747fl fr (MICH,NY, UB); 27088fr (MICH,NY, UB); 27954fr (MICH,NY,
UB); 28271 fr (MICH, NY, UB).
This species is readilydistinguishedfrom B. campestrisby its habitas a slender

tree, its smaller ovate to elliptic leaves with glands on the lateral veins but not
on the costa at the base, and its largerfruits, usually with a lateral wing on the
nut. It is distinguishedfrom B. andersonii by its wider and shorter pubescent
leaves which are not held erect, and its largerfruit.
Gates 361 and Anderson 11585 from the valley of Biribiri, near Diamantina,
have longer (up to 9.0 cm) lanceolate leaves which are only very sparsely pubescent below, and a laxer inflorescence.However, Gates 359, also from Biribiri
valley, is somewhatintermediatebetween these two specimens and "typical" arborea in its sparserpubescence, largerleaves and laxer inflorescence. Until more
is known about the variationwithinthis species, it would seem prematureto give
these almost glabrous-leavedindividualstaxonomic recognition.
31. BanisteriopsiscalcicolaB. Gates, sp. nov.
Frutex volubilis vel liana. Rami teretes, aureo-sericei,demumglabrati,cortice
brunnea,lenticellis dispersis punctato. Stipulaeusque 1 mm longae, lineares vel
lanceolatae, sericeae. Folia opposita lamina (3.3-)5.0-10.8 cm longa, (1.6-)2.95.5 cm lata, in inflorescentiaminori, elliptica vel in inflorescentiaorbiculari,basi
obtusa truncatave,apice acuta apiculatave, marginerevoluta, subtus basi prope
marginem2-4 glandes stipitates 0.4-0.5 mm diametro,stipite 0.6-1.0 mm longo,
in nervis lateralibusgerenti, supra breve velutina, pilis sessilibus, subtus laxe
velutina vel tomentosa, pilis stipitatis, pede 0.15-0.30 mm longo, trabecula0.30.6 mm longo, venis venulisque supra impressis, subtus prominentibus,petiolo
2-7 mm longo, sericeo, eglanduloso.Inflorescentiafloribusin umbellis4-floriferis
fasciculatis, umbellisin cymis vel dichasiis dispositis, sericea;bracteisbracteolisque 2.0-2.2 mm longis, anguste ovatis, abaxialitersericeis, adaxialiterglabris,
per anthesin caducis; pedicellis sessilibus, 11-14 mm longis, 0.8 mm diametro,
0.4-0.5 mm sine indumento, appresso-sericeis. Sepala 5, abaxialiter sericea,
adaxialiterminute velutina, apice glabra, elliptica, apice rotundaet interdumrevoluta; sepalumanticum3.2-3.8 mm longum, 2.0-2.4 mm latum, 4 lateralia3.84.4 mm longa, 2.5-3.8 mm lata, glandes 2.3-2.6 mm longas et 0.5-1.0 mm latas
gerentia, glandes 1.5-1.8 mm superantia.Petala 5, rosea?, fimbriata,4 lateralia
inter sepala patentia, ungue 1.0-1.8 mm longo, 0.4-0.6 mm lato, limbo 7.5-8.5
mm longo, 6.5-8.0 mm lato, orbiculari,2 anterioribusquam 2 posterioribusconcavioribus; petalum posticum ungue erecto 2.8-3.4 mm longo, usque 1.2 mm
diametro, apice constricto, limbo 4.5-6.5 mm longo, 3.5-5.0 mm lato, obovato,
retroflexo,plano convexove, fimbriisbasalibusglandulosis. Stamina 10, filamentis 1.5-4.2 mm longis, proximaliterconnatis, 3 posterioribusflexuosis sed non
inter stylos posticos inflexis; antherae loculis 0.6-1.2 mm longis, glabris, con-
105
nectivis petalis oppositis 0.3-0.8 mm longis, 0.4-0.6 mm latis, 0.2-0.6 mm profundis, non vel parum glandulosis, sepalis oppositis 0.5-1.9 mm longis, 0.4-1.0
mm latis, 0.3-1.0 mm profundis, 3 sepalis anterioribusoppositis glandulosis et
dilatatis, illis 2 sepalis antero-lateralibusoppositis loculos 0.9-1.3 mm superantibus. Ovarium1.0-1.2 mm altum, carpellis3, uniovulatis,liberis, toro pyramidali
adnatis, candido-sericeis;styli 3, aequales, 2.8-4.0 mm longi, recti parallelique,
glabri, stylo antico 0.4-0.5 mm lato, stylis posticis 0.3 mm latis, stigmatibus
usque 1.5 mm longo, 1.0 mm lato, nuce 7-9 mm alta, 6-8 mm longa, tuberculata
vel manifeste alulata, alulis usque 10 mm altis, appresso-pubescenti,ala 20-33
mm longa, 10-15 mm lata, oblonga vel late oblonga, appresso-sericea,pilis sessilibus persistentibus,T-formibus,trabeculaparumflexuosa 0.18-0.30 mm longa,
hebetato-carmesina.
Type. Anderson 9198, Brazil, Minas Gerais, 22 km by road W of Januaria,on
road to Serra das Araras, fr (holotype, UB; isotypes, MICH, NY).
Distribution.(Fig. 21). Found only in northeasternBrazil, in association with
limestone outcrops in northeastern Minas Gerais, Bahia, Piaui and Ceara, at
forest marginsor as a vine over trees in the cerrado, at altitudes of 500-650 m.
One collection, Gounelle s.n., is supposedly from Caragain Minas Gerais; it is
possible that B. calcicola occurs rarelyfarthersouth in the Planalto.
Collected in flower in Januaryand May, and in fruit in Marchto May.
BRAZIL. Piaui: Gardner 2071 fl fr (BM, K, W); Ule 7445 fl (G, K). Ceara: Allemao & Cysneiros
213 pp fl (R). Bahia: Anderson et al. 37133 fr (MICH, NY, UB). Minas Gerais: Gounelle s.n. fl (P).
State unknown: Sellow 1841 fl (W); Prinz von Neuwied s.n. fl (BR).
Banisteriopsiscalcicola is interestingin that it shares many characterswith the

B. campestris group, such as the form of its fruit and many aspects of its flower.
However, B. calcicola is a vine, and has greatly enlarged connectives on the
stamens opposite the antero-lateralsepals as in the B. muricata group, and the
stalked laminaglands, long petioles and lateralwinglets on the nut are characters
found also in B. adenopoda. Thus B. calcicola shares characterswith several
other species.
Banisteriopsis calcicola can be distinguished from B. campestris by its vining
habit, its enlargedconnectives on the antero-lateralsepals, stipitate leaf glands,

caducous bracts, and prominentlateral wings on the fruit nut; it can be distinguished from B. adenopoda by its ruguloseleaves, revolute leaf margins,straight
styles, glabrousantherlocules and dull red fruit-wing.
Where adequate field data are available, B. calcicola appears to occur in association with limestone outcrops, which is the basis for my choice of specific
epithet. It is interestingthat the supposed occurrence in southern Minas Gerais
is also from a limestone area.
The Banisteriopsis membranifolia group (species 32-34)
This group of three species are all vines occurringin the Atlanticcoastal forest
of SE Brazil and in Mato Grosso and Paraguay(Fig. 25).
They share the following characters(see Fig. 24 for illustrationof flower and
fruit type): vines; leaves with one to several pairs of glands on the lateral veins
near the margin;inflorescence of large dense cymes of four-floweredumbels;
bracts and bracteoles caducous before, during or immediately after flowering;
sepals rounded and often revolute at the apex, minutely pubescent throughout
adaxially, stamens with filamentsopposite the anteriorsepal and postero-lateral
petals longest, antherlocules pilose, connectives on stamens opposite the antero-
106
Flora Neotropica
lateral sepals enlarged and glandularand somewhat overtopping the locules;

styles stout, posterior styles somewhat longer than the anteriorstyle, diverging,
lyrate at the base; nut of the samarawith prominentwell-developedlateralwings.
32. Banisteriopsismembranifolia(Adr. Jussieu) B. Gates, comb. nov.
Fig. 24.
Banisteria membranifolia Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 39. 1832 [1833].
Banisteria membranifolia forma subglabrata Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem.: 23.
1900. Type. Glaziou 18950, Brazil, Rio de Janeiro, Alto Macah6, fl (holotype, B? destroyed;
isotypes, C, K, NY-fragment, P).
Liana or vining shrub, the young branchesflattened,the older branchesterete,

appressed-sericeous or velutinous. Stipules triangular, 0.5-1.0 mm long, appressed-sericeous. Leaves with the petiole 11-20(-27) mm long, appressed-sericeous or velutinous, bearing 1-2 pairs of stipitatepeltate glands, the lamina4.712.9 cm long, 2.7-6.3 cm wide, smaller in the inflorescence, elliptic to ovate,
obtuse at the base, acuminateat the apex, with the marginflat, sometimesbearing
1-2 pairs of peltate glands on the lateralveins near the marginabaxially, sparsely
appressed-sericeousto glabrateadaxially,appressed-sericeousabaxially,with the
venation prominulousto flat adaxiallyand prominentabaxially. Inflorescenceof
4-6-flowered umbels borne in cymes and forming large dense inflorescences,
sericeous or velutinous;bracts and bracteoles 1.5-2.2 mm long, narrowlyoblong,
sericeous abaxially, glabrousadaxially, caducous in bud. Pedicels sessile, 11-17
mm long, 0.6-0.8 mm wide, 0.6 mm without the hairs, appressed-sericeousto
velutinous. Sepals sericeous abaxially,minutelytomentose throughoutadaxially,
oblong, rounded at the apex, 3.0-3.5 mm long, 1.8-2.5 mm wide, the 4 lateral
sepals bearingglands 1.6-2.5 mm long, 0.8-1.0 mm wide, projecting1.5-2.0 mm
beyond the glands. Petals pale pink or white, becomingpale yellow in age, fimbriate, the 4 lateral petals reflexed between the sepals, the claw 0.9-1.2 mm long,
0.6-0.9 mm wide, the limb 7.0-10.0 mm long, 5.5-9.0 mm wide, orbicular,eglandular, the antero-lateralpetals more concave and largerthan the postero-lateral
petals, the posterior petal with the claw erect, 3.0-3.5 mm long, 1.2-1.5 mm
wide, not constricted at the apex, the limb 5.5-7.0 mm long, 4.5-6.0 mm wide,
broadlyobovate with the basal fimbriaegland-tipped.Stamens with the filaments
1.8-3.2 mm long, connate basally, the posterior 3 flexuous and inflexed between
the posterior styles, the locules pilose, 0.9-1.6 mm long, the connectives yellow,
those of the stamens opposite the petals 0.6-1.4 mm long, 0.6-0.9 mm wide, 0.20.6 mm deep, more or less glandular,those of the stamens opposite the sepals
0.8-1.8 mm long, 0.6-1.2 mm wide, 0.4-1.1 mm deep, all or only those opposite
the 3 anterior sepals glandularand enlarged, the connectives of the stamens
opposite the antero-lateralsepals projectingbeyond the connectives 0.2-0.8 mm.
Ovary 1.0-1.2 mm tall, white-sericeous, the anteriorstyle straight, 3.2-3.4 mm
long, 0.8 mm wide, the posteriorstyles straightand diverging,lyrate basally 3.63.8 mm long, 1.0 mm wide at the base, taperingto 0.3 mm wide at the apex, the
stigmascapitate. Samarawith the carpophoreup to 1.5 mm long, 1 mm wide, the
nut 7-9 mm tall, 5-7 mm long, alulate with the winglets up to 10 mm high,
appressed-pubescent,green, the wing 25-40 mm long, 10-18 mm wide, appressedsericeous, the hairs sessile, T-shaped, the wings of the posterior samarassomewhat rotated to lie more nearly parallelto the wing of the anteriorsamara.
Type. Saint Hilaire Catal. ?, 705, Brazil, Minas Gerais, As Bicas, near Itabira
de Mato Dentro, fl (holotype, P; isotypes, P-2 sheets).
Distribution.(Fig. 25). Atlantic coastal forest of Bahia, SE Minas Gerais, Es-
107
FIG. 24. Banisteriopsismembranifolia.A, flower bud x 4.2; B, flower x 4:2; C, stamens 1-6,
abaxialand lateralviews, x 8.4 (stamen I is opposite anteriorsepal, stamen6 is
semi-diagrammatic,
opposite posteriorpetal); D, gynoecium x 7; E, samara x 2. (A-D, Anderson11713;E, Gates 382.)
108
Flora Neotropica
pirito Santo, Rio de Janeiro and Sao Paulo, Brazil. Two collections from Amazonas, Brazil, may also belong to this taxon.
Collected in flower in Februaryto June and in fruit in Marchand April.
BRAZIL. Amaz6nas: Krukoff 8511 fl (BM, BR, F, G, GH, K, MICH, MO, NY, P, S, U, US);
8550 fl (BM, BR, F, G, GH, K, MICH, NO, NY, P, S, U, US). Bahia: Eupunino 255 fl (MICH);
Raimundo 1122 fl (MICH). Minas Gerais: Anderson 11732 fl fr (MICH); Araujo s.n. fl (R); Damazio
328 fl (G); Gates 380 fl (MICH); 381 fl (MICH); 382 fr (MICH); 383 fl fr (MICH); Glaziou 18948 fl
(R); Irwin 2681 fl (NY, F); Mexia 4396 fl (BM, F, G, GH, K, MICH, MO, NY, S, U, US); 4641 fr
(BM, F, G, GH, K, MICH, MO, NY, S, U, US); Rabello s.n. fl (R); Schwacke 7358 fl (RB); 7673
fl (NY, fragment); 10397 fl (P); Sellow III it.B1839c.1305 fl (NY, fragment). Espirito Santo: Anderson 11730 fl (MICH). Rio de Janeiro: Anderson 11713 fl (MICH); Duarte 4256 fl (RB); Frazdo s.n.
fl (RB); Glaziou 10357 fl (BR, X, F, G, K, NY, fragment); Sucre 2661/Braga 502 fl (HB, MICH,
UB); J. Vidal 11.6777 fl (R). Sao Paulo: Frazdo s.n. (RB).
There is some variationin this species with regardto leaf size and pubescence,
the pubescence in the inflorescence and flower size. Individuals segregated as
formasubglabratahave leaves which are abaxiallyglabrousor glabratecompared
with the sparse appressed-pubescenceof typical membranifolia. Since plants
showing these two types of pubescence grow sympatricallyand do not differ in
other respects, it seems unjustifiedto segregate the glabrate individuals, especially since density of pubescence is a characterwhich is frequently subject to
considerablevariation.The pubescence in the inflorescencecan be velutinous or
sericeous; there seems to be complete intergradationbetween the pubescence
types.
This species is closest to B. caapi, from which it is distinguishedby its less
coriaceous leaves, usually with denser pubescence, its longer pedicels and the
fruit with lateral wings. The two Krukoff collections from Amazonian Brazil
appearto belong to this taxon, but are unusualin the stouter and shorteranterior
style; the fruitis unknown.This representsa considerabledisjunctionin the range
of the species.
33. BanisteriopsispulchraB. Gates, sp. nov.
Frutex volubilisvel liana. Ramiteretes, appresso-sericei,demumglabrati.Stipulae minutae,usque 0.5 mm longae, triangulares,sericeae. Folia opposita, lamina
4.7-14.0 cm longa, 2.5-8.3 cm lata, late elliptica vel ovata, basi obtusa et parum
asymmetrica, apice apiculata caudatave, margine plana, subtus in nervis lateralibus2-4(-8) glandes stipitatasgerenti, supra sparsimappresso-sericeavel glabrata, subtus dense candidaappresso-sericeavel glabrata,pilis T-formibus,pede
0.04-0.09 mm longo, trabecula0.9-2.0 mm longa, venis venulisque supraparum
prominentibuset subtus prominentibus,petiolo 10-25 mm longo, appresso-sericeo, apice 2 glandes stipitatasgerenti. Inflorescentiafloribusin umbellis 4-floriferis fasciculatis, umbellis in cymis dispositis, appresso-sericea;bracteis bracteolisque 1.6-2.5 mm longis, lanceolatis, abaxialiterappresso-sericeis,adaxialiter
glabris, per vel post anthesin caducis; pedunculis 0-2 mm longis, pedicellis
(8-)12-17 mm longis, 0.9-1.2 mm diametro,0.6-0.8 mm sine indumento,appresso-sericeis. Sepala 5, abaxialitersericea, adaxialiteromnino minute tomentosa,
oblonga, apice rotundaet interdumrevoluta; sepalum anticum 4.5-5.0 mm longum, 2.0-2.5 mm latum, 4 lateralia4.5-5.8 mm longa, 2.5-2.8 mm lata, glandes
2.0-3.0 mm longas et 0.8-1.4 mm latas gerentiaet 2.5-3.0 mm superantia.Petala
5, rosea, aetate albicantia,fimbriata,4 lateraliainter sepala patentia, ungue 1.02.5 mm longo, 0.4-0.5 mm lato, limbo 7.5-11.0 mm longo, 7.0-10.5 mm lato,
orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribusconcavioribus;petalum posticum ungue erecto 3.5-4.2 mm longo, usque 1.4 mm diametro, apice
109
constricto, limbo 7.0-9.0 mm longo, 6.0-8.0 mm lato, oblongo, retroflexo,piano

convexove, fimbriisbasalibusdilatatisglandulosisque.Stamina10, filamentis2.05.0 mm longis, proximaliterconnatis et inter stylos posticos inflexis; antherae
loculis 0.8-1.6 mm longis, sparsimpilosis, interdumsolum basi et apice, connectivis canarinis,petalis oppositis 0.6-1.4 mm longis, 0.4-1.0 mm latis, 0.1-0.7 mm
profundis,solum 2 petalis anterioribusoppositisglandulosis,illis sepalis oppositis
0.6-2.2 mm longis, 0.4-1.6 mm latis, 0.3-1.2 mm profundis,omnibusglandulosis,
solum 3 sepalis anterioribusoppositis multumdilatatis, loculis 2 sepalis anterolateralibus0.2-1.4 mm supra superantibus.Ovarium1.0-1.4 mm altum carpellis
3, uniovulatis, liberis, toro pyramidaliadnatis, candido-sericeis; styli 3, stylo
antico rectum, 2.8-4.0 mm longo, 0.4 mm lato, stylis posticis 3.5-5.0 mm longis,
divergentibus,basi lyratis, stigmatibuscapitatis. Fructusex 3 samaris(vel abortu
1-2) constans; samara carpophorousque 3.5 mm longo, 0.8-1.0 mm lato, nuce
ovoidea 7-10 mm alta, 6-8 mm longa, appresso-pubescenti,viridi, tuberculata
vel manifestealulata, alulis usque 6 mm altis, ala 19-33 mm longa, 8-15 mm lata,
oblongaobovatave, viridi, appresso-sericea,pilis sessilibus T-formibus,trabecula
recta gracilis 0.4-0.5 mm longa, alis posterioribusparumparallelisad alam anteriorem, loculis interiusglabris.
Key to the Varieties of Banisteriopsis pulchra
1. Leaves densely appressed-sericeousbelow; inflorescencebranchessericeous. a. var. pulchra.
1. Leaves glabratebelow; inflorescencebranchesvelutinous.
b. var. glabrata.
33a. Banisteriopsispulchravar. pulchra

Leaves adaxially densely white-sericeous. Inflorescence branches sericeous.
Connectivesof stamens opposite the antero-lateralsepals globose, projecting0.20.6 mm beyond the locules. Anteriorstyle 3.4-4.0 mm long, posteriorstyles 4.05.0 mm long.
Chromosome number. n = 10. Voucher, Anderson 11789.
Type. Anderson 11789, Paraguay,50 km N of Rio Ypane, on road to Pedro

Juan Caballero,fl fr (holotype, MICH, isotypes K, NY).
Distribution.(Fig. 25). Vine in the cerradotrees and sometimesin galleryforest
of SW Goias and Mato Grosso in Brazil, extending southwardto Paraguay.
Collected in flower in March, April and May, and in fruit in April, May, June
and December.
BRAZIL. Mato Grosso: Hatschbach24328 fl (MICH,P); 38646 fl fr (MICH);Lindman2867 fl (F,
G, GH, NY, S, US); Santos s.n. fl (R); Commissdo Rondon (leg. Hoehne) 1581 fl (SP). Goias:
Anderson 9460 fr (MICH, NY, UB); Hatschbach & Kummrow 38336 fl (MICH); Irwin et al. 17805
fr (MICH,NY, UB). Stateunknown:Tamberliks.n. fl (W). PARAGUAY.Anderson11787fl (MICH);

11788 fl (MICH); Hassler 8218 fl (BM, G); 8317 pp fr (G); (leg. Rojas) 10303 fl (G, GH, K, NY, P,
W); 10303a fl (GH, MICH, NY, S); Schinini 9030 fr (CTES); Weddell 3278 fl (P).
This variety is distinguishedfrom B. membranifoliaby its densely white-sericeous leaves, the hairs with arms 0.4-1.0 mm long, largerflowers, with the fifth
petal apically constricted, and the 2 posteriorstyles longer than the anteriorone;
it is distinguishedfrom B. muricata by its caducousbracteoles, its longer, thicker,
sessile pedicels, stamens with the connectives opposite the antero-lateralsepals
not greatly enlarged, thicker styles, and fruit with broad lateral winglets.
33b. Banisteriopsis pulchra var. glabrata B. Gates, var. nov.
Folia lamina subtus glabrata.Inflorescentiavelutina. Stamina connectivis sepalis antero-lateralibusoppositis gibbosis loculis 1.0-1.4 mm superantibus.Stylus
anticus 2.8-3.2 mm longus, styli postici 3.5-3.6 mm longi.
Flora Neotropica
110
S'O.
I00
~ ~~
I40
'<I\;~~~~~~~~~~~~~~~~~~~~~X
^C^^-!
^J^^^^'^^'^r^^
""
^ .a.^.^^^.?^^ .-tjBi
W:.
FIG. 25. Distribution of Banisteriopsis membranifolia group, species 32-34. Triangle, B. membranifolia; closed circle, B. pulchra var. pulchra; open circle, B. pulchra var. glabrata; square, B.
adenopoda.
Type. Buchtien 1794, Bolivia, Mapiri, San Carlos, fl (holotype, NY; isotypes,
F, NY, fragment, US).
Distribution (Fig. 25). Known only from the type in Bolivia.

Collected in flower in November.
This specimen is very distinctive with its large glabrate leaves and large flowers
with greatly enlarged stamen connectives, and perhaps deserves recognition as
a species. However, in the absence of fruiting material, it seems better at this
time to recognise its affinity to B. pulchra var. pulchra, from which it chiefly
differs in its glabrate leaves, larger connectives on the stamens opposite the
antero-lateral sepals, and shorter styles.
34. Banisteriopsis adenopoda (Adr. Jussieu) B. Gates, comb. nov.
Banisteriaadenopoda Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 40. 1832[1833].
BanisteriasubcordataGardner,LondonJ. Bot. 2: 336. 1843.Type. Gardner338, Brazil, Rio de
Janeiro,fl (holotype, K; isotypes, BM, G, GH, K, NY, P, US, W).
Banisteria adenopoda var. subrotunda Niedenzu, Ind. Lect. Lyc. Brunsb.: 23. 1900. Type.
Mendonca242, Brazil, Sao Paulo, Jacarahy,fl (holotype, B? destroyed).
Liana, the branches terete, densely velutinous, eventually glabrate, the bark
dark-brown, smooth, with scattered minute lenticels. Stipules 1.0-1.5 mm long,
narrowly lanceolate, sericeous, sometimes glabrate. Leaves with the petiole 7-
111
18 mm long, velutinous, eglandular,the lamina(2.6-)5.8-14.9 cm long, (1.2-)2.97.3 cm wide, smaller in the inflorescence, elliptic, sometimes orbicular,obtuse
to cordate at the base, acuminateor more rarely apiculate at the apex, with the
margin flat to slightly revolute, bearing 1-2(-4) pairs of stalked glands on the
lateral veins near the marginabaxially, the stalk 0.6-1.5 mm long, adaxially velutinous, the hairs sessile, sometimes with inmixed golden-sericeoushairs, abaxially persistently velutinous, the hairs sessile, with the venation prominulousto
impressed adaxially, prominent abaxially. Infloresence of 4-flowered umbels
borne in large cymes on axillary branches; bracts and bracteoles 2.0-3.0 mm
long, linear to lanceolate, loosely sericeous abaxially, glabrous adaxially, caducous in bud; pedicels sessile, (9-)12-20 mm long, 0.6-0.8 mm wide, 0.6 mm
without the hairs, velutinous. Sepals sericeous abaxially, minutely velutinous
throughoutadaxially, oblong, roundedand sometimes revolute at the apex, 3.03.8 mm long, 1.5-2.1 mm wide, the glands green, 1.4-1.8 mm long, 0.5-0.6 mm
wide, the sepals projecting2.2-2.5 mm beyond the glands. Petals white, sometimes with pink veining, becoming yellow in age, fimbriate,the 4 lateral petals
reflexed between the sepals, the claw 1.0-1.5 mm long, 0.4-0.5 mm wide, the
limb 6.0-8.0 mm long, 5.0-6.5 mm wide, orbicular,eglandular,the antero-lateral
petals more concave than the postero-lateralones, the posterior petal with the
claw erect, 2.5-3.0 mm long, up to 0.8 mm wide, constricted apically, the limb
4.5-5.5 mm long, 4.0-4.5 mm wide, broadly obovate, reflexed, plane, with the
basal fimbriaegland-tipped.Stamens with the filaments 1.9-2.5 mm long, those
of the posterior 3 stamens flexuous and inbent between the posterior styles, the
locules densely pilose, 0.6-1.2 mm long, the connectives yellow, those of the
stamensopposite the petals 0.4-1.0 mm long, 0.4-0.8 mm wide, 0.1-0.5 mm deep,
only those opposite the antero-lateralpetals glandular,those opposite the sepals
0.6-1.5 mm long, 0.6-0.9 mm wide, 0.2-0.8 mm deep, those opposite the 3 anterior sepals with the connectives enlarged and glandular,the stamen opposite
the anterior sepal with the connective projecting 0.2 mm beyond the locules,
those opposite the antero-lateralsepals with the connectives projecting0.4-0.6
mm beyond the locules. Ovary0.8-1.0 mm tall, white-sericeous,the styles basally
pubescent, the anterior style straight, 2.2 mm long, 0.4-0.5 mm wide, the posterior styles diverging, lyrate basally, 2.4-2.5 mm long, tapering apically, the
stigmas capitate. Samarawith the carpophoreup to 2 mm long and 1.0 mm wide,
the nut 6-8 mm tall, 3-6 mm long, appressed-sericeous,cristate to alulate, the
winglets up to 8 mm high, the wing 22-37 mm long, 9-15 mm wide, appressedpubescent to glabrate, the hairs T-shaped, sessile, stiff and somewhat irritating
to the skin, the trabecula0.4-0.9 mm long, the wings on the 2 posterior samaras
somewhat rotated to lie parallelwith the wing of the anteriorsamara.
Type. Saint Hilaire Catal. D, 716, Brazil, Sao Paulo, Freguesia de Nossa
Senhora da Escadenha, fl (holotype, P; isotypes, P).

Distribution.(Fig. 25). A vine in the Atlanticcoastal forest of SE MinasGerais,
Rio de Janeiro, Sao Paulo and Parana.
Collected in flower Februaryto May, and in fruit April to August.
Representative collections. BRAZIL. Ceara: Allemdo & Cysneiros 215 pp fl (R). Minas Gerais:
Anderson 11618 (MICH); Damazio 864 fl (G, RB); Irwin et al. 30504 fl (MICH, NY); Regnell 11.27
fl fr (BR, C, F, K, M, MO, NY, P, R, S, U, US, W); L. Williams & Assis 6586 fl (GH, SP, US). Rio
de Janeiro: Glaziou 8582 fl (BR, C, G, K, NY, P, R); 9364 fl (BR, C, K, P); Occhioni 1215 fl (RB).
Sao Paulo: Brade 17812 fl (MICH, RB); Gehrt 5483 fl (SP); Glaziou 17488 fl (BM, C, G, IAN, P);
Martius 608 fl (M); Pereira 10756 fl (MBM). Parana: Dusen 8062 (BM, F, G, S, US); Hatschbach
666 fl (RB, SP); 6777 fl fr (HB); Kummrow 455 fl fr (MBM, MICH); Pereira 5415 fl (MICH, RB).
State unknown: Burchell 4689 (GH, GOET, K, NY, P).
Flora Neotropica
112
Local name. Brazil: "pragua".
This species shows some variationin the size and numberof the stalked leaf
glands, the size of the connectives on the stamens opposite the antero-lateral
sepals, and in the leaf pubescence, but seems to be well-defineddiscrete taxon.
One sheet of a Riedel collection from Cuiaba, Mato Grosso, is of this species;
the other sheets are Banisteriopsis confusa. It seems likely that the locality on
the sheet of B. adenopoda is incorrect.
35. Banisteriopsiscaapi (Spruce ex Grisebach)Morton, J. Wash. Acad. Sci. 21:
486. 1931.
Banisteriacaapi Spruceex Grisebachin Martius,Fl. bras. 12: 43. 1858.
BanisteriaquitensisNiedenzu, Ind. Lect. Lyc. Brunsb.:10. 1900.Type. Eggers 15485,Ecuador,
El Rosario, fl fr (holotype, B? destroyed, NY, fragment, probably of holotype; isotypes, F).
Banisteriopsis inebrians Morton, J. Wash. Acad. Sci. 21: 485. 1931. Type. Klug 1964, Colombia,
Putumayo,Umbria,fr (holotype, US; isotypes, BM, BR, ECON, F, GH, IAN, MICH,MO,
NY, P, RB, S, SP, US, W).
Banisteriopsisquitensis (Niedenzu)Morton,J. Wash. Acad. Sci. 21: 486. 1931.
Liana, the young branches sparsely appressed-sericeousto glabrate, the old

branches glabrous, terete, the bark becoming fissured into shallow corky splits
in age, with the wood sometimes conspicuously lobed. Stipules triangular,0.51.0 mm long, glabrousor appressed-sericeous.Leaves with the petiole 9-25 mm
long, sparsely appressed-sericeousor glabrate, eglandularor bearing a pair of
cupulate glands near the apex, the lamina (4.8-)8.2-15.9(-20.5) cm long, (2.5-)
3.5-7.7(-11.5) cm wide, smallerin the inflorescence,often quite coriaceous when
mature, broadly ovate to ovate, obtuse to truncate at the base, short- to longacuminateat the apex, flat to slightlyrevolute at the margin,bearingabaxially25 pairs of sessile glands near or at the margin and an additionalpair near the
midribat the base, glabrateadaxially, very sparsely appressed-sericeousto glabrate abaxially, the hairs T-shaped, sessile with the trabecula0.1-0.4 mm long,
with the primary veins prominulous adaxially and the reticulation sometimes
impressed, and the primaryand secondary veins prominentabaxially. Inflorescence of 4-floweredumbels arrangedin axillary cymes, subtended by very reduced leaves or the inflorescenceleaves deciduous before anthesis, sparsely tomentose to velutinous; bracts and bracteoles 1.0-1.8 mm long, triangularto
elliptic, appressed-pubescenceabaxially, glabrousadaxially, caducous before or
duringflowering, rarely immediatelyafter flowering. Pedicels sessile, 7-11 mm
long, 0.4-0.6 mm in diameter,0.3-0.5 mm without the hairs, appressed-sericeous
or tomento-sericeous.Sepals sericeous abaxially, minutelytomentose throughout
adaxially, elliptic, obtuse at the apex, 2.0-3.5 mm long, 1.5-2.0 mm wide, all
eglandularor the 4 lateral sepals biglandular,the glands 0.5-2.2 mm long, 0.41.2 mm wide, projecting1.2-2.0 mm beyond the glands;petals pale pink, becoming pale yellow in age, fimbriate,the 4 lateralpetals reflexed between the sepals,
the claw 1.0-1.5 mm long, 0.2-0.4 mm in diameter, the limb 5.0-8.5 mm long,
4.0-6.0 mm wide, the posterior petal with the claw erect, 2.5-3.0 mm long, up
to 1.0 mm wide, constricted at the apex, the limb 5.0-7.0 mm long, 2.5-4.5 mm
wide, broadly obovate with the basal fimbriaegland-tipped.Stamens with the
filaments2.0-4.0 mm long, connate basally, the posterior3 flexuous and inflexed
between the posterior styles, the locules sparsely pilose to glabrate,those of the
3 anterior stamens 0.7-1.2 mm long, those of the other 7 stamens 0.3-0.9 mm
long, the connectives of the 5 posterior stamens 0.2-0.7 mm long, 0.2-0.4 mm
wide, 0.1-0.2 mm deep, not glandular,those of the 5 anterior stamens 0.8-1.6
113
mm long, 0.5-1.0 mm wide, 0.3-1.0 mm deep, glandular, those opposite the

antero-lateral sepals enlarged and overtopping the locules by 0.5-1.0 mm. Ovary
1.0-1.2 mm tall, white-sericeous, the anterior style straight, 2.8-3.2 mm long, 0.2
mm in diameter, the posterior styles diverging and lyrate at the base, 3.0-4.0 mm
long, 0.15 mm in diameter, the stigmas capitate. Samara with the carpophore up
to 4 mm long and 0.4 mm wide, the nut 5-11 mm tall, 3-5 mm long, the abaxial
margin with a tooth at the base, appressed-pubescent to glabrate, the wing 18-42
mm long, 8-22 mm wide, appressed-pubescent soon glabrate, the wings of the
posterior samaras somewhat rotated to lie more nearly parallel to the wing of the
anterior samara, the locule of the nut hairy throughout within.
Chromosome number. n = 10. Baldwin, Bull. Torrey Bot. Club 73: 282.
Type. R. Spruce 2712, Brazil, Amazonas, Rio Vaupes, near Panure, fl fr (holotype, B? destroyed; isotypes, BM, BR, C, G, GH, K, NY, P, RB, W).
Distribution. (Fig. 27). Possibly native to, but widely cultivated by native peoples in, Bolivia, Peru, Ecuador, Colombia and Amazonian Brazil.
Collected in flower December to August and in fruit March to August.
Representative collections (specimens in flower or fruit). COLOMBIA. Valle: Cuatrecasas 14372
fl fr (F, US). Vaup6s: Garcia-Barriga 14250 fl fr (NY). Amazonas: Perez-Medena et al. Leticia 9
fl (US). VENEZUELA. Amazonas: Rutkis 209 fl (VEN). ECUADOR. Eggers 15494 fl (F, K, M, P,
US). PERU. Loreto: Rivier 3 fr (ECON). San Martin: Martin & Plowman 1805 fr (ECON, S); Velarde
Nunez 6577 fl (ECON); 6585 fl (ECON). BRAZIL. Amaz6nas: J. G. Kuhlmann s.n. fl (RB 60455,
US); Melin 96 fl (S). BOLIVIA. Pearce s.n. fl (BM, K); White 970 fl (K, MICH, NY, US). CULTIVATED IN BOTANIC GARDENS: Ducke 153 fl fr (F, GH, K, MO, NY, R, S, US); RB 25258
fl fr (P, RB, S, U, US); RB 25259 fl (GH, RB); RB 25260 fl (K, P, RB, S, U, US); RB 35599 fl
(G, K, MICH, P, R, S, U, US); Fennell 282 fl (ECON); Fr6es 28491 fl fr (GH, IAN); Garcia-Barriga
20035 fl (NY); Gillis 11237 fl (NY); Handro sn. SP 45735 fl (NY, SP); Pires 19 fl fr (ECON, IAN,
NY, P, US); N. T. Silva 59727 fl (IAN, MICH, NY).
Local names. Colombia: "yage," "'ayahuasca,"

"yage del monte," "yage sembrado." Ecuador: "ayahuasca," "natema," "nepe." Peru: "ayahuasca," "ayahuasca amarilla," "purga-huasca," "purga-huasca de los perros," "cielo ayahuasca"; "cuchi-ayahuasca," "ayahuasca negra," "shuri-fisopa." Brazil:
caapi," ya,"
"cauupuri mariri," mao de ona," "tiwaco-mariri."
This species is closely related to B. membranifolia, from which it chiefly differs
in its more coriaceous ovate leaves, shorter pedicels, smaller flowers, and fruit
without lateral wings on the nut and with the locule pubescent within. It is also
close to B. muricata, resembling this species in its small flower size, calyx with
or without glands, the shape of the posterior petal and the slender styles.
It is difficult to know where this species is native, since it is widely cultivated
throughout Peru, Ecuador, Colombia and Amazonian Brazil by native populations
for use in the preparation of an hallucinatory beverage; most of the collections
available to me were from cultivated plants according to their collection data,
and most of these specimens were sterile. After careful examination of all the
material I have concluded that the variation represented by them was best considered to be one taxon. The characters that show variation within this species
are the calyx glands, flower size, size and shape of the fruit wing and the size of
the inflorescence and the leaves associated with it. Flower size and inflorescence
size are very variable characters in all species, and the presence or absence of
calyx glands is also not significant taxonomically, glandular and eglandular forms
being present in the same population. The types of B. caapi and of B. inebrians
exhibit large fruits and very coriaceous leaves, as do most of the specimens
cultivated in Botanical Gardens in Brazil, whereas specimens from Bolivia, Peru,
Ecuador, and Colombia have smaller fruit wings and less coriaceous leaves. How-
114
Flora Neotropica
ever, both these characters show great variation on one plant and would not
appearto provide a sound basis for taxonomicsegregation.Apparentlythe native
populations who use this plant recognise many different kinds of caapi, with
differenthallucinogenicproperties;I consider these to be chemical variants.The
ease with which caapi can be vegetatively propagatedby stem cuttings makes it
possible for clones of such variantsto be maintained.
36. Banisteriopsisschwannioides(Grisebach)B. Gates, comb. nov.
Fig. 26.
Banisteria schwannioides Grisebach in Martius, Fl. bras. 12(1): 54. 1858.
Liana, the branchesterete, minutelyvelutinous, the barkbrown with scattered

prominulouslenticels. Stipules minuteor absent. Leaves with the petiole 3-7 mm
long, velutinous, eglandular,the lamina 2.2-8.3 cm long, 1.2-4.8 cm wide, reduced in the inflorescence, ovate to orbicular,cordate to truncate at the base,
apiculate to acuminateat the apex, with the marginrevolute, bearing abaxially
a pair of sessile glands 0.2-0.6 mm in diameterbeside the costa at the base and
sometimes 1-2(-5) pairs of minute stipitateglands, 0.1 mm in diameter,near the
margin, softly velutinous to glabrate adaxially, velutinous abaxially, with the
reticulationimpressed adaxially and prominentabaxially. Inflorescence of 4-6floweredumbels or short racemes borne in cymes on terminalor axillarybranches, velutinous; bracts and bracteoles 1.2-2.0 mm long, lanceolate, velutinous
abaxially, glabrousadaxially, caducous in flower or just after flowering;pedicels
sessile or stipitate, 11-24 mm long, 0.6-0.8 mm wide, 0.6 mm without the hairs,
minutely velutinous. Sepals broadly elliptic to ovate, rounded and sometimes
revolute at the apex, tomentose abaxially, tomentose throughoutadaxially, 3.05.0 mm long, 2.5-2.8 mm wide, the glands 2.0-3.0 mm long, 0.6-1.2 mm wide,
the sepals projecting1.8-2.0 mm beyond the glands, reflexedin fruit. Petals pink,
dentate, the 4 lateral petals reflexed between the sepals, the claw 1.2-2.5 mm
long, 0.4-0.8 mm wide, the limb 8.0-11.0 mm long, 6.0-9.5 mm wide, rotund,
eglandular,the antero-lateralpetals cochleate, the posterior petal with the claw
erect, 3.5 mm long, 0.8-1.0 mm wide, apicallyconstricted,rarelysericeous adaxially, the limb 6.0-6.5 mm long, 3.5-6.0 mm wide, rotund to obovate, the basal
teeth gland-tipped.Stamenswith the filaments0.5-1.6 mm long, connate basally,
the locules glabrous, 0.5-1.6 mm long, those of the stamens opposite the anterolateral sepals 0.6-0.8 mm long, the connectives of the 5 anteriorstamens 1.0-1.8
mm long, 0.8-1.1 mm wide, 0.5-1.5 mm deep, glandularand very enlarged,those
opposite the antero-lateralsepals projecting0.6-1.0 mmbeyond the locules, those
of the 5 posterior stamens 0.3-0.7 mm long, 0.3-0.6 mm wide, 0.1-0.4 mm deep,
not glandular.Ovary 1.0-1.4 mm tall, white-sericeous, the anteriorstyle 2.6-3.0
mm long, 0.6-0.9 mm wide, thick and straight, the posterior styles 1.5-1.8 mm
long, up to 0.5 mm wide and lyratebasally, taperingapically, the stigmascapitate.
Samarawith the carpophoreup to 2 mm long, 1.8-2.0 mm wide, the nut 7-8 mm
tall, 4-5 mm long, tuberculateto alulate, often asymmetric,appressed-pubescent,
the distal marginprolongedinto a tooth basally, the wing 27-30 mm long, 11-13
mm wide, appressed-sericeousto glabrate,the hairsT-shaped,sessile or stipitate,
the trabecula straight and stiff, 0.6-1.2 mm long, the wings of the 2 posterior
samarassomewhat rotated to lie parallelto the wing of the anteriorsamara.
Type. Spruce 1012, Brazil, Para, near Santarem,fl (lectotype, K; isolectotype,
K). Grisebach'soriginaldescriptionwas based on two collections, a fruitingcollection, Gardner 1484, and a flowering collection, Spruce 1012. I choose the
Spruce sheet as lectotype because the descriptionof the flowers is derived from
this collection, the descriptionof the leaves accords well with this plant, and the
115
*r,'' I' u
Di'
FI.
26
"
p' p
3.
(
.
4
.
A'
FIG. 26. Banisteriopsis prancei and B. schwannioides. A-C, B. prancei. A, flower x 4.2; B,
stamens, semi-diagrammatic, abaxial and lateral views, x 8.4 (stamen 1 is opposite anterior sepal,
stamen 6 is opposite posterior petal); C, gynoecium x 7. D-G, B. schwannioides. D, flower bud x
4.2; E, androecium and gynoecium x 6.3; F, stamens, semi-diagrammatic, abaxial and lateral
views, x 8.4 (stamen 1 is opposite anterior sepal, stamen 6 is opposite posterior petal); G, gynoecium x 7. (A-C, Anderson 990; D-G, Pires & Belem 12524.)
116
Flora Neotropica
epithet "schwannioides" seems more fittingfor a taxon with a greatly enlarged

anteriorstyle. The descriptionof the fruits is based on the Gardnersheet, which
is a galled individual of B. malifolia var. appressa.
Distribution.(Fig. 27). Collectedin the forests of Amazonasand Parain Brazil,

alt. 0-200 m.
Collected in flower in July to October, and in fruit August to October.
BRAZIL. Amaz6nas: Frdes 25130 fl (IAN, UB); 25491 fl fr (IAN, UB). Para: Black 47-1170 fl
(IAN); Ducke 2905 fl fr (MG); 8519 fl (MG); 11018 fl fr (MG); Frdes 31248 fl (IAN); Pires & Belem
12524 fl (IAN, MICH, RB).
This species is most closely relatedto B. caapi, but can be readilydistinguished

by its velutinous leaf pubescence, larger flowers with longer pedicels, glabrous
antherlocules, and most conspicuouslyby its reducedposteriorstyles and greatly
enlargedanteriorsytle.
Pires & Belem 12524 has sparse pubescence abaxially on the petals and adaxially along the claw and the middle of the limb of the posteriorpetals. However,
I do not feel that this interestingvariantwarrantstaxonomic recognition.
37. Banisteriopsisprancei B. Gates, sp. nov.
Fig. 26.
Frutex volubilis; ramiteretes, albi vel aureo-appresso-sericei,demumglabrati,

cortice fusco, in lamellis suberosis findenti. Stipulae nullae vel minutae. Folia
opposita, lamina 5.5-10.7 cm longa, 3.0-6.8 cm lata, in inflorescentiaminori,
ovata vel late ovata, basi obtusa truncatave, apice acuta vel apiculatavel acuminata,marginerevoluta, interdumsubtusprope marginem2-6 glandes stipitates
usque 0.4 mm diametrogerenti, supra molliter velutina, subtus sparsim et laxe
sericea, pilis T-formibus, stipitatis, trabecula 0.36-0.60 mm longa, venis venulisque supra planis, subtus prominentibus;petiolo 4-19 mm longo, 2-4 glandes
sessiles 1.0-1.5 mm diametroprope apicem vel in lamina basi gerenti, velutino.
Inflorescentiafloribus binatim vel in umbellis 4-floriferisfasciculatis, in thyrsis
dispositis, velutina; bracteis bracteolisque 0.8-1.0 mm longis, triangularibus,
abaxialitersericeis, adaxialiterglabris,post anthesincaducis;pedicellis sessilibus
9-12 mm longis, 0.6-0.8 mm diametro, 0.3-0.4 mm sine indumento, appressosericeis. Sepala 5, oblonga vel anguste ovata, apice rotunda et plus minusve
revoluta, abaxialitertomentosa, adaxialiterglabra vel basi minute velutina, in
fructu retroflexa, sepalum anticum 2.0-2.5 mm longum, 1.1-1.6 mm latum, 4
lateralia 2.5-3.2 mm longa, 1.4-2.0 mm lata, glandes 1.2-1.9 mm superantia.
Petala 5, rosea, dentata,4 lateraliainter sepala patentia,ungue 1.0-1.4 mm longo,
0.2-0.4 mm lato, limbo 5.0-7.0 mm longo, 4.0-6.0 mm lato, orbiculari, eglanduloso, 2 anterioribusquam 2 posterioribusparum majoribuset concavioribus,
petalumposticum ungue erecto 2.5-3.0 mm longo, usque 0.7 mm diametro,apice
constricto, limbo 4.5-6.0 mm longo, 3.5-5.5 mm lato, obovato, plano, dentis
basalibusglandulosis.Stamina10, filamentis1.4-3.8 mm longis, proximaliterconnatis; antherae loculis 0.6-1.2 mm longis, glabris, connectivis canarinis, illis
petalis oppositis 0.5-0.8 mm longis, 0.3-0.5 mm latis, 0.1-0.3 mm profundis, 2
petalis anterioribusoppositis glandulosis dilatatisque,illis sepalis oppositis 0.51.1 mm longis, 0.2-0.6 mm latis, 0.2-0.6 mm profundis, omnibus glandulosis
dilatatisque,2 sepalis antero-lateralibusoppositis loculos 0.3-0.4 mm superantibus. Ovarium0.8-1.2 mm altum carpellis 3, uniovulatis, liberis toro pyramidali
adnatis, candido-sericeis;styli 3, 1.8-2.8 mm longi, stylo antico quam 2 posterioribusparumbreviori, recti paralleliquevel parum divergentes, 0.15-0.20 mm
117
FIG. 27. Distribution of Banisteriopsis caapi, B. schwannioides, B. prancei and part of B. muricata group, species 35-37, 41, 43. Closed circle, B. caapi; open circle, B. schwannioides; closed
square, B. prancei; triangle, B. oxyclada; open square, B. goiana.
diametroversus apicem, stigmatibuscapitatis. Fructus ex 3 samaris (vel abortu

1-2) constans; samara carpophoro 1-2 mm longo, usque 0.6 mm lato, nuce 6-8
mm alta, 4.0-4.5 mm longa, rugosa tuberculatave,appresso-pubescenti,pilis sessilibus, T-formibus,trabecula flexuosa, ala 20-25 mm longa, 9-11 mm lata, appresso-sericea vel glabrata,pilis T-formibus,sessilibus, trabecularecta rigidave,
0.50-0.90 mm longa; loculo nucis interiusglabro.
Type. Prance et al. 14568, Brazil, Amazonas, Cachoeira Republica, Rio Curuquete, fl fr (holotype, INPA; isotypes, MG, MICH, NY).
Distribution.(Fig. 27). This species has been collected in western Mato Grosso
and southern Amazonas, Brazil, as a vine in open woods on drier sites, and at
forest margins, and in gallery forest, alt. 100-600 m.
Collected in flower from April to July, and in fruit in July.
BRAZIL. Mato Grosso: Anderson 9910 fl (MICH, NY, UB); J. G. Kuhlmann 2058 fl (R, RB);
Malme 3386 fl (S); Meader 10 fl (IAN, US); Ratter et al. R1156 fl (IAN, NY); R1470 fl (IAN, MO,
NY); Commissao Rondon (leg. Hoehne) 1891 fl (SP).
This species superficiallyresembles B. muricata with its dense inflorescences

of small pink flowers; however, in the details of its flower and the form of the
118
Flora Neotropica
fruit, B. prancei is closer to the B. malifolia group. Banisteriopsis prancei can be
distinguishedfrom B. muricata by its velutinous leaf pubescence, sepals with

roundedapex, the only slightlyenlargedconnectives on the stamens opposite the
antero-lateralsepals, the straightstyles, and the form of the fruit. It can be distinguishedfromthe B. malifoliagroupby its pink flowers, the constrictedclaw on
the fifth petal and its glabrousantherlocules, wings of the samarasnot rotated,
and nut locule glabrous.
The species is named after GhilleanT. Prance, who collected the type and has
made an inestimablecontributionto the knowledge of the Amazonianflora.
38. Banisteriopsismultifoliolata(Adr. Jussieu) B. Gates, comb. nov.
Banisteria multifoliolata Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 282. 1840.
Liana, the branches terete, persistently loosely sericeous, prominentlylenticellate. Stipules up to 0.5 mm long, triangular,sericeous. Leaves with the petiole
4-15 mm long, appressed-sericeous,eglandular,the lamina3.6-9.0 cm long, 2.46.0 cm wide, smaller in the inflorescence, elliptic to orbicular, truncate at the
base, apiculateat the apex, with the marginrevolute, bearing0-2 pairs of shortstalked glands on the basal lateral veins near the margin abaxially, sometimes
with an additional 1-2 pairs of glands distally, especially on the inflorescence
leaves, adaxially densely to sparsely appressed-hispido-sericeous,the hairs with
enlargedbases, abaxiallydensely and loosely hispido-sericeous,the hairs stalked,
T-shaped, with the lateral veins impressedadaxially, prominentabaxially. Inflorescence of 4-floweredumbels, the umbels borne in dense axillarycymes; bracts
and bracteoles 0.9-1.2 mm long, broadlytriangular,sericeous abaxially,glabrous
adaxially, persistent or deciduous in fruit; pedicels sessile, 9-15 mm long, 0.60.8 mm wide, 0.4-0.6 mm without the hairs, appressed-sericeous.Sepals loosely
sericeous abaxially, minutely tomentose or sericeous throughoutadaxially, with
the apex rounded and sometimes revolute, the anterior sepal 2.5-3.0 mm long
and 1.4-1.7 mm wide, elliptic, sometimes bearing 1-2 much reduced glands, the
posterior sepals 2.5-3.5 mm long, 1.5-2.8 mm wide, the glands 1.2-2.4 mm long,
0.4-1.2 mm wide, the sepals projecting 1.2-1.8 mm beyond the glands. Petals
pink, becoming white in age, lacerate to fimbriate,the 4 lateral petals reflexed
between the sepals, the claw 0.8-2.0 mm long, 0.4-0.5 mm wide, the limb 4.58.5 mm long, 4.0-8.0 mm wide, orbicular,eglandular,the antero-lateralpetals
more concave than the postero-lateralones, the posterior petal with the claw
erect, 2.0-3.0 mm long, up to 1.4 mm wide, not constricted apically, the limb
4.0-7.0 mm long, 3.5-5.0 mm wide, orbicular,reflexed, plane to convex, with the
basal fimbriaegland-tipped.Stamens with the filaments 1.5-3.5 mm long, those
of the posterior 3 stamens flexuous but not inflexed between the posterior styles,
those opposite the sepals longer than those opposite the petals, connate basally,
the locules densely pilose to glabrate, 0.8-1.2 mm long, the connectives of the
stamensopposite the petals 0.7-0.8 mm long, 0.4-0.6 mm wide, 0.1-0.4 mm deep,
only those opposite the antero-lateralpetals glandular,those opposite the sepals
0.8-1.4 mm long, 0.5-0.8 mm wide, 0.5-0.8 mm deep, globose, glandularand
enlarged, not projectingor projectinga little beyond the locules. Ovary 1.0-2.0
mm long, white-sericeous, the styles straight, divergingslightly, somewhat flattened laterally, 1.4-2.6 mm long, 0.3-0.5 mm wide, the stigmas capitate. Fruit
without carpophore?,the nut up to 8 mm tall and 6 mm long, laterallycorrugate,
appressed-sericeous,the hairs with the trabecula0.6-0.9 mm long, the wing up
to 32 mm long and 10 mm wide, appressed-sericeouslike the nut, the wings of
the 2 posteriorsamarasrotatedto lie at rightangles to that of the anteriorsamara;
locule of nut densely sericeous within.
119
Type. Vauthier 91, Brazil, Rio de Janeiro, Santa Thereza, fl (holotype, P;

isotypes, P, G).
Distribution. Most collections are from the tropical rain forest of the hills of
Rio de Janeiro, Brazil, but there is one collection from Bahia, and a fragment
from Minas Gerais.
Collected in flower in December, March to May, July, August and October,
and in fruit in April.
BRAZIL. Bahia: Mori et al. 10046 fl (MICH). Minas Gerais: Warming s.n. (GOET). Rio de Janeiro:
Frazdo s.n. fl (RB); Glaziou 1059 fl (BR, C, GOET, P); 8586 fl (C, G, P); 9363 fl (C, G, K, NY, R);
Pohl s.n. fl (BR); Schott s.n. fl (W); Widgren 990 fl (S); s.n. fl (S).
There are no recent collections of this distinctive species from the hills of the
city of Rio de Janeiro. It is possible that it has become extinct there, but more
probably it survives as a liana in the canopy of some of the steeper and more
inaccessible forests on the hills of Rio. However, there is a recent collection from
Bahia which in additionto extending the range of the species, provides the first
fruitingmaterialof this species.
Banisteriopsis multifoliolata is somewhat unusual in its combinationof characters; it shares certain characterswith the B. membranifoliagroup, such as its
habitas a vine, stipitatelaminarglands, sepals roundedat the apex, andpubescent
anther locules, but it has certain other characterssuch as persistent bracteoles,
stamens with connectives which are not very enlarged, straight styles, and
hairy nut locule, which make it distinct. It shares this last characterwith B. caapi
and the B. malifolia group.
39. BanisteriopsisquadriglandulaB. Gates, sp. nov.

Liana, rami pertinaciterappresso-sericei,pilis aureis, trabeculausque 0.3 mm
longa, vetustiores lenticellis brunneisprominentibusornati. Stipulaeminutaejugo
interpetiolarijunctae. Folia petiolo 8-13 mm longo, appresso-sericeo, eglanduloso, lamina 3.2-6.6 cm longa, 3.0-6.0 cm lata, late ovata vel orbiculari,basi
obtusa cordatave, apice emarginataapiculataque, margine plana, eglandulosa,
adaxialiter sparsim appresso-sericea, pilis trabecula usque 0.4 mm, abaxialiter
dense nitido-sericea,pilis argenteis,trabecula0.4-0.6 mm longa, adaxialiterlaevi,
venis lateralibus prominentibusabaxialiter. Inflorescentia floribus in umbellis
4-floriferisfasciculatis, umbellis in cymis vel dichasiis dispositis, appresso-sericea; bracteis bracteolisque 1.2-1.6 mm longis, late triangularibus,appresso-sericeis, involucratis,persistentibus;pedunculis0.5-2.0 mm longis, pedicellis 12-15
mm longis, 0.6-1.0 mm diametro,0.4-0.8 mm sine indumento,appresso-sericeis.
Sepala 5, abaxialitersericea, adaxialiterglabraet marginesericea, elliptica, apice
acuta et involuta, 3.0-3.5 mm longa, 1.5-2.0 mm lata, 2 postero-lateraliaglandes
roseas 1.0-2.4 mm longa, 0.6-0.8 mm latas gerentia, glandes 1.8 mm superantia.
Petala 5, rosea demumalba, lacerata, 4 lateraliainter sepala patentia,ungue 1.52.0 mmlongo, limbo7.0-10.0 mmlongo, 5.5-10.0 mmlato, late ellipticovel orbiculari, 2 anterioribusquam2 posterioribusmajoribuset concavioribus;petalumposticumungueerectousque3.0 mmlongoet 1.0-1.5 mmlato, apiceconstricto,limbo6.08.5 mm longo et 5.5-7.5 mm lato, late obovato, plano, dentibusbasalibusdilatatis
glandulosisque,ungue et parte basali limbi lutea. Stamina 10, filamentis2.0-4.0
mm longis, illis 3 sepalis anticis quam ceteris longioribus;antheraeloculis 0.81.6 mm longis, glabris, connectivis 3 sepalis anticis oppositis 1.0-1.4 mm longis,
1.0-1.2 mm latis, 0.6-0.8 mm profundis, obovatis, glandulosis, loculos non superantibus, ceteris 0.5-1.2 mm longis, 0.5-0.8 mm latis, 0.2-0.4 mm profundis,
oblongis, eglandulosisvel 2 petalis anticis oppositis interdumglandulosis. Ovar-
120
Flora Neotropica
ium 1.0-1.2 mm altum, carpellis.3, uniovulatis, liberis, toro pyramidaliadnatis,

candido-sericeis;styli 3, aequales, 2.0-3.0 mm longi, crassi usque 0.4 mm diametro, recti, parallelivel divergentes, stigmatibuscapitatis. Fructus ex 3 samaris
(vel abortu 1-2) constans; samara(solum unica suppetit) sine carpophoro,nuce
6 mm alta et 4 mm longa, rugulosa, appresso-sericea,ala usque 27 mm longa et
15 mm lata, appresso-sericea,pilis sessilibus trabecularecta 0.4 mm longa.
Type. J. A. de Jesus 393, Bahia, saida de Itirucu/Maracas,fl fr (holotype,
CEPEC; isotype, MICH) = T. S. Santos 442.
Distribution.Known only as a roadside vine in Bahia.

Collected in flower in April and May, and in young fruit in April.
BRAZIL. Bahia: Jesus 393/Santos 442 fl fr (CEPEC, MICH); Mori et al. 9965 fl (CEPEC, MICH).
This species is very interestingin that in its generalaspect, leaf size, shape and
pubescence, flower color, sepal shape and pubescence, it closely resembles B.
muricata. However, its androeciumand gynoecium are quite different. Instead
of the greatly enlargedconnectives on the stamens opposite the 3 anteriorsepals,
these connectives are similarin size to those of the other stamens, and the styles
are stout and straight,not slender with the posterior styles lyrate as in B. muri-
cata; in its androecium and gynoecium B. quadriglandula resembles B. multifoliolata. Banisteriopsis quadriglandula is further unusual in that only the 2 poste-
rior sepals bear glands, hence the name I have chosen for it.
The Banisteriopsis muricata group (species 40-43)
This group includes B. muricata, a vine found throughoutCentraland South

America from Mexico to Argentina, B. oxyclada, a vine of the cerrados of the
Planaltoof Brazil, B. pauciflora, a vine endemic to Cuba, and B. goiana, which
is a non-viningshrubfound only in SE Goias in Brazil. These species share the
following characteristics(see Fig. 28 for illustrationof flower and fruit type in
this group):bracts and bracteoles persistent;pedicels pedunculate;buds pointed
at the apex; sepals involute and acute at the apex; calyx glands pink;petals pink,
laciniate, the fifth petal with the proximal half yellow, with thick fleshy claw
apicallyconstricted,the limb broadlyobovate and with the basal laciniaeenlarged
and gland-tipped;stamens with locules glabrous, the filaments opposite the anterior sepal and postero-lateralpetals longest, those opposite the posteriorpetal
and postero-lateralsepals flexuous and incurvedbetween the posteriorstyles, the
connectives on the 5 anteriorstamensglandular,those opposite the antero-lateral
sepals greatly enlargedand overtoppingthe locules by ca. 1 mm, the locules on
these stamens 0.6-0.8 mm long; styles slender, up to 0.2 mm wide, diverging
(except B. goiana), the posterior styles longer and lyrate; fruit softly pubescent
throughout,the hairs sessile and persistent.
40. Banisteriopsismuricata(Cavanilles)Cuatrecasas,Webbia 13: 503. 1958.
Fig. 28.
Banisteria muricata Cavanilles, Tom. III, Nona Dissertatio Botanica: 423. 1790.
Heteropterys argentea Humboldt, Bonpland & Kunth, Nov. Gen. et Sp. PI. 5: 165. 1821 [1822].
Type. Humboldt & Bonpland s.n., Colombia, fl (holotype, P).
Banisteria argentea (Humboldt, Bonpland & Kunth) Sprengel, Syst. Veg.: 388. 1825.
Banisteria ? metallicolor Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 46. 1832 [1833]. Type.
Saint Hilaire Catal. Bl, 649 fl (holotype, P; isotypes, P).
Banisteria acanthocarpa Adr. Jussieu, Ann. Sci. Nat. Bot., Ser 2, 13: 281. 1840. Type. Dombey
s.n., Peru, Cochero, fr (holotype, P; isotypes, P).
121
Banisteria atrosanguinea Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 281. 1840. Type. Poeppig
Pl. exs. 1752, No. 99, Peru, fl (holotype, P; isotypes, F, G, NY, US).
BanisteriabenthamianaAdr. Jussieu,Ann. Sci. Nat. Bot., Ser. 2, 13: 281. 1840.Type. Mathews
2024, Peru, fl (holotype, P; isotypes, BM, G, K, NY).
Banisteriapruinosa Martiusex Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 281. 1840.Type.
Martiuss.n., Brazil, Piaui, Oeiras, fl (holotype, M).
Banisteria schomburgkiana Bentham, London J. Bot. 7: 129. 1848. Type. Schomburgk 1st. Coll.,
844, Brazil, Roraima,Rio Branco, fl (holotype, K; isotypes, BM, F, K, NY, P, US, W).
Banisteriaschlimii Turczaninow,Bull. Soc. Imp. NaturalistesMoscou 36: IL 584. 1863.Type.
Schlim 108, Colombia,near Ocafia,fl (holotype, CW?, n.v.; isotypes, BR, G, K, LE).
Heteropterys pirayuensis Morong, Ann. New York Acad. Sci. 7: 66. 1893. Type. Morong 672,
Paraguay,between Pirayuand Yaguaron,fr (holotype, NY; isotypes, F, MO).
Banisteria acanthocarpa var. glandulifera Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem.: 31. 1900.
Types. Ruiz s.n., Peru, fr (syntype, B? destroyed;isosyntypes, F, K). Poeppig 1214, Peru,
near Cuchero, fr (lectotype, B? destroyed; isolectotype, F, W). Poeppig 2433, Peru, fl (syntype, B? destroyed; isosyntypes, G, LE).
Banisteriaargentea var. acuminataNiedenzu, Ind. Lect. Lyc. Brunsb.p. hiem.: 30. 1900.Type.
J. D. Smith (leg. Heyde et Lux) 3703, Guatemala, Jalapa, Laguna de Ayarca, fl (lectotype,
M; isolectotypes, G, MICH, NY, P, US).

Banisteria argentea var. obtusiuscula Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem.: 30. 1900.
Type. Rusbv 511, Bolivia, Mapiri, fl (lectotype, W; isolectotypes, F, GH, MICH, MO, NY,
P, US).
Banisteriaargentea var. transiensNiedenzu, Ind. Lect. Lyc. Brunsb.p. hiem.: 30. 1900.Type.
Poeppig 1879, Peru, Maynas,fl (lectotype, W; isolectotypes, F, US, W).
Banisteriametallicolorvar. aurea Niedenzu, Ind. Lect. Lyc. Brunsb.p. hiem.: 29. 1900.Type.
Spruce 4530, Peru, Tarapoto, fl (holotype, B? destroyed; isotypes, BM, BR, C, G, K, NY,
W).
Banisteriametallicolorvar.falcata Niedenzu, Ind. Lect. Lyc. Brunsb.p. hiem.: 28. 1900.Type.

Balansa 2402, Paraguay, L'Assomption, fr (lectotype, GOET; isolectotypes, BM, BR, K,
NY, P, S).
Banisteria metallicolor var. pruinosa (Martius ex Adr. Jussieu) Niedenzu, Ind. Lect. Lyc.
Brunsb.p. hiem.: 28. 1900.

Banisteriametallicolorvar. sericea Niedenzu, Ind. Lect. Lyc. Brunsb.p. hiem.: 29. 1900.Type.
Blanchet 2737, Brazil, Bahia, Rio Sao Francisco, near Utinga, fr (lectotype, W; isolectotypes, BM, BR, F, G, K, NY, P).
Banisteria metallicolor var. subrotunda Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem.: 28. 1900.
Type. Bang 2813, Bolivia, fl fr (lectotype, M; isolectotypes, BM, C, F, G, GH, K, MICH,

MO, NY, US).
Banisteria metallicolor var. subsalicina Niedenzu, Ind. Lect. Lyc. Brunsb. p. hiem.: 29. 1900.
Type. Burchell6913, Brazil, Goias, fl (lectotype, GOET;isolectotype, K).

Banisteriopsis
Flora 25:
Banisteriopsis
1910.
Banisteriopsis
argentea (Humboldt, Bonpland & Kunth) Robinson in Small, North American

133. 1910.
schomburgkiana (Bentham) Robinson in Small, North American Flora 25: 133.
illustris Rusby, Bull. New York Bot. Gard. 8: 97. 1912. Type. R. S. Williams 62,
Bolivia, Apolo, fl (holotype, NY; isotypes, BM, K).
Banisteriopsis williamsii Rusby, Bull. New York Bot. Gard. 8: 97. 1912. Type. R. S. Williams
809, Bolivia, Mapiri,fl (holotype, NY; isotypes, BM, US).
Banisteria williamsii (Rusby) Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 453. 1928.
Banisteria illustris (Rusby) Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 451. 1928.
Banisteria atrosanguinea var. benthamiana (Adr. Jussieu) Macbride, Field Mus. Nat. Hist., Bot.
Ser. 8: 120. 1930.

Banisteriopsismetallicolor(Adr. Jussieu)O'Donellet Lourteig,Lilloa 9: 259. 1943.
Banisteriopsis metallicolor var. sericea (Niedenzu) O'Donell et Lourteig, Lilloa 9: 260. 1943.
Banisteriopsis metallicolor var. subrotunda (Niedenzu) O'Donell et Lourteig, Lilloa 9: 259. 1943.
Banisteria muricata var. atrosanguinea (Adr. Jussieu) Macbride, Publ. Field Mus. Nat. Hist.,
Bot. Ser. 13(3):833. 1950.
Vining shrub or liana. Young branches flattened, appressed-white- or goldensericeous or -velutinous, older branches terete, glabrate, the bark grey-brown to
dark brown with numerous punctiform lenticels. Stipules triangular, 0.5-1.0 mm
long, sericeous to glabrate. Leaves with the petiole 5-16 mm long, white- or
golden-appressed-sericeous or more rarely velutinous, eglandular or with 1-2
Flora Neotropica
122
'f
.*
:}
'
.
.: -
'tF
iC
..
..
-1
FIG. 28. Banisteriopsis muricata group. A-H, B. muricata. A, flower bud x 4.2; B, flower x
4.2; C, stamens, semi-diagrammatic x 8.4 (stamen 1 is opposite anterior sepal, stamen 6 is opposite
posterior petal); D, stamens 1-6 lateral view x 8.4; E, gynoecium lateral view x 7; F, gynoecium
anterior
view x 7; G and H, samaras
x 1.4. J-L,
B. goiana.
J, leaf x 0.7; K, samara x 1.4; L,
gynoecium x 7. (A-F, Anderson 11712; G, Gates 401; H, Woytkowski7615; J-L, Hatschbach &
38237.)
Kummrow 38237.)
123
pairs of cupulateglands 0.2-0.4 mm in diameternear the apex, the lamina(3.5-)

5.3-12.7(-16.5) cm long, (2.3-)4.8-7.5(-9.0) cm wide, smaller in the inflorescence, ovate, elliptic or rotund, cuneate to cordate at the base, acuminate to
apiculate at the apex, flat at the margin, eglandularor bearing 2-4(-5) pairs of
stipitate glands 0.4-0.6 mm in diameterabaxially, sparsely white- or golden-appressed-sericeous to glabrate adaxially, abaxially sparsely to densely silver- or
golden-appressed-sericeous,the hairs sessile with the trabeculastraight,0.2-0.6
mm long, with the venation scalariform,prominulousto flat adaxially,prominent
abaxially. Inflorescence of 4-flowered umbels in dense axillary and terminal
cymes with reduced leaves, often with several branches per node, silver- or
golden-appressed-sericeousor more rarelyvelutinous;bracts and bracteoles0.51.2 mm long, broadly triangular,sericeous abaxially, glabrousadaxially, persistent; peduncle 0.5-3.0(-7.0) mm long; pedicels 3-12 mm long, 0.6-0.8 mm in
diameter, 0.3-0.5 mm without the hairs, white- or golden-appressed-sericeous,
rarely velutinous. Sepals ovate, acute at the apex, involute at the margin,silveror golden-sericeous abaxially, adaxially sericeous near the margin, 1.5-3.0 mm
long, 1.2-2.5 mm wide, all sepals eglandularor the 4 lateral sepals each biglandular, the glands 1.3-2.0 mm long, 0.5-1.0 mm wide, the sepals projecting0.61.0 mm beyond the glands. Petals pink, becomingpaler in age, the fifthpetal with
the basal half yellow, the 4 lateral petals reflexed between the sepals, the claw
0.8-1.4 mm long, 0.2-0.5 mm wide, the limb 4.0-8.5 mm long, 3.5-8.0 mm wide,
laciniate, orbicular,eglandular,the antero-lateralpetals cochleate, the posterolateral petals plane, the posteriorpetal with the claw erect, 1.4-3.0 mm long, up
to 1.4 mm in diameter, apically constricted, the limb 4.0-7.0 mm long, 3.5-6.0
mm wide, obovate, fimbriatewith the basal fimbriaeenlarged and gland-tipped.
Stamens with the filaments 1.0-3.6 mm long, connate basally, those of the posterior 3 stamens flexuous and inbent between the posterior styles, the locules
0.6-1.2 mm long, glabrous, the connectives of the stamens opposite the petals
and opposite the postero-lateralsepals 0.4-1.0 mm long, 0.3-0.7 mm wide, 0.10.6 mm deep, those opposite the antero-lateralpetals slightly glandular,those
opposite the 3 anterior sepals 0.9-2.2 mm long, 0.7-1.6 mm wide, 0.6-1.2 mm
deep, glandularand enlarged, those opposite the antero-lateralsepals projecting
0.8-1.7 mm beyond the locules. Ovary 0.8-1.4 mm tall, wbite-sericeous, the
styles slender and diverging, the anterior style 2.2-2.8 mm long, up to 0.2 mm
wide, the posterior styles lyrate, 2.6-3.6 mm long, 0.15-0.20 mm wide at the
apex, the stigmas capitate. Samarawith the carpophoreup to 1.0 mm long, 0.81.0 mm wide, the nut 4-14 mm tall, 4-7 mm long, rugose, tuberculate,muricate
or alulate, appressed-pubescent,the wing (12-)20-34(-45) mm long, (9-)11-16
(-20) mm wide, appressed-sericeous,the hairs sessile, the armsstraightto slightly
flexuous 0.15-0.30 mm long.
Type. Joseph de Jussieu s.n., Peru, fr (holotype, P-JU; isotype, P).
Common names. Peru: "ayahuasca," "ayahuasca negro," "ayahuasca rosada," "ayahuasca de los brujos," "sarcello." Bolivia: "bejuco hoja de plata."
Argentina: "sombra de tora." El Salvador: "bejuco de casa," "pastora," "ala
de zompopo."
Distribution.A vine in tropicalforest and semideciduouswoodlandfrom Chiapas, Mexico, southwardsthroughoutSouth America to Argentina.
Collected in flower and fruit in every month of the year.
Representative collections. MEXICO. Chiapas: Breedlove & Raven 13487 fl (F, GH, MICH, US);
Matuda 598 fr (GH, MICH, MO, US); 1744 (F, GH, K, MO, NY, US); 17064 (F, MICH); Nelson
3278 (US). GUATEMALA. Bernoulli & Cario 3004 fl (GOET, K, NY); J. D. Smith (leg. Heyde et
Lux) 3703 fl (G, M, MICH, NY, P, US); 6309 fl (F, G, K, M, MICH, MO, NY, US); Standley 6214
fr (F, NY, US); Steyermark 30745 fl (F). HONDURAS. Hazlett 2025 fl (MO); Molina 13004 fl (F,
124
Flora Neotropica
NY, US); 13312 fr (F, NY); Rodriguez 1363 fl (F); Standley 23240 fl (F). EL SALVADOR. Calderon
68 (NY, US); 894 fl (NY); 931 fl (NY, US); Padilla 86 fl (US); Standley 2417 fl (F). NICARAGUA.
Cousin s.n. fl (BR); Levy s.n. fl (P); 386 fr (C, G, P, RB). COSTA RICA. Brenes 18949 fr (F, NY);
Daubenmire 203 fl (F); Heithaus 423 fl (MO); A. Jimenez M. 1327 fr (F, NY); Lankester 612 fl (BM,
C). PANAMA. P. Allen 2776 fl (G, S); Croat 12536 fl (F, MICH, NY); Duchassaing s.n. (GH, P);
Gentry & Dwyer 3544 fr (NY); Hayes 229 fl (BM, BR, K, P, W). COLOMBIA. Magdalena: C. Allen
701 fl (F, MO); Cuatrecasas & Castaneda 24980 fl fr (US); Haught 3733 fl (US); 3807 fl (GH, US,
U). Atlantico: Dugand 1056 fl (F, US); 1138 fl (F, US); 5330 fl (W). Bolivar: Killip & Smith 14325
fl (F, GH, NY, S); 14506 fr (F, GH). Norte de Santander: Cuatrecasas et al. 12094 fl fr (BM, F, GH,
MO, U, US); 13446 fr (US); Fassett 25992 (NY, US); Garganta 927 fr (F). Santander: Killip & Smith
14991 fr (F, GH, MICH, US); St. John 20729 fl (NY, US). Cundinamarca: Dawe 37 fl (K, US);
Holton s.n. fl (K, NY); Kalbreyer 2037 fl (BM, C, K); Pennell 2820 fl fr (F, GH, MO, NY, US);
Schneider 1063 fl (S). Boyaca: Garcia Barriga 18060 fl (COL); Lawrance 508 fl (BM, F, G, GH, K,
MO, NY, S, U, US). Tolima: Garcia Barriga 12037 fl (US); Haught 2368 fl (MO, US); 6348 fl (US);
Lehmann 2628 fl (F, K, NY, US); 6102 (F). Huila: Rusby & Pennell 1109 fl (NY). VENEZUELA.
Zulia: Pittier 10654 fl (G, GH, NY, US); Tejera 8 fl (GH, US). Yaracuy: Steyermark et al. 100363
fl (NY). Carabobo: Archer 3117 fr (US); Pittier 13398 fl (F, G, GH, MO, NY, SU). Aragua: Agostini
655 fl (NY); Chardon 167 fl (US); Lasser 819 fl (G); Vogel 1121 fl (M); Ll. Williams 10453 fl (F,
MICH). Distrito Federal: Bredemeyer 204 fl (W); Funck 377 fl (K); 497 fl (G, P); van Landsberge
263 fl (S); Pittier 12493 fl (G, GH, M, MO, NY, US). Miranda: Ll. Williams 10585 fr (F, MO); 13600
fl (F). Sucre: Tamayo 2160 fr (IAN). Monagas: Maguire 27302 fl (MICH, NY). Barinas: Breteler 4254
fl (G, IAN, NY, U). Portuguesa: Steyermark & Rabe 97422 fl (NY). Guarico: Davidse 4232 fr (MICH);
Ll. Williams 10712/Alston 108 fr (BM, F). Bolivar: Bernardi 7955 fl (VEN); Maguire et al. 35965 fl
(MICH, NY); Tovar 2 fr (VEN); Trujillo 5965 fr (MY). GUYANA. A. C. Smith 3176 fl (F, G, GH,
K, NY, P, U, US, W). FRENCH GUIANA. Serre s.n. fl (P). ECUADOR. Asplund 10323 fl (S);
Camp E6 fr (F, NY, US); E3773 fl (BR, G, GH, MICH, MO, NY, P, RB, S, W). PERU. Loreto:
Klug 3032 fl (BM, F, G, GH, MO, NY, S, US); Woytkowski 5762 fl (C, G, GH, MO); 7615 fr (F,
MO); Ule 6862 fl (G, MG). Cajamarca: Weberbauer 6160 fl (F, GH, US); Woytkowski 5588 bud (GH,
MO, US); 5674 fl (G, GH, K, MO, US); 5719 fl (G, GH, MO). Amazonas: Sagastegui 5879 fl (US);
Kavap 126 fl (MO); Woytkowski 5640 fl (MO). San Martin: Belshaw 3156 fl (F, MO, NY, US); Klug
3506 fl (F, G, GH, K, MO, NY, S, US); 4373 fl (BM, F, GH, MO, NY, S, U, US); Schunke 3863 fl
(F, G, GH, MO, NY); Woytkowski 5072 fr (F, MO, P). Huanuco: Mathias 3614 fl (US); Plowman
5966 fr (MICH); Schunke 2037 fr (F, G, NY); Woytkowski 5332 fl (BR, C, F, G, GH, MO, P, S, US);
7878 fl (GH, MO). Junin: Killip & Smith 23378 fr (F, NY, US); 23757 fr (F, NY, US); 25011 fr (F,
NY, US); 25069 fr (F, NY, US). Huancavelica: Tovar 4651 fr (US). Cuzco: Mexia 8027 fr (BM, F,
GH, K, MO, NY, S, US); 8029 fl (F, GH, K, MO, NY, US); C. Vargas C. 1842 fl (GH); Weberbauer
7935 fl (F, G, S, US). Ayacucho: Dudley 9085 fr (F). BRAZIL. Roraima: Black 51-13716 fl (IAN);
51-13781 fl fr (IAN, UB); J. G. Kuhlmann 732 fl (MICH). Acre: Steward et al. P12817 fl (MG, MICH,
NY); Ule 9480 fl (MG). Amaz6nas: Liitzelburg 20658 fl (M, R); 20686 fl (M, R). Para: Dahlgren &
Sella 122 fl (F, GH, S, US); Ducke 2873b fl fr (MG); Froes 27088 fl (IAN); A. Lima 53-1372 fl (IAN,
K). Maranhao: Hatschbach & Kummrow 38470 fl (MICH); Pires & Black 2588 fl (IAN, US). Ceara:
Allemdo & Cysneiros 214 fl (R); Ducke 2262 fl fr (MG); Gardner 844 fr (G); Huber 250 fr (MG);
Lofgren 481 fl (R, S). Rond6nia: Black et al. 52-14641 fr (IAN); Prance et al. 5177 fr (MG, MICH,
NY). Mato Grosso: Anderson 11234 fl (MICH); Commissdo Rondon (leg. Hoehne) 4758 fl (R, SP);
Malme s.n. fl (R, S); Patr. da Sa. Manso 325 (=Martius 325) fl (BR, P); N. Santos s.n. fl (R). Goias:
Burchell 8759 fr (GH, GOET, P); Gardner 3624 fl (BM, K); Oliveira 1566 fl (IAN, UB). Distrito
Federal: Gates 401 fl (MBM, MICH); Heringer 14400 fl (UB); Irwin et al. 14020 fl (MICH, NY, UB);
14027 fr (MICH, NY, UB). Bahia: Blanchet 2737 fr (BM, BR, F, G, K, NY, P, W); Castellanos
25121 fl (GUA, NY); Duarte 396 fl (MICH); Campos Porto 2434 fl fr (RB). Minas Gerais: Anderson
11606 fl (MICH); Carcerelli 16 fl (RB); Glaziou 16362a fl (P, R); Heringer 554 fl (SP); Regnell III.317
fl fr (BR, F, K, M, NY, P, R, S, US). Rio de Janeiro: Anderson 11712 fl (MICH); Frazdo s.n. fl
(RB); Glaziou 11816 fl (BR, C, G, K, NY, P, R); Miers 3460 fl (K, P); Peckolt 180 fl (W). Sao Paulo:
Anderson 11148 fl fr (MICH, NY); Eiten et al. 5918 fl (NY); M. Kuhlmann 2819 fl (SP); Mendes 245
fl (IAN, RB, SP); Mosen 1151 fl (C, R, S, US). Parana: Dusen 16622 fl (G, GH, MO, S); Hatschbach
21060 fl fr (MICH); Imaguire 1833 fl (MICH, RB); Lindeman & Haas 4409 fl (NY, U); Schwarz 7330
fl (S). BOLIVIA. Bang 1365 fl fr (BM, F, G, GH, K, M, MICH, MO, NY, US, W); Buchtien 913 fl
(F, GH, MICH, NY, P); Lorentz & Hieronymus 615 fr (GOET); Peredo s.n. (P, U, W); R. S.
Williams 192 fr (BM, K, NY, US). PARAGUAY. Fiebrig 2677 fl (BM, G, GH, GOET, K, M, NY,
P, S, U, US, W); Hassler 4089 fl fr (BM, F, G, GH, K, MICH, MO, NY, P, S, W); 12107 fl (BM,
C, G, GH, K, MO, NY, S, US); Jorgensen 4219 fl (F, GH, MO, NY, S, US); Woolston 790 fl (C,
K, NY, S, SP, U). ARGENTINA. Salta: Meyer 8497 fr (NY); Pierotti 1205 fr (NY, W); Rojas 11467
fl (U, US); Schreiter 11110 fl (F, GH, NY); Steinbach 7087 fl (BM, F, G, GH, GOET, MO, NY, S,
U). Misiones: Bertoni 3852 fl (BR); Ekman 1530 fl (GH, MO, NY, S, US); Meyer 5715 fl (GH, U);
125
Montes 15562 fr (G, W); Schwarz 7193 fl (P, S). Corrientes: Huidobro 2210 fl (S, W); Ibarrola 4030
fl (W); Pedersen 3883 fr (BR, C, G, GH, MO, NY, P, S, U, US).
This species is the most widespreadin the whole genus, occurringfrom Mexico
throughCentraland South Americato Argentina.Withinthis range, B. muricata
exhibits much variationin size and shape of leaves, type and density of pubescence, length of peduncle and pedicel, and in the fruit, size and shape of the wing
and surface of the nut. However, in the form of its flowers, B. muricata shows
great constancy (see Fig. 28). After examinationof all the specimens availableto
me I have concluded that any division of the variationwithin B. muricata would
be at best arbitrary,and of limited usefulness. All of the charactersused in the
past as a basis for segregatingtaxa within the B. muricata complex are represented in individualsfound in diverse parts of its range, and I was able to find
transitionalforms between previously segregated taxa, such that the variation
forms a continuum.I shall, however, describe the patternsof variation,and hope
that this will prove useful to individualsable to observe these plants in the field.
Perhaps a more detailed knowledge of the ecology of this species will suggest a
meaningfulsubdivisionin the future.
1. Leaf size and shape. Leaf size is a very variablecharacterin many species
in the genus; in vines, like B. muricata, small leaved specimens may represent
an artifact of collection, since the large leaves on the main branches are often
not collected. The leaf shape may be orbicularto ovate to lanceolate, with the
apex apiculateto acuminate;there seems to be no correlationbetween leaf shape
and geographicallocation, or with other characterswhich vary, and it would not
appearto be a good basis for taxonomic segregation.
2. Type and density of pubescence. Banisteriopsis muricata is characterised
by appressed pubescence on the abaxial surface of the leaves. Individualsfrom
Argentina,Paraguay,and southernBrazil exhibit sparse, very appressedpubescence, such that the leaves appeargreen but with a metallicsheen. However, this
type of pubescence is also commonin plants from Bolivia, Peru, CentralAmerica
and Mexico, and in some of these individualsfrom Bolivia and Peru the pubescence is golden. Most Venezuelan specimens have sparse pubescence, but the
hairs are not so closely appressedto the leaf surface, and in a very few individuals, the pubescence is subtomentose. These may have been young or regenerating individuals;seedlings which I grew had velutinous pubescence throughout.
Specimensfrom Colombia,Guyanaand northernBrazil, and also some specimens
from CentralAmericaand Peru, exhibit very dense appressedpubescence, giving
the leaves a white shiningappearance.
The pubescence of the inflorescences can be appressed or velutinous. The
formeris more common, but velutinouspubescence is found in some individuals
from Colombia, Venezuela, Ecuador, Peru, Brazil and Bolivia; some specimens
from Bolivia and Peru have golden-velutinouspubescence, and have been segregated as B. atrosanguinea or B. benthamiana in the past. Since in the same
areas individuals with golden-appressedand silver-velutinous pubescence are

found, I do not feel this recognitionis justified.
3. Peduncle and pedicel length. Throughoutmost of its range, B. muricatahas
peduncles which are three to four mm long; in Argentinaand Paraguayand Central America some individuals have peduncles only 0.5-1.0 mm long, and in
CentralAmerica and Mexico some individualshave peduncles up to seven mm
long. The pedicel also varies in length, with short-pedicelledindividuals(3-7 mm
long) being common in Peru. However, neitherof these charactersappearsto be
correlatedwith the other characterswhich vary such as leaf and stem pubescence.
126
Flora Neotropica
4. Fruit characters. Niedenzu represented most of the variation which I am

including under B. muricata in two species, B. metallicolor and B. argentea,
which he distinguishedchiefly on differences in their fruits. The former has the

nut of the samara rugose to tuberculate, whereas in the latter it is aculeate to
alulate. Niedenzu recognised that without fruits it was not possible to separate
the two species, and since there is considerable variation in the fruit on one
individual, and in some cases, even between the two sides of the same samara
(e.g., one side of the nut can be rugose, and the other side tuberculateto muricate), this seems to be an unsatisfactorycharacter for taxonomic recognition.
Specimensfrom the southernand easternpart of the rangeof B. muricatausually
have fruit nuts which are rugose to tuberculate, but this fruit type is found
throughoutthe range of the species. Similarly, individualswith large fruits and
the nut with long aculeate spines, previously segregatedas B. acanthocarpa, are
found in Peru and also in CentralAmerica. Vegetatively, these individualsare
indistinguishablefrom plants with rugose fruit nuts.
41. Banisteriopsisoxyclada(Adr. Jussieu) B. Gates, comb. nov.

Banisteriaoxyclada Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 282. 1840.
Banisteria albicans Grisebach, Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn:131.
1875.Types. Burchell7292, Brazil, Goias, fr (lectotype, GOET;isolectotype, K). Warming
s.n., Brazil, MinasGerais, Lagoa Santa, fl (syntype, C; isosyntypes, C, P). (I have used the
Burchellspecimen as lectotype, since it is in fruit, and Grisebachconsideredthe fruits of
the Brazilianspecimensto be different,and thus to warranttaxonomicrecognition.)
Banisteria oxyclada f. brevialata Niedenzu, Ind. Lect. Lyc. Brunsb.: 26. 1900. Type. O. Kuntze
s.n., Bolivia, Velasco, fr (holotype, B? destroyed;isotypes, F, NY).

Banisteria oxyclada var. rhombifoliaSkottsberg,Kongl. Svenska Vetenskapsakad.Handl. 35:
14. 1901.Type. Regnell II, 29, Brazil, MinasGerais, Caldas,fl (holotype, S; isotypes, W).
Liana; young branchesand inflorescencebranchesflattened,densely white- or

golden-tomentose, old branches terete, eventually glabrate, bark grey. Stipules
1-2 mm long, on old branchesborne on a conspicuous raised interpetiolarridge,
tomentose. Leaves with the petiole 14-25(-40) mm long, white-tomentose, canaliculate, bearing 1(-5) pairs of cupulate glands near the apex, the lamina 6.014.2(-18.0) cm long, 5.0-11.1(-14.8) cm wide, ovate, obtuse to cordate at the
base, apiculateat the apex, bearinga pair of glands basally beside the costa and
1-2 pairs of stipitateglands up to 0.4 mm in diameternear the marginabaxially,
adaxially densely velutinous to glabrate, abaxially densely white-tomentose to
sericeous, the hairsT-shaped,the trabecula0.4-0.8 mmlong, straightto flexuous,
with the venation prominentabaxially,and flat adaxially.Inflorescenceof 4-flowered umbels borne in large, dense, axillaryand terminalcymes, white-tomentose;
bracts and bracteoles 1.5-1.8 mm long, triangular,tomentose, persistent;peduncle 0-4 mm long, pedicels (8-)11-17 mm long, 1.0-1.5 mm in diameter, 0.7-0.9
mm without the hairs, tomentose to loosely sericeous. Sepals ovate, acute and
often inrolled at the apex, sericeous abaxially, sparsely sericeous near the apex
adaxially, 3.2-3.8 mm long, the anterior sepal 1.5-2.0 mm wide, the 4 lateral
sepals 2.5-3.0 mm wide, each biglandular,the glands 2.4-2.6 mm long, 0.8-1.0
mm wide, the sepals projecting 1.4-1.5 mm beyond the glands. Petals white to
pink, the 4 anterior petals reflexed between the sepals, the claws 0.6-1.5 mm
long, 0.5-0.8 mm wide, the limbs 7.5-13.0 mm long, 7.0-10.0 mm wide, laciniate,
eglandular,orbicular,the antero-lateralpetals cochleate,the posteriorpetalwiththe
claw erect, yellow, 3.0-3.5 mmlong, up to 1.3 mmwide, apicallyconstricted,thick
and fleshy, the limb 6.5-9.5 mm long, 4.5-8.5 mm wide, obovate, fimbriateto
laciniate, the basal fimbriaeenlargedand gland-tipped.Stamenswith the filaments
127
2.5-4.0 mm long, basally connate, those of the posterior 3 stamens flexuous and
inflexed between the posterior styles, the locules 0.6-1.2 mm long, glabrous, the
connectives of the stamens opposite the petals and opposite the postero-lateral
sepals 0.4-0.8 mm long, 0.3-0.8 mm wide, 0.1-0.6 mm deep, with only those
opposite the antero-lateralpetals glandular,those of the stamens opposite the 3
anteriorsepals glandularand enlarged, 1.2-1.8 mm long, 0.9-1.1 mm wide, 0.71.2 mm deep, those opposite the antero-lateralsepals projecting 1.0-1.2 mm
beyond the locules. Ovary 1.2-1.4 mm tall, white-sericeous, the anterior style
3.0-3.2 mm long, taperingto 0.2 mm wide apically, straight,the posterior styles
lyrate, diverging,3.6-3.8 mm long, taperingto 0.15 mm wide apically, the stigmas
capitate. Samarawith carpophoreup to 1.5 mm long, up to 1 mm wide, the nut
8-11 mm tall, 4.5-5.0 mm long, smooth to rugose, appressed-pubescent,the wing
(22-)28-40 mm long, 11-16 mm wide, appressed-sericeous,the hairs sessile, persistent, T-shaped, the trabeculastraight,0.2 mm long.
Chromosome number. n = 10. Voucher, Anderson 11144.
Type. d'Orbigny 804, Bolivia, Chiquitos, Santa Anna, fl fr (lectotype, P).

Claussen s.n., Brazil, Minas Gerais, fl (syntype, P). I have chosen the d'Orbigny
specimen as lectotype since the originaldescriptionincludes a descriptionof both
flowers and fruits, and this specimen has both.
Distribution.(Fig. 27). Large vine over trees and at gallery marginsthroughout
the Planaltoof Brazil, and apparentlyextending to Bolivia and Paraguay.
Collected in flower November to June and in fruit Marchto July.
Representative collections. BRAZIL. Para: Frdes 3002 fr (IAN); Pinheiro & Carvalho 677 fl (IAN).
Maranhao: Lisboa 2443 fr (MG, RB). Mato Grosso: Malme s.n. fl (S); 3414 fl (S). Goias: Anderson
7360 fl fr (MICH, NY); Glaziou 20755 fl (BM, BR, C, G, K, P); Hatschbach 36956 fr (MICH); Irwin
et al. 14976 fl (MICH, NY); Pohl 1695.d1533 fl (F, GH, K, NY, W). Distrito Federal: Gates 400 fl
(MICH); Martin 467 fr (GH); Santos 1633/Saco 1866 fl (MICH, NY); Sucre 667 fr (HB, IAN, NY,
RB). Minas Gerais: Assis 217 fl (GH, SP, US); Barreto 7705 fl (F, R, US); Claussen 548 fl (C, NY,
P, S, US); Glaziou 18945 fl (BM, BR, C, K, NY, P, R); Heringer 7483 fl (UB); Macedo 2260 fl
(MICH, S, SP); Pires 57945 fr (MICH, NY); Regnell 11.29 fl (BR, C, K, M, NY, P, R, S, U, US, W).
PERU? Hanke s.n. fl (M). PARAGUAY. Hassler 10189 fl (C, G, GH, K, MICH, MO, NY, P, RB,
S, US, W); s.n. fl (G).
Local names. Brazil: "Cipo folha de prata"; "cipo prata."
This species is very close to B. muricata, from which it is distinguishedby its

longer petioles, larger leaves with tomentose pubescence, the prominentinterpetiolar ridge, and largerflowers with thicker pedicels.
Throughoutmost of its range the abaxial leaf pubescence is tomentose, the
hairs very dense with flexuous trabeculae,giving the leaf a dense opaque white
appearance.However, in Goias, some collections have leaves which are shining
metallo-sericeousabaxially, the trabeculaestraightand aligned more or less parallel to each other. I do not consider this variation to have any taxonomic significance.
42. Banisteriopsispauciflora(Humboldt,Bonpland& Kunth)Robinsonin Small,
North AmericanFlora 25: 133. 1910.
Banisteria pauciflora Humboldt, Bonpland & Kunth, Nov. Gen. et Sp. P1. 5: 160. 1821 [1822].
Banisteria podadena Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 31: 392. 1858. Type. J.
Linden 2075, Cuba, Santiago, Nimanima, fl (holotype, CW or KW, n.v.; isotypes, BR, F,
G, K, LE, W).
Liana or vining shrub. Branches terete, appressed-sericeous,eventually glabrate, the bark pale brown with pale transverse lenticels in fissures in the bark.
Stipules absent. Leaves with the petiole 5-8 mm long, eglandular,sericeous, the
128
Flora Neotropica
lamina 1.3-6.8 cm long, 0.7-3.9 cm wide, narrowlyovate to elliptic to rotund,

with the base cordate to obtuse, the apex acute to apiculate,the marginrevolute,
bearing 1-2 pairs of stalked glands 0.2-0.3 mm in diameternear the marginabaxially, sparsely appressed-pubescentadaxially, appressed-sericeousabaxially, the
hairs T-shaped, the trabecula 0.2-0.5 mm long, sessile, with the veins slightly
impressed adaxially, the lateral veins prominent abaxially. Inflorescence of
4-floweredumbels on axillary branches, the umbels borne in cymes or dichasia,
appressed-sericeous;bracts and bracteoles 0.8-1.5 mm long, triangular,persistent; peduncle 0.5-3.5 mm long; pedicels (7-)11-18 mm long, 0.4-0.5 mm in
diameter,0.4 mm without the hairs, appressed-sericeous.Sepals sericeous abaxially, minutely sericeous adaxiallyat the apex, ovate, acute at the apex, 2.0-2.5
mm long, 1.5-1.8 mm wide, the 4 lateral sepals biglandular,the glands pink or
red, 1.0-1.8 mm long, 0.6-0.8 mm wide, the sepals projecting1.0-1.5 mm beyond
the glands. Petals pink or white, fimbriate,the 4 anteriorpetals reflexedbetween
the sepals, the claw 1.2-1.9 mm long, 0.4-0.5 mm wide, the limb 6.0-7.5 mm
long, 5.5-7.5 mm wide, orbicular,eglandular,the antero-lateralpetals more cochleate, the postero-lateralpetals plane to slightly convex, the posteriorpetal with
the claw erect, 2.5-3.8 mm long, up to 1.1 mm wide, apicallyconstricted,the limb
4-5 mm long, 4.0-4.5 mm wide, orbicular,the basal fimbriaeenlargedand glandular. Stamens with the filaments 0.8-3.8 mm long, those of the 3 posterior
stamens flexuous and inflexed between the posterior styles, connate basally, the
locules glabrous, 0.4-1.3 mm long, the connectives of the stamens opposite the
petals 0.3-0.9 mm long, 0.2-0.5 mm wide, 0.1-0.4 mm deep, not glandular,those
opposite the sepals 0.9-1.4 mm long, 0.4-1.0 mm wide, 0.5-0.9 mm deep, globose, glandularand enlarged,projectingbeyond the locules, those of the stamens
opposite the antero-lateralsepals projecting0.7-0.8 mm. Ovary 1.0-1.2 mm high,
white-sericeous, the anterior style straight, 2.4-2.6 mm long, 0.6 mm wide basally, taperingapically to 0.2 mm, the posterior styles diverging, slightly lyrate
basally, 2.5-2.6 mm long, 0.6 mm wide at the base, taperingapically to 0.1 mm.
Samarawith the carpophore1.0-1.5 mm long, 0.3-0.4 mm wide, the nut 5-9 mm
tall, 3.5-6.0 mm long, rugose, appressed-pubescent,the wing 16-23 mm long, 710 mm wide, appressed-sericeous.
Type. Bonpland s.n., Cuba, fl (holotype, P; isotype, P).
Distribution.Found only on the island of Cuba.
Collected in flower and in fruit in every month of the year.
Representative collections. CUBA. Alain 1608 fl (GH, US); Britton et al. 10323 fl (NY, US, U);
Clemente 2258 fl (GH, US); Regnell 111.1422 (leg. Ekman) fl (NY, S); 111.7639 fl (F, G, S); Howard
5810 fl fr (GH, MO, NY, US); Jack 8055 fl (F, GH, S, US); 8132 fl (F, GH, K, NY, S, US); Le6n
6380 fr (GH, NY); de la Sagra s.n. fl (P, W); Wright 95 fl (BR, F, G, GH, MO); 347 fl (GOET, NY);
1584 fl (GH, GOET, K, MO, P, S); 2158 fl (BM, G, K, 'MO,NY, P, S, US, W).
This species is most closely related to B. murcata. It can be distinguishedby

its smallerelliptic leaves with revolute margins,stipitateleaf glands, and smaller
fruit wing. It is the only species of Banisteriopsis thai grows in Cuba.
43. Banisteriopsisgoiana B. Gates, sp. nov.
Fig. 28.
Banisteria sailcijolia sensu Grisebachin MartiusFl. bras. 12(1): 53. 1858, et auctt. seq.; non
DC..
Frutex usque 2 m altus, non volubilis. Rami juniores complanati, appressosericei, ramivetustioresteretes, demumglabri,badii, lenticellispallidisnumerosis
punctati. Stipulae minutae vel nullae. Folia opposita vel ternata, lamina (1.9-)
4.9-8.9 cm longa, (0.4-)1.1-3.2 cm lata, lanceolata vel rarius obovata, in inflo-
129
rescentia minori et lineari, basi cuneata obtusave, apice acuta apiculatave, margine 2 glandes sessiles 0.2-0.3 mm diametrobasi in nervis lateralibusvel in costa
subtus gerenti, supra sparsimappresso-pubescenti,subtus dense metallo-sericea,
pilis T-formibus stipitatis, trabecula recta 0.6-1.2 mm longa, petiolo 3-7 mm
longo, sericeo, eglanduloso. Inflorescentiaramos terminans,floribusin umbellis
4-floriferisfasciculatis, umbellis in cymis dispositis, appresso-sericea;bracteis
bracteolisque 0.5-1.5 mm longis, triangularibus,abaxialitersericeis, adaxialiter
glabris, persistentibus;pedunculis0.5-6.5 mm longis, pedicellis 7-12 mm longis,
0.6-0.7 mm diametro, 0.3-0.4 mm sine indumento, appresso-sericeis. Sepala 5,
abaxialiter sericea, adaxialiterglabra et margine sericea, anguste ovata, apice
acuta et involuta, 2.5-3.5 mm longa, 1.0-2.0 mm lata, 4 lateraliaglandes roseas
1.6-2.5 mm longas, 0.6-1.0 mm latas gerentia,glandes 1.2 mm superantia.Petala
5, rosea, denticulata,4 lateraliainter sepala patentia, ungue 1.0-2.0 mm longo,
0.3-0.4 mm lato, limbo 6.0-7.0 mm longo, 4.5-6.0 mm lato, orbiculari,eglanduloso, 2 anterioribusquam 2 posterioribus concavioribus; petalum posticum
ungue erecto, 2.2-2.5 mm longo, usque 1.0 mm lato, apice constricto, limbo 5.56.0 mm longo, 3.0-4.0 mm lato, rotundo vel obovato, retroflexo, piano convexove, dentibus basalibus dilatatis glandulosisque.Stamina 10, filamentis 1.52.5 mm longis, proximaliterconnatis, 3 posterioribusflexuosis sed non inter stylos
posticos inflexis. Antheraeloculis 0.6-1.0 mm longis, glabris, connectivis canarinis, petalis oppositis 0.4-0.8 mmlongis, 0.4-0.6 mm latis, 0.2-0.6 mmprofundis,
eglandulosisvel 2 petalis anterioribusoppositis interdumglandulosis,illis sepalis
oppositis 0.5-1.9 mm longis, 0.4-0.9 mm latis, 0.2-0.9 mm profundis,glandulosis
dilatatisque,omnibus loculos superantibuset 2 sepalis anterioribusoppositis loculos usque 1.4 mm superantibus.Ovarium0.9-1.0 mm altum, carpellis 3, uniovulatis, liberis, toro pyramidaliadnatis, candido-sericeis;styli 3, aequales, 1.92.4 mm longi, recti parallelique,glabri, 0.2 mm diametro versus apicem, stigmatibus capitatis. Fructus ex 3 samaris (vel abortu 1-2) constans; samara carpophoro usque 0.3 mm longo, 0.4 mm lato, nuce ovoidea 6-7 mm alta, 3-4 mm
longa, rugosa, appresso-pubescenti,ala 11-17 mm longa, 8-12 mm lata, appressosericea vel glabrata,pilis sessilibus, T-formibus,trabecularecta rigidave, 0.250.35 mm longa.
Type. Hatschbach & Kummrow38237, Brazil, Goias, Mun. Morrinhos,Chapadao, fl fr (holotype, MBM; isotype, MICH).
Distribution.(Fig. 27). Found only in the cerrados of SE Goias.
Collected in flower December, January, March, August and October, and
in fruit January,March, August and October.
BRAZIL. Goias: Barroso et al. s.n. fl (RB 141454); s.n. fr (RB 141456); Burchell 6103 fl fr (GOET,
K, P); Heringer 12171 fl fr (UB); Heringer & Eiten 14068 fl (MICH); 14164 fl (MICH); Macedo 3530
fl (NY, RB, S, US); Weddell1996 fl fr (P).
This species is closely related to B. muricata, and is the only memberof this
group which is not vining. It can be readily distinguishednot only by its shrubby
habit, but also by its narrowly elliptic to lanceolate leaves, its straightparallel
styles, and the smallerfruit wing.
The Banisteriopsis martiniana group (species 44-54)
Membersof this largegroupof species occur in Amazonianand northernSouth

America, and in CentralAmerica. The species of this group share the following
characters:vines; leaves with scalariformsecondary veins; leaf marginusually
bearingglands which are often enlargedand cupulate, and sometimes developed
into cilia, especially on the inflorescenceleaves; pedicels sessile; bracts and brac-
130
Flora Neotropica
teoles triangular,persistent; calyx glandularor eglandular;corolla yellow, glabrous; fruit with stiff irritatinghairs; samarawith small nut with well-developed
filiformcarpophoreand without lateral winglets, wings held suberect.
The flower structureis not uniform in this group. Banisteriopsis martiniana
and B. velutinissima have more or less identical flower structure, which is like
that of B. muricata, with small flowers with enlargedconnectives which greatly
overtop the locules on the stamens opposite the antero-lateralsepals, the claw
of the posteriorpetal constricted,and the posteriorstyles longerand more slender
than the anteriorstyle, and lyrate. These species occur in lowland rain forest in
Amazonia and northwestern South America. Banisteriopsis elegans, B. wilburii,
B. padifolia, B. polygama, B. grandifolia and B. alternifolia have a more or less
similarflower structure,with the connectives on the stamensopposite the anterolateral sepals not greatly enlargedand not or a little overtoppingthe locules, the
claw of the posteriorpetal usually not constricted, and the styles stout, more or
less equal, and straightor slightlydivergingapically. These species occur around
the periphery of the Amazon basin along the Andes and their extension into
Venezuela, and north of the Andes into Central America. Banisteriopsis pulcherrimaand B. maguireihave flowers with enlargedconnectives on the stamens
opposite the three anteriorsepals, and longer posterior styles which are lyrate;
they are both endemic to the GuayanaHighlandin Venezuela and adjacentGuyana.
44. Banisteriopsismartiniana(Adr. Jussieu) Cuatrecasas,Webbia 13: 498. 1958.
Banisteria martiniana Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 284. 1840.
Banisteria leptocarpa Bentham, London J. Bot. 7: 129. 1848. Type. Martin s.n., French Guiana,
Cayenne, fr (lectotype, BM). Schomburgk 1st Coll., single specimen, Guyana (syntype, K
n.v.).
Banisteriopsis leptocarpa (Betham) R. O. Williams, Flora Trinidad & Tobago 1: 131. 1929.
Liana, young branches minutely tomentose, old branches glabrous, dark to

red-brownwith darkprominulouslenticels. Stipulesminute,triangular,pubescent
to glabrate. Leaves subcoriaceousto coriaceous, with the petiole 4-20 mm long,
glabrousor minutelytomentose at first, eglandular,the lamina4.5-17.0 cm long,
2.0-8.5 cm wide, those associated with the inflorescence smaller and often reduced to bracts, ovate to elliptic, obtuse to cordate at the base, obtuse to acute
to long acuminateat the apex, with the marginflat to slightly revolute, bearing
minute glands or large cupulateglands, more numeroustowards the apex, abaxially at the base of the leaf beside the midribbearinga pairof sessile or cupuliform
glands, glabrous adaxially, at first sparsely appressed-sericeous,abaxially very
soon glabrate or the midrib persistently pubescent, smooth or with the lateral
veins and reticulation prominulousto prominent on both sides. Inflorescence
axillary, the branches up to 40 cm long, minutely brown- or white-tomentose,
rarely glabrate, bearing greatly reduced leaves with a well developed pair of
glands at the base and more or less well developed glands or ciliate extensions
on the margin, of 4-flowered umbels borne in condensed dichasia of usually 3
umbels, the umbel-bearingbranches 1-8(-15) mm long; bracts and bracteoles
0.6-1.2 mm long, rounded at the apex, sparsely pubescent to glabrate, involucrate, persistent; pedicels sessile, 6-14 mm long, 0.3-0.6 mm in diameter, glabrous or sparselypubescent. Sepals sparsely appressed-pubescentabaxially,glabrous adaxially, 1.5-2.5 mm long and wide, rounded at the apex, eglandularor
the 4 lateralsepals biglandular,the glands 1.0-1.5 mm long and 0.6-1.2 mm wide,
the sepals enlargedand spreadingin fruit. Petals yellow, dentate to lacerate, the
131
4 lateralpetals reflexed between the sepals, with the claw 1.5-3.0 mm long, 0.30.4 mm wide, the limb 4.5-6.5 mm long and wide, orbicular,the antero-lateral
petals cochleate, the postero-lateralpetals plane or slightly concave, eglandular,
the posteriorpetal with the claw 2.5-3.0 mm long, up to 0.8 mm wide, thick and
fleshy, apicallyconstricted,the limb4.5-6.5 mm long, 2.5-3.5 mm wide, obovate,
reflexed, the narrowbasal part with 2-3 pairs of gland-tippedteeth. Stamenswith
the filaments 1.4-3.8 mm long, those opposite the 3 styles (i.e. opposite the
anteriorsepal and the postero-lateralpetals) longer than the others, those of the
three posterior stamens inflexed between the posterior styles; anthers with the
locules glabrous, 0.5-1.0 mm long, the connectives of the stamens opposite the
3 anteriorsepals 1.2-1.6 mm long, 1.0-1.3 mm wide, 0.6-1.0 mm deep, globose,
enlarged and glandular,those opposite the antero-lateralsepals overtoppingthe
locules 0.6-1.0 mm, those of the stamens opposite the petals and the posterolateral sepals 0.4-0.8 mm long, 0.3-0.5 mm wide, 0.1-0.2 mm deep, oblong, not
glandularor glandularat the apex. Ovary 1.0 mm tall, densely hairy throughout
or only apically, the anteriorstyle straight, erect or directed anteriorly,2.0-2.2
mm long, thicker than the posterior styles, the posterior styles divergingbasally,
2.6-3.0 mm long, slender, stigmas truncate. Fruit with filiformcarpophoreup to
3 mm long, the nut 3.0-4.0 mm tall and 2.0-3.0 mm long, appressed-pubescent,
the hairs stiff and irritating,with the trabecula0.6-1.5 mm long, smooth or with
a reticulatepattern of ridges laterally, the wing 16-32 mm long, 5-12 mm wide,
arisingfrom the upper1/3to /2 of the distal marginof the nut, appressed-pubescent
like the nut, to glabrate.
Key to the Varieties of Banisteriopsis martiniana
1. Leaves plane, up to 12.5 cm long; inflorescencebrancheswith dense minutelytomentose
ferrugineouspubescence;leaf marginbearingcupulateglands;inflorescenceleaves sparsely
a. var. martiniana.
sericeous.
1. Leaves falcate, up to 17.0 cm long; inflorescencebranchesminutelytomentose, the hairs
b. var. subenervia.
white; leaf marginbearingminuteglands;inflorescenceleaves glabrous.
44a. Banisteriopsismartinianavar. martiniana

Branches minutely puberulent,the hairs brown, soon glabrate. Leaves plane,
subcoriaceousto coriaceous, with the petiole 4-15 mm long, the lamina4.5-12.5
cm long and 2.0-6.9 cm wide, truncateto subcordateat the base, obtuse to shortacuminate at the apex, bearing cupulate glands on the margin, more numerous
towards the apex, sparsely appressed-sericeousat first, soon glabrate, on both
sides. Inflorescencebranches minutelytomentose, the hairs ferrugineous,rarely
glabrateor more loosely tomentose, bearingreducedleaves with a pair of glands
at the base beside the costa, and well developed cupulate marginalglands or
ciliate extensions of the marginapically, appressed-sericeouson both sides, the
branches subtendingthe umbels 4-8(-15) mm long.
Type. Martins.n. (92), Guyane Frangaise, Cayenne, fl yng fr (holotype, P).
Distribution.(Fig. 29). In the lowlandsof coastal NE South America, extending
up to elevations of 2000 m in the Guyana Highlands,and inland to Manaus.
Collected in flower in Februaryto December, and in fruit in April, and June
to December.
Representativecollections. VENEZUELA. Aragua: Gines 5720 fl (US). Terr. Delta Amacuro:
Bond et al. 76 fl (US); Steyermark 87415 fr (MICH, P). Bolivar: Bernardi 1627 fl (VEN); Cardona
2009 bud (VEN); Ferrari 1021 fl (MY, NY); Maguire 33888 fl (NY); 53523 fl fr (NY, VEN); Stey-
ermark1110 st (MICH,NY); 58559 fl (F, US, VEN); 59180 fr (F, NY, US); 59710 fl (F, NY, VEN);
132
Flora Neotropica
60807 fl (F, NY, VEN). TRINIDAD. Broadway 7172 fl (MO, US); Eggers (leg Rensch) 1145 fl (M,
P, W). GUYANA. Archer 2396 fr (US); De La Cruz 1718 fl (F, MO, NY, US); Fanshawe 5136 fl
(U, US); Irwin et al. 55415 fl fr (U, US); Maguire et al. 32499 fr (MICH, NY); Sandwith 143 fr (P,
U); Rich. Schomburgk 999 fl (GOET, NY, W); Robt. Schomburgk 2nd coll. 651 fl (C, F, G, P); Tillett
& Tillett 45387 fl (MICH, NY). SURINAME. Boerboom 9113 fl (U); Lanjouw 440 fl (U); Maguire
24615 fl (BR, F, G, GH, MO, NY, P, U, US); 24632 fl (NY, U, US); Schulz 8705 fl (U). FRENCH
GUIANA. Melinon s.n. fl (P); Oldeman B-1303 fl (P); Hb Sagot s.n. fl (NY). BRAZIL. Amaz6nas: Coelho s.n. fl (SP); Prance 23552 fl (MICH).
Local names. Venezuela: "chinak," "yukuna." British Guiana: "mabude hi."
This species is characterisedby its small flower with the posterior petal with
fleshy claw, apically constricted, the limb obovate with the basal teeth glandular,
and the connectives of the stamens opposite the anteriorsepals greatly enlarged
and overtoppingthe locules by up to 1.0 mm. This variety is distinguishedby its
ferrugineouspubescence on the inflorescence. The collections from the Guayana
Highlandare somewhat unusual, and have shorter petioles, smaller leaves, and
the inflorescence branches sometimes glabrate or with a sparse outer layer of
largerlooser hairs above the ferrugineouspubescence. There seems to be a continuous intergradationbetween the typical lowland form and the highlandform
so I have preferrednot to segregate them.
The Martincollection on which Jussieu based his descriptionis in flower and
young fruit, and bears the number92; Bentham based his descriptionof Banisteria leptocarpa in part on a Martincollection from Cayenne. However, the sheet
in the British Museumon which his descriptionis based is in fruit and is without
numberand so I consider the name legitimateratherthan a nomen superfluum.
44b. Banisteriopsis martiniana var. subenervia Cuatrecasas, Webbia 13: 501. 1958.
Banisteriopsis martiniana var. laevis Cuatrecasas, Webbia 13: 502. 1958. Type. Schultes &
Cabrera14569, Colombia,Amazonas-Vaup6s,rio Apaporis, Raudalde Jirijirimo,fl (holotype, US; isotype, A, F).
Branchesglabrous. Leaves falcate, coriaceous to fleshy, with the petiole 7-20

mm long, the lamina7.0-17.0 cm long and 4.1-8.5 cm wide, obtuse to cordate at
the base, short- to long-acuminateat the apex, bearing minute glands on the
margin,glabrouson both sides. Inflorescencebranchesminutelypuberulent,the
hairs white or grey, the reducedleaves of the inflorescenceglabrous,the branches
subtendingthe umbels 1-8 mm long.
Type. R. E. Schultes & I. Cabrera 16387, Colombia, Amazonas, Rio Miriti-
parana, canloGuacaya, fl (holotype, US; isotype, F, GH, U).

Distribution.(Fig. 29). In tropical rain forest of Amazonian Brazil and Peru,
the upper Vaupes in Colombia,and the upper Orinoco in Venezuela.
Collected in flower in January,Marchto June, and August to November, and
in fruit in June, August, and September.
Branchesglabrous. Leaves falcate, coriaceous to fleshy, with the petiole 7-20
mm long, the lamina7.0-17.0 cm long and 4.1-8.5 cm wide, obtuse to cordate at
the base, short- to long-acuminateat the apex, bearing minute glands on the
margin,glabrouson both sides. Inflorescencebranchesminutelypuberulent,the
hairs white or grey, the reducedleaves of the inflorescenceglabrous,the branches
subtendingthe umbels 1-8 mm long.
Type. R. E. Schultes & I. Cabrera 16387, Colombia, Amazonas, Rio Miriti-
parana, canioGuacaya, fl (holotype, US; isotype, F, GH, U).

Distribution.(Fig. 29). In tropical rain forest of Amazonian Brazil and Peru,
the upper Vaupes in Colombia,and the upper Orinoco in Venezuela.
133
. velutinissima; triangle, B. maguirei open

'
circle, . pucherrima. .
9+
+COLOMBIA
1-?
t+
++
.:
chus
&
674 ft (ECON,
(GH,NY, U, US); Soejarto & Cardozo
,
GH).
inUS).
~ Jue August,.. a:i Maguire
fri VENEZUELA.A
: 41638
. .....be
f (MIC,
17
Garcia-Barriga
N.
PER.
Lore:
n'
Cr
infruit in June,.A1ugust,and September.

FIG. 29.
. .Distribution
species4,44, 45, 53 and 54. Closed
Distribution of part of Banisteriopsis martiniana group, species
B. velutinissima; triangle, B. maguirei; open circle, B. pulcherrima.
Collected in flower in January, March to June, and August to November, and

in fruit in June, August, and September.
4184
fsquare (G,
martin.
NY);B.
Williams 971 f
19 closed
open (F,
square
US). San Martinan var. subenervia
13706 ft yng ft (NY, US); Schultes & Cabrera 13595 fl f (GH,,
,US)
114569t(;
circle,
ft (A,
f (M, R); Mo eSchultes 3836fl(MICH); RodrigSchultes

COLOAmazas:
45763fl
&
Luetzelburg&22128
67 Chagas
ffl (F, NY, US);
fr (GH, MICH);
Killip Smith 29922 fl (F, NY, US); Klug 136 fl (F, NY, US); 167
1175
fl
Revilla
364
fl
712
frt
63
Schunke
fl
(F, NY,
US); Soejarto& Cardozo
(MICH);674 fr (MICH);
(F, GH, NY, Peru:
US);
Tessman
(MICH);
Common(GH,
NY,
U, US);
Amazona
cia-Barriga
frpeyoo-wee(YukuGH). name).
fl yngfr (NY, US); Schultes& Cab 13595flfrgrey inflorescence pubescence.
Ths13706
The);
14569fl (A,
(SP,
(SP, US).
US).
leaveUS).
VENre probably almost fleshy when alive. The type of var. laevis has leaves
yng
Common names. Colombia. "'e-pe'-pe-yoo-wee" (Yukuna name). Peru: "nucha-wasca."
na-wasca.
are smooth above, but971
4184which
yng fr (F, US). SanMartinent
belytkowski
19 fl (F). BRAZ.lections
which
have attributed
to var. by
subenervia
fl (MICH);
va9198
4576
Rodrigues
Chagas
This Ivariety
is distinguished
its whiteexhibit
or
grey
inflorescence
pubescence.
The
flprominence
type of this variety has leaves which are quite smooth on both sides, and the
leaves were probably almost fleshy when alive. The type of var. laevis has leaves
which are smooth above, but with the veins prominent below. Other collections
which I have attributed to var. subenervia exhibit various degrees of prominence
of the leaf veins on both sides, but in all other respects are very similar. The
prominence of the leaf veins appears to be a very variable character; Maguire
41638 has leaves which are smooth above, or with the veins prominent, in the
same collection.
45. Banisteriopsis velutinissima B. Gates, sp. nov.
Liana, rami dense velutini, demum glabrati. Stipulae minutae. Folia petiolo 1015 mm longo, dense velutino, eglanduloso, lamina 5.5-10.1 cm longa, 3.6-8.1 cm
Flora Neotropica
134
lata, in inflorescentiaminori et saepe ad bracteas redacta, elliptica vel late elliptica, basi cuneata vel truncata, apice obtusa, marginerevoluta et glandibuscupulatis instructapraesertimversus apicem, abaxialiterbasi prope costam 2 glandibus sessilibus gerenti, adaxialitertomento-velutinademumglabrataintervenas,
abaxialitervelutina. Inflorescentiaaxillaris, rami usque 25 cm longi, dense velutina, folia redactaciliata gerens, composita umbellis4-floriferisin dichasiis dispositis, pedunculo communi 2-4 mm longo; bracteis bracteolisque 0.5-0.8 mm
longis apice rotundatis, glabratis abaxialiter, involucratis, persistentibus;pedicellis sessilibus, 4-9 mm longis, 0.3-0.4 mm diametro, sparsim appresso-pubescentibus. Sepala 1.4-1.6 mm longa latave, orbicularia,apice rotundata,sparsim
appresso-pubescentiavel glabrata abaxialiter, adaxialiter glabra, eglandulosa.
Petala lutea, longe fimbriata,4 lateraliainter sepala reflexa, ungue 1.5 mm longo,
limbo 3.0-5.0 mm longo latove, orbiculari,2 antero-lateralibusquam 2 posterolateralibusmajoribuset concavioribus, petalum posticum ungue erecto 2.5 mm
longo, 0.4 mm lato, ad apicem constricto, limbo usque 5.5 mm longo et 3.5 mm
lato, reflexo, obovato, parte angusta basali 2-3 paria laciniarumglandulosarum
gerenti. Staminafilamentis 1.2-2.8 mm longis, illis 3 stylis oppositis (i.e. sepalo
antico et petalis postero-lateralibusoppositis) longissimis, 3 posticis inter stylos
posticos inflexis;antheraeloculis glabris0.6-0.9 mm longis, connectivis petalis et
sepalis postero-lateralibusoppositis 0.3-0.6 mm longis, 0.3-0.4 mm latis, 0.1-0.2
mm profundis, oblongis, illis 3 sepalis anticis oppositis 1.1-1.4 mm longis, 0.91.0 mm latis, 0.7-1.0 mm profundis, obovatis, tumidis et loculos superantibus,
illis sepalis antero-lateralibusoppositis 0.6-0.8 mm loculos superantibus.Ovarium0.8 mm altum,pubescens, stylus anticus quamstylis posticis crassior,rectus
erectusque, 2.2 mm longus, styli postici graciles, divergentes basi, 2.0-2.2 mm
longi, stigmatatruncata.Fructus non suppetunt.
Type. J. Revilla 650, Peru, Loreto, Prov. Maynas, Distr. Iquitos, carretera
Nanta, caserio Pefia Negra, fl (holotype, MICH).
Distribution.(Fig. 29). Known only from two collections near Iquitos in lowland Peru.
Collected in flower in May and August, fruit unknown.
PERU. Loreto: Ducke 7573 fl (MG); Revilla 650 fl (MICH).
The flowers of this species are almost identical to those of Banisteriopsismartiniana, but the species is readily distinguishedby its velutinous pubescence
throughout.This feature is unique in this group of species.
46. Banisteriopsiselegans (Trianaand Planchon)Sandwith,J. Arnold Arbor. 24:
222. 1943.
Banisteria elegans Triana and Planchon, Prodromus Florae Novo-Granatensis: 320. 1862.
Banisteriopsis speciosa Small, North American Flora 25: 133. 1910. Type. H. von Tiirckheim II.
1785, Guatemala, Alta Verapaz, Coban, fl (holotype, NY; isotypes, BR, C, F, G, GH, M,
MO, S, W).
Banisteria elegans subsp. ovata Niedenzu, Verz. Vorles. Konig. Ak. Braunsb.: 16. 1912. Nomen
superfluum, = subsp. elegans.
Banisteria elegans subsp. ovata var. guatemalensis Niedenzu, Verz. Vorles. Konig. Ak.
Braunsb.: 16. 1912. Type. H. von Tiirckheim 11. 1785, Guatemala, Alta Verapaz, Coban, fl
(holotype B destroyed, NY, fragment; isotypes, BR, C, F, G, GH, M, MO, NY, S, W).
Banisteria elegans subsp. cordata Niedenzu, Verz. Vorles. Konig. Ak. Braunsb.: 16. 1912. Type
H. H. Smith 1509, Colombia, Magdalena, Santa Marta, fl (lectotype, MICH; isolectotypes
F, G, GH, MO, NY, P, US).
Banisteria elegans subsp. cordata var. ciliata Niedenzu, Verz. Vorles. K6nig. Ak. Braunsb.:
16. 1912. Type. H. H. Smith 1509, Colombia, Magdalena, Santa Marta, fl (holotype, B?
destroyed, fragment, NY; isotypes, F, G, GH, MICH, MO, NY, P, US).
135
Banisteriopsiselegans subsp. cordata (Niedenzu)Sandwith,J. ArnoldArbor. 24: 222. 1943.

Banisteriopsis elegans subsp. cordata var. ciliata (Niedenzu) Sandwith, J. Arnold Arbor. 24:
223. 1943.
Banisteriopsis scalariformis Schery, Ann. Missouri Bot. Gard. 7: 92. 1943. Type. H. von Wedel
2776, Panama,Bocas del Toro, WesternRiver, fl (holotype, MO;isotype, GH).

Banisteriopsismartiniana(Adr.Jussieu)Cuatrecasasvar. elegans (Trianaand Planchon)Cuatrecasas, Webbia13: 501. 1958.
Liana, branchesappressed-sericeousat first, soon glabrate,old branchessometimes with peeling papery bark. Stipules minute, triangular,up to 1 mm long.
Leaves with the petiole 4-10(-14) mm long, glabrous,eglandular,the lamina6.914.0(-20.7) cm long, 2.4-8.2(-9.1) cm wide, those associated with the inflorescence smallerand often reduced to bracts, lanceolate to ovate to broadlyelliptic,
obtuse to cordate at the base, acute to acuminateat the apex, with the margin
more or less revolute and glandular,the glands more numeroustowardsthe apex
and sometimes developed into short ciliate projections, bearingabaxially at the
base of the lamina beside the midriba pair of sessile glands, glabrous on both
sides or abaxiallysparsely appressed-sericeous,the trabeculaup to 0.5 mm long,
with the reticulationprominulousto prominentadaxiallyandprominentabaxially.
Inflorescenceaxillary, cymose, of 4 (rarely 6)-floweredumbels, the branchesup
to 35 cm long, glabrous to sparsely appressed-sericeous,the trabecula 0.2-0.5
mm, the side branchesbearing 1-5 umbels, the commonpeduncle 4-20 mm long;
bracts and bracteoles 0.8-1.4 mm long, triangular,acute at the apex, glabrous
abaxially or with a fringe of hairs apically, involucrate, persistent; pedicels sessile, 7-12(-14) mm long, 0.3-0.6 mm in diameter, glabrous. Sepals 2.0-3.0 mm
long, 1.5-2.8 mm wide, broadly elliptic, rounded at the apex, glabrous on both
sides, eglandularor the 4 lateral sepals biglandular,the glands 1.0-2.0 mm long,
0.6-1.0 mm wide. Petals yellow, lacerate to fimbriate, glabrous, the 4 lateral
petals reflexed between the sepals, the claw 1.0-2.5 mm long, 0.4-0.6 mm wide,
the limb 5.0-10.5 mm long, 4.5-9.0 mm wide, broadly elliptic to orbicular,the
antero-lateralpetals largerand more concave than the postero-lateralpetals, the
posteriorpetal with the claw erect, 2.0-3.4 mm long, up to 1.2 mm wide distally,
tapering at the base, not constricted, channelled distally, the limb 3.0-5.5 mm
long, 3.0-4.5 mm wide, broadly elliptic to broadly obovate, the basal fimbriae
sometimes gland-tipped.Stamens with the filaments 1.0-3.4 mm long, those opposite the 3 anteriorsepals longest, those opposite the postero-lateralpetals ca.
one-halfas long as the posteriorstyles; antherswith the locules glabrous,0.6-1.0
mm long, the connectives of the stamens opposite the sepals 0.6-1.0 mm long,
0.4-1.2 mm wide, 0.3-1.0 mm deep, obovate, those opposite the 3 anteriorsepals
usually largerthan those opposite the postero-lateralsepals and overtoppingthe
locules, those opposite the petals 0.5-1.0 mm long, 0.3-0.6 mm wide, 0.1-0.7
mm deep, oblong, not overtoppingthe locules. Ovary 1.0-1.8 mm tall, densely
sericeous, styles subequalor the posteriorstyles slightlylonger, 2.4-3.6 mm long,
straightand parallel, or the posterior styles divergingapically, stigmas truncate.
Fruit with filiformcarpophore3.5-4.0 mm long, the nut 4.5-5.0 mm tall, 2.5-3.5
mm long, sparsely sericeous to glabrate,the hairs stiff, sub-irritating,the trabecula straight,up to 2 mm long, the wing 23-33 mm long, 8-13 mm wide, sparsely
sericeous to glabratelike the nut, the hairs up to 3 mm long.
Type. Trianaet Planchon s.n., Colombia,El Valle, Buenaventura,yng fr (holotype, P).
Distribution.(Fig. 30). In coastal forests from sea level up to 1000m in Colombia and in CentralAmerica at altitudesup to 1500m.
Collected in flower in Februaryto June, and September,and in fruit in March,
May, June, and November.
136
Flora Neotropica
Representativecollections: GUATEMALA.A. Molina R. et al. 12233 fl (F, G, NY, W); J. D.

Smith 1698 fl (GH); Standley 91573 fl (F); 91589 fl (F, US); Tiirckheim 11. 1785 (BR, C, F, G, GH,
M, MO, NY, S, W). HONDURAS. A. Molina R. et al. 7205 fl (F). COSTA RICA. Brenes 3899 fl
(F); 4242 fr (F); Davidse & Pohl 1484 fl (MICH); Primack & Utley 716 fl (DUKE); Austin Smith
1829 fl fr (NY). PANAMA. Croat 33246 bud (MICH); Hunter & Allen 539 fl (F, G, GH, S, U); Mori
et al. 4599 yng fr (MICH); Porter et al. 4318 yng fr (F); von Wedel 2776 fl (GH); 2791 fl (GH).
COLOMBIA.Magdalena:H. H. Smith 1509fl (F, G, GH, MICH,MO, NY, P, US). Choc6: Forero
et al. 1189 fl (F, NY); Gentry & Fallen 17242 fl (MICH). El Valle: Alston 8625 fl (S, US); Cuatrecasas
17359 fl (F); 19990 fl (F, US); Killip & Cuatrecasas 28987 fr (F, GH, MO, NY, S, US); Triana s.n.
fl (G).
This species is characterisedby its glabrous or rarely sparsely appressed-sericeous inflorescence, flowers in four-flowered umbels, and usually glabrous
leaves. Smith 1509 is unusualin that the anthersof the five posteriorstamensare
smaller than the others, and the filamentsof the stamens opposite the posterolateralpetals are as long as those opposite the antero-lateralsepals, but in other
respects falls within the variationrepresentedby this species. Forero et al. 1189
and Alston 8625 have large fleshy leaves; both are apparentlylarge lianas and the
unusualsize and thickness of the leaves may reflect the part of the plant collected
or a response to some environmentalvariablesuch as light.
47. Banisteriopsispolygama(Niedenzu) B. Gates, comb. nov.
Banisteriapolygama Niedenzu, Fedde Rep. Spec. Nov. 26: 345. 1929.
Banisteriopsis martiniana var. popayanensis Cuatrecasas, Webbia 13: 500. 1958. Type. E. Perez
Arbaldez& J. Cuatrecasas6057, Colombia,Cauca, Popayan,La Capilla,fl (holotype, US;

isotype, F).
Liana, young branches terete, appressed-sericeous, old branches glabrate,

grey-brownwith numerousprominentdarklenticels. Stipules minute, triangular.
Leaves coriaceous with the petiole 3-8 mm long, appressed-sericeousto glabrate,
eglandular,the lamina3.8-9.4 cm long, 2.1-6.1 cm wide, broadlyellipticto ovate,
cuneate to obtuse at the base, acuminate to apiculate at the apex, the margin
plane, eglandular,or bearingminute glands on or just within the margin, sometimes bearinga pair of sessile glands on the laminabeside the midribat the base,
glabrouson both sides, smooth adaxiallyor with the veins prominulous,with the
lateral veins and the scalariformsecondary veins prominentabaxially. Inflorescence axillary, paniculate,the branchesup to 30 cm long, the side branches0.51.5 cm long and bearing 3 pairs of flowers, silvery-appressed-sericeous;bracts
and bracteoles 1.0-1.2 mm long, triangular,appressed-sericeousabaxially, involucrate, persistent; pedicels sessile, 7-11 mm long, 0.4-0.6 mm in diameter,
appressed-sericeous.Sepals 2.0-3.0 mm long, 1.5-2.2 mm wide, broadlyelliptic,
roundedat the apex, sparsely sericeous or glabrousabaxially,glabrousadaxially,
the 4 lateral sepals biglandularor more rarely eglandular,projecting 1.4-1.8 mm
beyond the glands, the glands 1.4-1.8 mm long, 0.8-1.0 mm wide. Petals lemonyellow, glabrous, fimbriate,the 4 lateral petals reflexed between the sepals, the
claw 1.5-3.0 mm long, the limb 4.0-6.5 mm long, 3.0-5.5 mm wide, broadly
elliptic to orbicular, the antero-lateralpetals larger and cochleate, the posterolateral petals plane, the posterior petal with the claw erect, 1.8-3.0 mm long,
wider, up to 0.8 mm, at the apex, the limb 3.5-5.0 mm long, 2.5-4.5 mm wide,
broadly obovate, with the basal fimbriaesometimes enlarged and gland-tipped.
Stamens with the filaments 1.8-4.0 mm long, those opposite the sepals longer
than those opposite the petals; anthers with the locules 0.6-1.0 mm long, glabrous, the connectives of the stamens opposite the sepals 0.8-1.2 mm long, 0.61.0 mm wide, 0.4-0.9 mm deep, projecting0.2-0.5 mm beyond the locules, those
opposite the petals 0.6-0.8 mm long, 0.3-0.6 mm wide, 0.2-0.6 mm deep, not
137
projectingbeyond the locules. Ovary 1.6-2.0 mm tall, densely sericeous; styles

2.6-3.5 mm long, subequal, straight and parallel, stigmas capitate. Fruit with
filiform carpophore up to 3 mm long, the nut 3-4 mm tall and 2-3 mm long,
sparsely sericeous to glabrate,the wing (16-)22-27 mm long, (6-)9-11 mm wide,
sparsely appressed-sericeous,the hairs irritatingwith stiff straighttrabecula1.01.5 mm long.
Type. Lehmann 8491, Colombia, Cauca, Popayan near Palace and Cajibio, fl
(holotype, B destroyed; isotypes, F, GH, K, S, US).
Distribution.(Fig. 30). In premontaneforests of the Andes in southernColombia, Ecuador and northernPeru.
Collected in flower from Januaryto Marchand June and July, and in fruit in
April and June.
COLOMBIA. Cauca: B. T. 403 (Herb. Lehmann) fl (GH, NY); 445 fl (GH, NY); 557 fl (NY);
Lehmann 8491 fl (F, GH, K, S, US); Pennell & Killip 6344 fl (GH, NY); Perez Arbeldez & Cuatrecasas 6057 fl (F, US); von Sneidern 5563 fl fr (US). ECUADOR. Lugo 179 fr (C, S). PERU. Amazonas: Woytkowski 8135 fl (MICH, Wurdack 1051 fl fr (F, US). San Martin: Klug 3329 fl (F, G, GH,
MO, NY, S, US).
This species is characterisedby its dense, appressed-sericeousinflorescence

pubescence, long narrowpanicles of flowers, its small, ellipticto ovate coriaceous
leaves with no or inconspicuous marginalglands, and the branches with very
numerousprominentblack or dark-brownlenticels. I am reluctantlyincludingin
this taxon Klug 3329, althoughI have not includedmeasurementsfrom this plant
in the description. The flowers of this plant are like B. polygama, and the inflorescence pubescence is appressed-sericeous.However, in many other characters
it is distinct, it has larger leaves (up to 14.8 cm long and 7.9 cm wide), longer
petioles (up to 14 mm long), and bears prominentglands or glandularteeth on
the acumen; the flowers are borne in loose panicles, the side branches of which
bear 10-16 flowers, and the pedicels are glabrous. I hope that future collections
will resolve the problem of this plant's affinities.
48. Banisteriopsispadifolia(Niedenzu) B. Gates, comb. nov.
Banisteriapadifolia Niedenzu, Ind. Lect. Lye. Brunsb.:7. 1900.
Banisteria parviflora var. boliviensis Niedenzu, Fedde Rep. Spec. Nov. 26: 345. 1929. Type.
Guiinther
(leg Buchtien) 1809, Bolivia, Mapiriregion, Sarampiuninear San Carlos, fl (holotype, B destroyed;isotype, US).
Liana, young branchesterete, appressed-sericeous,the hairs with the trabecula

straightor flexuous, older branchesglabrate,darkbrown, with short longitudinal
fissures and prominentlenticels. Stipules minute, triangular.Leaves, only young
newly expanded ones available, with the petiole 5-7 mm long, appressed-sericeous to glabrate, bearing 1, rarely 2, pairs of sessile glands at the apex, the
lamina4.6-10.6 cm long, 1.8-4.6 cm wide, narrowlyovate to lanceolate, obtuse
to truncate at the base, acute to acuminateat the apex, with the marginplane
and bearing 3-5 pairs of cupulate glands laterally, glabrous adaxially, at first
densely tomento-sericeousabaxially, soon glabrous or the midribsparsely sericeous, smooth or with the lateralveins prominulouson both sides. Inflorescence
axillary, paniculate, the branches up to 15 cm long, tomento-sericeous,the side
branches up to 3.0 cm long and bearing 6-8 flowers, the inflorescence leaves
reduced, with prominentcupulateglands along the margin;bracts and bracteoles
1.0-1.4 mm long, triangular,sparsely tomento-sericeous abaxially, involucrate,
persistent; pedicels sessile, 6-10 mm long, up to 0.6 mm in diameter, tomentosericeous. Sepals 2.0-2.5 mm long, 1.8-2.0 mm wide, elliptic, rounded at the
138
Flora Neotropica
apex, sparsely sericeous abaxially,glabrousadaxially, the 4 lateralsepals biglandular or more rarely eglandular,projecting 1.0-1.2 mm beyond the glands, the
glands 1.6-1.8 mm long, 0.8-1.0 mm wide. Petals yellow, glabrous,fimbriate,the
4 lateralpetals reflexed between the sepals, broadlyelliptic to orbicular,the claw
1.6-2.0 mm long, the limb of the antero-lateralpetals 6.0-7.5 mm long, 5.5-6.0
mm wide, cochleate, that of the postero-lateralpetals 4.0-4.5 mm long and wide,
plane to slightly convex, the posterior petal with the claw erect, up to 2.4 mm
long, flaringand channelleddistally, the limb 4.0-4.5 mm long and wide, broadly
obovate, with the basal fimbriaeenlarged and gland-tipped. Stamens with the
filaments 1.6-2.6 mm long, subequal, that opposite the anterior sepal slightly
longer, and those of the 3 posterior stamens slightly shorter, than the others;
anthers reflexed, the locules 0.5-0.8 mm long, glabrous, the connectives of the
stamens opposite the sepals 0.8-1.0 mm long, 0.5-0.8 mm wide, 0.4-0.6 mm
deep, obovate, projectinga little beyond the locules, those opposite the petals
0.5-0.7 mm long, 0.4-0.5 mm wide, 0.2-0.4 mm deep, obovate, not projecting
beyond the locules. Ovary 1.4-1.6 mm tall, densely sericeous apically;styles 2.02.6 mm long, subequalor the posterior styles somewhat longer than the anterior
style, straightand parallelor divergingslightlyat the apex, stigmascapitate. Fruit
unknown.
Type. Poeppig 1288, Peru, Huanuco, near Cuchero(holotype, W; isotypes, F,
NY, W).
Distribution.(Fig. 30). In the Andes of Peru and Bolivia.
Collected in flower in July and August in Peru and in November and December
in Bolivia.
PERU. Huanuco: Poeppig 1288 fl (F, NY, W). Cuzco: C. Vargas C. 4571 fl (F); 12970 fl (US).
BOLIVIA. Buchtien 1809 fl (US); Weddell 4207 fl (P).
This species is most closely related to Banisteriopsispolygama, from which

it can be distinguishedby its larger, narrowly ovate to lanceolate leaves with
prominentmarginalglands, tomento-sericeousinflorescencepubescence, and by
bearingits flowers when the plant is leafless or the new leaves are just emerging.
49. BanisteriopsiswilburiiB. Gates, sp. nov.
Liana, rami brunneo-tomentosi,demum glabrati, pilis trabecula 1.0-1.8 mm
longa, interdumpilis basifixis usque 1.5 mm longis interspersis. Stipulae usque
2 mmlongae, triangulares,sparsimappresso-sericeaevel glabratae.Folia coriacea
petiolo 8-11 mm longo, sparsim tomentoso, eglanduloso, lamina 4.7-15.1 cm
longa, 2.4-8.4 cm lata, in inflorescentiaminori,et ad bracteasredacta, lanceolata
vel ovata vel late elliptica, basi cordata,apice acuta vel breve acuminata,margine
parum revoluta et glandis instructa praesertim versus apicem, abaxialiterbasi
prope costam 2 glandes sessiles gerenti, adaxialiter tomentosa mox glabrata,
abaxialitersparsimtomentosa, pilis flexuosis, trabeculasessili 0.6-1.8 mm longa,
venis venulisqueprominulisadaxialiteret prominentibusabaxialiter.Inflorescentia axillaris, umbellis 4-floriferiscomposita, in cymis dispositis, ramis usque 20
cm longis, sparsim tomento-hispidis,pilis brunneis, pede usque 0.8 mm longo,
trabecula 1.5-2.5 mm longa, pedunculo communi usque 25 mm longo; bracteis
bracteolisque usque 1.0 mm longis, triangularibus,apice rotundatis,glabris vel
marginepubescenti, involucratis,persistentibus;pedicelli sessiles, 12-18 mm longi, 0.6-0.7 mm diametro, glabri. Sepala 2.8-3.2 mm longa, 2.0-2.5 mm lata,
elliptica, apice rotundata,utrinqueglabra,4 lateraliabiglandulosa,glandibus1.52.6 mm longis et 0.8-1.4 mm latis. Petala citrina,laciniata,4 lateraliainter sepala
139
+.
id,9"?
fCi?
*' ---- . -?:
*'.
' '....
0
?f
?.
t;
:.
.....:..
:7
'
': !*
.._.;......
FIG. 30. Distribution of part of Banisteriopsis martiniana group, species 46-52. Open circle, B.
elegans; open square, B. polygama; closed square, B. padifolia; closed circle, B. wilburii; triangle,
B. pubescens; open diamond, B. grandifolia; closed diamond, B. alternifolia.
patentia, ungue 1.0-3.0 mm longo, 0.4-0.6 mm diametro, limbo 6.0-8.5 mm longo,

5.0-8.0 mm lato, orbiculari vel rotundato, antero-lateralibus quam postero-lateralibus majoribus et concavioribus, petalum posticum ungue erecto, 3.0 mm longo,
dilatato et usque 1.0 mm lato ad apicem, saepe canaliculato distaliter, limbo 5.57.0 mm longo, 4.0-5.0 mm lato, late elliptico vel late obovato, laciniis basalibus
glanduliferis. Stamina filamentis 2.2-3.6 mm longis, illis sepalis oppositis quam
illis petalis oppositis longioribus, praesertim 3 anticis; antherae loculis 0.8-1.0
mm longis, glabris, connectivis sepalis oppositis 1.0-1.2 mm longis, 0.7-1.0 mm
latis, 0.6-0.9 mm profundis, late obovatis, apice glandulosis, loculos 0.2-0.6 mm
superantibus, illis petalis oppositis 0.8-1.0 mm longis, 0.4-0.7 mm latis, 0.2-0.6
mm profundis, oblongis, illis 4 petalis lateralibus oppositis apice parum glandulosis et loculos usque 0.2 mm superantibus. Ovarium 1.6-2.0 mm altum, dense
sericeum; styli recti, aequales, apice parum divergentes, 2.4-3.0 mm longi, crassi,
usque 0.4 mm diametro, stigmata capitata. Fructus ex 3 samaris (vel abortu 1-2)
constans, carpophoro filiformi usque 3 mm longo, nuce usque 6 mm alta, 3-5 mm
longa, appresso-sericea vel glabrata, pilis rigidis, trabecula 1.5-2.0 mm longa, ala
21-26 mm longa, 8-13 mm lata, sparsim appresso-sericea nucis similis.
Type. R. L. Wilbur & J. A. Teeri 13615, Panama, slopes of Cerro Jef beyond
Cerro Azul, ca. 5.5 miles on road to La Eneida, fl fr (holotype, DUKE; isotype,
F).
Distribution. (Fig. 30).. In high forest in Boyac and Antioquia in Colombia and
on mountain ridges in Panama and Costa Rica.
Collected in flower in December to May, July, and September and in fruit in
January to May and July.
Flora Neotropica
140
COSTA RICA. Almeda et al. 2943 fl (MICH); 2978 (MICH). PANAMA. Almeda 3453 fr (MICH);
Croat 33790 fl (MICH); Dwyer et al. 5040 fl (NY); Mori 3604 fl (MICH); 6084 fl fr (MICH); 7111 fl
(F, MICH); Nee 10517 fl (MICH); Wilbur et al. 11371 fl (DUKE); 13615 fl fr (DUKE, F); 19468 fl
(DUKE). COLOMBIA. Antioquia: Fosberg 21625 fr (NY, US); Toro 585 fl (NY); 606 fl (NY).
Boyaca: A. E. Lawrance 639 fl (G, GH, MO, S, US, W).
This species is distinguishedfrom Banisteriopsis elegans by its largerflowers

with large calyx glands, its longerpedicels, tomentose pubescence on the branches and inflorescence, and sparse pubescence on the leaves abaxially;in addition,
B. elegans is a lowland plant whereas B. wilburiioccurs at higherelevations (up
to 2200 m) in pre-montanewet forest.
This species is namedfor R. L. Wilbur,a student of the flora of CentralAmerica, where he has done much collectingincludingsome very complete collections
of this species.
50. Banisteriopsispubescens(Niedenzu) Cuatrecasas,Webbia 13: 503. 1958.
Banisteria elegans subsp. cordata var. pubescens Niedenzu, Verz. Vorles. K6nig. Ak. Braunsb.:
17. 1912.
Liana, branches glabrous. Stipules minute. Leaves with the petiole 2-7 mm
long, glabrous, eglandularor apically biglandular,the lamina 7.0-15.0 cm long,
6.0-10.0 cm wide, smallerand often reducedto bractsin the inflorescence,broadly elliptic to rotund, strongly cordate at the base, apiculateat the apex, with the
marginflat and bearing small glands, bearingabaxially at the base beside or on
the midrib a pair of sessile glands, glabrous adaxially, sparsely appressed-sericeous, abaxially, the hairs with the trabecula0.5-2.0 mm long, the reticulation
prominulous adaxially and prominent abaxially. Inflorescence axillary, the
branches up to 20 cm long, loosely brown-tomentose;the hairs with the trabecula 1.2-1.6 mm long, cymose, of 4-floweredumbels, the common peduncle
up to 24 mm long, bracts and bracteoles up to 1.2 mm long, acute at the
apex, abaxially sparsely tomentose, involucrate, persistent; pedicels sessile,
10-14 mm long, 0.6 mm in diameter, loosely tomentose. Sepals 2.0-3.0 mm
long, 2.0-2.2 mm wide, elliptic, rounded at the apex, abaxially densely sericeous, adaxially glabrous, the 4 lateral sepals eglandular or biglandular,the
glands up to 1.2 mm long and 0.8 mm wide. Petals yellow, glabrous, lacerate, the 4 lateral petals reflexed between the sepals, the claw 1.0-2.5 mm
long, 0.2-0.4 mm wide, the limb 5.0-9.0 mm long, 5.0-8.5 mm wide, rotund
to orbicular,the antero-lateralpetals larger and more concave than the posterolateral petals, the posterior petal with the claw erect 3.0-3.5 mm long, 0.5-1.0
mm wide distally, not constricted, the limb 4.0-5.5 mm long, 4.0-4.5 mm
wide, obovate to orbicular,eglandularor with 1-2 glands at the base. Stamens
with the filaments 1.8-3.0 mm long, those opposite the sepals longer than those
opposite the petals; antherswith the locules 0.6-1.2 mm long, glabrous,the connectives of the stamens opposite the 3 anteriorsepals 1.0-2.0 mm long, 0.8-1.2
mm wide, 0.8-1.2 mm deep, obovate, glandularapically and projectingup to 0.8
mm beyond the locules, those of the stamens opposite the petals and the posterolateral sepals 0.4-1.2 mm long, 0.4-0.6 mm wide, 0.2-0.6 mm deep, oblong,
eglandular,not overtoppingthe locules. Ovary 1.0-1.4 mm tall, densely sericeous
apically; styles equal, 3.0-3.4 mm long, straight and parallel throughouttheir
length or hooked at the apex, stigmastruncate. Fruitunknown. Sepals enlarging,
up to 5 mm long, after flowering.
Type. Lehmann5159, Colombia,Narifio,Tuquerres,fl (holotype, B destroyed,
NY fragment;isotype, F).
141
Distribution.(Fig. 30). Known only from the type locality Narifio, Colombia,
and two collections from adjacentEcuador.
COLOMBIA. Narino: Lehmann 5159 fl (F, NY fragment ex B). ECUADOR. Madison et al. 5138
fl (MICH); Gentry & Shupp 26586 (MICH).
In its flowers and inflorescencethis species is similarto Banisteriopsiselegans,

but can be distinguishedby its short petiolate leaves which are strongly cordate
at the base, its tomentose pedicels and sericeous sepals. Additionalcollections
would no doubtfurtherincrease the size rangesfor leaf and flowermeasurements.
51. Banisteriopsisgrandifolia(Niedenzu) B. Gates, comb. et stat. nov.
Banisteria elegans var. grandifolia Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 412. 1928.
Banisteriopsis martiniana var. grandifolia (Niedenzu) Cuatrecasas, Webbia 13: 499. 1958.
Liana, young branchesbearingbasifixedhairs up to 5 mm long, older branches

glabratewith pale flakybark. Stipules minuteor linearup to 1.5 mm long. Leaves
very coriaceous, with the petiole 5-13 mm long, glabrous or bearing sparse basifixed hairs, eglandular,with the lamina 5.9-17.7 cm long, 4.2-9.6 cm wide,
muchreducedor bract-likein the inflorescence,ovate, cordateat the base, obtuse
to acute at the apex, the marginslightly revolute, with minute glands along the
leaf edge abaxially 1-2 mm from the margin,bearingabaxially on the midribat
the base a pair of large stalked glands, adaxially sparsely appressed-sericeous,
soon glabrate, abaxially sparsely tomento-sericeous,the hairs flexuous, more or
less sessile, soon glabrate except near the base, with 3-5 pairs of lateral veins
arising in the proximal half of the leaf, prominenton both sides. Inflorescence
axillary, the branches up to 20 cm long, minutely velutinous, of racemes of up
to 12 pairs of flowers, congested distally, suboppositeto alternateproximally,the
inflorescenceleaves up to 12 mm long and wide, orbicular,apiculateat the apex,
with the marginlong-ciliateor eglandular;bracts and bracteoles0.6-0.8 mm long,
acute at the apex, glabrousabaxially, involucrate,persistent;pedicels sessile, 610mmlong, up to 0.6 mmin diameter,loosely sericeous,the hairswiththe trabecula
1.0-1.5 mmlong. Sepals 2.0-2.2 mmlong, 1.8-2.0 mmwide, rotund,roundedat the
apex, glabrouson both sides, or sparselysericeous abaxially,the 4 lateralsepals biglandular,the glandsup to 1.2 mm long and 0.7 mm wide. Petals yellow, glabrous,
dentate, the 4 lateral petals reflexed between the sepals, the claw 1.2-1.5 mm
long, the limb 6.0-6.5 mm long and 5.0 mm wide, rotund,the antero-lateralpetals
cochleate, the postero-lateralpetals concave to plane, the posterior petal with
the claw erect, up to 3.0 mm long, flaringand canaliculatedistally, up to 1.0 mm
wide, the limb 5.0 mm long and wide, orbicular,plane. Stamenswith the filaments
2.5-3.4 mm long, those opposite the styles (i.e. opposite the anteriorsepal and
the postero-lateralpetals) longest, longer than those opposite the antero-lateral
sepals; anthers with the locules 0.6-1.0 mm long, glabrous, the connectives opposite the 3 anterior sepals 0.8-1.0 mm long, 0.8 mm wide, 0.6-0.7 mm deep,
obovate and glandularapically, projectingup to 0.3 mm beyond the locules, those
opposite the petals and postero-lateralsepals 0.5-0.8 mm long, 0.4-0.5 mm wide,
0.2-0.4 mm deep, oblong. Ovary 1.2-1.4 mm tall, densely sericeous apically;
anteriorstyle straight,up to 3.0 mm long and 0.2 mm in diameter,posteriorstyles
up to 3.5 mm long and 0.15 mm in diameter,divergingbasally. Fruitof 3 samaras
with filiformcarpophoreup to 3 mm long, the nut 5.0-5.5 mm tall, 3.0-3.5 mm
long, striate, densely sericeous to glabrate, the hairs stiff and irritating,sessile,
the trabecula 1.2-2.4 mm long with the point of attachmentvery eccentric, the
wing 23-30 mm long, 7-10 mm wide, sparsely sericeous to glabrate.
Flora Neotropica
142
Type. Pittier 6117, Venezuela, Miranda,El Cedral de Las Ajuntas near Los
Teques, fl (holotype, B destroyed?;isotypes, NY, US).
Distribution.(Fig. 30). Known only from the mountainsnear Caracas, Venezuela, at altitudes of 1000-1800m, in cloud forest.
Collected in flower in April, October, and December, and in fruit in July,
October and December.
VENEZUELA. Yaracuy: Steyermark 100303 yng fr (NY). D.F.: Pittier 8299 fl fr (GH, US); Rohl
34 fl (US); Steyermark (leg Lasser) 55102 fr (F); 97497 fl fr (NY, US); LI. Williams 10918 fl (F).
Miranda: Pittier 6117 fl (NY, US). Falcon: Steyermark 99089 fr (NY).
This species is characterisedby its large fleshy leaves with few lateral veins,
its basifixed hairs on the young branches, minutely velutinous inflorescence
branches and flowers borne in racemes.
52. Banisteriopsisalternifolia(Steyermark)B. Gates, comb. nov.
Banisteria alternifolia Steyermark, Fieldiana, Bot. 28: 280. 1952.
Liana, the branches sparsely appressed-sericeousat first, eventually glabrate,

stipules minute, triangular.Leaves alternate, coriaceous, with the petiole 8-15
mm long, sparselyappressed-sericeousto glabrate,bearinga pair of sessile glands
apically, with the lamina 4.6-10.6 cm long, 2.3-6.4 cm wide, ovate, obtuse to
truncate at the base, obtuse to acuminateat the apex, with the marginbearing
minute glands apically, adaxiallyglabrous, abaxially at first sparsely appressedsericeous, the hairs sessile, with the trabecula0.2-0.5 mm long, soon glabrate,
with the reticulationprominenton both sides. Inflorescenceaxillary,the branches
up to 30 cm long, silvery appressed-sericeous,the hairs with the trabecula0.150.25 mm long, bearingbract-likeleaves up to 5 mm long subtendingracemes of 910 flowers borne alternately; bracts and bracteoles subtending the pedicels
0.6-1.2 mm long, acute at the apex, glabrous, involucrate, persistent; pedicels
sessile, 5-10 mm long, 0.4-0.6 mm wide. Sepals glabrouson both sides, 2.6-3.0
mm long, 2.0-2.2 mm wide, roundedat the apex, the 4 lateralsepals biglandular,
the glands 1.2-1.3 mm long, 0.7-0.9 mm wide. Petals yellow, laciniate, the 4
lateralpetals reflexed between the sepals, with the claw 1.5-2.0 mm long and up
to 0.4 mm wide, the limb 5.0-6.0 mm long and 4.5-6.0 mm wide, orbicular,the
antero-lateralpetals somewhat largerand more concave than the postero-lateral
petals, the posterior petal with the claw erect, 3.0 mm long and up to 0.6 mm
wide, canaliculate at the apex, the limb up to 4.5 mm long and 4.0 mm wide,
broadlyobovate, with the basal laciniaegland-tipped.Stamens with the filaments
2.2-3.6 mm long, those opposite the 3 anterior sepals and the postero-lateral
petals longest, those opposite the postero-lateralpetals almost the same length
as those opposite the antero-lateralsepals, those of the 3 posterior stamens inflexed between the posterior styles, the anthers with the locules glabrous, 0.81.0 mm long, the connectives of the stamens opposite the 3 anteriorsepals 1.01.2 mm long, 0.6-0.8 mm wide and 0.6-0.8 mm deep, broadlyobovate, overtopping the locules 0.2-0.4 mm, those of the stamens opposite the petals and the
postero-lateralsepals 0.6-0.9 mm long, 0.4-0.6 mm wide, 0.2-0.6 mm deep, oblong, not overtoppingthe locules. Ovary 1.0 mm tall, sparselypubescent, anterior
style straightand erect, 2.6-2.8 mm long, the posterior styles diverging,2.8-3.0
mm long, stigmas capitate. Fruit unknown.
Type. J. A. Steyermark61914, Venezuela, Monagas, Cerro de la Cueva de
Dona Anita, fl (holotype, F).
143
Distribution.(Fig. 30). Known only from two collections, one in Trinidadand

one in Monagas, Venezuela.
Collected in flower in Marchand April.
VENEZUELA. Monagas: Steyermark 61914 fl (F). TRINIDAD. R. Williams 10121 fl (NY).
This is the only species in the genus with alternateleaves, and the flowers are
also borne alternately.It is possible that this is a developmentalabnormalityand
does not warrantspecific recognition,but I preferfor the moment to maintainits
specific identity as it is such a unique species in this respect. It is probablymost
closely related to Banisteriopsis polygama with which it shares the following
characteristics:relatively small coriaceous leaves; silvery-appressedpubescence
in the inflorescence;connectives of the stamensopposite the three anteriorsepals
only a little largerthan the others; styles subequal.
53. BanisteriopsismaguireiB. Gates, Brittonia31: 108. 1979.
Fig. 31.
Liana, branchesgolden-appressed-sericeous,sometimes glabrate. Stipules minute. Leaves opposite or subopposite,very coriaceous, with the petiole 5-14 mm
long, appressed-sericeous,eglandular,with the lamina4.5-12.2 cm long, 2.3-10.0
cm wide, those associated with the inflorescence much smaller and usually reduced to bracts, ovate to orbicular,cordate at the base, obtuse to apiculate at
the apex, with the margin revolute and bearing minute glands on or near the
margin and a single pair of sessile glands at the base of the lamina beside the
midribabaxially, glabrousadaxially, sparsely appressed-sericeousabaxially, the
hairs with the trabecula0.2-1.0 mm long, with the reticulationprominulousadaxially and prominentabaxially. Inflorescence composed of large axillary cymes,
the branchesup to 30 cm long, densely to sparsely appressed-sericeous,the hairs
with the trabecula 0.5-1.3 mm long, with the reduced bract-like leaves of the
inflorescencebearinga well-developedpair of glandsat the base beside the midrib
and no or tiny marginalglands, the flowers borne in racemes of up to 7 pairs of
flowers or rarely 4-flowered umbels; bracts and bracteoles up to 1.0 mm long,
acute at the apex, sparsely sericeous; pedicels sessile, 8-12 mm long, 0.4-0.6
mm in diameter, sparsely to densely appressed-sericeous.Sepals sparsely sericeous abaxially, glabrous adaxially, 2.0-3.0 mm long and 2.0-2.5 mm wide,
roundedat the apex, the 4 lateral sepals biglandular,the glands 1.4-2.0 mm long
and 1.0-1.4 mm wide. Petals yellow, long-fimbriate,the claws sometimes red or
orange, the 4 lateralpetals reflexedbetween the sepals, with the claw 1.5-2.5 mm
long, 0.3-0.4 mm wide, the limb 4.0-7.0 mm long and wide, orbicular,sometimes
with orange median area, the antero-lateralpetals largerand more concave than
the postero-lateralpetals, the posteriorpetal with the claw erect, 2.5-2.8 mmlong,
0.7-1.0 mm wide distally, often canaliculate,the limb 3.5-5.0 mm long, 3.0-4.0
mm wide, broadly ovate to orbicular,the basal fimbriaegland-tipped.Stamens
with the filaments 1.8-3.6 mm long, all nearlythe same length, with the filaments
opposite the styles (i.e. opposite the anterior sepal and postero-lateralpetals)
somewhat longer than the others; antherswith the locules glabrous, 0.6-1.0 mm
long, the connectives opposite the 3 anteriorsepals 1.0-1.2 mm long, 0.8-1.0 mm
wide and 0.6-0.7 mm deep, enlargedand glandular,obovate, projecting0.2-0.5
mm beyond the locules, those of the stamensopposite the petals and the posterolateral sepals 0.4-1.0 mm long, 0.4-0.6 mm wide, 0.2-0.6 mm deep, oblong, not
glandularor glandularat the apex. Ovary 1.0-1.4 mm tall, anteriorstyle 3.0-3.2
mm long, straight and erect, somewhat thicker than the posterior styles, the
144
Flora Neotropica
FIG. 31. Banisteriopsis maguirei. a, flowering branch x 0.5; b, detail of the inflorescence x 5;
c, flower x 5; d, androecium,with posteriorpetal for orientation,x 7.5; e, gynoecium, the anterior

carpel in center, x 7.5; f, samara x 1.5. (Drawnby KarinDouthit;a-e, Steyermark105918;f, Maguire 32850.)
posterior styles 3.2-3.6 mm long, straight and subparallel or diverging, slender,

stigmas capitate. Fruit with filiform carpophore up to 3 mm long, the nut 4-5 mm
tall, 2-3 mm long, sericeous to glabrate, the hairs stiff and irritating with the
trabecula 0.5-1.0 mm long, ridged, with the wing 18-25 mm long, 8-10 mm wide
distally, 2-3 mm wide at the base, appressed-sericeous to glabrate, the hairs with
the trabecula 1.0-1.3 mm long, the wing arising from the upper one-third of the
distal margin of the nut.
145
Type. Maguire & Politi 28323, Venezuela, Amazonas, Cerro Sipapo, fl (holotype, MICH; isotypes, MICH, NY).
Distribution(Fig. 29). Known only from the GuayanaHighlandin Venezuela
at altitudes of 1300-1800m.
Collected in flower in December to May, and in fruit in December, February
and May.
VENEZUELA. Amazonas: Barnes 50 fl (VEN); Maguire 29807 fl (NY); 30010 fl (NY); 30096 fr
(MICH, NY); 30244 fr (NY); 31699 fr (NY); 35163 fl (NY); 35285 fl (NY); 35495 fl (NY); Steyermark
105918 fl (NY). Bolivar: Maguire 32850 fr (NY); Steyermark 109672 at (MICH).
This species is distinguishedby its very coriaceous leaves which are usually
appressed-sericeousabaxially, its appressed-sericeouspubescence in the inflorescence, and the flowers usually borne in racemes.
It is named for Bassett Maguirewho has contributedmore to the knowledge
of the flora of the Guayana Highland, where this species is endemic, than any
other individual.
54. Banisteriopsispulcherrima(Sandwith)B. Gates, Brittonia31: 109. 1979.
Fig. 32.
Banisteriopsis elegans var. pulcherrima Sandwith, J. Arnold Arbor. 24: 223. 1943.
Shrub or vine on ground, branches glabrous. Stipules minute. Leaves very

coriaceous, with the petiole 2-6 mm long, glabrous, eglandular,with the lamina
5.2-14.5 cm long, 3.5-11.8 cm wide, those associated with the inflorescence
smaller and often reduced to bracts, ovate to broadly ovate, strongly cordate at
the base, acute to short-acuminateat the apex, with the margin more or less
revolute and bearing apically glandularteeth or ciliate extensions, abaxially at
the base of the leaf beside the midrib bearing a single pair of sessile glands,
glabrouson both sides, with the lateralveins and the scalariformsecondaryveins
interconnectingthem prominenton both sides. Inflorescenceaxillaryor terminal,
cymose, of 4-flowered umbels, the branches up to 45 cm long, glabrous, the
reduced or bract-likeleaves of the inflorescence with ciliate projectionsup to 6
mm long aroundthe margin,subtending1-3 umbels or 3-4 pairs of flowers, the
umbel-bearingbranches 15-25 mm long; bracts and bracteoles 0.6-0.8 mm long,
glabrous; pedicels sessile, 12-18 mm long, 0.4-0.5 mm in diameter, glabrous.
Sepals glabrouson both sides, 2.0-2.5 mm long and wide, rounded at the apex,
the 4 lateralsepals biglandular,the glands 1.5-2.0 mm long and 1.2-1.5 mm wide.
Petals yellow, lacerate, the 4 lateralpetals reflexed between the sepals, with the
claw 3.5-4.0 mm long, 0.6-0.7 mm wide, the limb 8.0-11.0 mm long and wide,
orbicular, the antero-lateralpetals larger and more concave than the posterolateralpetals, the posteriorpetal with the claw erect, 3.5 mm long and up to 1.2
mm wide at the apex, the limb 8.5-9.5 mm long and wide, reflexed, broadly
obovate, often veined with red, the basal teeth gland-tipped.Stamens with the
filaments 2.6-4.0 mm long, those opposite the 3 anterior sepals longest, those
opposite the postero-lateralpetals almost as long as those opposite the anterolateral sepals, those of the 3 posterior stamens inflexed between the posterior
styles; antherswith the locules glabrous,0.7-1.0 mm long, the connectives of the
stamens opposite the 3 anteriorsepals 1.2-1.6 mm long, 1.4-1.8 mm wide, 1.01.4 mm deep, globose, enlargedand glandular,overtoppingthe locules, those of
the stamens opposite the petals and the postero-lateralsepals 0.5-1.0 mm long,
0.4-0.8 mm wide, 0.4-0.6 mm deep, oblong, not overtoppingthe locules. Ovary
1.2-1.4 mm tall, densely hairy, anteriorstyle straightand erect, 2.8-3.0 mm long,
Flora Neotropica
146
..
~~~~~~~~~~~~~~~~~~,.
%?
i.. -
FIG. 32. Banisteriopsis pulcherrima. a, flowering branch x 0.5; b, flower x 1.5; c, androecium
and gynoecium, side view, the enlargedanther to the left opposite the eglandularanteriorsepal, 1
large anther removed, x 5; d, part of androecium,the leftmost anther opposite the anteriorsepal,
the rightmostopposite the posterior petal, x 5.5; e, gynoecium, the shorter style anterior, x 6; f,
fruit, the middlesamaraanterior, x 1. (Drawnby KarinDouthit;a, Maguire& Fanshawe 23125; bf, Maguire et al. 43889.)
147
0.2-0.3 mm in diameter, the posterior styles divergingat the base and directed
posteriorly, 3.8-4.0 mm long and 0.10-0.15 mm wide, stigmascapitate. Fruitwith
filiformcarpophoreup to 3 mm long, with the nut 4-6 mm tall and 3-4 mm long,
ridged, appressed-sericeousto glabrate,the hairs sessile, with the trabeculastiff
and irritating, 1.0-1.5 mm long, the wing 24-35 mm long, 10-13 mm wide, appressed-sericeousto glabratelike the nut, arisingfrom the upper one-thirdof the
distal marginof the nut.
Type. N. Sandwith 1430, Guyana, Kaieteur Savannah, fl fr (holotype, K;
isotype, U).
Distribution.(Fig. 29). In open places in the GuayanaHighlandat altitudes of
400-2130 m.
Collected in flower in December to September,and in fruit in December, February, and May to October.
VENEZUELA. Bolivar: Bunting 2964 fl (MY); 3042 fl (F, MY); Ferndndez 1120 fl (MY); Kunhardt
2 fl (NY); 21 fl (NY); Maguire 53561 fl (NY, VEN); 53669 fl (NY, VEN); Steyermark & Nilsson 83
fl (NY, VEN); 834 fl (VEN); Steyermark 58689 fl (F, NY); 92094 fl (F, NY, VEN); 11290 fr (MICH);
Trujillo3541 fl (MY). GUYANA. Cowan & Soderstrom 1815 fr (NY, US); 2030 fl (NY, US); Hollister
s.n. fl (NY); Jenman 243 fl (US); Maguire 23125 fl fr (BR, F, GH, MO, NY, P, U, US); 43889 fl fr
(MICH, NY); Tillett 44905 fl (MICH, NY); 45597 fr (NY); Tutin 694 fl (US).
This is a distinctive and beautiful species with its large flowers arrangedin
showy inflorescences, its coriaceous subsessile leaves, and long-ciliateinflorescence bracts.
The Banisteriopsis nummifera group (species 55-59)
This group consists of one widespreadand very variable species, B. nummifera, and four species with more restricteddistributions.The species in this group
share the following characters:habit of a liana (in open habitatsthey will sometimes vine upon themselves);leaves long-petiolatewith a largepairof protuberant
glands at the apex of the petiole; flowers borne in large showy panicles, with 620 flowers borne in racemes on the ultimate branches; fruits with appressed,
irritatingpubescence, the hairs sessile with the trabeculastraight,up to 1.5 mm
long.
55. Banisteriopsisnummifera(Adr. Jussieu) B. Gates, comb. nov.
Banisteria nummifera Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 281. 1840.
Banisteria blanchetiana Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 285. 1840. Type. Blanchet
2576, Brazil, Bahia, Serra de Jacobina, fl (holotype, P, isotypes, BR, F, G, GH, NY, P, W).
Banisteria sellowiana var. blanchetiana (Adr. Jussieu) Niedenzu, Ind. Lect. Lyc. Brunsb.: 8.
1900.
Banisteria gardneriana var. nummifera (Adr. Jussieu) Niedenzu, Ind. Lect. Lyc. Brunsb.: 9.
1900.
Banisteria gardneriana var. nummifera f. luschnathiana Niedenzu, Ind. Lect. Lyc. Brunsb.: 9.
1900. Types. Luschnath s.n., Brazil, Bahia, Ilheos, fl (lectotype, BR; isolectotype, BR).
Schiich s.n., Brazil, Rio de Janeiro, fl (syntypes, BR).
Banisteria gardneriana var. nummifera f. blanchetiana Niedenzu; Ind. Lect. Lyc. Brunsb.: 9.
1900. Types. Blanchet 3174 et 3174a, Brazil, Bahia, fl (holotype, B destroyed?; isotypes, F,
G, P, W).
Banisteria cinerea Rusby, Bull. New York Bot. Gard. 4: 335. 1907. Type. M. Bang 2489, Bolivia,
Unduavai, fl (holotype, NY; isotypes, F, GH, MICH, MO, NY, US).
Banisteria sphaerandra Rusby, Mem. New York Bot. Gard. 7: 276. 1927. Type. G. S. McCarty
(Mulford Biol. Explor. of the Amazon Basin) 126, fl (holotype, NY).
Banisteria sellowiana var. blanchetiana f. grandifolia Niedenzu in A. Engler, Das Pflanzenreich
IV, 141: 409. 1928. Type. Steinbach 6530, Bolivia, Dep. Santa Cruz, Prov. Sara, Buenavista,
fl (holotype, B destroyed, NY fragment; isotype, GH).
148
Flora Neotropica
Liana, branches sparsely appressed-sericeousor glabrous. Stipules minute,

triangular.Leaves with the petiole 7-22 mm long, sparsely appressed-sericeous
to glabrous, apically biglandular,the lamina4.3-19.0 cm long, 2.0-8.8 cm wide,
narrowly ovate to ovate or elliptic, obtuse to cordate at the base, obtuse to
acuminateat the apex, with the marginslightly revolute, glabrousadaxially, appressed-sericeousto glabrousabaxially,with the reticulationprominentadaxially
and the lateral veins prominentabaxially. Inflorescenceof large axillarypanicles
bearingvery reduced leaves, appressed-sericeousto glabrate, the side branches
bearing6-12 flowers borne alternatelyor in pairs; bracts and bracteoles 0.8-1.4
mm long, triangular,sparsely appressed-sericeousto glabrate, involucrate, persistent; pedicels sessile or short-stalked,5-10 mm long, 0.3-0.6 mm in diameter,
sparsely appressed-sericeousto glabrate. Sepals 2.2-3.4 mm long, 1.6-2.8 mm
wide, elliptic, rounded and often revolute at the apex, sparsely appressed-sericeous or glabrous abaxially, glabrousadaxially, the 4 lateral sepals biglandular,
projecting 1.0-2.0 mm beyond the glands, the glands 1.0-2.0 mm long, 0.6-1.4
mm wide. Petals yellow, glabrous,dentateto lacerate, the 4 lateralpetals reflexed
between the sepals, the claw 1.5-2.0 mm long, the limb 6.0-11.0 mm long and
5.0-9.0 mm wide, elliptic to orbicular,the antero-lateralpetals larger and more
concave than the postero-lateralpetals, the posterior petal with the claw erect,
2.5-4.0 mm long, constricted at the apex, the limb 5.0-8.0 mm long and 4.0-7.5
mm wide, broadlyobovate, plane. Stamens with the filaments2.0-3.6 mm long,
those opposite the sepals longer than those opposite the petals, or those opposite
the postero-lateralpetals as long as those opposite the sepals; anthers with the
locules 0.6-1.2 mm long, glabrous, the connectives of the stamens opposite the
3 anterior sepals 0.8-1.3(-1.8) mm long, 0.7-1.0(-1.8) mm wide, 0.5-0.9 mm
deep, obovate, sometimes swollen and glandularapically and projecting0.2-0.6
(-1.0) mm beyond the locules, those opposite the petals 0.5-1.0(-1.4) mm long,
0.4-0.7(-0.9) mm wide, 0.2-0.4 mm deep, oblong, not projecting beyond the
locules. Ovary0.8-1.4 mm tall, sericeous or glabrous,the styles 2.2-3.0 mm long,
stout and subequal, divergingfrom the base, stigmas capitate. Fruit with carpophore up to 2.5 mm long and 0.6 mm wide, the nut 6-8 mm tall, 3-5 mm long,
rugulose, appressed-sericeousto glabrate, the hairs stiff and irritating,the trabecula 1.0-1.6 mm long, the distal marginwith a prominenttooth, the wing 2232 mm long, 11-15 mm wide, oblong, appressed-sericeousto glabrate like the
nut, with the sepals enlargingin fruit.
Type. Salzmann pl. Bras. exs. 92, Brazil, Bahia, fl yng fr (holotype, P-JU;
isotype, G).
Distribution.(Fig. 33). In rain forest in Amazonia and Bolivia, in the Atlantic
coastal forest of SE Brazil and also in gallery forest in the Planaltoof Brazil.
Collected in flower in Januaryand from May to November, and in fruit from
August to November.
FRENCH GUIANA. de Granville 1265 fr (NY, P). PERU. Loreto: Ule 6350 fl (G, HG).
BRAZIL. Amazbnas: Chagas s.n., INPA 2016 fl fr (INPA, MG, SP); s.n., INPA 4037 fl fr (IAN,
INPA, MG, SP); Coelho 5809 fr (SP); Jobert & Schwacke 450 fl (P, R); J. G. Kuhlmann 150 fl (RB);
Mello & Coelho s.n., INPA 4189 fl (IAN, MG); Prance et al. 14357 bud (NY). Para: Bockermann
213 fl (GH, IAN, SP). Rond6nia: Prance et al. 5891 fl (MICH, NY). Mato Grosso: Archer & Gehrt
127 fl (SP, US); Argent in Richards 6663 fl (NY); Harley & Souza 10034 fl fr (NY); Robert 523 fl
(NY). Goias: Hatschbach 38771 fl (MICH); Lane s.n. fl (SP). Bahia: Blanchet 2576 fl (BR, F, G,
GH, NY, P, W); 3174 fl (F, G, P, W); 3174A fl yng fr (G); Eupunino 264 fl (MICH); 283 fl (MICH);
Harley 18078 fl (K); Huber 164 fl (MG); Luschnath s.n. fl (BR); Martius s.n. fl (P); Salzmann 92 fl
yng fr (G, P). Minas Gerais: Glaziou 12484 fl (BR, G, GH, NY, P); Macedo 2512 fl fr (IAN, NY, S,
SP, US). Rio de Janeiro: Glaziou 9676 fl fr (P); Schiich s.n. fl (BR). Sao Paulo: Frazdo s.n. fl (RB);
Gehrt s.n. fl (SP); M. Kuhlmann 986 fl (SP); Lofgren 101 fl (C); 1351 fl (P, SP); Mosen 3967 fl (S);
Novaes s.n. fl (GH, SP). BOLIVIA. Bang 2489 fl (F, GH, MICH, MO, NY, US); Buchtien 669 fl
149
(GH); 3797 fl (NY, US); Cdrdenas5692 fl (US); Krukoff10857fl (F, G, GH, MICH, MO, NY, S, U,
US); 11184fl fr (G, GH, MICH, MO, NY, S, U); McCarty126 fl (NY); Rusby 852 fl (MICH, NY);
1033 fl (NY); Steinbach6495 fl (F, G, GH, NY); 6530 fl (G, NY); Troll2693 fl (M).
There is probably more than one taxon represented by the variation I have
includedin B. nummifera,but I failed to find any consistent basis for subdivision.
The characterwhich shows most obvious variationis the abaxialleaf pubescence;
some individualsare densely and persistently appressed-sericeous,others are at
first appressed-sericeous, and soon or eventually glabrate, whereas some individuals are always glabrous. However, this characterdoes not seem to correlate
with geographyor other charactersthat vary, such as leaf size and shape, inflorescence pubescence, number of flowers on the side branches of the inflorescence, flower size, and degree of enlargementof the stamen connectives.
Specimens of Banisteriopsis nummiferawith sericeous leaves can be distinguished from B. gardneriana by their three stout equal styles. Glabrousspecimens can be distinguishedfrom B. sellowiana by their largerflowers and larger
leaves.
56. Banisteriopsissellowiana(Adr. Jussieu) B. Gates, comb. nov.
Banisteriasellowiana Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 419. 1843.
Liana, branchesglabrous, young branches smooth, terete, older branches minutely fissured with numerousprominulouslenticels. Stipules up to 1 mm long,
triangular,glabrous,joined by an interpetiolarridge. Leaves with the petiole 510 mm long, glabrous,eglandularor apicallybearinga pair of protuberantglands,
the lamina4.1-11.4 cm long, 2.2-5.0 cm wide, elliptic, obtuse at the base, acute
to short-acuminateat the apex, with the marginslightly revolute, usually bearing
a gland at the base of the leaf beside the midribon each side, glabrouson both
sides, with the reticulation prominent on both sides. Inflorescence axillary or
terminal, paniculate, minutely appressed-sericeousto glabrate, bearing reduced
leaves bearing a pair of prominentglands at the base and abaxially appressedsericeous, the side branches bearing 3-5 pairs of flowers; bracts and bracteoles
0.6-1.0 mm long, triangular,glabrous,involucrate,persistent;pedicels sessile or
borne on a peduncle up to 1.5 mm long, 5-7 mm long, 0.2-0.4 mm in diameter,
sparsely appressed-sericeousto glabrate. Sepals 1.8-2.6 mm long, 1.4-2.0 mm
wide, broadly elliptic, rounded at the apex, glabrouson both sides, the 4 lateral
sepals bearing glands 0.6-1.2 mm long and 0.5-0.7 mm in diameter, projecting
1.0-1.5 mm beyond the glands. Petals yellow, irregularlydentate,glabrous,eglandular, the 4 lateral petals reflexed between the sepals with the claw 1.0-2.0 mm
long, the limb 5.0-7.0 mmlong, 4.5-6.0 mm wide, rotund,the antero-lateralpetals
largerand more concave than the postero-lateralpetals, the posteriorpetal with
the claw erect, up to 2.8 mm long, the limb 4.5-5.5 mm long and 3.5-4.0 mm
wide, obovate, plane. Stamenswith the filaments1.2-3.2 mm long, those opposite
the 3 styles longest, those of the 3 posterior stamens shortest; anthers with the
locules 0.4-1.0 mm long, glabrous, the connectives of the stamens opposite the
3 anterior sepals 1.0 mm long, 0.6-0.9 mm wide, 0.6-0.8 mm deep, obovate,
glandularapically, the others 0.4-0.8 mm long, 0.2-0.4 mm wide, 0.1-0.3 mm
deep, elliptic, eglandular.Ovary 1.0-1.4 mm tall, sericeous, the styles slender
and subequal,2.4-2.8 mmlong, the anteriorstyle directedanteriorlywith the apex
bent inwards, the posterior styles divergingfrom the base, hooked at the apex,
stigmas truncate. Fruit of 3 samaraswith filiformcarpophoreup to 1.5 mm long,
the nut up to 5 mm tall and 3 mm long, reticulate,appressed-sericeousto glabrate,
the hairs stiff and straightwith the trabeculaup to 1 mm long, bearinglaterally
150
Flora Neotropica
on the upper distal margin of the nut a triangular crest 2-3 mm along its base and
1.5-3.5 mm tall, the wing 20-24 mm long, 9-11 mm wide, narrower towards the
base, appressed-sericeous to glabrate like the nut. Sepals enlarging, up to 4 mm
long, in fruit.
Type. Sellow II it. 234 (a. 1815), Brazil, Rio de Janeiro, between Rio and
Campos, fl (holotype, B destroyed, fragment NY).
Distribution. (Fig. 33). Only known from restingas near Rio de Janeiro, Brazil.
Collected in flower in October and in fruit in October.
BRAZIL. Rio de Janeiro:Gardner18 fl (GH, W); Glaziou8587 fl fr (BR, C, G, P, R, S); Lund
1058 fr (C); Mikan s.n. fl (W); Porto s.n. fl (MICH); Pereira 4363 fl (MICH, RFA); Riedel 1037 (X,
1832) fl (C, F, NY, P, US, W); Sellow II it. 234 (a. 1815) fl (NY fragment ex B); Vauthier s.n. fl (P).
Banisteriopsis sellowiana is distinguished from most members of the B. nummifera group by its glabrous leaves. It can be distinguished from glabrous-leaved
individuals of B. nummifera by its inflorescence with shorter side branches (up
to 1.5 cm) bearing fewer smaller flowers with small calyx glands and slender
styles, and its fruit with a smaller nut bearing a lateral winglet, and with a more
slender wing. Superficially, B. sellowiana resembles the B. martiniana group,
but the reticulate secondary veins easily distinguish it from this group.
57. Banisteriopsis anisandra (Adr. Jussieu) B. Gates, comb. nov.
BanisteriaanisandraAdr. Jussieuin Saint Hilaire, Fl. bras. mer. 3: 47. 1832[1833].
BanisteriaclaussenianaAdr. Jussieu,Ann. Sci. Nat. Bot., Ser. 2, 13: 285. 1840.Type. Claussen
s.n. (121), Brazil, MinasGerais, Caxoeirado Campo,fl (holotype, P; isotypes, G, P, P-JU).
Banisteriopsisclausseniana(Adr. Jussieu)Andersonet Gates, Contr.Univ. Mich. Herb. 11: 53.
1975.
Liana, young branches flattened, densely and minutely tomentose, older

branches terete, eventually glabrate, grey-brown. Stipules up to 1.5 mm long,
glabrate, their bases joined by a prominent interpetiolar ridge. Leaves opposite
or subopposite, the petiole 7-20(-32) mm long, white-tomentose, ridged abaxially,
bearing a pair of large sessile glands apically, the lamina 5.0-16.7(-22.0) cm long,
2.2-8.1(-11.8) cm wide, elliptic or rarely narrowly elliptic, obtuse to truncate at
the base, obtuse to apiculate or rarely emarginate at the apex, with the margin
slightly revolute, adaxially tomentose, soon glabrate except the midrib, abaxially
densely white-tomentose, with the reticulation prominent adaxially and the lateral
veins prominent abaxially. Inflorescence of large axillary panicles bearing reduced
leaves, the flowers borne in pairs on the lateral branches, densely tomentose to
tomento-velutinous; bracts and bracteoles 1.2-2.5 mm long, narrowly oblong,
rounded or obtuse at the apex, sparsely tomentose abaxially, involucrate, persistent; pedicels 7-10(-12) mm long, 0.5-0.7 mm in diameter, golden-tomentose
sessile or pedunculate, the peduncle up to 3 mm long. Sepals 2.0-4.0 mm long,
2.0-3.0 mm wide, ovate to oblong, rounded and often revolute at the apex, sparsely tomentose abaxially, glabrous adaxially, the 4 lateral sepals biglandular, the
glands 1.0-1.6 mm long and 0.6-1.3 mm wide, the sepals projecting 1.5-2.5 mm
beyond the glands. Petals yellow, glabrous, dentate, the 4 lateral petals reflexed
between the sepals, the claw 2.0-3.0 mm long, the limb 7.0-10.0 mm long, 6.59.0 mm wide, rotund to orbicular, the antero-lateral petals slightly larger and
more concave than the postero-lateral petals, the posterior petal with the claw
erect, 3.0-3.5 mm long, 0.5-0.6 mm in diameter, the limb 6.0-7.0 mm long and
5.0-5.5 mm wide, oblong, plane. Stamens with the filaments 2.0-3.4 mm long,
that opposite the anterior sepal longer, and that opposite the posterior petal shorter than the others, the others subequal, those of the 3 posterior stamens inflexed
151
\?
-'.'..'' .,i
' .":.:?
'
'' .
-..
.'.
FIG. 33. Distribution of Banisteriopsis nummifera group, species 55-59. Open circle, B. nummifera; closed circle, B. sellowiana; triangle, B. anisandra; open square, B. gardneriana; closed
square, B. lyrata.
between the posterior styles; antherswith the locules 0.8-1.2 mm long, glabrous,
the connectives of the stamens opposite the 3 anterior sepals 1.0-1.4 mm long,
0.8-1.1 mm wide, 0.7-1.0 mm deep, obovate, glandularand enlarged apically,
projecting a little above the locules, those opposite the petals and the posterolateral sepals 0.5-0.9 mm long, 0.4-0.6 mm wide, 0.1-0.6 mm deep, oblong,
eglandularor those opposite the antero-lateralpetals and postero-lateralsepals
glandular apically, not projecting beyond the locules. Ovary 0.8-1.2 mm tall,
152
Flora Neotropica
apically sericeous, the styles equal or subequal, 2.4-3.2 mm long, diverging,

stigmas capitate. Fruit with carpophoreup to 2.5 mm long and 0.8 mm wide, the
nut 4-8 mm tall and 3-4 mm long, rugulose, appressed-sericeousto glabrate,with
a "tooth" at the base of the lower margin, the wing 18-28 mm long, 8-13 mm
wide, narrower at the base, the lower margin arcuate, appressed-sericeousto
glabrate, the hairs stiff and straight, the trabecula 1.0-1.5 mm long, bearing a
shallow rounded appendageat its base up to 2.5 mm long and 1 mm tall. Sepals
enlarged, up to 6 mm long in fruit.
Type. A. de Saint-Hilaire C?-549, Brazil, Minas Gerais, Rio Manso, fl (holotype, P; isotypes, P).
Distribution. (Fig. 33). In cerrado in Minas Gerais, Goias and the Distrito
Federal of Brazil, and occasionally collected in Sio Paulo and Bahia.
Collected in flower from August to October, and in fruit in September and
October.
Representativecollections. BRAZIL. Goias: Glaziou 20754 fl (G, NY, P); 20756 fl (G, P, R);
Hatschbach & Kummrow 37273 fl (MICH); Irwin et al. 34359 old fr (NY); Ule 144 fl (NY, P, R).
D.F.: L. Duarte & R. S. Santos 105 fl (HB, NY, U); Heringer 103 fl (MICH); 11868 fl (NY); Irwin
& Soderstrom 6035 fl (MICH, NY); Prance & Silva 59071 fl (F, G, GH, NY, P, S, U, US); Ratter
R 3759 fr (E). Bahia: R. S. Santos s.n. fl (NY). Minas Gerais: Barreto 864 fl (RB); 7695 fl (R);
Claussen 86A fl (BR, W); 1538 fl (F, GH, P, RB, US); Fr6es 33317 fl (IAN); Glaziou 18942 fl (C, P,
R); Goodland 830 fl (MICH); Lund s.n. fr (GOET); Oliveira 1145 fl (IAN, US); R. S. Santos s.n.
(HB 28461)fr (MICH);Semiret al. 4373 fl (E); Warming353 fl (M); s.n. fl fr (C, US). Rio de Janeiro:
Riedel s.n. (IX. 1824) fl (C, P, US, W). Sao Paulo: Bicalho 27 fl (SP); J. Campos 47 fl (NY, US); M.
Kuhlmann 3062 fl (SP); Lofgren 862 fl (C).
Common names. Brazil: "cipo de prata."
This species is most closely relatedto Banisteriopsisgardneriana,from which

it can be distinguishedby its dense, white, tomentose abaxial leaf pubescence,
the reticulationprominenton the adaxialleaf surface, its small calyx glands, and
three stout equal styles. It has the habit of a liana when growing in forest, and
will vine upon itself formingdense thickets when growingin open cerrado.Unlike
most cerrado species of Banisteriopsis which flower in the rainy season, B. anisandra flowers in the dry winter season. Few fruiting collections have been
made, perhapsbecause it producesfew fruits, or perhapsbecause the fruits mature rapidlyand are immediatelydispersed.
58. Banisteriopsisgardneriana(Adr. Jussieu) Anderson & Gates, Contr. Univ.
Mich. Herb. 11: 54. 1975.
BanisteriagardnerianaAdr. Jussieu, Arch. Mus. Paris3: 421. 1843.
BanisteriamonostylaNiedenzu, Arb. Bot. Inst. Ak. Braunsberg8: 61. 1926.Types. Commissdo
Rondon (leg Hoehne) 1898, Brazil, Mato Grosso, fl (syntype, B, destroyed?);1909, Brazil,
Mato Grosso, Juruena,fl (syntype, B, destroyed?).
Liana, branches minutely tomento-velutinous, eventually glabrate, the bark

minutelyfissuredwith numeroustiny lenticels. Stipules 1.0-3.0 mm long, tomentose to glabrate, their bases often joined by a prominent interpetiolarridge.
Leaves coriaceous, with the petiole 5-16 mm long, densely tomentose or velutinous, bearinga pair of large sessile glands apically, the lamina 5.2-13.9(-18.1)
cm long and 3.2-6.9(-11.1) cm wide, ovate to elliptic, obtuse to cordate at the
base, obtuse to acute or apiculate,rarelyemarginate,at the apex, with the margin
slightly revolute, adaxially at first sparsely tomentose, soon glabrateexcept the
midribat the base, shining,abaxiallydensely appressed-sericeous,adaxiallywith
the primary and secondary veins prominent in young leaves, prominulousor
smooth in older leaves, the lateral veins prominentabaxially. Inflorescence of
153
large axillarypanicles, bearingreduced leaves, the side brancheswith 6-14 flowers borne alternatelyor in pairs, minutely white-tomento-velutinous;bracts and
bracteoles 1.0-2.0 mm long, ovate to lanceolate, roundedto obtuse at the apex,
sparsely tomentose abaxially, involucrate, persistent; pedicels 7-13 mm long,
0.6-1.0 mm in diameter,white-tomento-velutinoussessile or pedunculate,the peduncle up to 5 mm long. Sepals 2.6-4.0 mm long, 1.6-3.4 mm wide, elliptic,
rounded at the apex, sparsely appressed-sericeousabaxially, glabrousadaxially,
the 4 lateral sepals biglandular,projecting 1.0-2.4 mm beyond the glands, the
glands 1.4-2.6 mm long, 0.8-1.8 mm wide. Petals yellow, glabrous, dentate, the
4 lateral petals reflexed between the sepals, the claw 1.5-3.0 mm long, the limb
7.0-12.0 mm long and wide, orbicular,the antero-lateralpetals largerand more
concave than the postero-lateralpetals, the posterior petal with the claw erect,
3.0-4.0 mm long, up to 1.4 mm wide, articulatedat the apex, the limb 5.0-9.5
mm long and wide, orbicularto broadly obovate, plane. Stamens with the filaments 1.8-3.8 mm long, those opposite the sepals usually longer than those opposite the petals, sometimes those opposite the postero-lateralpetals as long as
those opposite the sepals; anthers with the locules 0.6-2.0 mm long, glabrous,
the connectives of the stamens opposite the 3 anteriorsepals 1.2-2.4 mm long,
0.9-2.2 mm wide, 0.9-1.6 mm deep, obovate or globose, projectingup to 0.4 mm
beyond the locules, those opposite the petals and postero-lateralsepals 0.5-1.6
mm long, 0.4-1.0 mm wide, 0.2-1.0 mm deep, elliptic, not projectingbeyond the
locules. Ovary 1.0-1.8 mm tall, sericeous throughoutor only apically,the anterior
style 2.8-3.6 mm long, stout and erect, up to 0.5 mm in diameter, the posterior
styles shorter and more slender than the anterior style, or absent, 1.0-3.0 mm
long, up to 0.2 mmin diameter,divergingfromthe base, stigmasstronglycapitate.
Fruit with carpophore2.0-4.0 mm long and 1.0-2.0 mm wide, the nut 7-13 mm
tall and 5-8 mm long, rugulose, appressed-sericeousto glabrate,the hairsgolden,
the trabeculastiff, straight, 1.0-1.5 mm long, the distal marginwith a prominent
"tooth," the wing 20-33 mmlong, 10-21 mm wide, oblong, the lower marginmore
or less parallel to the upper margin, sparsely appressed-sericeousto glabrate,
without a basal appendage.Sepals not enlargedin fruit.
Type. G. Gardner2502, Brazil, Piaui, Paranagoa,fl (holotype, G; isotypes,
GH, MO, P, W).
Distribution.(Fig. 33). In the northernpartof the Planaltoof Brazil, in northern
MinasGerais, Goias and the DistritoFederal,extendingN to Piauiand Maranhao,
and W to Mato Grosso.
Collected in flowerfrom Aprilto Septemberand in December, and in fruitfrom
July to September.
BRAZIL. Maranhao: Fr6es 27796 fl (IAN). Piaui: Ducke 390 fl (MG); 806 fl fr (MG); Gardner 2502
fl (G, GH, MO, P, W). Mato Grosso: Argent in Richards 6478 fl (IAN, NY); 6646 fl (IAN, NY);
Commissdo Rondon (leg Hoehne) 1892 fl (R); Maguire et al. 56430 fl (NY); 56489 fl fr (NY); Ratter
R 32 fl fr (NY). Goias: Andrade 342/Emmerich 334 fl (HB); Estabrook 17 old fr (MGM, MICH);
Gemtchujuicov s.n. fl (SP); Haas & Belem 370 fl (MICH); Heringer 15885 fl (MICH); 15890 fl
(MICH); Liitzelburg 1582 fl (M, NY); Macedo 3692 fl (MO, RB, S, US); 3899 fl (IAN, RB); 5382 fl
(US). Distrito Federal: Heringer 14654 fl (MICH); Irwin et al. 7838 fl (MICH, NY); 8377 fl fr (NY,
US); Ratter R3418 fl (E, MICH); R3509 fl (E, MICH). Bahia: Liitzelburg 151 fl (M); 1824 fl (M).
Minas Gerais: Davis et al. 2355 fl (E); Glaziou 12484 fl (R); Magalhdes 4688 fl fr (IAN); Santos &
Castellanos 24197 fl (NY); Sobrinho 208 fl (GUA, MICH, NY); L. O. Williams 8196 fr (GH). Rio de
Janeiro:Glaziou 12489fl (BR, C, G, P).
Common names. Brazil: "crista de galo," "cipo prata."
Banisteriopsis gardneriana is a common and variable species, showing variation in leaf size and shape, type and density of the inflorescencepubescence, and
154
Flora Neotropica
size of the posterior styles. Typical B. gardnerianafrom the eastern part of the
rangeof the species has posteriorstyles which are slenderbut only a little shorter
than the anterior style. Collections from Goias, D.F. and Mato Grosso include
plants in which one or both posterior styles are aborted, and those plants were
segregated by Niedenzu as Banisteria monostyla. However, since all intermediates between well-developedposteriorstyles and no posterior styles are found,
sometimes on the same inflorescence, I am not maintainingB. monostyla as a
separatetaxon. The collections with one style from Mato Grosso have relatively
shorter petioles and longer side branches in the inflorescence than collections
from other parts of the range, but these differences do not seem sufficient to
warranttaxonomic recognition.
Associated with the reductionof the posterior styles there is a reductionin the
size of the posterior stamens and an increase in the size of the anthers of the
stamens opposite the three anteriorsepals so that these flowers are very markedly
zygomorphic. It is interesting that B. schwannioides, which exhibits a similar
loss of the posteriorstyles, shows this same asymmetryin the size of the stamens.
This species also apparentlyoccurs in a diversity of habitats;it has been collected in dry forest, galleryforest, cerradoand on rocky outcrops. Whenit occurs
in forest it is a liana, but in the absence of trees to climb on, it will vine upon
itself formingthickets like its close relative Banisteriopsis anisandra. Most collections of B. gardneriana in flower were made from July to September, during
the dry season. A few floweringcollections were made in April to June. Perhaps
these collections were made in unusuallydry years which stimulatedthe plants
to flowerearly. Thereare relativelyfew collections of this species in fruit;perhaps
this species produces few fruits or is inconspicuous in fruit, or matures and
disperses its fruits very early, soon after flowering.
Banisteriopsis gardneriana can be distinguished from Banisteriopsis anisandra
by its appressed-sericeousleaf pubescence, large calyx glands, and slender, more

or less reduced posterior styles.
59. Banisteriopsislyrata B. Gates, Brittonia31: 108. 1979.
Fig. 34.
Liana, young branches appressed-sericeous,older branches glabrate. Stipules
minute, triangular,appressed-sericeousto glabrate.Leaves with the petiole 6-10
(-20) mm long, golden-tomento-sericeous,with a pair of prominentglands apically, the lamina 6.0-12.8 cm long, 2.9-6.5 cm wide, narrowlyelliptic to elliptic
or ovate, cordate at the base, obtuse to acute at the apex, with the marginplane,
adaxially glabrous, abaxially densely appressed-sericeous,more or less shining
and golden, with the lateralveins prominenton both sides. Inflorescenceaxillary,
paniculate, the lateral branchesbearing3-4 pairs of flowers, sparsely to densely
tomento-sericeous,the hairs minutewith the trabeculaup to 0.4 mm long, bracts
and bracteoles 1.0-1.5(-2.0) mm long, sparsely sericeous to glabrate, broadly
triangular,acute at the apex, involucrate, persistent; pedicels sessile, 6-10 mm
long, 0.4-0.6 mm in diameter, sparsely to densely sericeous. Sepals 2.2-3.0 mm
long, 1.6-2.0 mm wide, rounded at the apex, sericeous throughoutabaxially or
glabrous distally, glabrous adaxially, the 4 lateral sepals biglandular,projecting
0.8-1.7 mm beyond the glands, the glands 0.8-2.0 mm long, 0.6-1.4 mm wide.
Petals yellow, long-fimbriate,glabrous, the 4 lateral petals reflexed between the
sepals, the claw 1.0-2.0 mm long, the limb 6.0-10.0 mm long and 4.5-7.0 mm
wide, broadly elliptic, plane, the antero-lateralpetals somewhat larger than the
postero-lateralpetals, the posterior petal with the claw erect, 3.0 mm long and
0.6-1.0 mm wide, apically constricted,the limb 5.0-8.0 mm long and 3.5-5.0 mm
FIG.. 34.
155
Banisteriopsis lyrata. a, leafy shoot and inflorescence on older leafless stem x 0.5; b,
flower and posterior(flag)petal x 5; c, androeciumand gynoecium, lateral-posteriorview, x 7.5; d,

stamen opposite antero-lateralsepal x 7.5; e, gynoecium, anteriorstyle in center, x 5. (Drawnby
KarinDouthitfrom Prance & Ramos 6960.)
156
Flora Neotropica
wide, obovate, with 2-3 prominentgland-tippedteeth proximally. Stamens with

the filamentsopposite the 3 anteriorsepals and the postero-lateralpetals 3.4-4.4
mm long, those opposite the postero-lateralpetals longest, those opposite the
other petals and the postero-lateralsepals 1.8-2.6 mm long, those of the 3 posterior stamens inflexed between the posterior styles; anthers with the locules
glabrous,0.6-1.0 mm long, the connectives of the stamensopposite the 3 anterior
sepals 0.9-1.3 mm long, 0.7-1.1 mm wide, 0.5-0.8 mm deep, broadly obovate,
glandularand enlarged, overtoppingthe locules 0.2-0.7 mm, those opposite the
petals and the postero-lateralsepals 0.4-0.8 mm long, 0.3-0.7 mm wide and 0.20.4 mm deep, oblong. Ovary 1.0-1.5 mm tall, sparsely sericeous apically, the
anteriorstyle straightand erect, 2.4-3.4 mmlong, the posteriorstyles lyrate, much
longer than the anteriorstyle, 3.0-4.2 mm long, slender, stigmas capitate. Fruit
unknown.
Type. G. T. Prance & J. F. Ramos 6960, Brazil, Rondonia, vicinity of Santa
Barbara, 15 km E of km 117 on Porto Velho to Cuiaba highway, fl (holotype,

INPA 23719;isotypes, MICH, NY).
Distribution.(Fig. 33). In forest on terrafirme on both sides of the Amazon
basin, to the northin the foothills of the GuayanaHighlandin Bolivar, Venezuela
and in northernPara, Brazil, and to the south in Rondonia, Brazil.
Collected in flower in February,March,June, August, November and December. Not known in fruit.
VENEZUELA. Bolivar: Blanco 676 fl (MY); 715 fl (NY); LI. Williams 11627 bud (F, US, VEN).
BRAZIL. Para: Oliveira 3501 fl (IAN, MICH, NY); N. T. Silva 2859 fl (IAN). Rond6nia: Black 5214867 fl (IAN); Prance & Ramos 6960 fl (INPA, MICH, NY).
This species is named for its lyrate posterior styles, which are unusualin this
group where the styles are usually subequal or the posterior styles are shorter
than the anteriorstyle, or even lacking. The Black collection from Rondoniais
unusualfor its longer petioles and largerbracts and bracteoles.
The Banisteriopsis parviflora group (species 60-64)
The five species in this group occur only in the Atlantic coastal forest of SE
Brazil. They share the following characters: vining habit; stipitate glands on
the leaf marginnear the base or on the basal lateralveins; the flowers are borne
in four-floweredumbels which are borne in racemes; the bracts and bracteoles
are broadly triangularand papery;the flowers are small, and the posterior styles
are shorterand more slenderthan the anteriorstyle, directedposteriorly,parallel
throughouttheir length and often connate apically.
60. Banisteriopsisparviflora(Adr. Jussieu) B. Gates, comb. nov.
BanisteriaparvifloraAdr. Jussieu, Ann. Sci. Nat. Bot., S6r. 2, 13: 284. 1840.
?Banisteriaviridis Niedenzu, Verz. Vorles. Ak. Braunsberg:14. 1912.Type. Zietz s.n., Brazil,
Sao Paulo, near Sao Paulo, fr (holotype, B destroyed, n.v.) ex descriptione.
Liana, young branchesterete, smooth, glabrousor with sparse appressedhairs

or sparsebasifixedhairs, old branchesglabrouswith paperypeelingbark. Stipules
up to 1 mm long, triangular.Leaves opposite or ternate, with the petiole 8-18
mm long, glabrous, eglandular,the lamina 3.6-16.8 cm long, 1.3-6.4 cm wide,
elliptic to narrowlyelliptic, cuneate to obtuse at the base, acute to acuminateat
the apex, with the marginslightly revolute and bearinga stipitategland on either
side of the midribnear the base, glabrouson both sides or more rarely sparsely
sericeous abaxially, with the reticulationprominenton both sides. Inflorescence
157
axillary, of "racemes" of 4-floweredumbels, glabrous or with a few appressed

hairs, the inflorescenceleaves reduced to linear bracts, the bracts and bracteoles
0.6-1.0 mm long, broadly triangular,papery, glabrous, involucrate, persistent;
pedicels sessile, 5-10 mm long, 0.4 mm in diameter,glabrous.Sepals 1.8-2.2 mm
long, 1.2-1.8 mm wide, broadlyelliptic to rotund, roundedat the apex, glabrous
on both sides, eglandularor the 4 lateral sepals biglandular,the glands 1.0-1.3
mm long, 0.9-1.0 mm wide, elliptic to round. Petals white, becomingpale yellow
in age, short-fimbriate,glabrous, the 4 lateralpetals reflexed between the sepals,
the claw 1.8-2.2 mm long, the limb 3.0-4.5 mm long, 2.5-4.5 mm wide, the
antero-lateralpetals orbicular,cucullate, the postero-lateralpetals broadly elliptic, plane, somewhat smaller than the antero-lateralpetals, the posterior petal
withthe claw erect, 2.5-3.0 mmlong, constrictedapically,the limb3.0-3.5 mmlong,
2.5-3.0 mm wide, broadly elliptic, plane. Stamens with the filaments glabrous,
those opposite the sepals 1.8-3.0 mm long, those opposite the petals 1.3-1.8 mm
long, subequal;anthers with the locules 0.6-1.2 mm long, glabrous, the connectives opposite the 3 anteriorsepals 0.8-1.1 mm long, 0.6-1.2 mm wide, 0.7-0.8
mm deep, tumid, glandular,obovate, projectingup to 0.6 mm beyond the locules,
those opposite the petals and the postero-lateralsepals 0.5-0.7 mm long, 0.4-0.6
mm wide, 0.2-0.4 mm deep, broadly elliptic, not glandularor sometimes those
opposite the antero-lateralpetals with a glandulararea in the middle, not projecting beyond the locules. Ovary 1.0-1.2 mm tall, sericeous, the anteriorstyle
2.6-2.8 mm long, 0.2-0.3 mm in diameter,directedanteriorly,the posteriorstyles
2.0-2.4 mm long, more slenderthan the anteriorstyle, paralleland almost touching throughouttheirlength, directedposteriorly;stigmascapitate. Fruitwith short
broad carpophore, the nut 6-7 mm tall and 4 mm long, sparsely sericeous to
glabrate, bearingon each side a short winglet projectingup to 3 mm, the dorsal
wing 20-24 mm long, 8-9 mm wide, sparsely sericeous to glabrate,the hairs stiff,
sessile, the trabecula0.8-1.0 mm long.
Types. C. Gaudichaud s.n. (Herbier Imperial du Bresil 580), Brazil, fl (lecto-
type, P). Sellow IV it 5933, Brazil, Sao Paulo, fl (syntype, B destroyed, NY fragment ex B).
Distribution.(Fig. 35). In the Atlantic coastal forests of Parana,Sio Paulo and
Rio de Janeiro.
Collected in flower in Februaryand April to June and in fruit in April.
BRAZIL. Rio de Janeiro: A. Ab. 605 fl (HB); Dionisio & Otavio 270 fl (RB); A. F. Coimbra Filho
II fl (GUA, MICH, NY); Glaziou 10365 fl (BR, C, G, K, NY, P); Occhioni 4836 fl (HB, MICH,
RFA); 4891 fl (MICH, RFA); H. Velloso 432 fl (R); Wawra 39 fl (F, W). Sao Paulo: Constantino 7
st (RB); Handro s.n. fl (SP); Hoehne s.n. fl (GH, SP); s.n. fl (M, SP 3965); Sellow IV it 5933 fl (NY
ex B). Parana: Dusen 7915 fl (S); 8326 fl fr (G, S); Hatschbach 5543 (leg Stellfeld) fl (P); 6034 fl (HB
16927, MICH, US).
This species is most closely related to B. salicifolia from which it can be

distinguishedby it' glabrousleaves and inflorescence, and its fruits with lateral
winglets on the nut and a shorterdorsal wing.
I have included Banisteria viridis in the synonymy for this species as the descriptionfits this species very well: the leaves are quite glabrous,the flowers are
borne in umbels, the calyx is sometimes eglandular,and the fruit nut bears a
winglet on each side; only the size of the appendageon the nut is unus: -' ~ -his
species.
61. Banisteriopsissalicifolia(de Candolle)B. Gates, comb. nov.

Banisteria salicifolia de Candolle, Prodromus 1: 590. 1824 (non salicifolia sensu Jussieu et auctt.
seq.).
158
Flora Neotropica
BanisteriaparvifloraAdr. Jussieuvar. tomentosaNiedenzu, Verz. Vorles. Konig. Ak. Braunsb.:
17. 1912.Type. Schwacke10433,Brazil, MinasGerais, Serrade OuroPreto, fl fr (holotype,
B destroyed,fragmentNY).
Liana, branchesdensely tomento-sericeous.Stipulesup to 1 mm long, tomentosericeous. Leaves subopposite, opposite or ternate, the petiole 10-16 mm long,
tomento-sericeous, eglandular,the lamina 5.3-10.0 cm long, 2.5-4.1 cm wide,
elliptic to ovate, the base cuneate to obtuse, acute to short-acuminateor rarely
apiculateat the apex, with the marginslightlyrevolute, bearinga pair of stipitate
glands beside the midribat the base, glabrous adaxially, densely tomento-sericeous abaxially, with the reticulationprominentadaxially, and the lateral veins
prominent abaxially. Inflorescence of axillary racemes of 4-flowered umbels,
densely tomento-sericeous; bracts and bracteoles up to 1.0 mm long, broadly
triangular,glabrous, papery, involucrate, persistent; pedicels sessile, 8-9 mm
long, 0.4-0.5 mm in diameter, sparsely tomento-sericeous. Sepals 1.6-2.0 mm
long, 1.5-1.8 mm wide, broadly elliptic, rounded at the apex, glabrous on both
sides or sparselysericeous abaxially,eglandularor the 4 lateralsepals biglandular,
projecting 0.6-1.0 mm beyond the glands, the glands 1.0-1.2 mm long and 0.8
mm wide. Petals probably white, becoming pale yellow in age, denticulate, glabrous, the 4 lateralpetals reflexedbetween the sepals, the claw 1.2-2.0 mm long,
the limb 3.5-5.0 mm long and 2.5-5.0 mm wide, the antero-lateralpetals orbicular
and cucullate, the postero-lateralpetals broadly elliptic, plane, slightly smaller
than the antero-lateralpetals, the posterior petal with the claw erect, up to 2.5
mm long, constricted at the apex, the limb up to 3.5 mm long and 2.5 mm wide,
obovate, the basal teeth gland-tipped.Stamens with the filamentsopposite the 3
anteriorsepals 1.8-2.6 mm long, those of the other 7 stamens subequal, 1.2-1.6
mm long; antherswith the locules 0.7-1.2 mm long, glabrous, the connectives of
the stamens opposite the 3 anteriorsepals 1.0-1.3 mm long, 0.9-1.0 mm wide,
0.6-0.8 mm deep, enlarged and glandular,projectingup to 0.8 mm beyond the
locules, those opposite the petals and postero-lateralsepals 0.6-0.9 mm long,
0.5-0.6 mm wide, 0.2-0.4 mm deep, broadly elliptic, not projectingbeyond the
locules. Ovary 1.2-1.4 mm tall, sericeous; anteriorstyle 2.4-2.6 mm long, erect,
posteriorstyles 1.6-1.8 mm long, straightand parallelthroughouttheirlengthand
joined apically, directed posteriorly, stigmas capitate. Fruit with broad carpophore up to 3 mm long, the nut 8-9 mm tall and 5 mm long, smooth, sparsely
appressed-sericeous,the wing up to 40 mm long and 16 mm wide, sparsely appressed-sericeous, the hairs golden, the trabeculastiff and straight, 1.0-1.5 mm
long, with a shallow appendageat the base up to 1.5 mm tall.
Type. Herbariumde Candolle, Prodromus1, 590, fl (holotype, G-DC.).
Distribution.(Fig. 35). In forests in the vicinity of OuroPreto in MinasGerais.
Collected in fruit in August.
BRAZIL. Minas Gerais: Magalhdes Gomes 2632 fr (RB); Schwacke 10433 fl fr (NY ex B).
This species has been very rarely collected, and it is possible that it is now
extinct due to the destructionof the forests near Ouro Preto. It is readily distinguished from Banisteriopsisparviflora by its tomento-sericeousleaves and inflorescence, and its fruit with smooth nut and large fruit wing.
62. Banisteriopsisscutellata(Grisebach)B. Gates, comb. nov.
Banisteriascutellata Grisebach,Vidensk. Meddel. Dansk. Naturhist.Foren. Kjobenhavn:131.
1875.
Banisteria scutellata f. obtusa Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1900. Type. Glaziou
159
16362, Brazil, Rio de Janeiro, Ayuruocato Pico do Papageio, fl (holotype, B destroyed?;

isotypes, C, K, P).
Liana, branches densely velutinous. Stipules up to 1 mm long, broadly triangular, sparsely sericeous. Leaves with the petiole 18-38 mm long, densely goldenvelutinous, eglandular,the lamina 4.2-12.7 cm long, 1.9-7.3 cm wide, ovate,
truncate to cordate at the base, short- to long-acuminateat the apex, with the
marginslightly revolute, bearing 1(-2) stipitate gland(s) on the basal lateral vein
on each side near the margin,adaxiallyvelutinous, abaxiallytomento-velutinous,
with the reticulationprominentadaxially and the lateral veins prominentabaxially. Inflorescenceaxillary,of 4-floweredumbels, bornein shortaxillaryracemes,
inflorescenceleaves usually reducedto linearbracts, common peduncle 4-8 mm,
velutinous; bracts and bracteoles broadly triangular1.2-2.0 mm long, glabrous,
papery, involucrate, persistent; pedicels sessile 3-7 mm long, 0.2-0.3 mm in
diameter,sparselyvelutinous. Sepals 1.6-2.0 mm long, 1.6-1.8 mm wide, broadly
elliptic, roundedat the apex, sparsely sericeous abaxially,glabrousadaxially,the
4 lateral sepals biglandularand projecting 0.8-1.2 mm beyond the glands, the
glands 1.0-1.2 mm long, 0.8-1.0 mm wide, broadly elliptic to orbicular.Petals
white?, becoming pale yellow in age?, fimbriate,eglandular,the 4 lateral petals
reflexedbetween the sepals, the claw 1.5-2.0 mm long, the limb 3.5-5.0 mm long
and wide, orbicular,the antero-lateralpetals largerthan the postero-lateralpetals
and strongly cucullate, the postero-lateralpetals a little concave, the posterior
petal with the claw erect, 2.5 mm long and up to 1.0 mm wide, constricted
apically, the limb up to 4.0 mm long and wide, orbicular. Stamens with the
filaments opposite the 3 anteriorsepals 2.2-2.8 mm long, the other 7 subequal,
1.2-1.8 mm long; anthers with the locules 0.4-1.0 mm long, sometimes very
reduced or absent on the stamens opposite the antero-lateralsepals, glabrous,
the connectives of the stamens opposite the antero-lateralsepals 1.0-1.2 mm
long, 1.0-1.2 mm wide, 0.6 mm deep, glandular,obovate, projectingup to 0.6
mm beyond the locules, the connectives opposite the petals and the posterolateralsepals 0.4-1.0 mm long, 0.4-0.6 mm wide, 0.3-0.4 mm deep, not glandular,
elliptic, not projecting beyond the locules. Ovary 1.0 mm tall, sericeous, the
anterior style erect, 2.6 mm long, the posterior styles directed posteriorly, 2.2
mm long, parallelthroughouttheir length and fused at the apex, stigmascapitate.
Fruitwith shortbroadcarpophore,the nut 6-8 mm tall and 4-5 mm long, smooth,
appressed-sericeousto glabrate, the hairs golden with straightstiff trabeculaup
to 2 mm long, sometimes with a short lateralwinglet or crest, the wing 23-30 mm
long, 9-13 mm wide, glabrous, with a triangularappendageat the base up to 2.5
mm tall.
Type. Glaziou 2512, Brazil, Rio de Janeiro, Tijuca, fl (hblotype, GOET, fragment; isotypes, BR, C, P).
Distribution.(Fig. 35). Known only from the Atlantic coastal forest in Rio de
Janeiro, Brazil.
Collected in flower from Januaryto March, and in fruit in March.
BRAZIL. Rio de Janeiro:Brade 11282 fl (R); Gates s.n. yng fr (MICH); Glaziou 2512 fl (BR,
GOET,P); 8583 fl (K, P); 10370fl (C, G, K, NY, P); 11812fl (G, K, P, R); 16362fl (C, K, P); Martins
121 fl (GUA, MICH, NY); Peckolt & Freire 482 fl (R); Pereira 4266 fl (HB, RFA, MICH); Ule 127
fl fr (R); Vidal 11-304 fl (R).
This distinctive species has few modern collections, perhaps because it is inconspicuous in flower and apparentlyproducesfew fruits. It is distinguishedfrom
Banisteriopsis basifixa by its velutinous pubescence throughout, and sparsely
pubescent pedicels.
Flora Neotropica
160
J*t
3C%E..,
.--%.3
0
0.
5., o
i4'
40
4
&U
' 35
FIG. 35. Distribution of Banisteriopsis parviflora group, species 60-64, and B. pseudojanusia and
B. magdalenensis (southeastern Brazil). Open circle, B. parviflora; closed circle, B. salicifolia; closed
square, B. scutellata; open square, B. basifixa; closed triangle, B. parviglandula; star, B. pseudojanusia; open triangle, B. magdalenensis.
63. Banisteriopsis basifixa B. Gates, sp. nov.

Liana, rami sparsim hispidi, pilis Y-formibus, interdum uno brachiorum brevissimo vel nullo, usque 2.5 mm longis, vetustiores glabrati, grisei. Stipulae usque
1 mm longae, triangulares. Folia petiolo 10-20(-35) mm longo, dense aureo-hispido-velutino, eglanduloso vel rarius 2 glandes parte superiore gerenti, lamina
3.6-15.0 cm longa, 1.8-6.9 cm lata, ovata vel rarius juniore lobata, basi cordata,
apice acuta vel acuminata, margine parum revoluta, venis lateralibus basi 1-2
glandes stipitatas gerenti, adaxialiter hispida, pilis Y-formibus, brachiis 0.3-0.5
mm longis, abaxialiter hispido-tomentosa, pilis T-formibus, trabecula 1.0-1.5 mm
longa, pede 0.2-0.3 mm, venis venulisque prominentibus adaxialiter, venis lateralibus prominentibus abaxialiter. Inflorescentia axillaris, floribus in umbellis
4-floriferis fasciculatis foliis redactis usque ad bracteas lineares subtentis, umbellis in racemis dispositis, sparsim aureo-velutina, pedunculo umbellae 2-3 mm longo, glabro; bracteis bracteolisque 0.8-1.0 mm longis, late triangularibus, glabris,
membranaceis, persistentibus, involucratis; pedicellis sessilibus, 4-9 mm longis,
0.3 mm diametro, glabris. Sepala 5, 1.8-2.0 mm longa, 1.4-1.8 mm lata, late
elliptica, apice rotundata, abaxialiter sparsim sericea, adaxialiter glabra, eglandulosa vel 4 lateralia 2 glandes 1.0-1.4 mm longas et 0.6-1.0 mm latas gerentia,
glandes 0.7-1.2 mm superantia. Petala 5, orbicularia, eglandulosa, fimbriata, luteola?, 4 lateralia inter sepala patentia, ungue 1.5-2.0 mm longo, limbo 3.0-4.0
mm longo et lato, antero-lateralia cochleata et quam postero-lateralia majoria,
postero-lateralia concava, petalum posticum ungue erecto usque 2.5 mm longo,
apice constricto, limbo 3.0 mm longo latoque. Stamina filamentis 3 sepalis anticis
161
oppositis 2.0-2.7 mm longis, ceteris 0.8-1.8 mm longis; antheraeloculis petalis

oppositis 0.6-0.8 mm longis, sepalo antico opposito 1.2-1.3 mm longo, sepalis
antero-lateralibusoppositis redactis vel nullis, sepalis postero-lateralibusoppositis 0.3-0.4 mm longis, glabris, connectivis 3 sepalis anticis oppositis 0.9-1.2 mm
longis, 0.8-1.0 mm latis, 0.8 mm profundis,glandulosis, obovatis, loculos superantibus, ceteris 0.5-0.9 mm longis, 0.3-0.7 mm latis, 0.2-0.4 mm profundis, ellipticis, illis sepalis postero-lateralibusoppositis loculos superantibus,illis petalis
oppositis loculos non-superantibus.Ovarium0.8-1.0 mm altum, carpellis3, uniovulatis, liberis, toro pyramidaliadnatis, sericeis; styli 3, anticus usque 2.6 mm
longus, erectus, postici 1.8-2.0 mm longi, recti parallelique,liberi, quamanticum
graciliores, stigmatibuscapitatis. Fructusex 3 samaris(vel abortu 1-2) constans;
samaracarpophorousque 2 mm longo et 1 mm lato, nuce ovoidea 5-7 mm alta,
3-5 mm longa, sparsim sericea, rugulosa, ala 18-26 mm longa, 12-15 mm lata,
sparsim sericea, pilis trabecularecta 1.0-2.0 mm longa, basi appendicularotundata usque 2 mm alta munita.
Type. Hoehne & A. Gehrt 17711, Brazil, Sao Paulo, Sao Jose do Barreiro,fl
(holotype, SP 17711;isotype, GH).
Distribution. (Fig. 35). In the Atlantic coastal forest of Rio de Janeiro and
adjacent Sao Paulo and Minas Gerais, Brazil.
Collected in flower from Marchto May and in July, and in fruit in June.
BRAZIL. Minas Gerais: Riedel & Langsdorff 355 pp fl (US). Rio de Janeiro: Anderson 11201 st
(MICH); Glaziou 2512 pp fl (R57963); 3894 fl (C, P); 11811 fl (C, K, P); Goes & Dionisio 368 fl
(RB 138860); 378 fl (RB 138858); 662 fr (RB51463). Sao Paulo: Hoehne & Gehrt 17711 fl (GH, SP17711).
This species is most closely related to Banisteriopsisscutellata, from which it

can be distinguishedby its hispid stem pubescence with basifixed hairs, its glabrous pedicels, strongly cordate leaf base, and the fruit with shorter broader
wing. The two species appearto be sympatricfor most of their range. There are
very few modern collections of either species, but it appears that there is a
separation between the species in their time of flowering: this species flowers
from Marchto July, and B. scutellata from Januaryto March.
The name refers to the basifixedhairs on the stems.
64. BanisteriopsisparviglandulaB. Gates, sp. nov.
Liana, ramiminutegriseo-velutini,pilis basifixis2.0-3.5 mmlongis interspersis,
vetustiores glabrati. Stipulae minutae. Folia opposita vel ternata, petiolo 3-12
mm longo, aureo-velutino, pilis T-formibustrabecula 1.0-1.5 mm longa interspersis, eglanduloso, lamina 3.1-9.7 cm longa, 0.9-3.2 cm lata, lanceolata, basi
cordata, apice acuta, marginerevoluta, venis lateralibusprope costam par glandium stipitatarumgerenti, adaxialiterpilis Y-formibus vel basifixis, abaxialiter
appresso-sericea,trabecula0.6-1.5 mm longa, adaxialiterplano, abaxialitervenis
prominentibus.Inflorescentiaaxillaris,floribusin umbellis4-floriferisfasciculatis
foliis usque ad bracteas lineares redactis subtentis, umbellis in racemis brevibus
vel dichasiis dispositis, aureo-appresso-sericea;bracteis bracteolisque 0.8-1.0
mmlongis, late triangularibus,glabris,membranaceis,involucratis,persistentibus;
pedicellis sessilibus 6-8 mm longis, 0.3 mm diametro,glabris. Sepala 5, 1.4-1.5
mm longa lataque, orbicularia, apice rotundata, utrinque glabra, 4 lateralia 2
glandes 0.4-0.6 mm longas et latas gerentia, glandes 0.9 mm superantia.Petala
5, lutea, orbicularia,fimbriata,eglandulosa,4 lateraliainter sepalapatentia,ungue
1.5-2.5 mm longo, limbo 3.0-3.5 mm longo latoque, antero-lateraliacochleata,
postero-lateraliaplana;petalumposticum ungue erecto 2.5 mm longo, apice constricto, limbo 3.0-3.5 mm longo latoque. Staminafilamentis3 sepalis anticis op-
162
Flora Neotropica
positis 2.4-2.6 mm longis, ceteris 1.3-1.8 mm longis; antheraeloculis 0.6-1.0 mm

longis, glabris, connectivis 3 sepalis anticis oppositis 1.0-1.1 mm longis, 0.9-1.3
mm latis, 0.6-0.9 mm profundis,obovatis, glandulosis, loculos superantibus,ceteris 0.5-0.7 mmlongis, 0.4-0.6 mmlatis, 0.2-0.3 mmprofundis,ellipticis, loculos
non superantibus.Ovarium 1.0-1.2 mm altum, carpellis 3, uniovulatis, liberis,
toro pyramidali,adnatis, sericeis; styli 3, anticus 2.8-3.0 mm longus, erectus,
postici 2.2-2.5 mm longi, recti parallelique,apice connativel liberi, quamanticum
graciliores, stigmatibuscapitatis. Fructus non suppetunt.
Type. Vasco Gomes 1215, Brazil, MinasGerais, Esta9ao Experimentaldo Cafe
Coronel Pacheco, fl (holotype, RB) = E. P. Heringer 1215 (isotype, SP).
Distribution.(Fig. 35). In woodland in SE Minas Gerais in Brazil, apparently

extending south into the state of Rio de Janeiro.
Collected in flower from April to June.
BRAZIL. Minas Gerais: V. Gomes 1215 fl (RB); Heringer 1215 fl (SP); J. G. Kuhlmann 2300 fl
(RB); H. Magalhaes 1190 fl (R); Mexia 4630 bud (F, G, GH, MO, US). Rio de Janeiro: Anderson
11727 st (MBM, MICH).
This species is most closely related to Banisteriopsis scutellata and B. basifixa,
from which it can be distinguishedby its short petioles, smallerlanceolate leaves

with sericeous pubescence, shorterinflorescencesof 3-5 umbels, and small calyx
glands. In leaf size, shape and pubescence, B. parviglandularesembles B. pseudojanusia.
The name refers to the very small calyx glands.
65. Banisteriopsispseudojaunsia(Niedenzu) B. Gates, comb. nov.

Banisteria pseudojanusia Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 408. 1928.
Liana, branchestomento-sericeouswith mixed, sessile T- and Y-shapedhairs,

the arms unequal. Stipules up to 1.5 mm long. Leaves with the petiole 7-11 mm
long, golden-tomentose, biglandular,the lamina 2.2-7.5 cm long, 0.8-2.2 cm
wide, lanceolate, truncate at the base, acute to apiculate at the apex, with the
margin revolute, adaxially sparsely appressed-sericeous, abaxially sericeous,
smooth adaxially and with the lateral veins prominentadaxially. Inflorescence
axillary, of single 4-floweredumbels, appressed-sericeous,the commonpeduncle
bibracteate, up to 4 cm long; bracts and bracteoles 0.8-1.5 mm long, broadly
triangular,papery, glabrous, involucrate, persistent;pedicels sessile, 12-15 mm
long, 0.4-0.5 mm in diameter, glabrous. Sepals 1.8-3.0 mm long, 1.8-2.5 mm
wide, broadly elliptic, roundedat the apex, glabrouson both sides, the 4 lateral
sepals biglandular,projecting0.6-1.4 mm beyond the glands, the glands 1.2-2.6
mm long, 0.9-1.3 mm wide, elliptic. Petals yellow, glabrous, fimbriate, eglandular, orbicular,the 4 lateralpetals with the claw 0.8-2.0 mm long, the limb 4.07.5 mm long, 4.0-7.0 mm wide, the antero-lateralpetals largerand more concave
than the postero-lateralpetals, the posterior petal with the claw erect, 2.2-3.0
mm long, 0.8-1.2 mm in diameter, constricted at the apex, the limb 4.0-6.0 mm
long and wide. Stamens 10, or sometimes 9 by abortion, with the filaments 1.63.2 mm long, those opposite the sepals somewhat longer than those opposite the
petals; anthers with the locules 0.8-1.2 mm long, glabrous, the connectives 0.71.0 mm long, 0.5-0.7 mm wide, 0.2-0.6 mm deep, elliptic, not overtoppingthe
locules. Ovary0.8-1.4 mmtall, sparselysericeousto glabrous,the styles subequal,
2.0-2.4 mm long, divergingapically, stigmas capitate. Fruit unknown.
Types. Dusen 9079, Brazil, Parana, Turma, fl (lectotype, S). Dusen s.n. (17
Feb 1911), Brazil, Parana,Desvio Ribas, fl (syntype, S).
163
Distribution.(Fig. 35). In woods in southernParanaand Santa Catarina,Brazil.

Collected in flower in Januaryand February.
BRAZIL. Parana: Dusen 9079 fl (S); s.n. fl (S). Santa Catarina: Reitz & Klein 12424 fl (P).
This species resembles Banisteriopsisparviglandula in the size and shape of

its leaves, but can be readily distinguishedby its inflorescencesof single umbels,
and its much largerflowers with large calyx glands, and stamens and styles subequal. It resembles Janusia superficiallyin the form of its leaves and its inflorescences of single axillaryumbels.
66. BanisteriopsismagdalenensisB. Gates, sp. nov.
Frutex?, rami juniores complanati, appresso-sericei, vetustiores teretes, glabrati. Stipulae minutae, triangulares,glabrae. Folia petiolo usque 3 mm longo,
sparsimappresso-sericeo,eglanduloso,lamina2.7-10.0 cm longa, 1.8-5.0 cm lata,
ovata ellipticave rotundave, basi cordata vel amplexicauli,apice mucronatavel
emarginata,margineparumrevoluta, 2 glandes sessiles usque 1.5 mm diametro
ad costam basi gerenti, adaxialitersparsim appresso-sericea, abaxialiterdense
appresso-sericeavel rariusglabrata,venis venulisque prominentibusadaxialiter,
venis lateralibusprominentibusabaxialiter.Inflorescentiaaxillaris,floribusin umbellis 4-floriferisfasciculatis, umbellis in dichasiis dispositis, ramis complanatis
sparsim appresso-sericeis, bracteis bracteolisque 1.0-1.4 mm longis, late triangularibus, glabris, membranaceis,involucratis, persistentibus;pedicellis sessilibus 19-22 mm longis, 0.5-0.6 mm diametro,glabris. Sepala 5, 1.8-2.2 mm longa
lataque, late depresso-ovata,apice rotundata,sparsimappresso-sericeaabaxialiter, glabraadaxialiter,4 lateralia2 glandes 1.0-2.0 mm longas et 0.7-2.0 mm latas
gerentia. Petala 5, lutea, dentata, 4 lateraliainter sepala patentia, antero-lateralia
ungue 2.5 mm longo, limbo 11.0-12.0 mm longo lataque, concavo, orbiculari,
postero-lateraliaungue 1.5-2.0 mm longo, limbo 10.5-11.0 mm longo, 9.5-11.0
mm lato, late elliptico, plano, petalum posticum ungue erecto, usque 4.5 mm
longo, 0.6 mm lato, limbo 8.0-8.5 mm longo latoque, late obovato. Staminafilamentisglabris, illis 3 sepalis anticis oppositis 3.0-3.4 mm longis, ceteris 1.6-2.4
mm longis, basi connatisusque 1 mm;antheraeloculis 0.7-1.1 mm longis, glabris,
connectivis 3 sepalis anticis oppositis 1.0 mm longis, 0.7-1.0 mm latis, 0.7-1.1
mm profundis, obovatis, glandulosis, tumidis, ceteris 0.6-0.8 mm longis, 0.5-0.7
mm latis, 0.3-0.5 mm profundis, ellipticis, eglandulosis. Ovariumusque 1.4 mm
altum, setosum, carpellis 3, univolatis, liberis, toro pyramidaliadnatis. Styli 3,
erecti paralleliquevel divergentes ad apicem, stylo antico 2.2-2.4 mm longo,
0.25-0.30 mm diametro,stylis posticis 2.0-2.2 mm longis, 0.2 mm diametro,basi
setosis, stigmatibuscapitatis. Fructus ignotus.
Type. Santos Lima 350, Brazil, Rio de Janeiro, Santa MariaMagdalena,Pedra
de Abreusinho,fl (holotype, RB 28982).
Distribution.(Fig. 35). Known only from 3 collections in the vicinity of Santa
MariaMagdalenaat elevations of 1200-1700m.
Collected in flower in February,Marchand May.
BRAZIL. Rio de Janeiro: S. Lima 350 fl (RB 28982); S. Lima & Brade 13281 fl (RB 26260); 14239
bud (RB 26439).
This species shares several characterswith Banisteriopsisparviflora, such as

its papery bracts and bracteoles, enlarged connectives on the stamens opposite
the three anterior sepals, and its prominentleaf reticulation.In many other respects, it is quite distinct: it is apparentlya shrub;the leaves are almost sessile,
164
Flora Neotropica
with large basal glands; the flowers are large, with erect subequal styles; the
umbels are borne in dichasia. Santos Lima & Brade 14239 differsfrom the other
2 collections in that its leaves are quite glabrousabaxially. In all other respects
it is very similarto the other collections. It was apparentlycollected at a higher
altitude and perhaps the absence of hairs is correlatedwith this, or perhaps the
presence or absence of leaf pubescence in this species is a very variablecharacter.
Banisteriopsissubgenus 3. Pleiopterys(Niedenzu) B. Gates, comb. nov., species
67-92.
Banisteria subgenusPleiopterys Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1901.
Leaf glands marginalor on the lamina, or a single pair on the petiole, sessile
or rarely stipitate. Flowers borne in condensed axillaryracemes or 4-6-flowered
umbels, the umbels sometimes borne in cymes. Bracts and bracteoles persistent,
pedicels sessile, often thickerin fruit. Calyx 8-glandularor eglandular,the glands
borne on the receptacle below the free part of the sepal. Petals yellow, entire to
dentate to lacerate. Stamens with glandularconnectives, the glandulartissue in
the form of projectingconvex cells or papillae. Styles equal or the posteriorstyles
differentfrom the anteriorstyle, hairyat least at the base and for up to 2 of their
length. Ovary bearinglong hairs. Fruit nut ca. orbicular,smooth or on each side
with several ridges radiatingfrom the areole and terminatingin crests or winglets,
or with 1-3 winglets parallel to the areole, bearing a prominent appendage on its upper edge, nut locule glabrouswithin.
Lectotype species. Banisteriopsis pubipetala (Adr. Jussieu) Cuatrecasas as
Banisteria pubipetala.
Banisteriopsis subgenus 3. Pleiopterys section 1. Sciurostylis (Skottsberg) B.

Gates, comb. nov., species 67-73.
Banisteria subgenus Sciurostylis Skottsberg,Kongl. Svenska Vetenskapsakad.Handl. 35: 15.
1901.
Banisteria subsection Psilopetalum Niedenzu, Ind. Lect. Lyc. Brunsb.: 16. 1901. Lectotype
species. Banisteriopsis lutea (Grisebach) Cuatrecasas as Banisteria nitrosiodora.
Leaves with 1 (rarely2) pairs of protuberantglands on the petiole, and sometimes with 1-2 stalked glands on the marginof the lamina near the base. Inflorescence axillaryor terminal,of 4-floweredumbels, the umbels often arrangedin
cymes or dischasia. Pedicels slender, not thickeningin fruit. Sepals eglandular,
often reflexed and somewhat enlargedin fruit. Petals glabrous(except B. valvata
and B. heterostyla). Styles slenderand sigmoid, densely hairyfor up to half their
length (except B. valvata). Samaraswith carpophoreup to 2 mm long, probably
functional;hairs on samarastiff, sessile, irritatingto the skin, the trabecula 1-3
mm long.
Type species. Banisteriopsis lutea (Grisebach) Cuatrecasas as Banisteria hassleriana Chodat.
67. Banisteriopsisvalvata Anderson & Gates, Contr. Univ. Mich. Herb. 11: 51.
1975.
Fig. 36.
Liana, young branches flattened, brown-sericeous, older branches terete, sericeous to glabrate. Stipules up to 1.0 mm long, often dissected at the apex and
joined by an interpetiolarline. Leaves with the petiole 10-37 mm long, sericeous
165
--"
e~~~~~~~''';.r
FIG. 36. Banisteriopsisvalvata. a, habit and inflorescence,with largervegetative leaf, x 0.5; b,

flowerbud x 2.5; c, flower, with posterior(flag)petal to right, x 1.5; d, androeciumand gynoecium,
posteriorview with shortest stamen(that opposite the posteriorpetal) in foreground,x 5; e, androecium and gynoecium, lateral view, x 5; f, gynoecium x 5; g, young fruits enclosed by accrescent
sepals x 0.5; h, immaturefruit, lateralview, x 1.5. (Drawnby KarinDouthitfrom Andersonet al.
36981.)
166
Flora Neotropica
to glabrate, bearing 2 stalked or sessile protuberantglands near the apex adaxially, the lamina5.0-15.0 cm long, 4.0-14.5 cm wide, smallerin the inflorescence,
orbicularto broadly ovate, truncate to subcordate at the base, obtusely shortacuminateat the apex, with the marginplane, eglandularor bearing 1-2 sessile
or stipitate glands near the base, pubescent and soon glabrate except on the
nerves adaxially, tomentose abaxially, the hairs white, T-shaped, the trabecula
straight or flexuous 0.7-0.9 mm long, the stalk 0.1 mm long, with the venation
obscure adaxially and prominentabaxially with the lateral veins connected by
parallelsecondaryveins. Inflorescenceaxillary,paniculate,of 4-floweredumbels,
the branches of the inflorescence bearing a single umbel or a dichasium of 3
umbels, brown-sericeous;bracts and bracteoles 1.5-2.0 mm long, triangularor
ovate, abaxially sericeous, involucrate,persistent;pedicels sessile or subsessile,
9-19 mm long, 0.8-1.0 mm in diameter, brown-sericeous. Sepals 6.5-8.0 mm
long, 5.5-6.5 mm wide, connate basally for 2-3 mm, eglandular,broadly ovate,
cordate or subcordateat the base, obtuse at the apex, exduplicate-valvate,abaxially brown-sericeous, adaxially near the marginminutely white-pubescent, accrescent in fruit, up to 16 mm long and 12 mm wide in young fruit. Petals yellow,
laciniate, eglandular,abaxially sparsely white-sericeous, the 4 lateral petals reflexed between the sepals, the claw 4.0-5.0 mm long, 0.5-0.6 mm wide, the limb
10.0-14.0 mm long and wide, orbicular,plane, the antero-lateralpetals somewhat
larger than the postero-lateralpair, the posterior petal with the claw erect, 4.04.5 mm long, 1.1-1.7 mm wide, constricted at the apex, the limb 8.0-10.0 mm
long, 6.0-9.0 mm wide, broadly elliptic, obovate or orbicular, often wrinkled.
Stamens with the filamentsconnate at the base for 0.3-0.7 mm, that opposite the
anteriorsepal 4.5-5.5 mm long, that opposite the posteriorpetal 2.0-3.0 mm long,
the 8 lateral filaments 3.0-4.0 mm long; anthers subequal, reflexed, the locules
1.0-1.5 mm long, glabrous,the connectives swollen, papillate,projecting0.4-0.8
mm beyond the locules, those of the stamens opposite the sepals 1.2-1.4 mm
long, 1.0-1.2 mm wide, 0.6-0.8 mm deep, obovate, those opposite the petals 1.01.4 mm long, 0.8-1.0 mm wide, 0.4-0.6 mm deep, oblong. Ovary 1.5 mm tall,
velutinous; styles straightand diverging, subequal 4.0-5.0 mm long, pubescent
at the base, the hairs white or brown, straightor flexuous, the armsvery unequal,
the longerarm 1-2 mm long, stigmascapitate.Immaturefruitwithoutcarpophore,
densely appressed-sericeous,with stiff sub-basifixedhairs 1.5 mm long and shorter medifixed hairs, nut up to 1 cm tall and 1 cm long, bearing 2-3 winglets on
each side parallelto the areole, up to 16 mm along the base and projectingup to
8 mm from the nut, sometimes confluentbeneath the nut, the wing up to 20 mm
long and 9 mm wide, bearinga triangularappendageon its upperedge at the base
5 mm tall and 5 mm long.
Type. Anderson, Stieber & Kirkbride 36981, Brazil, Bahia, ca. 13 km S of
Cocos and 3 km S of Rio Itaquari,fly yng fr (holotype, UB; isotypes, F, IAN,

K, MICH, MO, NY, P, RB, SP, U, US).
Distribution.(Fig. 37). In scrubby "caatinga"thickets on the eastern slopes of
the Espigao Mestre in Bahia, extendingnorthwardinto Maranhao.
Collected in flower in Marchand April and in young fruit in March.
BRAZIL. Maranhao: Eiten & Eiten 4457 fl (NY). Bahia: Anderson et al. 36981 fl yng fr (F, IAN,
K, MICH, MO, NY, P, RB, SP, U, UB, US); Hatschbach 42115 fl (MICH).
The large valvate calyx of this species is unique in the genus; in its fruit and
leaf glands, B. valvata resembles B. heterostyla, but the androeciumand gynoecium are like those of B. hypericifolia in section Anisopterys.
167
68. Banisteriopsis heterostyla (Adr. Jussieu) Cuatrecasas, Webbia 13: 491.

1958.2
Fig. 38.
Banisteria heterostyla Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840.
Banisteria jasminellum Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840. Type. Watts
s.n., Colombia, Bolivar, Cartagena, fl (holotype, K; isotype, P).
Liana, young branchestomentose, soon glabrate,barkred-brown,old branches

glabrouswith thick corky ridges. Stipules minute. Leaves with the petiole (3-)520 mm long, tomentose to glabrate,bearinga pair of protuberantglands near the
apex adaxially,the lamina1.9-9.5 cm long and 1.4-4.8 cm wide, ellipticto broadly
elliptic, cuneate to obtuse at the base, acute to apiculate at the apex, with the
marginmore or less revolute and sometimesbearinga pair of stalkedglands near
the base, adaxially glabrate, abaxially sparsely white-tomentose, with the veins
prominulous adaxially, prominent abaxially. Inflorescence on short axillary
branches bearing reduced leaves and terminatingwith 1-3 4-flowered umbels;
bracts and bracteoles 1.0-1.5 mm long, broadly triangular,tomentose abaxially,
involucrate,persistent;pedicels sessile, 12-20 mm long, 0.4-0.5 mm in diameter,
sparsely tomentose. Sepals 2.6-3.0 mm long and wide, orbicular,roundedat the
apex, eglandularor the 4 lateral sepals each bearing a pair of minute orbicular
glands at the base, abaxially tomentose, adaxially glabrous, reflexed and somewhat enlarged in fruit. Petals yellow, denticulate, plane, eglandular,abaxially
sparsely tomentose, the four lateral petals reflexed between the sepals, with the
claw 2.5-3.0 mm long, the limb 8.5-10.0 mm wide, orbicular,the posteriorpetal
with the claw erect, 3.5-4.2 mm long, 0.8-1.0 mm wide, the limb 7.0-8.0 mm long
and 6.0-6.5 mm wide, rotund, veined with red. Stamens with the filamentsslender, 2.0-5.2 mm long, those opposite the 3 styles (i.e., opposite the anteriorsepal
and postero-lateralpetals) longest, those opposite the antero-lateralsepals and
posterior petal shorterthan the others, those of the 3 posterior stamens inflexed
between the posterior styles, the other 7 erect; anthers with the locules 0.9-1.2
mm long, glabrous, diverging distally on the stamens opposite the sepals, the
connectives of the stamens opposite the sepals 0.9-1.2 mm long, 0.8-1.0 mm
wide, 0.5-0.7 mm deep, obovate, papillose apically, those opposite the petals
0.6-1.0 mm long, 0.4-0.6 mm wide, 0.3-0.4 mm deep, oblong. Ovary 1.0-1.5 mm
tall, minutely white-sericeous, styles slender, sigmoid and diverging from the
base, the anteriorstyle 4.8-5.5 mm long, hairy adaxiallyin the middle, the posterior styles 4.0-4.5 mm long, glabrous, stigmas truncate. Fruit with the carpophore up to 2 mm long and wide, the nut up to 6 mm tall and 4 mm long, sparsely
sericeous, bearingon each side 3 winglets up to 10 mm long and projectingup to
3 mm, parallelto the areole, sometimes confluentventrally, the dorsal wing 2035 mm long, 9-14 mm wide, sparsely sericeous, the hairs stiff and irritating,
sessile, the trabeculastraight,0.6-1.5 mm long, bearingon its upper edge at the
base an appendage4.5-8.0 mm tall and 5-8 mm along the base.
Type. Bertero2201, Colombia,Magdalena,Santa Marta,fl fr (holotype, P-JU;
isotypes, M, NY fragmentex B, P).
2
It was only after this treatment went to press that I was sent a specimen of Banisteriopsis
caduciflora with eglandular sepals. Such eglandular specimens would key out here. They can be
readily distinguished from Banisteriopsis heterostyla by their glabrous leaves with 3-5 pairs of glandular areas on the lamina.
Flora Neotropica
168
Distribution.(Fig. 37). In thickets on sandy hillsides in the arid coastal belt of

the Caribbeancoast of Colombia.
Collected in flower in Januaryto May and in fruit in Februaryto April.
COLOMBIA. Atlantico: Dugand 751 fl (F, NY); 1060 fl fr (F, US); 2320 st (US); 5025 fl (US);
5102 fl fr (NY, US); 6704 fl fr (US); Bro. Elias s.n. fl (NY); 387 fl (US); 492 fl (US); 697 fl (NY, US);
1198 fl (F, US); 1288 fl (F, G, US); 1442 fl fr (F, US); P!owman 3521 fl (GH). Magdalena: Funck 308
fl (G, P); Goudot s.n. fl (GOET); Haught 3996 fl (F, NY, US); Killip & Smith 21087 fl (GH, NY,
US); H. H. Smith 1524 fl fr (BR, F, G, MO, MICH, NY, P, US). Bolivar: Bro. Heriberto 302 fr
(US); 306 fl (US); Pennell 4734 fl (NY). No other locality: Bro. Ariste-Joseph A 883 fl (US); Dugand
339/767 fl fr (F, NY); 547 fl (F); Schultze 298 fr (NY fragment ex B); Stiibel 22 fl (NY fragment ex
B).
Common names. Colombia: "bejuco colorado," "coloradito," "bejuco golandrina. "
In its androeciumand gynoecium, Banisteriopsisheterostyla resembles the B.

lutea group, but differs in its externally hairy petals, sepals which are glabrous
adaxially, and the lateral winglets on the fruit nut parallelto, not radiatingfrom,
the areole.
The Banisteriopsis lutea group (species 69-70)
One of the two species of the group, B. lutea, is widely distributedto the west,
south, and east of the Amazon basin, whereas B. cristata has a more restricted
distributionto the north of the Amazon basin in Venezuela, Guyana, and Surinam. These species share the following characters:flowers producedwhen plant
is leafless, in axillaryfour-floweredumbels; flowers very similar,with eglandular
sepals which are tomento-sericeousadaxially, glabrouslong-clawedpetals, long
exserted stamens and styles, the stamens opposite the styles with very long filaments, the three posterior stamens inflexed between the posterior styles, the
styles sigmoid and slender, with the anterior style bearded for up to 2/3 of its
length; fruit with lateral crests or winglets radiatingfrom the areole.
69. Banisteriopsis lutea (Grisebach) Cuatrecasas, Ciencia (Mexico) 23: 141.
1964.
Fig. 38.
Banisterialutea Grisebach,Linnaea22: 15. 1849.
Banisteriapraecox Grisebach,Vidensk.Meddel.Nat. For. Kjobenhavn:133. 1875.Type. Warming 31, Brazil, MinasGerais, Lagoa Santa, fl (holotype, C).
BanisterianitrosiodoraGrisebach,Abh. Konigl. Ges. Wiss. G6ttingen24: 65. 1879.Type. Lorentz & Hieronymus486, Argentina,Salta, Oran, fl (holotype, GOET;isotypes, G, GOET,
NY).
Banisteria hassleriana Chodat, Bull. Herb. Boissier 6, Appendix 1: 16. 1898. Types. Hassler
812, Paraguay,near Ypacaray,fl (lectotype, G; isolectotypes, G, NY). Hassler 812a, Paraguay, fr (syntypes, G, NY, P).
Banisteriahispida Niedenzu, Verz. Vorles. Ak. Braunsb.:21. 1912. Type. Fiebrig 4193, Paraguay, near Centurion,between rivers Apa and Aquidaban,fr (holotypeG; isotype, G).
Jubistylis mollis H. H. Rusby, Mem. New York Bot. Gard. 7: 274. 1927. Type. H. H. Rusby
732, Bolivia, Bopi river, fl (holotype, NY; isotypes, GH, MICH).
Banisteriopsisnitrosiodora(Grisebach)O'Donell & Lourteig,Lilloa 9: 260. 1943.
Vine, rarely a subshrubor shrub with vining branches, young branches greytomentose, soon glabrate, the bark pale to reddish brown with prominentcorky
lenticels, old branches sometimes with prominent corky ridges. Stipules triangular, up to 1 mm long. Leaves with the petiole 5-22 mm long, tomentose to
glabrate,bearinga pair of protuberantglands near the apex adaxially, the lamina
4.0-10.1 cm long, 2.6-8.8 cm wide, ovate to orbicular,obtuse to truncateat the
base, acute to apiculateat the apex, with the marginplane, at first tomentose on
169
both sides, usually soon glabrateadaxiallyexcept the marginand midribnear the

base, abaxially usually persistently sparsely tomentose, sometimes glabrate, the
hairs T-shaped, stalked, with the trabecula more or less flexuous, 0.6-2.0 mm
long, with the adaxialsurface smooth and the lateralveins prominulousabaxially.
Inflorescence of 4-flowered umbels, 1-2(-6) borne in the axils of leaf scars of
leaves of previous seasons, the branchesup to 7 mm long, tomentose, bracteate;
the floriferousbracts and bracteoles 1.0-2.0 mm long, lanceolate to linear, abaxially tomentose to glabrate,deciduous duringor after flowering;pedicels sessile,
(10-)14-22(-28) mm long, 0.4-0.6 mm in diameter, sparsely tomentose to glabrate. Sepals 1.5-3.0 mm long, 1.2-2.5 mm wide, ovate to orbicular,roundedat
the apex, sparsely tomentose in the middle abaxially, tomento-sericeousadaxially, eglandular,enlargingin fruitup to 5 mm long. Petals yellow, laciniate,plane,
eglandular,glabrous, the 4 lateral petals reflexed between the sepals, with the
claw 2.0-3.0 mm long, the limb 5.0-9.0 mm long and 5.0-8.0 mm wide, orbicular
to broadly elliptic, the posteriorpetal with the claw erect, 3.0-4.0 mm long, 0.60.8 mm wide, the limb 4.0-5.0 mm long, 3.0-5.0 mm wide, broadly obovate.
Stamens with the filaments opposite the 4 lateral petals and the sepals 2.8-6.0
mm long, that opposite the anteriorsepal somewhatlonger than the others, those
opposite the antero-lateralsepals somewhatshorterthan the others, that opposite
the posteriorpetal 2.2-3.0 mm long, those of the 3 anteriorstamens erect, those
opposite the antero-lateralsepals and postero-lateralpetals directed posteriorly,
those of the 3 posterior stamens inflexed between the posterior styles; anthers
with the locules 0.7-1.2 mm long, glabrous, the connectives of the stamens opposite the sepals 0.8-1.0 mm long, 0.4-0.9 mm wide and 0.4-0.6 mm deep, obovate, papillateapically, the locules divergingdistally, those opposite the petals
0.6-1.0 mm long, 0.3-0.6 mm wide, 0.2-0.4 mm deep, oblong. Ovary 1.0-2.0 mm
tall, densely sericeous, the styles slender, sigmoid, and tapering apically, the
anterior style 4.4-5.8 mm long, hairy for one-half to two-thirds of its length,
directed anteriorly,the posteriorstyles 3.2-4.8 mm long, hairy at the base for up
to one-thirdof their length, directedposteriorly, stigmas truncate. Fruit with the
sepals enlarged and diverging, the carpophoreup to 1.5 mm long and 0.5 mm
wide, the nut 4-6 mm tall and 3-4 mm long, globose or laterally compressed,
bearingon each side 3 crest-likewings up to 3 mm high and 5 mm long extending
diagonallyfrom the areole, sparsely sericeous, the wing 19-26 mm long and 7-12
mm wide, sparsely appressed-sericeous,the hairs T-shaped, the trabeculastiff,
straight, irritating, 1.0-1.5 mm long, bearing on its upper marginat the base a
roundedappendage2-5 mm tall and 5-8 mm along the base.
Type. Ruiz s.n. (33/64), Peru, in nemoribus Andium, fl fr (holotype, MA
n.v.; isotype, F).
Distribution.(Fig. 37). In Argentina,Paraguayand Bolivia, extendingwest and
northup the Andes in Peru, and extendingeast and norththroughsouthernBrazil
to Maranhao.
Collected in flower and in fruit from June to December.
PERU. Cajamarca: A. Sagdstegui-A. 5859 fl yng fr (US); Woytkowski 6801 fl (MO). San Martin:
Spruce 4199 fl (BR, G, MG, NY, W). Junin: Woytkowski 5917 fl (G, GH, MO, US); 6499 fl (MO).
Cuzco: Marin 1651 fl (US); C. Vargas C. 1144 fl (GH); 2044 fl (GH); West 7191 fl (GH, MO).
BRAZIL. Maranhao: Ducke 793 fl fr (MG); Eiten & Eiten 4627B fl (NY); 4875 fl (NY, SP); Lisboa
2484 fl (MG, RB); Pires & Black 1640a (IAN, US). Piaui: Castellanos & Duarte 571 fl (MICH).
Ceara: Allemao & Cysneiros 221 fl (R); Cutler 8079 fl (IAN, R, US); 8410 (F, US); Dahlgren 814 fl
(F); Drouet 2522 fl fr (F, MICH, MO, NY, R, US, W); A. P. Duarte 1257 yng fr (RB); L. Duarte 483
fl (MICH); Ducke 2373 fr (IAN, UB); Eugenio 741 fl (RB); Gardner 1481 fl fr (G, P, W). Rio Grande
do Norte: Luetzelburg 11216 fl (M). Paraiba: Dobermir 45 fl (RFA); Luetzelburg 12019 fl (M, NY);
J. Colho de Moraes 1977 fl (IAN). Pernambuco: Chase 7687 fl (US); Falcao et al. 1059 fl (RB);
170
Flora Neotropica
TROPIqAL AERA
s
,e ,e-----
.-----
--l.
-. ,-
----
FIG. 37. Distributionof Banisteriopsissection Sciurostylis, species 67-73. Open square, B. val-
vata; closed square, B. heterostyla; open circle, B. lutea; closed circle, B. cristata; closed triangle,
B. leiocarpa; diamond, B. parvifolia; open triangle, B. populifolia.
Pickel 133 fl (F); 275 fl (GH, MICH, NY, P, US); Tovares 558 yng fr (US). Rond6nia: Black &
Cordeiro 52-15188 fl (IAN); Prance et al. 5152 fl fr (MICH, NY). Mato Grosso: Commissdo Rondon
(leg Hoehne) 169 fl (SP); 170 fl (R). Goias: Burchell 7325 fl (GOET, P); Eiten & Eiten 5580 fl (UB,
US); Gottsberger 25-23771 fl yng fr (MICH). Minas Gerais: Heringer 5691 fl (UB); Irwin & Soderstrom 5485 fl (NY, US); Macedo 415 fl (SP); 1140 fl (SP); 2500 fl fr (S, P); Martius s.n. fl (M); Pires
58019 fl (UB); Regnell 111.149 fl (S, US). Sao Paulo: J. M. de Freitas Campos 35 fl (NY, US); Gehrt
s.n. fl yng fr (SP); Gottsberger 13-26970 fl (NY). BOLIVIA. Bang 2457 fl (MO); Brooke 5568 fl (F,
NY); Cardenas 5634 fl (US); Herzog 1182 fl (G, S, W); Steinbach 1601 fl (GH); 6386 fl (GH); 6438
fl (G, GH); 7603 fl (F, GH); 8188 bis fl (GH); Troll 30 fl (M); Werdermann 2375 fl (MO, NY); White
255 fl (GH, NY); R. S. Williams 796 fl fr (NY, US). PARAGUAY. Chodat 3779 fl fr (G); Fiebrig 56
fl (F, G, GH, M); Hassler 3274 fl (F, G, GH, NY, P, W); 4451 fl (G, NY); 5118 fl (G); 7172 fl (F, G,
GH, MICH, MO, NY, P, S, W); 7355 fl (G, NY); 7470 fl (G, NY); (leg Rojas) 10537fl (G, GH, NY,
P, W); Lindman A2175 fl (F, G, GH, S, US); Osten 8254 fl (GOET, S); 9043 fl (S); Rojas 12917A fr
(S, W); 13552 fl (S, W); 13629 fl fr (MO, U). ARGENTINA. Jujuy: Cabrera & Fabris 15977 fr (P);
15992 fl (P); Eyerdam et al. 22274 fl (GH); Fries 520 fl (G, S); Venturi 5190 fl (BM, GH, US). Salta:
Inst. Miguel Lillo 3078 fl (GH); Rodriguez 1112 fl (NY); Schreiter 11429 fl (F, GH); Schulz 5490 fl
(MICH, W); Steinbach 1712 fl (GH). Tucuman: Venturi 1426 fl (GH).
Common names. Peru: "huillca bejuco." Brazil, Rio Grande do Norte: "Cipo
de Sdo Joao."
This species, like Banisteriopsis cristata, usually flowers when leafless in the
171
dry season, or as the new leaves are expandingat the beginningof the wet season.
Consequently,leaves are often not availablefor comparison,and the great variation exhibitedin leaf size and pubescence on collections with leaves is probably
at least in part due to the relative age of the leaves. Most specimens with large
leaves exhibit persistent, sparse, tomentose pubescence on the abaxial leaf surface, but some collections from lowland Paraguayand Maranhaoare almost glabrous. Until more collections with leaves are made and better data on variation
in leaf size and pubescence with age on one individual are collected it seems
preferableto include all the variationin one large taxon, since in flower and fruit
the specimens are very similar.Banisteriopsislutea can be distinguishedfrom B.
cristata by its tomentose pubescence on the leaves and inflorescence, and its
smallerfruit wing. Some leafless specimens from the cerradosof Brazil with the
habit of a subshrub,and densely tomentose stems with many flowers, have been
segregatedin the past as B. praecox. Probablythese plants represent young or
regeneratingindividuals,and young or regeneratingbranches are often different
in pubescence from branches on matureundamagedplants.
70. Banisteriopsiscristata (Grisebach) Cuatrecasas, Ciencia (Mexico) 23: 141.
1964.
Fig. 38.
Banisteriacristata Grisebach,Linnaea22: 16. 1849.
Banisteria orbicularis Niedenzu, Fedde, Rep. Spec. Nov. 33: 69. 1933. Type. E. G. Holt & W.
Gehriger155, Venezuela, Anzoategui,near Soledad, fl (holotype, B destroyed?;isotypes,

G, NY, US).
Liana, or shrub with vining branches, branches terete, pale to dark brown,
glabrous, lenticels numerous, more or less prominent,old branches with prominent corky ridges. Stipules triangular,up to 1 mm long. Leaves with the petiole
5-15 mm long, sparsely appressed-sericeousto glabrate, bearing a pair of protuberantglands adaxially near the apex, the lamina 2.7-9.0(-12.0) cm long, 2.55.9(-8.0) cm wide, ovate to rotund, cuneate to obtuse or rarely truncate at the
base, acute to acuminateor apiculateat the apex, with the marginplane, at first
appressed-sericeouson both sides, soon glabrousexcept the marginand midrib,
the hairs sessile, the trabeculastraight, 1.0-1.5 mm long, with the lateral veins
prominulouson both sides. Inflorescence in the axils of leaf scars of leaves of
previous seasons, of 4-flowered umbels, 1-several umbel-bearingbranches in
each axil, the branchesup to 6 mm long, sericeous; bracts and bracteoles 1.0-1.5
mm long, triangular,appressed-sericeousabaxially, involucrate,persistent;pedicels sessile, (10-)14-19(-21) mm long, 0.4-0.5 mm in diameter, sparsely appressed-sericeous.Sepals 1.7-2.3 mmlong, 1.4-2.2 mm wide, oblongto orbicular,
largerin fruit, appressed-sericeousabaxially,tomento-sericeousadaxially,eglandular. Petals yellow, laciniate, eglandular,glabrous, the 4 lateral petals reflexed
between the sepals, with the claw 2.0-3.5 mm long, the limb 5.0-11.0 mm long
and 4.5-8.0 mm wide, the petal outermost in bud largest, broadly elliptic, the
posteriorpetal with the claw erect, 2.5-3.5 mm long and up to 0.8 mm wide, the
limb 4.0-6.0 mm long, 2.5-5.0 mm wide, broadly obovate. Stamens with the
filamentsopposite the 4 lateralpetals and the anteriorand postero-lateralsepals
3.6-5.4 mm long, that opposite the anterior sepal longest (4.6-5.4 mm), those
opposite the antero-lateralsepals 2.6-3.6 mm long, that opposite the posterior
petal 1.6-3.0 mm long, those of the 3 anteriorstamens erect, those opposite the
antero-lateralsepals and postero-lateralpetals directed posteriorly, those of the
3 posteriorstamensinflexedbetween the posteriorstyles; antherswith the locules
0.6-1.0 mm long, glabrous, the connectives of the stamens opposite the sepals
172
Flora Neotropica
0.7-1.0 mm long, 0.6-0.8 mm wide, 0.3-0.6 mm deep, broadlyobovate, with the

locules divergingdistally, papillate, those opposite the petals 0.4-0.7 mm long,
0.3-0.4 mm wide, 0.2-0.4 mm deep, oblong. Ovary 1.0-1.3 mm tall, sericeous,
the styles slender and taperingapically, hairy for part of their length, the hairs
T-shaped, sessile, with unequal arms, the longer arm 1.0-2.0 mm long, the anterior style 4.5-5.4 mm long, curved and directed anteriorlydistally, hairyfor up
to half of its length, the posterior styles 3.8-4.2 mm long; sigmoid, directed
posteriorlydistally, hairyfor one-fourthto one-thirdof their length, stigmastruncate. Fruit with the carpophoreup to 1.5 mm long and 0.5 mm wide, the nut 45 mm tall and 4-5 mm long, globose, bearingon each side 3 to 7 crest-likelateral
wings radiatingfrom the areole, sparsely sericeous to glabrate, the wing 25-34
mm long, 9-11 mm wide, sparsely appressed-sericeous,the hairs stiff, irritating,
the trabeculastraight, 1.0-1.5 mm long, bearingon its upper marginat the base
an appendage6-7 mm tall and 5-7 mm along its base.
Type. Richard Schomburgk594, Guyana, fl fr (holotype, GOET).
Distribution.(Fig. 37). In deciduous woodland (savannah)in Venezuela, Guyana and Surinam.
Collected in flower in January,Marchto July and November throughDecember, and in fruit from May to July.
VENEZUELA. Zulia: Curran 149 fl (NY). Yaracuy: Pittier 11762 fl (G, GH, NY, US). Carabobo:
Pittier 8867 fl (GH, US). Aragua: Pittier 11346 fl (G, GH, NY, US); Vogl 471 fl (M, S). D.F.: Morillo
& Manara 1083 fr (VEN); 1141 fl (MICH); 1165 fl (MICH); Pittier 10307 fl yng fr (GH, NY, US);
10307a fl (GH); 10417 fl yng fr (GH, NY, US); 12591 fl (G, GH, NY, US). Guarico: Aristeguieta
6104 fl (NY); Pittier 11798 fl (G, GH, NY, US). Anzoategui: Holt & Gehriger 155 fl (G, NY, US);
Pittier 15180 fl (US); F. D. Smith 180 fl (US); 253 fl (US). GUYANA. Rich. Schomburgk 594 fl fr
(GOET); A. C. Smith 3152 fl (MO, NY, US). SURINAM. Schulz 8962 fl (GH, NY).
This species is distinguishedfrom Banisteriopsis lutea by its glabratemature

leaves, and the appressed-sericeouspubescence on the leaves and inflorescence.
Usually B. cristata flowers in a leafless condition so that few collections have
matureleaves. The type of B. orbicularishas matureleaves althoughthe flowers
are not borne in their axils, and this specimenwas collected in November. Pittier
11346also has matureleaves, and was collected in December. Perhapsthese two
specimens represent individualswhose floweringresponse was triggeredin the
wet season, perhapsin response to anomalousweather conditions.
The Banisteriopsis leiocarpa group (species 71-73)
The three species of this group have very restricteddistributionsin the Andes
of Peru. They share the following characters:calyx eglandular;petals small, all
strongly reflexed between the sepals; stamens similar, erect, projectingbeyond
the sepals; styles equal, slender, as long as the stamens, hairy at the base, incurved apically;fruit nut laterallycompressed, smooth, with appressedirritating
hairs.
71. Banisteriopsisleiocarpa(Adr. Jussieu) B. Gates, comb. nov.
Fig. 38.
Banisteria leiocarpa Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 395. 1843.
Vine or viningshrub,young branchesappressed-grey-sericeous,olderbranches

glabrate, the bark brown with pale lenticels. Stipules minute, triangular.Leaves
with the petiole 9-24 mm long, tomentose, eglandularor apicallybiglandular,the
lamina 3.0-7.3 cm long and 2.0-7.4 cm wide, sometimes smaller in the inflorescence, ovate to orbicular,obtuse to truncateat the base, apiculateto emarginate
173
#//
--
,...,
I>
I'
N
!?
''
:
f-I,
Ki
FIG. 38.
38. Banisteriopsis
B. lutea,
B. cristata
B. leiocarpa.
FIG.
cristata and
and B.
B. leiocarpa.
lutea, B.
A-D, B.
A,
Banisteriopsisheterostyla,
heterostyla,B.
leiocarpa. A-D,
leiocarpa. A,
flower x 3.5; B, stamen x 7.5; C, gynoecium x 5; D, fruit x 1.5. E, F, B. heterostyla. E, bud x 3;

F, fruit x 1.5. G-L, B. lutea. G, bud x 3; H, flower side view x 4; J, androeciumx 5 (the stamen
to the left is opposite the anteriorsepal); K, anther, adaxialview, x 7.5; L, gynoecium x 5. M, B.
fruit x 1.5.
1.5. (A-D,
Plowman 5558;
Prance et
et al.
al. 5152;
cristata,
5558; E,
cristata, fruit
E, F,
F, Dugand
(A-D, Plowman
G-L, Prance
M,
3391767;G-L,
5152; M,
Dugand 339/767;
Morillo& Manara
Manara1083.)
Morillo
1083.)
at the apex, with the margin plane and often bearing a pair of glands near the
midrib at the base, adaxially sparsely appressed-sericeous to glabrate, the hairs
with the trabecula 1.0-1.5 mm long, abaxially densely and loosely tomentose, the
trabecula 1.0-2.0 mm long, plane or with the reticulation prominulous adaxially
174
Flora Neotropica
and with the 4-7 pairs of lateral veins prominent abaxially. Inflorescence of cymes
of 1-5 4-flowered umbels in the axils of reduced leaves on short lateral branches,
appressed-sericeous; bracts and bracteoles 1.0-1.8 mm long, lanceolate, appressed-sericeous to glabrate abaxially, involucrate, persistent; pedicels sessile,
6-11 mm long, 0.4-0.6 mm in diameter, appressed-sericeous. Sepals 1.5-2.0 mm
long and 0.9-1.4 mm wide, elliptic, appressed-sericeous abaxially, tomentose
adaxially, erect, eglandular. Petals yellow, glabrous, strongly reflexed between
the sepals, the posterior petal scarcely distinguishable from the 4 lateral petals
but a little smaller, the claw 1.2-1.5 mm long, the limb 3.5-5.0 mm long, 3.5-4.5
mm wide, rotund, denticulate, concave. Stamens with the filaments 2.4-3.6 mm
long, erect, those opposite the petals slightly longer and thicker than those opposite the sepals, anthers with the locules 0.7-1.0 mm long, glabrous, the connectives subequal, 0.6-0.8 mm long, 0.4-0.6 mm wide, 0.2-0.4 mm deep, elliptic
to oblong, papillose. Ovary 1.0-1.4 mm tall, densely sericeous; styles 2.6-3.2 mm
long, diverging, hairy for half their length, subequal or the posterior styles a little
shorter and more slender than the anterior style, tapering and strongly incurved
apically, stigmas truncate. Fruit with the carpophore up to 2 mm long and 1 mm
wide, the nut 5-6 mm tall and 3-4 mm long, smooth, laterally compressed, with
appressed irritating hairs, the trabecula 1.2-1.5 mm long, soon glabrate, with an
appendage on its upper edge 1-2 mm tall and 5-6 mm along the base, the wing
16-20 mm long, 10-14 mm wide, oblong, appressed-sericeous, soon glabrate, the
hairs irritating, the trabecula straight, 1.5-2.0 mm long.
Type. Mathews 2126, Peru, Amazonas, Chachapoyas, fl fr (holotype, P; isotypes, BR, G, K).
Distribution. (Fig. 37). Known only from Amazonas in Peru at altitudes of
1500-1900 m.
Collected in flower in February and April, and in fruit in April.
PERU. Amazonas: Ferreyra 7133 fl fr (F, US); Mathews 2126 fl (BR, G, K, P); Plowman 5558 fl
(GH, MICH).
This species can be distinguished from Banisteriopsis parvifolia by its larger

leaves, flowers and fruits, and from B. populifolia by its tomentose leaf pubescence.
72. Banisteriopsis parvifolia (Niedenzu) B. Gates, stat. et comb. nov.
Banisteria leiocarpa var. parvifolia Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 451. 1928.
Vine or vining shrub, branches grey-tomento-sericeous, soon glabrate, the bark

brown with pale lenticels. Stipules minute, triangular. Leaves with the petiole 510(-15) mm long, tomento-sericeous, eglandular, the lamina 1.4-4.5 cm long and
0.9-2.5 cm wide, narrowly ovate to ovate, obtuse to cuneate at the base, acute,
apiculate or emarginate at the apex, with the margin plane, bearing a pair of glands
near the midrib at the base, adaxially sparsely appressed-sericeous to glabrate,
the trabecula up to 1.0 mm long, abaxially densely and loosely tomento-sericeous,
the trabecula 1.0-1.5 mm long, plane or with the reticulation prominulous adaxially and the 4-7 pairs of lateral veins prominent abaxially. Inflorescence axillary,
of cymes of 1-9 4-flowered umbels, appressed-sericeous; bracts and bracteoles
1.0-1.4 mm long, triangular, sparsely appressed-sericeous to glabrate abaxially,
involucrate, persistent: pedicels sessile, 3-9 mm long, 0.3-0.4 mm in diameter,
densely sericeous. Sepals 1.6-1.8 mm long, 0.6-1.2 mm wide, narrowly elliptic
to elliptic, appressed-sericeous abaxially, sparsely tomentose adaxially, erect,
eglandular. Petals yellow, glabrous, strongly reflexed between the sepals, the
175
posteriorpetal scarcely distinguishablefrom the 4 lateralpetals, but a little smaller, the claw 0.8-2.0 mm long, the limb 2.0-4.5 mm long, 1.5-4.0 mm wide, rotund,
denticulate, cochleate. Stamens with the filaments 1.8-4.0 mm long, erect, those
opposite the petals slightly longer and thicker than those opposite the sepals;
anthers with the locules 0.6-1.0 mm long, glabrous, the connectives subequal,
0.5-0.8 mm long, 0.3-0.6 mm wide and 0.2-0.3 mm deep, elliptic to obovate,
papillose. Ovary 0.6-1.2 mm tall, densely sericeous; styles 2.0-3.4 mm long,
divergingdistally, hairy for half their length, subequal, or the posterior styles a
little shorterand more slender, taperingand hooked at the apex, stigmastruncate.
Fruit with the carpophoreup to 2 mm long and 1 mm wide, the nut 3-4 mm tall
and up to 3 mm long, smooth, laterally compressed, appressed-sericeous,soon
glabrate, the hairs irritating,the trabecula straight, 1.0-1.5 mm long, with an
appendageon its upperedge up to 1 mm tall and 3-4 mm along the base, the wing
8-12 mm long and 7-8 mm wide, oblong, appressed-sericeous,soon glabrate,like
the nut.
Type. Weberbauer6063, Peru, Piura, Huancabambafl fr (lectotype, NY fragment ex B; isolectotypes, F, GH, US).
Distribution.(Fig. 37). Known only from three collections in Andean Peru at
altitudes of 1900-2000m.
Collected in flower in March, April and June, and in fruit in April.
PERU. Piura: Weberbauer 6063 fl fr (F, GH, NY fragment ex B, US); Fosberg 27700 fl (MO).
Huancavelica: Weberbauer 6466 fl (syntypes, F, GH, US).
The two collections of this species from Piura differ in their pubescence and
flower size fromthe collection from Huancabamba,exhibitinglooser more tomentose pubescence, and much smallerflowers with narrowlyelliptic sepals. Other
collections of this species will give a better indicationfor the range of variation
within it. Banisteriopsis parvifolia can be distinguished from B. leiocarpa by its
smaller leaves and smallerfruits.
73. Banisteriopsispopulifolia(Niedenzu) B. Gates, comb. nov.

Banisteria populifolia Niedenzu in Engler, Bot. Jahrb. 40: 281. 1908.
Liana, branchesappressed-silvery-sericeous,eventuallyglabrate,the barkwith

prominentred-brownlenticels. Stipulesminute. Leaves with the petiole 7-14 mm
long, densely white-sericeous, eglandularor apicallybiglandular,the lamina3.38.4 cm long, 1.7-6.1 cm wide, ovate, obtuse to truncateat the base, apiculateat
the apex, with the marginplane and often bearinga pair of glands near the midrib
at the base, adaxiallysparselyto densely persistentlyshining-sericeous,the hairs
golden with the trabecula0.8-1.2 mm long, abaxially densely shining-sericeous,
the hairs T-shaped, very short-stalked,the trabecula 1.0-1.5 mm long, adaxially
plane, abaxially with the 7-9 pairs of lateral veins prominent.Inflorescence axillary, of 4-floweredumbels in cymes of 3-7 umbels, appressed-sericeous;bracts
and bracteoles0.8-1.2 mm long, lanceolate, appressedsericeous;bractsand bracteoles 0.8-1.2 mm long, lanceolate, appressed-sericeousto glabrate abaxially,
involucrate,persistent;pedicels sessile, 3-16 mm long, 0.5-0.6 mm in diameter,
appressed-sericeous.Sepals 1.5-2.2 mm long, 1.2-1.7 mm wide, elliptic to broadly elliptic, appressed-sericeousabaxially, loosely tomento-sericeous adaxially,
erect, eglandular.Petals yellow, strongly reflexed between the sepals, the posterior petal scarcely distinguishablefrom the 4 lateral petals but a little smaller,
the claw 1.0-1.5 mm long, the limb 4.0-5.5 mm long, 4.0-5.0 mm wide, rotund,
denticulate, concave. Stamens with the filaments 2.0-4.0 mm long, erect, those
176
Flora Neotropica
opposite the petals slightly longer and thicker than those opposite the sepals;
anthers with the locules 0.8-0.9 mm long, glabrous, the connectives subequal,
0.6-0.8 mm long, 0.4-0.6 mm wide, 0.1-0.3 mm deep, elliptic to oblong, papillate.
Ovary 1.0-1.2 mm tall, densely sericeous; styles subequal, 3.0-3.6 mm long, hairy
for up to half their length, diverging, incurved apically, stigmas truncate. Fruit
(only immature known) with the nut up to 3 mm tall and 2 mm long, smooth,
densely sericeous to glabrate, the hairs stiff and irritating, the trabecula 1.0-1.5
mm long, bearing an appendage on its upper edge up to 1 mm tall and 2 mm along
the base, the wing up to 7 mm long and 5 mm wide, appressed-sericeous to
glabrate like the nut.
Type. Weberbauer 4794, Peru, Amazonas, "in declivibus saxosis apricis in
valle fl. Maranon prope Tupen," fl (holotype, B destroyed, fragment NY; isotype,
G).
Distribution. (Fig. 37). In the valley of the river Marafi6n in Peru at altitudes
of 500-800 m.
Collected in flower in March and May.
PERU. Cajamarca: Weberbauer 6230 fl (F, GH, NY, fragment ex B). Amazonas: Ferreyra 14196
fl (US); Weberbauer 4794 fl (G, NY, fragment ex B).
This species can be distinguished from Banisteriopsis leiocarpa by its ovate

leaves which are densely shining sericeous abaxially, and from B. parvifolia by
its larger leaves and larger flowers.
Banisteriopsis subgenus 3. Pleiopterys section 2. Anisopterys (Grisebach) B. Gates,
comb. nov., species 74-92.
Banisteria section Anisopterys Grisebach, Linnaea 13: 201. 1839.
Banisteria subsection Octadenia Niedenzu, Ind. Lect. Lyc. Brunsb.: 18. 1901. Lectotype
species. Banisteriopsis pubipetala (Adr. Jussieu) Cuatrecasas as Banisteria pubipetala.
Banisteria subsection Anadenia Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1901. Lectotype species.
Banisteriopsis patula Gates as Banisteria eglandulosa Adr. Jussieu.
Banisteria section Actinoctenia Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1901. Lectotype species.
Banisteriopsis lucida (Richard) Small, as Banisteria lucida.
Banisteria subsection Sericopetalum Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1901. Lectotype
species. Banisteriopsis lucida (Richard) Small, as Banisteria lucida.
Banisteria series Orthanthele Niedenzu, Ind. Lect. Lyc. Brunsb.: 12. 1901. Lectotype species.
Banisteriopsis hypericifolia (Adr. Jussieu) Anderson et Gates as Banisteria hypericifolia.
Banisteria series Cyrtanthele Niedenzu, Ind. Lect. Lyc. Brunsb.: 14. 1901. Lectotype species.
Banisteriopsis lucida (Richard) Small as Banisteria lucida.
Leaves with the petiole eglandular, the lamina with many minute glands on or
just within the margin (except B. caduciflora with 2-4 pairs of glandular areas on
the lamina). Inflorescence axillary, of 4- to 6-flowered umbels or condensed axillary racemes of up to 8 pairs of flowers. Pedicels often becoming thicker and
woody in fruit. Sepals bearing glands below the free part of the sepal on the
receptacle, eglandular in B. patula and B. sepium, not reflexed or enlarging in
fruit. Petals densely sericeous externally (except B. caduciflora). Styles stout and
equal or the posterior styles more slender, hairy basally or for up to half the
length of the anterior style. Samaras with carpophore short and broad, up to 1
mm long and 2 mm wide, or absent; hairs on samaras sessile, not irritating to the
skin, the trabecula straight or flexuous, 0.2-1.0 mm long.
Type species. Banisteriopsis pubipetala (Adr. Jussieu) Cuatrecasas as Banisteria pubipetala.
177
The Banisteriopsislucida group (species 74-83)

This group includes one widespreadspecies, B. lucida, and nine species with
more restricted distributions.They share the following characters:leaves with
marginalglands, often more numeroustowards the apex; flowers borne in fourfloweredumbels or condensedracemesof up to 12flowers, axillary;petals yellow,
externallysericeous;fruitwith the nut bearingnumerouslateralcrests, sometimes
developed into winglets, radiatingfrom the areole.
74. Banisteriopsislucida (Richard)Small, North Amer. Flora 25: 133. 1910.
Fig. 39.
Banisterialucida Richard,Actes Soc. Hist. Nat. Paris 1: 109. 1792.
Heteropterys appendiculata DC., Prodr. 1: 592. 1824. Type. Lambert s.n., St. Vincent, fl fr
(holotype, G; istotype, P-JU). Probablycultivated.

Banisterialobulata E. Meyer, Nov. ActorumAcad. Caes. Leop.-CarolXII, 2: 813. (1825?)Type.
E. Meyers.n. (813), Surinam,fr (holotype, B destroyed;isotype, G fide Kostermans,n.v.)
Banisteria divaricata Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 284. 1840. Type. Richard
s.n., FrenchGuiana,fl (holotype, P; isotype, P).

Banisteriaappendiculata(DC.) Adr. Jussieu, Arch. Mus. Hist. Nat. 3: 158. 1843.
Banisteriacalocarpa Miquel,Linnaea18: 53. 1844.Type. Focke 329, Surinam,fr (holotype, U).
Banisteriapolita Miquel, Natuurk. Verh. Holl. Maatsch. Wetensch. HaarlemII, 7: 80. 1851.
Type. Kappler ed. Hohenacker 1900, Surinam,river Marowini,fl fr (holotype, U; isotype, P).
Stigmaphyllon nigrescens 0. Kuntze, Rev. Gen. pl. 1: 89. 1891. Type. O. Kuntze s.n., West
Indies, PuertoRico (holotype, NY). Probablycultivated.
Diplopterys marshalliana Riley, Kew Bull.: 135. 1925. Type. L. A. M. Riley 94, Trinidad, Arena
Reserve, fl (holotype, K; isotype, NY).
Liana or vining shrub, young branches appressed-sericeousto glabrate, old

branches glabrate, smooth. Stipules minute, 0.5-1.0 mm long, triangular,joined
by an interpetiolarline. Leaves falcate, the petiole 4-10 mm long, sparsely appressed-sericeous to glabrate, eglandular,the lamina (3.6-)6.0-13.3(-16.4) cm
long, (2.2-)3.2-6.0(-7.5) cm wide, elliptic, cuneate at the base, with the apex
short- to long-acuminate,the marginplane or rarely slightlyrevolute and bearing
minute glands, more numeroustowards the apex, adaxially at first sparsely appressed-sericeous, soon glabrateexcept near the midvein at the base, abaxially
sparsely appressed-sericeous, sometimes glabrate, the hairs with the trabecula
0.2-0.3 mm long, with 4-6 pairsof mainlateralveins, the reticulationprominulous
adaxially and prominentabaxially. Inflorescenceaxillary, of 2-4 4-floweredumbels in each axil, the branches 6-12 mm long, bibracteate, densely brown-sericeous; bracts and bracteoles up to 1.0 mm long, cymbiform, densely brownsericeous, involucrate, persistent; pedicels sessile, 13-20 mm long, 0.6-0.7 mm
in diameter, brown-sericeous,glabratein age. Sepals 1.8-2.2 mm long, 1.4-2.0
mm wide, broadly elliptic, roundedat the apex, densely golden-sericeous, the 4
lateral sepals biglandular,projecting0.6-1.0 mm beyond the glands, the glands
1.0-1.8 mm long, 0.8-1.2 mm wide. Petals yellow, externally sericeous, the 4
lateral petals reflexed between the sepals, with the claw 1.0-1.5 mm long, the
limb dentate to lacerate, plane, broadly elliptic, that of the antero-lateralpetals
7.0-10.0 mm long and 6.0-8.5 mm wide, that of the postero-lateralpetals 5.0-7.5
mm long and 4.0-6.5 mm wide, the posterior petal with the claw erect, 3.0-3.5
mm long, 0.4-0.6 mm wide, constrictedapically, the limb 4.0-5.0 mm long, 3.04.0 mm wide, obovate, lacerate. Stamens with the filaments glabrous, 1.4-4.4
mm long, those opposite the styles (i.e. opposite the anteriorsepal and the pos-
178
Flora Neotropica
tero-lateralpetals) longest, those of the 3 anteriorstamenserect with that opposite

the anterior style inflexed apically, those opposite the antero-lateralsepals and
postero-lateral petals directed posteriorly and connate for one-third of their
length, those of the 3 posterior stamens inflexed between the posterior styles;
anthers with the locules 0.9-1.4 mm long, pubescent, diverging distally on the
stamens opposite the 4 lateralsepals, the connectives of the stamens opposite the
sepals 0.7-1.2 mm long, 0.6-1.0 mm wide, 0.3-0.7 mm deep, obovate, swollen
distally, those of the stamens opposite the petals 0.6-1.0 mm long, 0.2-0.5 mm
wide, 0.1-0.2 mm deep, oblong, with the locules parallel.Pollen (33-)35-40(-45)
utmin diameter. Ovary 1.0-1.4 mm tall, densely hairy, the anteriorstyle 2.4-4.8
mm long, stout, apicallyinflexed,hairyfor up to one-halfits length, the 2 posterior
styles 2.0-3.4 mm long, slender, divergingfrom the base, directed towards the
posteriorpetal. Samarawith slendercarpophoreup to 1 mm long, the nut globose,
5-10 mm tall and long, ridged or rarely cristate, the ridges radiatingfrom the
areole, sparsely appressed-sericeous,bearingalong its upper edge an appendage
up to 10 mm tall and up to 5 mm along its base, the wing 36-63 mm long, 14-25
mm wide distally, usually much narrowerat the base, sparsely appressed-sericeous.
Type. Leblonds.n., French Guiana,Cayenne, fl (holotype, P; isotypes, G (no.

49), P, P-LAM).
Distribution.(Fig. 40). In tropical rain forests of eastern South America, extending westward throughAmazonian Brazil to lowland Bolivia and Peru. The
two type collections from Puerto Rico and St. Vincent probablywere cultivated;
there are no other collections from the West Indies.
Representative collections. COLOMBIA. Magdalena: Cuatrecasas et al. 25359 fr (US); 25395 bud
yng fr (US); Kirkbride 1979 fl (MICH, NY). Antioquia: Romero-Casteneda 10091 fr (NY). VENEZUELA. Zulia: Steyermark 105613 fl (NY). Aragua: Gines 4843 fl (US); 5018 fr (US); Pittier 14137
fl (F, G, US). D.F.: Delgado 74 fr (F, US). Bolivar: Bernardi s.n. fl (NY); Brewer 134 fl (VEN);
Cardona 1178 fl (US, VEN); Steyermark 89510 fl (VEN). Amazonas: Holt & Blake 689 fl (US); Lizot
77 fl (VEN); Maguire 34721 fr (MICH, NY); 34748 fl fr (MICH, NY); Medina 384 fl (VEN); Spruce
3276 fl (BR, G, GH, MG, NY, P, W); LI. Williams 15719 fr (F, US, VEN); Wurdack 43130 fl (MICH,
NY); 43650 fr (MICH, NY). TRINIDAD. Baker s.n. st (U); Britton et al. 603 fl fr (NY); Broadway
4032 fl (F, G, M, NY, S, U); 6910 fr (F, MO, S, US); Cowan 1182 fr (GH, NY, P, S, US); A. C.
Smith 10076 fl (GH, US). GUYANA. Cowan 2093 fl (F, NY); De La Cruz 1386 fl (F, GH, MO, NY,
US); 3081 fr (F, GH, MO, NY, US); Forest Dept 6967 fl (MICH, P, S); Macguire 23354 fr (F, GH,
MO, NY, U, US). SURINAM. Berthoud-Coulon 295 fl (F, MICH, MO, P); Hostmann & Kappler
1027 fl (C, F, G, GOET, M, MO, P, U, US); Kramer & Hekking 2753 fl (C, F, NY, U); Lanjouw &
Lindeman 3465 fr (NY, U); Wessels Boer 977 fl (U); Wullschlaegel 1387 fl (BR). FRENCH GUIANA.
Melinon 245 fl fr (NY, P, US); Oldeman B-1362 fr (P); Sagot 96 fl (BR, C, GH, GOET, P,
W). PERU. Amazonas: Ancuash 284 fl (MICH). San Martin: Ll. Williams 7274 fl (F); 7776 fl (F, US).
Cuzco: Gay s.n. fl (P). BRAZIL. Amaz6nas: Cardona 1422 fr (US, VEN); 1428 fl (US). Roraima:
Prance et al. 10740 yng fr (MICH, NY); 11056 fr (MG, MICH, NY); 11203 fr (MG, MICH, NY).
Amapa: Fr6es 25770 fl (IAN); Irwin et al. 47437 bud fr (IAN, MG, MICH, NY). Para: Black 48-3406
fl (IAN); E. Oliveira 3298 bud (IAN); A. Silva 133 yng fr (IAN, NY, US). BOLIVIA. Steinhach 6817
yng fr (F, G, GH, MO, NY, fragment ex B).
Common names. Venezuela: "mune-yek" (Arekuna). Brazil: "totorinmo"

(Uaica-Mucajai); "guarand-rana macho."
Banisteriopsislucida is a widespreadspecies, but has been most often collected

in eastern Venezuela, Guyana and Surinam. It is characterizedby bearing its
flowers in axillaryfour-floweredumbels, by small calyx glands, a hooked anterior
style much longer than the posterior styles, and the fruit with its globose, ridged
nut without lateral wings, and large dorsal wing which is much narrowerat the
base. Throughoutmost of its range B. pubipetala is easily distinguishedfrom B.
lucida, not only in fruitbut also in flower, as its flowers are borne in 6-8-flowered
179
axillary racemes, have large calyx glands and an anteriorstyle which is curved
but not inflexed at a sharp angle apically. However, in eastern Colombia and
western Venezuela specimens of B. pubipetala in flower are very difficultto distinguish from B. lucida, as the flowers are borne in 4-floweredumbels, and the
anteriorstyle is more or less hooked apically;they can be distinguishedin flower
only by the smaller calyx glands and smallerpollen of B. lucida.
Cardona 1422 from Amazonian Brazil is unusual in having fruits with a relatively large nut and in this respect approachesB. peruviana.
75. Banisteriopsisnutans (Niedenzu) B. Gates, comb. nov.
Banisterianutans Niedenzu, Ind. Lect. Lyc. Brunsb.: 14. 1901.
Banisteriopsispanamensis Cuatrecasas& Croat,Ann. MissouriBot. Gard.67: 861. 1981.
Liana, branches appressed-sericeous,soon glabrate. Stipules triangularup to

1 mm long, sparsely sericeous. Leaves falcate with the petiole (3-)5-9 mm long,
sparsely appressed-sericeousto glabrate, eglandular,the lamina 4.2-12.7(-16.2)
cm long, 1.9-5.7(-7.3) cm wide, elliptic or rarely obovate, cuneate at the base,
short- to long-acuminateat the apex, with the marginplane to slightly revolute
and bearing minute glands, glabrous adaxially, sparsely appressed-sericeousto
glabrateabaxially, the hairs 0.2-0.6 mm long, plane adaxially, with the reticulation prominulousto prominentabaxially.Inflorescenceaxillary, 1-3 branchesper
leaf axil, each branch usually bearing a single 4-floweredumbel, rarely a cyme
of 3 umbels, appressed-sericeous,the hairs brown to golden; bracts and bracteoles 1.0-1.6 mm long, broadlytriangular,appressed-sericeous,involucrate,persistent; pedicels sessile, 12-22(-25) mm long, 0.6-0.8 mm in diameter,appressedsericeous, the hairs brown with the trabecula0.3-0.5 mm long. Sepals 2.0-2.8
mm long, the anterior sepal 1.2-2.2 mm wide, the 4 lateral sepals 2.0-2.8 mm
wide at the base, narrower at the apex, broadly ovate, rounded at the apex,
appressed-sericeousabaxially,glabrousadaxially,the 4 lateralsepals biglandular,
projecting0.2-0.8 mm beyond the glands, the glands 2.0-3.8 mm long, and 1.22.0 mm wide. Petals yellow, externally sericeous, lacerate to fimbriate, the 4
lateral petals reflexed between the sepals, broadly elliptic, the claw 1.0-2.0 mm
long, the limb of the antero-lateralpetals 8.5-11.0 mm long and 7.5-9.0 mm wide,
concave, the limb of the postero-lateralpetals 6.0-8.0 mm long and 5.0-7.0 mm
wide, plane, the posterior petal with the claw erect, 3.0-4.0 mm long, up to 1.0
mm wide distally, constrictedat the apex, the limb 5.0-6.5 mm long and 3.5-5.0
mm wide, obovate, plane, the basal laciniae sometimes gland-tipped.Stamens
with the filaments 1.0-4.0 mm long, that opposite the anteriorsepal longest, that
opposite the posterior petal shortest, those opposite the sepals usually longer
than those opposite the petals, or those opposite the postero-lateralpetals as long
as or longer than those opposite the antero-lateralsepals, those of the 3 posterior
stamens sometimes inflexed between the posterior styles, those of the 3 anterior
stamens connate basally, those of the other 7 stamens connate for up to half their
length; antherswith the locules 0.9-1.8 mm long, sparselyhairy, the connectives
of the stamens opposite the sepals 0.9-1.6 mm long, 0.7-1.0 mm wide, 0.6-0.8
mm deep, obovate or elliptic, those opposite the petals 0.8-1.2 mm long, 0.4-0.8
mm wide and 0.3-0.6 mm deep, narrowlyoblong or narrowlyelliptic. Ovary 1.01.4 mm tall, densely hairy, the styles subequal, 2.2-2.6 mm long, 0.2-0.3 mm in
diameter,the anteriorstyle sometimesa little stouterandlongerthanthe posterior
styles, diverging, glabrous or basally hairy, stigmas capitate. Fruit with short
broad carpophore,the nut 6-8 mm tall and 5-7 mm long, with a series of parallel
ridges or crests radiatingfrom the areole laterally,tomento-sericeousto glabrate,
180
Flora Neotropica
the hairs sessile with the trabeculaflexuous and 0.2-0.4 mm long, the wing 3047 mm long, 10-19 mm wide, with the upper and lower margins more or less
parallel, appressed-sericeousto glabrate,the hairs sessile with the trabecula0.20.4 mm long, bearing an appendageon the upper edge at the base 2.5 mm tall
and 4-7 mm along the base.
Type. Poeppig 1497, Peru, Huanuco, Cuchero, bud (holotype, B? destroyed;
isotypes, F, W).
Distribution.(Fig. 40). In the Andes in Peru and Bolivia, with one collection
from the northernmostend of the Andes in Venezuela, and two collections from
Panama.
Collected in flower in February, April, May, August, and November, and in
fruit in May, August and September.
PANAMA. Gentry 6159 fr (MICH); Mori & Kallunki 1836 fr (MICH). VENEZUELA. Lara: Mocquerys 6 fr (P); Tamayo 253 fl fr (US). PERU. Huanuco: Poeppig 1497 bud (F, W). Cuzco: Cook &
Gilbert 982 fr (US); Hunnewell 16039 fl (GH). BOLIVIA. Buchtien 1821 fl (US); 1822 fl (US); 1924
fl (US).
This species differs from Banisteriopsislucida in its largerflowers with large

calyx glands, its stout, subequal, diverging styles, and its shorter, stouter fruit
wing which is not constricted at the base. There is considerablevariationin the
leaves and fruits shown by the specimens I have included in this taxon. The
specimens from Venezuela differ from the Peruvianspecimens in their smaller,
more coriaceous leaves with short petioles (3-5 mm) but in flower and fruit are
very similar;the Bolivian specimens are unusualin their very large calyx glands,
large coriaceous leaves, and subequal, erect stamen filaments, and in this last
characterthey approachB. nigrescens. The two collections from Panamaare like
the Venezuelan specimens in leaf size and shape, but the fruit nut is alulate, with
the winglets developing from ridges radiatingfrom the areole but superficially
appearingparallel to the areole. This development of the lateral crests on the
fruit into winglets is also found in B. lucida and I have not considered it to be
taxonomicallysignificant.With the winglets appearingparallelto the areole, the
Panamanianspecimens resemble B. platyptera and these two species are almost
indistinguishablein flower, althoughthe mode of developmentof the lateralwings
is apparentearly in fruit and makes their separationeven in young fruit simple.
76. Banisteriopsisperuviana(Niedenzu) B. Gates, comb. nov.
Banisteria peruviana Niedenzu, Ind. Lect. Lyc. Brunsb.: 15. 1901.
Liana (?), young branches compressed, sparsely appressed-pubescent, old

branchesterete, glabrous. Stipules triangular,up to 1 mm long, their bases joined
by a more or less prominentinter-petiolarridge. Leaves with the petiole up to 8
mm long, sparsely appressed-pubescent,the lamina7.7-12.2 cm long and 4.0-6.5
cm wide, elliptic, cuneate at the base, acuminate at the apex, with the margin
slightlyrevolute and bearingminuteglands, adaxiallyglabrous,abaxiallysparsely
appressed-pubescent,the hairs with the trabecula0.3-0.6 mm long, with the 68 pairs of lateral nerves prominulousadaxially and prominentabaxially. Inflorescence axillary, of 4-floweredumbels, appressed-pubescentto glabrate,bracts
and bracteoles 1.2-1.6 mm long, broadlytriangular,involucrate,persistent. Pedicels sessile, 17-22 mm long, 0.6-0.7 mm in diameter, much stouter in fruit.
Sepals 2.0-2.2 mm long and wide, rounded at the apex, abaxially brown-sericeous, the 4 lateral sepals.biglandular,the glands 2.0-3.0 mm long and 1.2-1.5
mm wide. Petals yellow, externally sericeous. Stamens with the filaments gla-
181
brous, erect, that opposite the anteriorsepal inflexed apically, those opposite the
sepals and the postero-lateralpetals 3.6-4.2 mm long, those opposite the other
petals 2.8-3.2 mm long, the locules 1.0-1.4 mm long, pubescent, the connective
opposite the anteriorsepal short and oblong, those opposite the other sepals and
the petals longer, those opposite the sepals obovate and those opposite the petals
narrowlyoblong. Ovaryup to 2.0 mm tall, densely brown-pubescent,the anterior
style 3.0-3.2 mm long, stout, incurved apically, the posterior styles 2.4-2.6 mm
long, straightand diverging.Fruits without carpophore,the nut up to 20 mm tall
and 15 mm long, globose, bearinglaterally5-7 ridge-likecrests radiatingfrom the
aerole, up to 3 mm tall and sometimes interconnected;apically with a rounded
appendage 14 mm long and up to 10 mm tall, the dorsal wing up to at least 45
mm long and 17 mm wide, narrowertowards the nut, very thickened along its
upper edge, appressed-sericeous.
Type. Const. de Jelski 343, Peru, Cajamarca,Tambillo, (fl) fr (holotype, B?
destroyed; isotypes, US, W).
Distribution.(Fig. 40). Known only from the type in Cajamarca,Peru.
Collected in fruit in September.
PERU. Cajamarca: Jelski 343 fr (US, W).
The fruit of this species is so unusual with its large fruit nut and prominent
ridge-likecrests radiatingfrom the areole, that I have maintainedits specific rank.
There are no additionalcollections in fruit or flower; the flower measurementsI
have included in the description were taken from flowers in the packet on the
sheet from Vienna. Niedenzu states in his descriptionthat he did not have flowers, so it is possible that these flowers do not belong to Jelski 343. Indeed, the
flowers are almost identicalto those of the type of Banisteriopsisnutans. Perhaps
it is difficultto distinguishB. peruviana and B. nutans in flower; only additional
collections will resolve this problem.
77. BanisteriopsiswoytkowskiiB. Gates, sp. nov.
Fig. 39.
Liana, ramijuniores appresso-aureo-sericei,canaliculati,rami vetustiores glabrati. Stipulae usque 1.0 mm longae, triangulares,jugo interpetiolarijunctae.
Folia falcata, petiolo 4-10 mm longo, appresso-brunneo-sericeo,adaxialitercanaliculato,eglanduloso,lamina4.5-17.5 cm longa, 1.6-8.2 cm lata, lanceolatavel
ovata, basi obtusa vel truncata,apice acuminata,margineparumrevolutaet glandulis minutis instructa, adaxialiter sparsim tomento-sericea, mox glabrata, abaxialitertomento-velutina,pilis sessilibus brachiis0.4-0.5 mm longis, abaxialiter
venis appresso-sericeis et prominentibus,adaxialiterprominulis. Inflorescentia
axillaris umbellae quadrifloraecomposita, rariusin cymis 3-5 umbellarumfasciculata, in axilla 2-3 ramos floriferes gerens, appresso-aureo-sericea;bracteis
bracteolisque 0.8-1.2 mm longis, late triangularibus;pedicellis sessilibus 10-22
mm longis, 0.6-0-8 mm diametro,brunneo-sericeis,in fructu incrassatis. Sepala
2.0-2.2 mm longa, 1.5-2.2 mm lata, rotunda, apice rotundata, abaxialiterappresso-aureo-sericea,adaxialiterglabra,4 sepalis lateralibus2 glandesgerentibus,
glandes 0.6 mm superantibus,glandibus 2.0-3.4 mm longis, 1.0-2.4 mm latis.
Petala lutea, extus sericea, lacerata vel longe fimbriata,rotundavel orbicularia,
4 lateraliainter sepala patentiaungue 1.5-2.0 mm longo, limbo antero-lateralium
7.0-9.0 mm longo, 6.0-8.0 mm lato, concavo, illo postero-lateralium5.0-6.0 mm
longo et 4.0-5.0 mm lato, plano, petalo postico ungue erecto usque 2.5 mm longo,
apice constricto, limbo 3.5-4.0 mm longo latoque, orbiculari,fimbriisbasalibus
glanduliferis. Stamina filamentis 1.4-3.0 mm longis, illo stylo antico opposito
182
Flora Neotropica
S.
I,a.
FIG. 39.
Banisteriopsis lucida, B. woytkowskii, B. hypericifolia, B. amplectens and B. rondon-
iensis. A-F, B. hypericifolia. A, inflorescence branch with bud x 1.5; flower x 2.5; C, androecium x 5 (the stamen at the left is opposite the anteriorsepal); D, adaxial view of stamen opposite
postero-lateralsepal x 5; E, gynoecium x 5; F, fruit x 1. G-K, B. lucida. G, inflorescencebranch
with bud x 1.5; H, androeciumx 5 (the stamen at the left is opposite the anteriorsepal); J, adaxial
view of stamen opposite postero-lateralsepal x 5; K, gynoecium x 5. L, B. woytkowskii,fruit x 1.
M, N, B. amplectens. M, leaf x h; N, fruit x 1. 0, B. rondoniensis, inflorescence branch with
fruits x 1. (A-E, Gates 349; F, Gates 366; G-K, Wurdack & Adderley 43130; L, Woytkowski 7328;
Cordeiro 903.)
0, Cordeiro
34693; O,
903.)
M, N, Hatschbach 34693;
183
longissimo, illo petalo postico opposito brevissimo, ceteris subaequalibus.Antherae loculis 1.0-1.3 mm longis, glabris, connectivis sepalis oppositis 0.8-1.2
mm longis, 0.5-0.8 mm latis, 0.4-0.6 mm profundis,obovatis, illis sepalis anterolateralibus oppositis quam ceteris majoribus,illis petalis oppositis 0.8-0.9 mm
longis, 0.2-0.4 mm latis, 0.2-0.3 mm profundis, anguste oblongis. Ovarium 1.0
mm altum, dense brunneo-sericeum;stylus anticus 2.4-3.5 mm longus, crassus,
apice argute inflexus, plus quam medium pubescens, styli postici 2.0-3.0 mm
longi, graciles, divergentes et petalum posticum versus spectantes, basi pubescentes; stigmatibustruncatis. Fructus ex 3 samaris (vel abortu 1-2) constans,
samara carpophorobrevi latoque usque 1.0 mm longo, nuce globosa, 8-11 mm
alta, 8-10 mm longa, porcata, porcis ex areola radiantibus, sparsim tomentosericea, pilis sessilibus, trabeculaflexuosa 0.4-0.6 mmlonga, ala 51-67 mm longa,
21-25 mm lata, sparsim tomento-sericea nucis similis, marginibussuperiore et
inferiore subparallelisvel parumdivergentibus,appendicula4-10 mm alta et 512 mm longa basi in margineinstructa.
Type. Woytkowski7328, Peru, Pasco, Villa Rica, fr (holotype, MO; isotypes,
F, NY).
Distribution.(Fig. 40). Known from two collections in the Andes of Peru at
1500m, and a single collection from AmazonianPeru at 220 m.
Collected in fruit in June and December.
PERU. Loreto: C. Diaz et al. 788 fr (MICH). Pasco: Woytkowski 7328 fr (F, MO, NY). Junin: C.
Schunke 306 fl (F).
This species is characterisedby its tomento-velutinousabaxialleaf pubescence,

and large fruit. The floweringcollection, Schunke306, shows the characteristic
leaf pubescence but has much smaller leaves than the two fruiting collections,
and the calyx glands and styles are smaller also. Additionalcollections will provide further measurementsfor this species which should indicate whether the
inclusion of Schunke306 in this species is justifiable.
This species is namedfor Felix Woytkowski,whose collections from Peruhave
done much to document the modernflora of that country.
78. Banisteriopsisnigrescens(Adr. Jussieu) B. Gates, comb. nov.
Banisteria nigrescens Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 44. 1832 [1833].
Banisteriafimbriata Grisebach (non Vellozo), Linnaea 13: 191. 1839. Type. Sellow VI it, 304 et
306, Brazil, Sao Paulo, fl (holotype, B destroyed, fragment NY).
Vine, branches appressed-golden-sericeous,glabrous in age; stipules up to 1

mm long, triangular,sparsely sericeous to glabrate.Leaves falcate, with the petiole 7-10 mm long, the lamina 3.9-15.5 cm long and 1.7-5.7 cm wide, often
smallerassociated with the inflorescence, elliptic to narrowlyelliptic, cuneate at
the base, long-acuminateat the apex, with the marginslightly revolute and glandular,the glands minute, numerous,adaxiallysparselyappressed-sericeous,soon
glabrate,abaxially sparsely appressed-sericeous,the trabecula0.3-0.6 mm long,
with the reticulation prominulous adaxially and prominent abaxially. Inflorescence of 4- to 6-floweredumbels borne in the axils of full-sized leaves or in the
axils of reduced leaves borne on short axillarybranches, appressed-golden-sericeous; bracts and bracteoles 1.0-2.0 mm long, broadlyovate, roundedor obtuse
at the apex, sparselyappressed-sericeousto glabrate,abaxiallyerect or appressed,
persistent;pedicels sessile, 15-22 mm long, 0.6-0.7 mm in diameter,very sparsely appressed-golden-sericeous.Sepals 1.8-2.0 mm long and 1.5-2.0 mm wide,
rotund, sparsely sericeous abaxially, glabrous adaxially, the 4 lateral sepals biglandular,projecting1.0-1.5 mm beyond the glands, the glands 1.5-3.0 mm long,
Flora Neotropica
184
TROPAL
A MER
J.""l"_
t.
......|~-
-F
--
.,
.'- ,.
..n
FIG. 40. Distributionof part of Banisteriopsislucida group, species 74-77. Circle, B. lucida;
triangle, B. nutans; closed square, B. peruviana; open square, B. woytkowskii.
1.0-1.8 mm wide, free apically. Petals yellow, fimbriate, sparsely sericeous externally, the 4 lateral petals reflexed between the sepals, with the claw 1.5-2.0
mm long, the limb 7.0-9.0 mm long, 5.0-7.0 mm wide, orbicular, concave, the
antero-lateral petals slightly larger and more concave than the postero-lateral
petals, the posterior petal with the claw suberect, 2.0-2.5 mm long, up to 0.9 mm
wide distally, the limb 6.0-8.0 mm long, 5.0-5.5 mm wide, obovate, plane. Stamens with the filaments connate at the base, erect, those opposite the sepals
subequal, 2.8-3.0 mm long, those opposite the petals subequal, 2.0-2.5 mm long,
thicker than those opposite the sepals, the locules glabrous, 1.2-1.4 mm long, the
connectives papillose apically, those opposite the sepals 1.1-1.4 mm long, 0.60.8 mm wide, 0.6-0.8 mm deep, obovate, those opposite the petals 1.0-1.2 mm
long, 0.4-0.5 mm wide, 0.3-0.5 mm deep, oblong. Ovary 1.3-1.8 mm tall, densely
hairy, the hairs with unequal arms, with the longer arm up to 1.5 mm long; styles
equal, straight and parallel or diverging apically, glabrous or basally hairy, 2.42.8 mm long, 0.2-0.4 mm wide, stigmas capitate. Fruit with short broad carpophore, the nut 7.0-8.5 mm long and 5.0-6.5 mm wide, globose, laterally bearing
numerous ridges radiating from the areole, appressed-golden-sericeous, with a
shallow appendage apically up to 1 mm tall and 3-4 mm along the base, the wing
185
20-25 mm long, 13-16 mm wide, oblong, sparsely appressed-sericeous,the hairs

with the trabecula0.3-0.4 mm long.
Type. A. de Saint Hilaire, Catal. C2, 1186, Brazil, Sao Paulo, near Sao Paulo,
fl (holotype, P; isotypes, F-fragment, P).

Distribution.(Fig. 41). In the Atlantic coastal forest of Sao Paulo and Rio de
Janeiro, Brazil.
Collected in flower in August, November and December, and in fruit in December.
BRAZIL. Rio de Janeiro: Glaziou 3884 fl (C, P); 12483 fl (LE); Herb. Imp. Brazil (leg Gaudichaud)
583 fl (P); 855 (P). Sao Paulo: Brade 12328 fr (R); Burchell 8729-2 fl (GOET); Hoehne s.n. fl (SP);
Martius 517 fl (M).
This species resembles several species in the Banisteriopsislucida groupin its

leaf shape, petiole length and abaxial leaf pubescence. However, it is readily
distinguishedin fruit by its shorter wing with a much shallower appendage. In
flower, B. nigrescens resembles B. nutans in the B. lucida group, although it can
be distinguishedby its sparser, grey, pedicel pubescence, sparser petal pubescence, its erect filaments with those opposite the petals subequal and always
shorterthan those opposite the sepals, and its equal styles.
There are no modern collections of this species. The forest around Sao Paulo
has been destroyed, but perhaps this species persists in the forests of Rio de
Janeiro.
79. BanisteriopsiscachimbensisB. Gates, sp. nov.
Frutex usque 1.5 m, rami dense luteo-tomento-sericei.Stipulaeminutae. Folia
opposita, suboppositavel ternata, petiolo 6-9 mm longo, dense aureo-tomentosericeo, eglanduloso, lamina 4.4-9.6 cm longa, 1.9-4.9 cm lata, interdumin inflorescentia parviore, ovata vel anguste elliptica, basi cordata, apice breviacuminata, margineplana et glandulis minutis instructa praesertimin acumine,
adaxialiter sparsim appresso-sericea, pilis aureis, trabecula 1.0-1.2 mm longa,
abaxialiter dense albo- vel luteo-tomentosa, venis venulisque prominulis vel
prominentibusadaxialiter, venis lateralibusprominentibusabaxialiter.Inflorescentia axillaris in umbellis 4- vel 6-floriferiscomposita, dense luteo-tomentosericea; bracteisbracteolisque1.5-2.2 mm longis, angusteovatis vel ovatis, apice
acutis, carinatis, dense tomentosis abaxialiter, involucratis, persistentibus;
pedicellis sessilibus, 15-20 mm longis, 0.8-1.0 mm diametro,laxe fulvo-tomentosericeis, pilis T-formibus,trabecula0.7-1.0 mm longa. Sepala 2.0-2.3 mm longa
lataque, late obovata, apice rotundataet inflexa, abaxialiterdense aureo-sericea,
adaxialiterglabra, 4 lateralia 2 glandes gerentia et 1.0-1.2 mm superantia,glandibus 2.0-2.5 mm longis et 1.0-1.4 mm latis. Petala lutea, extus sericea, lacerata
denticulatave, 4 lateraliainter sepala patentia, ungue 2.0-2.2 mm longo, limbo
orbiculariantero-lateralium10.0-12.0 mm longo latoque, cochleato, illo posterolateralium7.0-9.0 mm longo latoque, plano, petalo postico ungue erecto 1.8-2.0
mm longo, usque 1.0 mm lato ad apicem, limbo usque 8.0 mm longo et 7.5 mm
lato, late obovato. Staminafilamentiscrassis erectis, illis sepalis oppositis 3.03.2 mm longis, illis petalis oppositis 2.2-2.5 mm longis; antheraeloculis 1.3-1.6
mm longis, pilosis, connectivis ad apicem papillosis, illis sepalis oppositis 1.01.4 mm longis, 0.8-0.9 mm latis, 0.6-0.7 mm profundis, obovatis, illis petalis
oppositis 1.0-1.2 mm longis, 0.4-0.6 mm latis, 0.4-0.5 mm profundis, anguste
oblongis. Ovarium 1.5-1.6 mm altum, dense sericeum; styli aequales recti vel
parum divergentes ad apicem, 2.0-2.2 mm longi, usque 0.3 mm lati; stigmata
capitata. Fructus non suppetit.
Flora Neotropica
186
Type. J. H. Kirkbride, Jr. & E. Lleras 2957, Brazil, Para, Serra do Cachimbo,
along BR-163, Cuiaba-Santaremroad, km 823.5, fl (holotype, INPA; isotype,

MICH).
Distribution.(Fig. 41). Known only from the Serrado Cachimboon white sand
campina, Para, Brazil, and adjacentMato Grosso.
Collected in flower in Februaryand June.
BRAZIL. Para: Kirkbride & Lleras 2886 fl (MICH); 2957 fl (INPA, MICH); Sick s.n. fl (HB). Mato
Grosso: Prance 25282 bud (MICH).
This species is distinguishedby its hairy, ovate to narrowlyelliptic leaves with

the petiole up to 9 mm long.
80. Banisteriopsishypericifolia(Adr. Jussieu) Anderson & Gates, Contr. Univ.
Mich. Herb. 11: 54. 1975.
Fig. 39.
BanisteriahypericifoliaAdr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 42. 1832[1833].

Banisteriarigida Adr. Jussieu, Ann. Sci. Nat. Bot., S6r. 2, 13: 284. 1840.Type. Vauthier448,
Brazil, MinasGerais, Tejuco, fl (holotype, P-JU; isotypes, F, fragment,G, P).
Much-branchedshrub to 1 m, or shrublet with wand-like stems, sometimes

with twiningtips, to 2 m; young branchesterete, at first sparsely sericeous, soon
glabrous, green, drying black, old branches pale brown, glabrous. Stipules minute, triangular.Leaves opposite or 3 or 4 at a node, the petiole 1-2 mm long,
glabrous, eglandular,the lamina (1.0-)2.5-6.6(-9.0) cm long, (0.5-)1.2-6.4(-7.5)
cm wide, ovate to rotund, truncateto cordate at the base, short acuminateat the
apex, with the marginrevolute and bearingnumerous small glands towards the
apex, glabrous on both sides, with the veins prominenton both sides. Inflorescence of axillary4- or 6-floweredumbels or condensed racemes, glabrous;bracts
and bracteoles 1.2-2.0 mm long, triangular,glabrous, involucrate, persistent;
pedicels sessile, 15-25(-30) mm long, 0.6-0.8 mm in diameter,glabrous. Sepals
2.2-3.0 mm long, 1.6-2.0 mm wide, elliptic, sparselysericeous abaxially,glabrous
adaxially, the 4 lateral sepals biglandular,projectingup to 2.0 mm beyond the
glands, the glands 1.2-2.7 mmlong, 1.0-1.6 mm wide, free apically.Petals yellow,
externally sericeous, lacerate, orbicular,the 4 lateralpetals reflexedbetween the
sepals, the claw 2.0-4.0 mm long, the limb 7.5-11.0 mm long and wide, plane,
the antero-lateralpetals slightly largerthan the postero-lateralpetals, the posterior petal with the claw erect, 3.0-3.4 mm long, up to 0.8 mm wide, fleshy, the
limb 6.5-9.0 mm long and wide. Stamens with the filamentserect, those opposite
the sepals 2.6-3.8 mm long, subequal,those opposite the petals 2.0-2.6 mm long,
subequal, those opposite the postero-lateralpetals slightly longer than those opposite the other petals; anthers reflexed with the locules glabrous, 1.2-1.8 mm
long, the connectives papillate, those opposite the sepals 1.0-1.4 mm long, 0.60.8 mm wide, 0.5-0.7 mm deep, obovate, those opposite the petals 0.9-1.2 mm
long, 0.3-0.6 mm wide, 0.3-0.6 mm deep, oblong. Ovary 1.0-1.4 mm tall, densely
hairy apically, styles straightand parallelor slightly diverging,2.2-3.0 mm long,
equal, stigmas capitate. Fruit of 3 samaras,the carpophoreup to 2 mm long and
1 mm wide, the nut 5-7 mm tall, 4-6 mm long, globose, sparsely sericeous, with
a series of ridges radiatingfrom the areole forming crests distally, bearing a
shallow appendageon the upperedge up to 2 mm tall and 5-7 mm along the base,
the wing 16-29 mm long, 8-16 mm wide, sparsely sericeous to glabrate, oblong,
with the base of the wing curving in to meet the nut one-thirdof the perimeter
from the base of the areole.
Type. A. de Saint-Hilaire Catal. B], 1691, Brazil, Minas Gerais, Sobrado, fl
fr (holotype, P; isotypes, F, fragment,P).
187

,
TROFP
CALEC
*w--_
_.,
,,a o.mapn
,: ,-?
M
-OKfMm
Ul--- ll
1l
'.
m.! m--l--M
; 4- F
-"--
FIG. 41. Distribution of part of Banisteriopsis lucida group, species 78-83. Open square, B.
nigrescens; closed square, B. cachimbensis; closed circle, B. hypericifolia; open circle, B. virgultosa;
open triangle, B. amplectens; closed triangle, B. rondoniensis.
Distribution. (Fig. 41). On white sand or crystalline outcrops in the Serra do

Espinhago in Bahia and Minas Gerais, Brazil.
Collected in flower in January to June, and in fruit in January to April, and
August.
BRAZIL. Bahia: Fryxell 1960 fl fr (NY); Hatschbach 39499 fl (MICH). Minas Gerais: Anderson
8532 fl fr (MICH, NY); 11533 (MICH); 11548 fl (MICH); 11550 fl fr (MICH); 11603 fl (MICH); 35190
fl (MICH, NY); Gates 349 fl (MGM, MICH); 366 fl fr (MGM, MICH); Glaziou 18935 fl fr (BR, C,
G, NY, P); 18936 fl fr (C, P); Irwin et al. 21824 fl fr (MICH, NY); 22288 fl (MICH, NY); 28319 fl
(MICH, NY); Maguire et al. 49165 fl fr (MICH, NY); Mexia 5804 fl (MC); Santos & Castellanos
24204 fl fr (NY); Schwacke 8377 fl (NY); Tschudi s.n. fl (W).
Common name. Brazil: "margadin."

This species occurs in two different habits: that of a low, much-branched bushy
shrub with small leaves and few flowers, distinguished in the past as B. rigida,
and that of a subshrub, with unbranched, wand-like stems up to 2 m tall, often
twining at the tip, with larger leaves, often ternate, and many flowers. This is the
form of B. hypericifolia which is most commonly collected as it is very showy,
and the type is of this form. Personal observations in the field in 1975 in Minas
188
Flora Neotropica
Gerais, Brazil, near Diamantina,with Dr. W. Anderson revealed that the 'hypericifolia' form is the aspect of this species when regeneratingafter damageby
cutting or burning;in a few specimens the wand-like branches were actually
arising within the burnt skeleton of a "rigida"-type plant, from the same rootstock. If undisturbedsubsequently, side branches develop from the bases of the
wand-likestems and by successive branchinggive rise to the 'rigida' form after
a few years.
81. Banisteriopsisvirgultosa (Adr. Jussieu) Anderson & Gates, Contr. Univ.
Mich. Herb. 11: 54. 1975.
BanisteriavirgultosaAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 284. 1840.
Banisteria pubipetala var. virgultosa (Adr. Jussieu) Grisebach in Martius, Fl. bras. 12(1): 45. 1858.
Liana, young branches appressed-sericeous, old branches glabrate, greyishbrown with numerous raised pale lenticels. Stipules minute, triangular.Leaves
coriaceous, with the petiole 1-3 mm long, appressed-golden-sericeous,eglandular, the lamina 3.5-7.5 cm long, 2.9-5.4 cm wide, sometimes smaller on vining
shoots, orbicularto broadlyovate, cordateat the base, obtuse to short-acuminate
at the apex, with the marginrevolute and bearing small glands, more numerous
on the acumen,glabrouson both sides or sometimessparselyappressed-sericeous
on the midrib and apex on both sides, the hairs golden, with the reticulation
prominenton both sides. Inflorescence axillary, of 4- or 6-floweredumbels, appressed-sericeous, the hairs golden; bracts and bracteoles 1.0-1.5 mm long, narrowly triangular,cymbiform,appressed-sericeousabaxially, erect or appressed,
persistent. Pedicels sessile, 12-23 mm long, 0.6-0.8 mm in diameter, sparsely
appressed-sericeous.Sepals 1.8-2.5 mm long, abaxiallygolden-sericeous, adaxially glabrous, the anterior sepal 1.6-1.8 mm wide, the 4 lateral sepals 2.2-2.8
mm wide, biglandular,projecting0.8-1.2 mm beyond the glands, the glands 2.03.0 mm long, 1.6-1.8 mm wide, free apically. Petals yellow, externallysericeous,
the 4 lateral petals reflexed between the sepals, with the claw 2.0-2.5 mm long,
the antero-lateralpetals with the limb 9.0-11.0 mm long, 8.0-9.5 mm wide, elliptic, cochleate, denticulate, the postero-lateralpetals with the limb 6.0-7.5 mm
long and wide, orbicular,plane, lacerate, the posteriorpetal with the claw erect,
up to 4 mm long and 0.8 mm wide distally, fleshy, the limb up to 7.5 mm long
and wide, obovate, lacerate, plane, the basal teeth sometimesglandular.Stamens
with thick erect filamentsjoined basally, those opposite the sepals 3.2-3.6 mm
long, those opposite the petals 2.2-2.6 mm long; anthersreflexed with the locules
1.2-1.4 mm long, glabrous,the connectives papillose apically, those opposite the
sepals 1.1-1.3 mm long, 0.7-0.8 mm wide and 0.7-0.8 mm deep, obovate, those
opposite the petals 1.0-1.2 mm long, 0.6-0.7 mm wide, 0.4-0.5 mm deep, oblong.
Ovary 1.4-1.6 mm tall, densely hairy, the styles 2.6-2.8 mm long, straight or
slightly diverging, 0.2-0.3 mm thick, glabrous or basally strigose, stigmas capitate. Fruit of 3 samaras, each with short broad carpophore,the nut 5-6 mm tall
and long, globose, smooth to reticulate,densely appressed-golden-sericeous,with
the appendageon the upper edge 1-5 mm tall and 3-5 mm along the base, the
wing 22--30 mm long, 12-18 mm wide, red, sparsely appressed-sericeous,the
hairs sessile, with the trabecula0.4-0.6 mm long.
Type. Martius 1761, Brazil, Goias, Chapadade Paranan,near R. Formosa, fl
(holotype, M).
Distribution.(Fig. 41). Known only from a few collections in northernGoias
and adjacentBahia, Brazil.
189
Collected in flower in March,August and Septemberand in fruit in Marchand

August.
BRAZIL. Goias: Martius 1761 fl (M); Prance & Silva 58533 fl fr (MICH, NY). Bahia: Anderson
et al. 36742 fl fr (MICH, NY).
This species is distinguishedfrom Banisteriopsis hypericifoliaby its habit as

a liana, its sericeous pedicels, stems and petioles, and the smooth nut of the
samara.
82. BanisteriopsisamplectensB. Gates, sp. nov.
Fig. 39.
Frutex pauciramosus,rami glabri, lenticellis numerosis instructi. Stipulae minutae. Folia coriacea, sessilia, lamina 5.5-10.6 cm longa, 5.2-9.2 cm lata, interdum minorein inflorescentia,orbicularivel late ovata, basi valde cordatavel amplexicauli, apice brevi-acuminata,margine plana et glandulis minutis instructa
praesertimin acumine, utrinqueglabraet venis venulisqueprominentibus.Inflorescentia axillarisin umbellis 4- vel 6-floriferiscomposita, glabra;bracteis bracteolisque 1.2-1.5 mm longis, anguste triangularibus,glabris, involucratis,persistentibus; pedicellis sessilibus, 15-20 mm longis, 0.4-0.6 mm diametro, glabris.
Sepala 2.0-2.2 mm longa, 1.5-2.0 mm lata, late elliptica, apice rotundata, abaxialitersericea, adaxialiterglabra,4 lateralia2 glandes gerentiaet usque 1.2 mm
superantia, glandibus 2.0-2.5 mm longis, 1.0-1.5 mm latis. Petala lutea, extus
sericea, dentata, 4 lateralia inter sepala patentia, ungue usque 2.5 mm longo,
limbo 9.0-9.5 mm longo, 7.5-8.0 mm lato, rotundato, cochleato, petalo postico
ungue erecto 3.0-3.5 mm longo, usque 0.8 mm lato distaliter,ad apicem constricto, limbo 7.0 mm longo latoque, orbiculari,plano, reflexo. Stamina filamentis
crassis, erectis, illis sepalis oppositis 2.8-3.0 mm longis, subaequalibus,illis petalis oppositis 2.0-2.2 mm longis, subaequalibus;antheraereflexae, loculis sparsim pilosis, 1.2-1.6 mm longis, connectivis ad apicem papillosis, illis sepalis oppositis 1.2-1.3 mm longis, 0.6-0.7 mm latis, 0.5-0.6 mm profundis,obovatis, illis
petalis oppositis 1.0-1.1 mm longis, 0.6 mm latis, 0.2-0.3 mmprofundis,oblongis.
Ovarium1.5-1.8 mm altum, dense sericeum, styli recti, 2.6-3.0 mm longi, usque
0.3 mm lati, stylus anticus erectus et postici divergentes;stigmatacapitata.Fructus ex 3 samaris(vel abortu 1-2) constans, carpophoro1-2 mm longo usque 0.5
mm lato, nuce usque 9 mm alta et 8 mm longa, globosa, caperata, sparsim appresso-sericea vel glabrata, pilis trabecula usque 1.0 mm; ala (17-)21-28 mm
longa, (10-)12-17 mm lata, glabra,appendiculain marginesuperiorebasi 1-2 mm
alta et 1-2 mm longa secus basim.
Type. G. Hatschbach34693, Brazil, Mato Grosso, CorregoRancho(Mun. Alto
Araguaia)fl fr (holotype, MBM; isotype, MICH).
Distribution.(Fig. 41). Known only from a few collections in the cerrado of
Mato Grosso.
Collected in flower in May and July and in fruit in July and September.
BRAZIL. Mato Grosso: Hatschbach 34693 fl fr (MBM, MICH); 35026 fr (MICH); Occhioni 7298
fl (MICH).
This species is distinguishedby its large leaves with amplexicaulbase and its
large fruit.
83. BanisteriopsisrondoniensisB. Gates, sp. nov.
Fig. 39.
Frutex volubilis vel liana usque 10 m, ramijuniores dense luteo-tomento-sericei, ramivetustiores demumglabrati,griseo-brunnei.Stipulaeminutae.Folia sub-
190
Flora Neotropica
sessilia, petiolo usque 2 mm longo, tomentoso, eglanduloso, lamina 4.6-7.3 cm

longa, 2.7-5.4 cm lata, in inflorescentiaparviore, elliptica vel late elliptica vel
rotundata, basi cordata, apice brevi-acuminata,margine plana et glandulis instructapraesertimin acumine, adaxialitersparsimaureo-tomentosavel glabrata,
pilis flexuosis, pagina polita, abaxialiterdense tomentosa, venis venulisque prominulis adaxialiter, venis lateralibus abaxialiter prominentibus. Inflorescentia
axillaris, in racemis 6- usque 12-floriferisparibusoppositis composita, aureo-tomento-sericea;bracteis bracteolisque 1.0-2.5 mm longis, anguste ovatis, apice
acutis, carinatis, involucratis, persistentibus, sparsim vel dense tomentosis abaxialiter; pedicellis sessilibus, 14-22 mm longis, 0.5-0.6 mm diametro, aureotomento-sericeis. Sepala 2.0-2.2 mm longa, 1.5-2.0 mm lata, abaxialiteraureosericea, adaxialiter glabra, omnia biglandulosavel interdum sepalum anticum
uniglandulosum,usque 1.0 mm superantiaglandes, glandibus 1.8-2.3 mm longis
et 0.8-1.2 mm latis. Petala lutea, extus sericea, fimbriata,4 lateraliainter sepala
patentia, ungue 1.5-2.0 mm longo, limbo 8.5-9.5 mm longo, 6.5-8.0 mm lato,
rotundata, cochleato vel concavo, 2 anterioribusquam 2 posterioribusparum
majoribuset concavioribus,petalumposticum ungue erecto usque 2.5 mm longo
et 0.8 mm lato ad apicem, limbo usque 7.5 mm longo et 5.5 mm lato, elliptico,
fimbriisbasalibus interdumglandulosis. Stamina filamentis crassis, erectis, illis
sepalis oppositis 2.4-2.8 mm longis, subaequalibus,illis petalis oppositis 1.4-2.4
mm longis, subaequalibus,illis petalis postero-lateralibusoppositis quam ceteris
majoribus;antherae reflexae, loculis 1.3-1.6 mm longis, pilosis, connectivis ad
apicem papillosis, illis sepalis oppositis 1.1-1.2 mm longis, 0.5-0.6 mm latis, 0.40.6 mm profundis, obovatis, illis petalis oppositis 1.0 mm longis, 0.3-0.4 mm
latis, 0.3-0.4 mm profundis, auguste oblongis. Ovarium 1.5 mm altum, dense
sericeum; styli recti paralleliquevel divergentes apicem versus, stylus anticus
2.0-2.6 mm longus, usque 0.2 mm latus, postici 1.6-2.0 mm longi usque 0.2 mm
lati; stigmata capitata. Fructus ex 3 samaris (vel abortu 1-2) constans, samara
carpophorousque 1.5 mm longo et 0.2-0.4 mm lato, nuce 4.0-5.0 mm alta, 3.03.5 mm longa, globosa, caperata, dense aureo-appresso-sericea,ala 14-18 mm
longa, 6-9 mm lata, sparsim aureo-appresso-sericea,pilis sessilibus, trabecula
0.3-0.4 mm longa, appendiculain marginesuperiorebasi 1-2 mm alta et 3-4 mm
longa secus basim.
SubbaseProjecto
Type. M. R. Cordeiro903, Brazil, Rondonia,Guajara-Mirim,
RADAM, Estrada do Palheta FronteiraBrasil-Bolivia, Ponto 23, SC-20-VB, fr
(holotype, IAN; isotype, MICH).
Distribution.(Fig. 41). Known only from Rondoniain Brazil.
Collected in flower in Januaryand April and in fruit in April.
BRAZIL. Rond6nia: Cordeiro 902 fl (MICH); 903 fr (IAN, MICH); Kuhlmann 2059 fl (R, RB).
This species is distinguishedby its tomentose leaf pubescence and by its small
fruits and flowers borne in axillaryracemes of 6-12 flowers, not "umbels" of 46 flowers. Banisteriopsiscachimbensis in this group also has tomentose leaf pubescence, but its leaves are petiolateand not subsessile. Banisteriopsisrondoniensis is unusualin that the anteriorsepal is usually glandular.
The Banisteriopsis pubipetala group (species 84-88)
This group consists of one widespread,very variablespecies, and four species

with more restricted distribution.The species of the group share the following
characters:they are all vines; leaves with minute glands on or near the margin;
flowers axillary, borne in four-floweredumbels or condensed racemes of up to
191
16 flowers, axillary;petals yellow, externallysericeous, fruit with the nut bearing

two or three lateral winglets parallelto the areole.
84. Banisteriopsispubipetala(Adr. Jussieu) Cuatrecasas, Ciencia (Mexico) 23:
142. 1964.
Fig. 42.
Banisteria pubipetala Adr. Jussieu in Saint Hilaire, Fl. bras. mer. 3: 41. 1832 [1833].
Banisteria spruceana Grisebach in Martius, Fl. bras. 12(1): 45. 1858. Types. Spruce s.n. (=Pl.
Spruc. 240), Brazil, Caripi, near Para (=Belem), fl (lectotype, GOET; isolectotypes, C, G,
GH, M, MG, NY, P (=no. 156), W). Gardner 2499, Brazil, Piaui, Urucuai, fr (syntypes,
BM, GH). Martius s.n., Brazil, near Ega, fl (syntype, M).
Banisteria constricta Grisebach in Martius, Fl. bras. 12(1): 46. 1858. Type Spruce s.n. (=PI.
Spruc. 1727), Brazil, Amaz6nas, Barra, fr (holotype, GOET; isotypes, F, G, GH, M, P, W).
Banisteria pubipetala Adr. Jussieu var. oblongata Grisebach, Vidensk. Meddel. Dansk. Naturhist. Foren. Kj0benhavn: 129. 1875. Type. Warming s.n. (1022), Brazil, Minas Gerais, Lag6a
Santa, fl (holotype, C).
Banisteria pubipetala Adr. Jussieu var. scandens Grisebach, Vidensk. Meddel. Dansk. Naturhist.
Foren. Kj0benhavn: 129. 1875. Type. Warming s.n., Brazil, Minas Gerais, Lagoa Santa, fl
(holotype, C? n.v.)
Banisteria pubipetala Adr. Jussieu f. constricta (Grisebach) Niedenzu, Ind. Lect. Lye. Brunsb.:
20. 1901.
Banisteria pubipetala Adr. Jussieu f. spruceana (Grisebach) Niedenzu, Ind. Lect. Lye. Brunsb.:
20. 1901.
Banisteria pearcei Rusby, Bull. New York Bot. Gard. 4: 334. 1907. Type. Bang 2811 pp., Bolivia,
fl fr (holotype, NY; isotype, NY). (Most collections of this number are Banisteriopsis muricata (Cav.) Cuatr.)
Banisteria pubipetala Adr. Jussieu f. cordata Niedenzu, in A. Engler, Das Pflanzenreich IV,
141: 447. 1928. Types. Hassler (leg. Rojas) 10562, Paraguay, Sierra de Amambay, fl (lectotype, G; isolectotypes, G, GH, MO, NY, P, S). Hassler 10833, Paraguay, Sierra de Amambay, fl (syntypes, G, GH, NY, P).
Banisteria pubipetala Adr. Jussieu var. constricta (Grisebach) Macbride, Field Mus. Nat. Hist.,
Bot. 13: 837. 1950.
Liana, or shrub with vining branches, young branches terete, sparsely sericeous, soon glabrate, older branchesglabrous, red-brownor pale, minutelyfissured with numerous eruptive lenticels. Stipules 0.5-1.2 mm long, triangular,
sparsely sericeous or glabrous, sometimes inflexed apically, and situated on a
more or less conspicuous interpetiolarridge. Leaves with the petiole 2-8(-12)
mm long, sparsely sericeous to glabrous, eglandular,the lamina (2.5-)5.1-13.0
(-15.5) cm long, (1.1-)2.1-6.0(-8.3) cm wide, narrowlyto broadly elliptic or obovate, cuneate to cordate at the base, short- to long-acuminateat the apex, with
the marginflat to slightly revolute, bearing minute glands on or just within the
margin,more numeroustowards the apex, glabrouson both sides with 4-8 pairs
of main lateral veins, the reticulationprominulousto prominenton both sides.
Inflorescence axillary, of 2-3 condensed racemes of 2-3(-5) pairs of flowers or
cymes of 3-5 such condensed racemes, sparsely appressed-sericeousto glabrate;
bracts and bracteoles 0.8-1.6(-2.0) mm long, triangular, sparsely sericeous
to glabrateabaxially, erect or appressed, persistent, pedicels sessile, 10-22(-26)
mm long and 0.3-1.0 mmin diameter,wider distally, sparselyappressed-sericeous
to glabrate, rarely tomento-sericeous. Sepals broadly elliptic, rounded at the
apex, 1.4-3.2 mm long, 1.0-2.5 mm wide, golden-sericeous,the 4 lateral sepals
biglandular,the glands 1.5-3.0 mm long, 0.8-2.0 mm wide, free apically, with the
sepals projecting0.4-1.4 mm beyond the glands. Petals yellow, externally sericeous, the 4 lateralpetals reflexedbetween the sepals, the claw 1.0-2.0 mm long,
the limb 7.0-12.0 mm long, 6.0-11.0 mm wide, laciniate, the antero-lateralpetals
largerand more cochleate than the postero-lateralpetals, the posteriorpetal with
Flora Neotropica
192
H''
X,-..n,
--~..-..
:'""':'~
~,--~. ~i:.i-,,.
-~-
....
-'~-''' ~"~
~
~'~~
.~~
?.
*I
FIG.
loer,sd
I?Y/G
42. Ban~~~~~~~~iseipipuieaaB.shneanB.spu.AEB.pbpta.,ifor-
,cecebrnc x2.; wthbu

righ isopst
h neirspl;D
scuk
i.
,la
B.~~~~
4635.)/,
yocu
;H
ri
;E
tmna
h
.;C
nreimx5(h
ri
,F .spu,futx1
.(-,Gts32;F
eer
iw
,H
86 ,$hneV
the
clweet
hc n lsy .-. mln,aial
osrce,telm
fmrit,hebslfmrasotms
3.-.mlnanwide
oricua,
fiaetgaros1.8 -5
gland-tipe.Sae
sw
.tth
mmln,thsp
poit
te tle
(.e ppsiehateio
spl
ndth
oser-ltra
ptas
FIG. 4. Banisteriops. pubi

.
a B. schu i ad , B. s,
infloresp
cence branch with bud x 2.5; B, flower, side view, x 2.5; C, androeciumx 5 (the stamen at the
rightis opposite the anteriorsepal); D, gynoecium x 5; E, fruit x 1; F, B. sepium, fruit x 1; G, H,
B. schunkei. G, leaf x h; H, fruit x 1. (A-E, Gates 323; F, Pereira 5866; G & H, Schunke Vigo
4635.)
the claw erect, thick and fleshy, 3.0-5.0 mm long, apically constricted, the limb
3.5-8.5 mm long and wide, orbicular, fimbriate, the basal fimbriae sometimes
gland-tipped. Stamens with the filaments glabrous, 1.8-5.0 mm long, those opposite the styles (i.e. opposite the anterior sepal and the postero-lateralpetals)
193
and the postero-lateralsepals longest, that opposite the posterior petal shortest,
those of the 3 posterior stamens usually inflexed between the posterior styles,
with the locules 0.8-2.0 mm long, sparselyto densely pilose, those of the stamens
opposite the 4 lateral sepals divergingdistally, the connectives yellow, those of
the stamens opposite the sepals 0.8-1.8 mm long, 0.5-1.0 mm wide and 0.5-0.8
mm deep, obovate to narrowly obovate, papillate apically, those opposite the
petals 0.7-1.4 mmlong, 0.3-0.7 mm wide, and 0.2-0.6 mm deep, narrowlyoblong.
Ovary 1.0-1.5 mm tall, apically densely pilose, the hairs white and straightin
fresh material,brown and flexuous in dry material,with the arms very unequal,
one arm up to 0.5 mm long and the other up to 3.0 mm long, the styles 3.0-5.0
mm long, the anterior style usually longer and thicker than the posterior ones,
directed anteriorly, sometimes inflexed apically, bearinghairs like those on the
ovary for up to half of its length, the posterior styles diverging, directed posteriorly, lyrate at the base, hairy in the basal third, stigmas capitate. Samarawith
carpophoreup to 2.0 mm long and 1.0 mm wide, the nut 3.5-6.0(-11.0) mm tall
and 3.5-5.0(-9.0) mm long, bearing2-3 lateralwings on each side parallelto the
areole 1-3(-5) mm high, glabrate, bearing apically an appendage 1.5-4.0(-7.0)
mm high, and 2.5-4.0(-9.0) mm wide at the base, with the locule of the nut
sphericaland smooth inside, the dorsal wing 20-40(-65) mm long, 9-16(-23) mm
wide, often narrowerat the base, glabrate,the hairs appressedwith the trabecula
0.4-0.6 mm long, brightred or yellow.
Type. A. de Saint Hilaire, Catal. Cl, 762, Brazil, Goias, Villa Boa, fl (lectotype, P; isolectotypes, P). Saint Hilaire s.n., Brazil, Minas Gerais, Formigas, fl
(syntypes, P). Saint Hilaire, Catal. B1, 1617, Brazil, Minas Gerais, near Chapada, fr (syntypes, F, fragment,P).
Distribution.WidespreadthroughoutBrazil, extendingnorthto Colombia,west
to Peru and Bolivia, and south to Paraguay.
Collected in flower and fruit in every month of the year.
Representative collections. COLOMBIA. Norte de Santander: Cuatrecasas 13301 fl (F, US). Valle:
Cuatrecasas 15043 fl (F, US). Meta: Hermann 11115 fr (COL, US); Idrobo & Schultes 1329 fr (US);
Philipson et al. 1719 fr (F, NY, US). Vaupes: Zarucchi et al. 1817 fr (MICH). Putumayo: Ewan
16717 fr (NY, US). No other locality: Mutis 2365 fr (US); 2368 fl (US); 2396 fl (US); 2411 fl (US);
2418 fl (US); 2523 fr (US). VENEZUELA. Tachira: de Bruijn 1364 yng fr (MICH). PERU. San
Martin: Spruce 4272 fr (BR, G, GH, MG, NY, W); Ule 6439 fr (G, MG); LI. Williams 5426 fl (F, G);
6157 fr (F); Woytkowski 7259 fr (MICH). Junin: Woytkowski 5581 fl (C, G, MO); 5879 fl (G, MO);
5909 fl fr (G, MO); 7392 fl (M). BRAZIL. Roraima: Black 51-13610 fl (IAN). Amaz6nas: Chagas 1437
fl (IAN, MG), Frbes 20638 fr (IAN, NY); Litzelburg 22822 fl (M, R); Prance et al. 14615 fl (MG,
MICH, P, U); Ule 8892 fl (G, MG). Para: Black 52-15550 fl fr (IAN); Ducke 8587 fl fr (MG); Egler
827 fl (IAN, MG); Prance et al. 1628 fr (IAN, MICH, NY, U); N. T. Silva 2726 fl (IAN). Maranhao:
Black 54-16659 fl (IAN); Devereux 15 fl fr (US); Eiten & Eiten 5402 fl (SP); 5325 fl (SP); Lisboa 2517
fl (MG); Prance & Silva 58607 fl (MICH, U). Piaui: Jobert 958 fl (P); 1185 fl (P); Lutzelburg 336 fl
(M); Snethlage 615 fl (F, NY, fragment ex B). Ceara: Dahlgren 916 fl fr (F, P); 959 fl (F); Drouet
2523 fr (F, GH, S); Ducke 2484 fl fr (IAN); Lofgren 270 fr (R, S). Rio Grande do Norte: Wurdack
B-156 fl fr (GH, NY). Rond6nia: Prance et al. 5422 fl (MG, MICH, NY, U); Rodrigues, Coelho 4312
fl (IAN, SP); J. F. Silva 67 fr (IAN). Mato Grosso: Anderson 11308 fl fr (MICH); Argent in Richards
6448 fl (IAN, NY); A. Ferreira et al. R1181 fr (IAN, MO, NY); Fonseca 1064 fl (NY, UB); Hatschbach 21853 fl (MICH, P); Irwin et al. 6591 fl (IAN, S, SP); Martius 658 (=P. Manso 119) fl (BR, G,
M, MO, P, W); Prance et al. 59447 fr (MICH, U); Ratter et al. R161 fl fr (MO, NY); Regnell (leg
Malme) II. 1915 fl (G, S). Goias: Burchell 7339 fl (GOET, P); Glaziou 20758 fl (BR, G, P, NY,
fragment ex B); Irwin et al. 10838 fl (MO, NY, S, SP); Macedo 3901 fl (MO, SP), Pohl 1516d. 2330
fl (F, W). Distrito Federal: Duarte et al. 104 fl (UB); 113 fr (HB, U); Heringer 7930 fl (SP); Irwin et
al. 6167 fl fr (MICH); 9284 fl (IAN, MO, NY, S, SP, U). Bahia: Anderson 11751 fr (MICH); 36484
fl (NY); 37150 fl (MICH); Irwin et al. 30773 fl fr (MICH, UB); 31371 fl (MICH). Minas Gerais:
Barreto 7724 fl fr (F); Gardner 4475 fl (BM); Hatschbach 28066 fl fr (C, P); Macedo 2505 fl fr (S,
SP); Regnell III. 312 fl (BR, C, M, P, R, U, US, W); Warming s.n. fl (C); L. O. Williams & Assis
194
Flora Neotropica
7535 fl (GH, MO, US). Rio de Janeiro: Armond 80 fr (R); 82 fl (R); Glaziou 7679 fl (C, P); 10383 fl
fr (C, P). Sao Paulo: Felippe 111 fl (SP); Hatschbach & Kummrow 34814 fl fr (MICH); Hemmendorff
145 fr (S); Lofgren 504 fl (S); Mosen 365 fl (S); Raben 141 fl (C). Parana: Braga & Lange 127 fl
(MICH, NY, P); 128 fl fr (P, S); Hatschbach 16987 fl (P, US); 21584 fl fr (MICH, P); Lindeman &
Haas 986 fr (U). BOLIVIA. Bang 2457 fl fr (F, NY); Brooke 5703 fl (F, NY); Buchtien 3798 fl (GH,
NY); Krukoff 10048 fl (F, G, GH, MO, S, U); 11234 fl fr (F, G, GH, MO, NY); Rusby 515 fr (F,
NY); Steinbach 2819 fl (F, G, NY-fragment ex B); 3018 yng fr (GH); 7305 fr (MO, S); R. S. Williams
1511 fl fr (NY). PARAGUAY. Hassler 10562 fl (G, GH, MO, NY, P, S); 10562a fr (G, GH, P); 10833
fl (G, GH, NY, P).
Common names. Brazil: "Cipo preto," "cipo de rego," "crista de gallo."
This is an enormously variable species, showing variation in the form and

pubescence of the inflorescence, the size of the bracts and bracteoles, flower
size, petiole length, leaf shape and size, size of the fruit nut, and size and shape
of the fruit wing. However, the variationdoes not appearto correspondwell to
geography. For example, the type of Banisteriaspruceana Grisebachfrom Para,
Brazil, has smallleaves and flowers, but individualswith smallleaves and flowers
have been collected from Sao Paulo and Mato Grosso in Brazil, and from Bolivia.
In some instances most of the leaves of a collection will be small but if older
leaves are representedin the collection these are often much larger, and small
leaf size may be an artifact of collection. The collections from Paraguayhave
sessile leaves with a cordate base and were recognized by Niedenzu as forma
cordata. However, these plants are indistinguishablein other respects and some
specimens from adjacent Mato Grosso and Parana in Brazil also have sessile
leaves with cordate base. There appears to be a continuous gradationbetween
this form and the more typical form, and individualswith cordate leaf base have
been collected in other parts of the range of this species. For these reasons I
prefer not to maintainthe forma.
Some individuals from Bolivia (Brooke 5703; Steinbach 2819; 3018; 7305) show
tomentose pubescence in the inflorescence, and racemes of up to ten flowers; in

these charactersthey approachthe Bolivian specimens of Banisteriopsislongialata. However, in their flower size, leaf size, petiole length, and fruit characters
they resemble typical B. pubipetala. Individualswith largefruits similarto those
characteristicof B. platyptera and B. longialata have been collected in Amazonian Brazil and Colombia. These large-fruitedcollections are furtherunusual
in that their leaf glands lie within and not on the margin.However, since other
collections from these places have smallerfruits whose size lies within the usual
range for B. pubipetala, I prefer not to segregate these individuals.
Most of the collections for this species have been made in the Planaltoregion
of Brazil, and I think that this reflects the relative ease of collecting plants in the
more open vegetation of this area compared with collecting in tropical forest.
With more collections from the other parts of the range of this species and with
additionaldata on habit and habitat,perhapsthe variationwithinthis species will
be seen to meaningfullyreflect the geographyor ecology of the plants.
85. Banisteriopsisplatyptera(Grisebach)Cuatrecasas,Webbia 13: 494. 1958.
Banisteria platyptera Grisebach, Linnaea 22: 17. 1849.
Liana, branchesterete, glabrous. Stipules up to 1 mm long, triangular.Leaves

chartaceous with the petiole 3-7 mm long, glabrous or sparsely appressed-sericeous, eglandular,the lamina 4.7-10.4 cm long and 1.8-4.5 cm wide, elliptic or
obovate, cuneate at the base, acuminate at the apex, with the margin slightly
revolute and bearingminuteglands, glabrouson both sides, with the lateralveins
prominulousadaxiallyand prominentabaxially. Inflorescenceaxillary, of 4-flow-
195
ered umbels or 6-floweredracemes, usually 2-3 per leaf axil, brown-appressedsericeous, the bractsand bracteolesup to 1.0 mm long, broadlytriangular,brownappressed-sericeousabaxially, glabrous adaxially, involucrate, persistent; pedicels sessile, 17-25(-36) mm long, 0.6-0.8 mm in diameterin flower, much thicker
(up to 1.5 mm) in fruit, brown-appressed-sericeous.Sepals 2.0-2.4 mm long, 1.82.3 mm wide, roundedat the apex, densely brown-sericeousabaxially, glabrous
adaxially, the 4 lateral sepals each biglandular,the glands 1.8-2.4 mm long and
1.0-1.4 mm wide, the sepals overtoppingthe glands up to 1.2 mm. Petals yellow,
externally sericeous, orbicularto broadly elliptic, dentate, the 4 lateral petals
reflexed between the sepals, the claw 1.0-2.0 mm long, the limb 7.0-9.0 mm long
and 6.0-8.0 mm wide, the antero-lateralpetals largerand more concave than the
postero-lateralpetals, the posteriorpetal with the claw erect, up to 4.0 mm long,
the limb up to 6.5 mm long and 5.5 mm wide, plane. Stamens with the filaments
glabrous, erect, those opposite the sepals and the postero-lateralpetals 2.4-3.0
mm long, those opposite the antero-lateraland posteriorpetals 1.8-2.2 mm long,
the locules 1.0-1.4 mm long, pubescent, the connectives opposite the sepals 0.91.2 mm long, 0.6-0.7 mm wide, 0.4-0.5 mm deep, obovate to broadly oblong,
those opposite the petals 0.8-1.2 mm long, 0.3-0.4 mm wide and 0.2-0.3 mm
deep, narrowly oblong. Pollen 45-50 gtm in diameter. Ovary 1.4-1.5 mm tall,
densely brown-pubescent,the hairs sessile, with very unequalarms, the long arm
up to 1.5 mm long; styles subequal, 2.4-3.0 mm long, stout, divergingfrom the
base, hairy for half their length, stigmasterminal.Fruit with short broad carpophore up to 2.0 mm long and 1.5 mm wide, the nut 11-15 mm tall, 9-11 mm long,
sparsely appressed-sericeous,bearing2 lateral wings on each side parallelto the
areole 3-5 mm high and an appendageon its upper edge 7-12 mm long and 4-10
mm tall, the wing 45-65 mm long and 20-28 mm wide, sparsely appressed-sericeous, the hairs with the trabecula0.2-0.6 mm long.
Type. H. Karsten (4), 55, Venezuela, Perija, fl fr (holotype, GOET;isotypes,
NY, fragmentex B, S).
Distribution. (Fig. 43). Known from only a few collections from wet forest
along the Atlantic coast of South America.
Collected in fruit in October, December, and January.
PANAMA. Gentry et al. 13826 fr (MICH). VENEZUELA. Distrito Federal: Bailey 425 fr (US);
Steyermark 91740 yng fr (F, NY). BRAZIL. Pernambuco: Pickel 2906 fr (NY, fragment ex B).
This species has been seldom collected and the only floweringmaterialis that
of the type. Its flowers appearto be very similarto those of Banisteriopsisnutans
in the B. lucida group, althoughin fruit the two species are easily distinguished.
Perhapsadditionalfloweringcollections of B. platyptera will make the separation
of these two species in flower easier. Banisteriopsisplatyptera is characterized
by its large samaras with a large alulate nut and large robust wing. It is distinguished from B. longialata which has similarlarge samaras,by its small leaves,
short petiole, flowers usually in four-floweredumbels and its stout straightsubequal styles. It is distinguishedfrom B. pubipetala by its large samaras, dense
rust-brownpubescence in the inflorescence,flowers usually in four-floweredumbels, and its stout, straight,subequalstyles.
86. Banisteriopsislongialata(Niedenzu) B. Gates, comb. nov.
Banisteria longialata Niedenzu, Ind. Lect. Lyc. Brunsb.: 18. 1901.
Banisteria rusbyana Niedenzu, Ind. Lect. Lyc. Brunsb.: 18. 1901. Type. H. H. Rusby 2171,
Bolivia, Beni River, fl (holotype, B destroyed, NY fragment; isotypes, F, GH, MICH, MO,
NY, US).
196
Flora Neotropica
Banisteria bopiana Rusby, Mem. New York Bot. Gard. 7: 274. 1927. Type. H. H. Rusby 663,
Bolivia, Bopi River valley, fr (holotype NY; isotype, MICH).

Heteropterysbopiana (Rusby)Niedenzu, in A. Engler, Das PflanzenreichIV, 141: 377. 1928.
Banisteriopsisrusbyana(Niedenzu)Morton,J. Wash. Acad. Sci. 21: 486. 1937.
Liana, branchesglabrous, stipules minute. Leaves coriaceous, with the petiole

(6-)10-27 mm long, glabrous or sparsely appressed-sericeous, eglandular,the
lamina (8.0-)12.0-25.5 cm long and (2.4-)4.5-12.0 cm wide, smaller associated
with the inflorescence, elliptic to obovate, cuneate to obtuse at the base, longacuminate to cuspidate at the apex, with the marginplane to slightly revolute,
bearing minute glands on or rarelyjust within the marginabaxially, the glands
more numerous on the acumen, glabrous on both sides, the reticulationprominulous to prominenton both sides. Inflorescenceaxillary,of panicles or racemes,
1-3 per leaf axil, the ultimatebranchesbearing4-8 pairs of flowers, sparsely to
densely appressed- or tomento-sericeous, the hairs brown or golden, the bracts
and bracteoles 1.0-2.5 mm long, broadly triangular,involucrate, persistent, appressed-sericeous abaxially, glabrous adaxially; pedicels sessile or subsessile,
12-22 mm long, 0.5-0.8 mm in diameter,densely pubescent. Sepals 2.0-2.5 mm
long, 1.8-2.2 mm wide, rounded at the apex, densely golden-sericeous, the 4
lateral sepals biglandular,overtoppingthe glands 0.6-1.2 mm, the glands 1.2-2.5
mm long and 0.7-1.2 mm wide. Petals yellow, densely sericeous externally, dentate, the 4 lateral petals reflexed between the sepals, the claw 1.0-2.0 mm long,
the limb 7.0-12.0 mm long and 6.0-11.0 mm wide, orbicular,the antero-lateral
petals largerand more concave than the postero-lateralpetals, the posteriorpetal
with the claw erect, 3.0-4.0 mm long, up to 0.8 mm wide, apically constricted,
the limb 4.5-5.5 mm long and 4.0-5.5 mm wide, orbicularto obovate, plane.
Stamenswith the filaments1.5-4.5 mm long, glabrous,those opposite the 3 styles
longest, that opposite the anteriorsepal curved inwards apically, those of the 3
posterior stamens inflexed between the posterior styles, the locules 0.9-1.4 mm
long, sparsely to densely pubescent, the connectives opposite the sepals 0.8-1.4
mm long, 0.5-1.0 mm wide, 0.4-0.8 mm deep, obovate, those opposite the 4
lateral sepals with the locules divergingapically, those opposite the petals 0.71.0 mm long, 0.3-0.5 mm wide, 0.2-0.4 mm deep, oblong. Ovary 1.0-1.8 mm tall,
densely hairy, the anterior style 3.0-5.0 mm long, stout, apically inflexed, the
posterior styles 2.4-4.0 mm long, shorter than the anterior style, divergingand
directedposteriorly. Fruitwith shortbroad carpophore,the nut 7-13 mm tall and
7-10 mm long, sparsely appressed-pubescent,bearingtwo lateral wings on each
side up to 8 mm high, and an appendagealong its upper edge up to 10 mm long
and up to 8 mm tall, the wing (38-)44-55 mm long and 15-20 mm wide, sparsely
appressed-pubescent,more or less parallel-sided,not constricted at the base.
Type. Ruiz & Pavon a. 1784, Peru, Huanuco, Pozuzo, fr (holotype, B destroyed; isotype, F).
Distribution.(Fig. 43). In the Andes in Bolivia and Peru, extending north to
Colombia.
Collected in flower in April, July and October, and in fruit from July to September.
COLOMBIA.Caqueta:Cuatrecasas8985 fl (COL, F, US). PERU. Loreto: Torres110 fl (ECON).
Amazonas: Woytkowski 8066 fr (MICH). Huanuco: Ruiz & Pavon a. 1784 fr (F). BOLIVIA. Krukoff
10022 fl fr (F, MICH, MO, NY, US); 10590 fl fr (F, MICH, MO, NY, US); Rusby 663 fr (MICH,
NY); 2171 fl (F, GH, MICH, MO, NY, US).
This species is distinguishedfrom Banisteriopsispubipetala by its large leaves

with long petiole, and its large samaraswith a large nut and a stout wing. There
197
is some variationamong the collections that I have assigned to this taxon. The
type has appressed,brownpubescence in the inflorescenceand the ultimateflower-bearingbranchesare short condensed racemes of four to five pairs of flowers.
The inflorescences of the other Peruvianspecimens and of the Colombianspecimen are similarto that of the type. However, the Colombianspecimenis unusual
in bearing the minute leaf glands within the marginon the abaxial leaf surface
and not on the margin.The Bolivian specimens exhibit loose tomentose pubescence in the inflorescence, and the ultimateflower-bearingbranches are longer,
ratherlax, racemes of six to eight pairs of flowers.
87. Banisteriopsiserianthera(Adr. Jussieu) B. Gates, comb. nov.
BanisteriaeriantheraAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 284. 1840.
BanisteriaplatypteraGrisebachf. grandifoliaNiedenzu, Verz. Vorles. Konig. Ak. Braunsb.:21.
1912. Type. Ule 6705, Peru, Loreto, S. Antonio de Cumbaso, fr (holotype, B destroyed;
isotypes, G, MG).
Liana, young branches flattened, appressed-sericeous,the hairs golden, older

branchesterete, sparsely sericeous to glabrate.Stipules up to 2 mm long, goldensericeous abaxially, projectingperpendicularto the stem, inflexed and glabrate
apically, with their bases greatly widened and sometimes fusing laterallyto form
an interpetiolarridge. Leaves chartaceous, with the petiole (7-)12-20(-27) mm
long, appressed-sericeous,the hairs golden, eglandular,the lamina 8.2-23.2 cm
long and 4.3-12.2 cm wide, elliptic, obtuse at the base, long-acuminateat the
apex, with the margin flat, bearing minute glands, the glands more numerous
towardsthe apex, adaxiallyglabrateexcept the midrib,abaxiallysparselygoldensericeous, the hairs sessile with the trabecula 0.1-0.2 mm long, with the veins
more densely sericeous, with 8-18 pairs of lateral veins extending almost to the
marginbefore formingloops, with the primaryand secondary veins prominulous
adaxially and prominentabaxially. Inflorescence densely golden-sericeous, axillary, of 4- to 6-floweredumbels, 2 to 3 per leaf axil, the umbel-bearingpeduncles
7-15 mm long, bracts and bracteoles 1.0-1.5 mm long, obtuse at the apex, cucullate, densely golden-sericeousabaxially, glabrousadaxially, involucrate,persistent, the pedicels sessile, 13-26 mm long, up to 1.0 mm in diameter, and
thicker, up to 2.0 mm in fruit, densely golden-sericeous,glabratein fruit. Sepals
golden-sericeous abaxially, 2.5-3.0 mm long, 2.0 mm wide, the 4 lateral sepals
biglandular,projecting 0.6-0.8 mm beyond the glands, the glands 2.0-3.0 mm
long and 1.2-2.0 mm wide, free apically. Petals yellow, densely sericeous abaxially, the 4 lateralpetals reflexed between the sepals, the claw 1.5-3.0 mm long,
the antero-lateralpetals up to 12.5mm long and 12.0mm wide, orbicular,lacerate,
cochleate, the postero-lateralpetals up to 9.0 mm long and 6.0 mm wide, elliptic,
lacerate, plane to slightly cochleate, the posterior petal with the claw erect, up
to 4.0 mm long and 1.0 mm wide, the limb up to 7.0 mm long and 4.4 mm wide,
obovate, fimbriate,plane. Stamens with the filaments 1.8-5.0 mm long, that opposite the anteriorsepal longest and incurvedapically, that opposite the posterior
petal shortest, those opposite the styles thicker than the others, those opposite
the antero-lateralsepals shorter than those opposite the lateral petals, those of
the 3 posterior stamens inflexed between the posterior styles, the anthers with
the locules densely pilose, 1.0-1.8 mm long, those of the 5 anteriorstamens 1.51.8 mm long, those of the 5 posterior stamens 1.0-1.4 mm long, the connectives
opposite the sepals 1.0-1.6 mm long, 0.6-0.8 mm wide and 0.5-1.0 mm deep,
obovate, those opposite the petals 0.6-1.2 mm long, 0.2-0.4 mm wide and 0.10.4 mm deep, narrowlyoblong. Ovary 1.5 mm long, apically pubescent, anterior
198
Flora Neotropica
style 4.0-6.0 mm long, hairy for up to half its length, apically inflexed, posterior
styles 4.0-5.0 mm long, basally pubescent, diverging at the base and inflexed
apically. Fruit with short broad carpophore,the nut 15-17 mm long and 12-13
mm wide, bearing3 lateral wings on each side parallelto the areole up to 5 mm
high, and an appendageapically 17-18 mm along its base and 10-14 mm high,
the wing 63-75 mm long, 20-30 mm wide, greatly thickenedalong its upperedge,
sparsely appressed-pubescent.
Type. Martiuss.n., Brazil, Amazonas, Japura,fl (holotype, M; isotype, M).
Distribution.(Fig. 43). In tropicalrain forest of AmazonianBrazil and Peru at
low elevations.
Collected in flower in December and in fruit in March.
PERU. Loreto: Ule 6705 fr (G, MG). BRAZIL. Amaz6nas: Fr6es 23731 fl (IAN); 23754 fl (IAN);
Krukoff 6444 bud (BR, G, MICH, MO, U). Rond6nia: Prance et al. 8872 yng fr (MG, MICH, NY).
This species is easily distinguishedby its large leaves with numerousparallel

lateral veins and persistent abaxialpubescence, and its very large fruits.
88. BanisteriopsisschunkeiB. Gates, sp. nov.
Fig. 42.
Liana, ramis glabris. Stipulis minutis. Folia opposita; petiolo (6-)12-17 mm
longo, glabro, eglanduloso, adaxialitercanaliculato;lamina (9.0-)13.5-23.0 cm
longa et (3.5-)7.4-13.0 cm lata, obovata vel elliptica, basi cuneata, apice longe
acuminata,margineplana, subtus prope marginemnumerosasareas glandulosas
circularesellipticasvegerenti, utrinqueglabra,venis venulisquesupraprominulis,
subtusprominentibus.Inflorescentiaaxillaris,floribusin racemis4- vel 6-floriferis
fasciculatis, binis in axilla, sparsimappresso-sericea,pilis trabecularecta 0.2-0.4
mm longa; bracteis bracteolisque0.8-1.2 mm longis, late triangularibus,sparsim
sericeis abaxialiter,glabrisadaxialiter,involucratis,persistentibus;pedicellis sessilibus vel subsessilibus, 15-17 mm longis, 1.0 mm diametro,glabris. Flores non
suppetunt. Sepala 5, abaxialitersparsim sericea, 4 lateralia 2 glandes gerentia.
Fructus ex 3 samaris constans; samaracarpophorousque 1 mm longo et 2 mm
lato, nuce 13-17 mm alta, 11-13 mm longa, glabra, lateraliterutrinque2 alulas
usque 8 mm altas ad areolamparallelasgerenti, et appendicerotundata10-13 mm
longa secus basim, usque 10 mm alta in marginesuperioreinstructa,ala (56-)6073 mm longa, 20-28 mm lata, glabra.
Type. J. Schunke V. 4635, Peru, San Martin, Prov. Mariscal Caceres, Dept.
Tocache Nuevo, Instituto Agropecuariade Tocache, fr (holotype, F; isotypes,
G, GH, MO, NY).
Distribution. (Fig. 43). Known only from the type from high forest in the
groundsat the Instituto de Agropecuariade Tocache, San Martin,Peru.
Collected in fruit in January.
PERU. San Martin: Schunke 4635 fr (F, G, GH, MO, NY).
Common name. Peru: "auca ayahuasca."
This species is distinguishedby its large chartaceousleaves bearing"smeary"

glands withinthe margin,and its largeglabrousfruit. It is namedfor Jose Schunke
Vigo, who has madeby his collections such an importantcontributionto the knowledge of the Peruvianflora.
89. Banisteriopsiscaduciflora(Niedenzu) B. Gates, comb. nov.
Banisteria caduciflora Niedenzu, Ind. Lect. Lyc. Brunsb.: 21. 1901.
199
?.TROPICA...E..CA
TI
"
?--'
n ' rr;~
,,
---,
- ....?!- ' ' T'L'k
i.
..1no.
.-
. ?
"
.........
'
--
marn
'"
'-'- .'
amp no!1_,
FIG. 43. Distribution of Banisteriopsis pubipetala group (excluding B. pubipetala), and of B.

caduciflora, B. krukoffii, B. patula and B. sepium (species 85-92). Closed circle, B. platyptera; open
square, B. longialata; closed triangle, B. erianthera; closed square, B. schunkei; open circle, B.
caduciflora; open triangle, B. krukoffii; half circle, B. patula; closed star, B. sepium.
Liana or vining shrub, branches terete, glabrous. Stipules up to 1.0 mm long,

deltate. Leaves with the petiole 3-7 mm long, glabrous, eglandular,the lamina
4.6-11.4 cm long and 1.3-3.6 cm wide, narrowly elliptic, cuneate at the base,
long-acuminateat the apex, with the margin plane, with 2-4 or more pairs of
glandularareas in the lamina V-2-6 of the distance to the marginabaxially, glabrous on both sides, plane adaxiallyand with the reticulationprominentabaxially.
Inflorescenceaxillary, of up to 3 branchesper axil, of racemes of up to 4 pairs of
flowers, golden-appressed-sericeous;bracts and bracteoles 0.6-1.0 mm long, triangular,sparsely sericeous, erect or appressed, persistent;pedicels sessile, 9-14
mm long, 0.4 mm in diameter,glabrous. Sepals oblong, roundedat the apex, 1.42.0 mm long and 1.0-1.4 mm wide, all eglandularor the anteriorsepal eglandular
or bearing 1 gland and the 4 lateral sepals biglandular,overtopping the glands
0.2-0.6 mm, the glands 1.0-1.6 mm long and 0.6-0.8 mm wide. Petals yellow,
externally glabrous or sparsely sericeous, the 4 lateral petals reflexed between
the sepals with the claw 1.0-2.0 mm long, the limb 4.5-6.0 mm long and 3.0-6.0
mm wide, orbicular, lacerate, with the antero-lateralpetals more concave than
200
Flora Neotropica
the postero-lateralpetals, the posterior petal with the claw erect, up to 3.0 mm
long, with the limb 4.0-4.5 mm long and 3.0-3.5 mm wide, obovate with the basal
teeth gland-tipped.Stamens with the filaments 1.4-3.2 mm long, those of the 3
anterior stamens erect, the next 2 pairs directed posteriorly and those of the 3
posterior stamens inflexed between the posterior styles, antherswith the locules
0.5-0.8 mm long, hairy, the connectives opposite the sepals 0.6-0.8 mm long,
0.5-0.6 mm wide and 0.3-0.4 mm deep, broadlyoblong, those opposite the petals
0.4-0.6 mm long, 0.3-0.6 mm wide and 0.1-0.3 mm deep, oblong. Ovary 1.0 mm
tall, densely hairy, the styles subequaland up to 2.2 mm long, divergingbasally
and inflexed apically, inconspicuouswithinthe dense mass of hairs on the ovary,
the ovary hairs up to 3 mm long, stigmas truncate. Fruit unknown.
Type. Poeppig 2474, Peru, Loreto, Maynas, woods near Yurimaguas,fl (holotype, B destroyed?;isotypes, F, W).
Distribution.(Fig. 43). Known from two collections from the upper reaches of
the Amazon in Peru, and a single collection from the basin of the Rio Madeira
in Rondonia, Brazil.
Collected in flower in July.
PERU. Loreto: Poeppig 2474 fl (F, W). Junin: Killip & Smith 26082 bud (F, NY, US). BRAZIL.
Rond6nia: Vieira et al. 392 fl (MICH).
This species is unusual in its smeary leaf glands. This type of leaf gland is
found only in this species and in Banisteriopsisschunkei. The inflorescenceand
leaves of Banisteriopsis caduciflora most closely resemble those of the B. pu-
bipetala group. However, in its small anthers and almost glabrouspetals B. caduciflora is different from that group. Examinationof fruiting material of this
species would make its affinitiesclearer. The presence of individualswith glandular or eglandularcalyx within the same species is unknown elsewhere in the
subgenus Pleiopterys, though it occurs in the subgenus Hemiramma.
90. BanisteriopsiskrukoffiiB. Gates, sp. nov.
Liana, rami dense aureo-sericei, demum glabrati. Stipulae minutae, triangulares, basi per jugum interpetiolarejunctae. Folia opposita; petiolo 6-14 mm
longo, appresso-sericeo,pilis aureis, supracanaliculato,ad apicem biglanduloso;
lamina 5.1-14.3 cm longa, 2.8-7.9 cm lata, falcata, coriacea, elliptica, basi cuneata obtusave, apice longe acuminata,marginerevoluta et glandes minutasgerenti, supra glabra, subtus sparsimappresso-sericea,pilis trabecula0.2-0.3 mm
longa, venis venulisque supra prominulis subtus 6-8 venis lateralibusutroque
costae latere prominentibus.Inflorescentiaaxillaris, floribus in umbellis 4-floriferis fasciculatis, 1-5 paribusumbellae in racemis 3-11 mm longis dispositis, 13 in axilla, appresso-sericea,pilis aureis; bracteis bracteolisque 1.0-1.6 mm longis, spathulatis,apice rotundatis,abaxialiterappresso-sericeis,adaxialiterglabris,
divergentibus,persistentibus;pedicellis sessilibus, 5-12 mm longis, 0.4-0.6 mm
diametro, in fructu incrassatis, dense appresso-sericeis, pilis aureis. Sepala 5,
1.6-2.0 mm longa, 1.0-1.4 mm lata, elliptica, apice rotundata, abaxialiterappresso-sericea, adaxialiterglabra, 4 lateraliaglandes 1.2-1.6 mm longas et 0.50.6 mm latas gerentia,glandes0.6-1.2 mm superantia.Petala 5, lutea, extus dense
sericea, denticulata fimbriatave, eglandulosa, 4 lateralia inter sepala patentia,
ungue 0.5-1.0 mm longo, limbo 4.5-7.0 mm longo et 4.0-5.0 mm lato, late obovato, 2 anterioribusquam 2 posterioribusmajoribusconcavioribusque,petalum
posticum ungue erecto 2.0-2.5 mm longo, limbo 3.5-4.0 mm longo latoque, orbiculari, plano. Stamina 10, filamentis 1.4-3.8 mm longis, glabris, illo sepalo
201
antico opposito quam ceteris longiore et apice incurvato, illis sepalis antero-lateralibus et petalis postero-lateralibusoppositis versus petalum posticum inclinatis et in dimidio inferiore connatis, 3 filamentisposticis inter stylos posticos
inflexis; antheraeloculis 0.8-1.2 mm longis, glabris, connectivis sepalis oppositis
0.8-1.0 mm longis, 0.6-0.7 mm latis, 0.4-0.5 mm profundis,obovatis, illis petalis
oppositis 0.6-0.8 mm longis, 0.3-0.4 mm latis, 0.2-0.3 mm profundis, anguste
oblongatis. Ovarium 1.0 mm altum, densius appresso-pubescens, pilis aureis,
trabecula 0.2-0.4 mm longa, stylo antico 3.6 mm longo, crasso, ad apicem incurvato, dimidio inferiorepubescenti, pilis flexuosis 2 mm longis, stylis posticis
3.0 mm longis, sigmoideis,basi divergentibuset pubescentibus,stigmatibuscapitatis. Fructus ex 3 samaris(vel abortu 1-2) constans; samaracarpophorousque
1 mm longo, nuce usque 7 mm alta et 6 mm longa, appresso-pubescenti,pilis
trabecula 0.2-0.4 mm longa, lateraliterutrinquealula usque 8 mm longa ad areolam parellela instructa, et appendice rotundatausque 3 mm alta in margine
superioreinstructa,ala (?immatura?)usque 11 mm longa et 9 mm lata, appressopubescenti.
Type. B. Krukoff6791, Brazil, Amazonas, Mun. Humayta, near Livramento
on Rio Livramento,fl (holotype, RB; isotypes, A, BR, F, G, MICH, MO, NY,
S, U, US).
Distribution.(Fig. 43). Known only from two collections in AmazonianBrazil
on terrafirme, and one collection from Venezuela.
Collected in flower in May and October and in fruit in May.
VENEZUELA. Amazonas: Liesner 7375 (MICH). BRAZIL. Amaz6nas: Fr6es 22331 fr (IAN
38874);Krukoff6791 fl (BR, F, G, GH, MO, MICH, NY, RB, S, U, US).
This species differs from the Banisteriopsispubipetala group in its diverging

spatulatebracts and bracteoles, and in the fruit, with a reduced dorsal wing and
only one lateralwing on each side of the nut. It is possible that the reducedfruit
wing is an artifact;perhapsthe fruits are immatureor have been damagedduring
development. Most species of the B. pubipetala grouphave glabrousleaves; only
B. erianthera exhibits sparse appressed pubescence abaxially like that of B.
krukoffii.In the form of its bracts and bracteoles and its leaf pubescence B.
krukoffiiresembles Diplopterys. It can be easily distinguishedfrom that genus in
fruit, and in flower B. krukoffiican be distinguishedby its petals, which are
densely sericeous and denticulateor short-fimbriate,and its styles, of which the
anteriorstyle is longer, pubescent for half its length and incurved at the apex.
This species is named for Dr. Boris Krukoff whose Amazonian expeditions
produced many species never before or since collected.
91. Banisteriopsis patula B. Gates, nom. nov.
BanisteriaeglandulosaAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840,non DC.
Liana, young branches sometimes hairy, the hairs stiff with the trabecula 1.02.0 mm long and the stalk up to 0.2 mm long, old branches glabrate. Stipules
minute. Leaves chartaceous or coriaceous, with the petiole 9-20(-36) mm long,
appressed-sericeous,canaliculateabove, eglandularor apically biglandular,the
lamina 3.0-11.4 cm long and 1.8-6.8 cm wide, ovate, obtuse to truncate at the
base, acute to acuminateat the apex, with the marginplane to slightly revolute
and bearing minute glands, the one nearest the base on each side sometimes
enlarged,adaxiallysparsely sericeous to glabrate,abaxiallysparsely sericeous or
tomentose, rarely glabrate, the hairs with the trabecula 0.3-1.0 mm long, with
202
Flora Neotropica
the reticulation smooth or prominulous adaxially, the lateral veins prominent

abaxially. Inflorescence of 4-floweredumbels, 1-9 umbels borne in racemes, on
short axillary branches up to 3.5 cm long with very reduced leaves or bracts,
appressed-golden-sericeous;bracts and bracteoles 1.0-2.0 mm long, appressedgolden-sericeousabaxially, narrowlyoblong and roundedat the apex, spreading,
persistent; pedicels sessile or subsessile, 9-15 mm long, 0.4-0.6 mm wide, appressed-golden-sericeous.Sepals 1.4-2.5 mm long and wide, rotund, roundedat
the apex, appressed-sericeousabaxially, glabrous adaxially, eglandular.Petals
yellow, externallysericeous, subentireto denticulate,the 4 lateralpetals reflexed
between the sepals, with the claw 1.5-3.0 mm long and 0.4-0.5 mm wide, the
limb 7.0-9.0 mm long and 5.5-7.5 mm wide, broadly elliptic, concave, the posterior petal with the claw erect, 2.5-3.0 mm long and 0.7 mm wide, apically
constricted, the limb 4.0-7.5 mm long and 2.5-5.0 mm wide, obovate to broadly
elliptic, plane, reflexed. Stamens with filaments 1.6-3.6 mm long, that opposite
the posterior petal much shorter than the others, the other 9 almost the same
length except those opposite the antero-lateralsepals sometimes shorter, those
of the 3 posterior stamens inflexed between the posterior styles, the locules 0.51.5 mm long, those of the 3 posterior stamens shorterthan the others, glabrous,
the connectives opposite the 3 anteriorsepals 0.8-1.6 mm long, 0.7-1.2 mm wide,
0.6-0.9 mm deep, obovate, glandularand enlarged, apically papillate, with the
locules divergingdistally, those opposite the petals and the postero-lateralsepals
0.4-1.0 mm long, 0.3-0.8 mm wide and 0.2-0.6 mm deep, narrowlyoblong, only
those opposite the antero-lateralpetals apicallypapillate. Ovary 1.0-1.4 mm tall,
densely pubescent, styles 2.0-3.0 mm long, subequal, divergingfrom the base,
stigmas capitate. Fruit without carpophore,the nut 7-12 mm tall and 6-10 mm
long, laterally compressed, sparsely appressed-golden-sericeous,the hairs with
the trabecula 1.0-1.5 mm long, bearinglaterally on each side 2 winglets up to 8
mm high, and a rounded appendageon its upper edge up to 8 mm tall and 7-12
mm long, the wing 35-56 mm long and 10-18 mm wide, narrowlyoblong, sparsely
appressed-pubescentto glabrate.
Type. Blanchet Pl. exs. 1632, Brazil, Bahia, fl (holotype, G; isotypes, BM, F).
Distribution.(Fig. 43). In the Atlanticcoastal forest of Rio, EspiritoSanto, and
Bahia in Brazil.
Collected in flower in March, April, August, and October, and in fruit in September.
BRAZIL. Bahia: Blanchet 1632 fl (BM, F, G). Minas Gerais: Heringer 401 fl (RB); Schwacke 10864
fr (NY fragment ex B, P). Espirito Santo: Belem 1521 fl (IAN, NY, UB). Rio de Janeiro: Bunbury
I fl (BR); Burchell 2808 fl (GH, GOET, P); Kuhlmann s.n. fl (RB 41444); Peckolt 235 fl (BR); Sastre
& Emmerich 1218 fl (P).
The variationrepresentedby these collections may correspondto more than

one taxon. I prefer not to subdivide the species at this stage because of the
paucity of material,and especially of fruitingmaterial. Maybe additionalcollections will fill in the apparentdiscontinuitiesin the variation. However, I shall
detail the variation I have observed. The type is characterisedby chartaceous
leaves with sparse appressed pubescence abaxially, the hairs with straighttrabecula 0.2-0.6 mm long. Its sepals are short, up to 1.6 mm long, the lateralpetals
are short-clawed, with the claw 1.5-2.5 mm long, and the limb of the posterior
petal is small (4.0-5.0 mm long and 2.5-3.0 mm wide). The only fruitingcollection
of this species (Schwacke 10864), and Peckolt 235 are similarto the type in these
characters. Three collections (Kuhlmann s.n.; Burchell 2808; Bunbury 1) also
have chartaceous leaves, but the abaxial leaf pubescence is more dense, the
203
hairs with the trabecula flexuous and longer (0.5-1.0 mm). In addition, the
sepals are larger, 2.0-2.6 mm long, the claw of the lateral petals is longer
(2.5-3.0 mm), and the limb of the posterior petal is larger (6.5-7.5 mm long,
and 4.0-5.0 mm wide). Belem 1521 is unusual in its very condensed axillary
inflorescences and its coriaceous leaves which are glabrate, although the hairs
are short and appressedlike those of the type: in its sepal size, claw length of the
lateral petals, and limb size of the posterior petal, it also resembles the type.
Heringer401 also has coriaceous leaves, but its petioles are very long, up to 3.5
cm; other collections of this species have a maximumpetiole length of 2 cm. In
its sepal size and abaxialleaf pubescence this collection resembles the Kuhlmann
collection, but its lateral petals are short clawed and the posteriorpetal is small
like the type.
This species is unusualnot only for its eglandularcalyx, but also for its spreading, lingulate bracts and bracteoles; only Banisteriopsis krukoffiihas similar
bracts and bracteoles, which are like those of Diplopterys. Banisteriopsis patula
can be readilydistinguishedfrom Diplopterysin flowerby its entireto denticulate

petals which are densely sericeous externally, and its eglandularcalyx; in fruit,
B. patula exhibits a well-developeddorsal wing typical of Banisteriopsis.
The specific epithet refers to the spreadingfloriferousbracts and bracteoles.
92. Banisteriopsissepium(Adr. Jussieu) B. Gates, comb. nov.
Fig. 42.
Banisteriasepium Adr. Jussieu, Ann. Sci. Nat. Bot., Ser 2, 13: 284. 1840.
Vine, young branchesflattened, appressed-golden-sericeous,old branchesglabrate. Stipules minute, joined by an interpetiolarline. Leaves very coriaceous,
with the petiole 4-9 mmlong, densely appressed-sericeous,eglandular,the lamina
3.3-9.0 mm long, 1.6-4.4 cm wide, cuneate at the base, obtuse to emarginateat
the apex, the marginthickenedand more or less revolute, bearingminuteglands,
the gland nearest the base on each side sometimes enlarged, new leaves at first
sparsely appressed-sericeouson both sides, mature leaves quite glabrous, with
the reticulation prominent on both sides. Inflorescence axillary, of 4-flowered
umbels, appressed-golden-sericeous;bracts and bracteoles 2.0-3.0 mm long, triangular, carinate, acute at the apex, sparsely appressed-sericeousto glabrate
abaxially,glabrousadaxially, erect or appressed, persistent;pedicels sessile, 2125 mm long, 1.0 mm in diameter,thicker, up to 2.0 mm, in fruit, glabrous. Sepals
up to 3.0 mm long and 2.0 mm wide, elliptic, rounded at the apex, appressedgolden-sericeous, eglandular.Petals yellow, externally sericeous, fimbriate,the
4 lateral petals reflexed between the sepals, the claw 3.0-4.0 mm long, the limb
7.0-11.0 mm long, 7.0-9.0 mm wide, rotund, concave, the posterior petal with
the claw erect, up to 4.5 mm long and 1.0 mm wide, apicallyconstricted,the limb
up to 9.0 mm long and 7.0 mm wide, broadly obovate, plane, reflexed. Stamens
with the filaments2.8-3.8 mm long, those opposite the styles thick, those opposite
the postero-lateralpetals and sepals longest, those of the 5 anterior stamens ?
the same length, those of the 3 posterior stamens inflexed between the posterior
styles; anthers with the locules 1.2-1.5 mm long, glabrous, connectives of the
stamens opposite the sepals up to 1.4 mm long, 1.0 mm wide and 0.8 mm deep,
obovate, those opposite the petals up to 1.0 mm long, 0.8 mm wide and 0.4 mm
deep, oblong. Ovary 1.5 mm tall, densely pubescent, styles ca. 3 mm long, divergingat least in bud, the anteriorstyle inflexed apically, stigmascapitate. Fruit
without carpophore, nut up to 32 mm tall and 27 mm long, smooth, glabrous,
with a single shallow winglet on each side parallelto the areole up to 7 mm high,
sometimes confluent beneath the nut, the wing reduced and forming a deltate
204
Flora Neotropica
projection up to 20 mm tall and extending forward as a shallow crest to just

behind the style, glabrous.
Type. Martius s.n., Brazil, Piaui, fl (holotype, M).
Distribution. (Fig. 43). In dry thickets in Piaui and Bahia.
Collected in fruit in June and October.
BRAZIL. Piaui: Martius s.n., fl (M). Bahia: Belem 1219 fr (NY); Pereira 58661 A. Lima 61-3935
fr (HB).
This species exhibits the largest fruit in the genus; with its massive nut and
relatively small fruit wing it is perhaps adapted for water dispersal.
Excluded Taxa
Banisteria aculeata Miller =non Malpighiaceae fide Adr. Jussieu.
B. albida Willdenow ex Steudel =Gaudichaudia albida Chamisso & Schlechtendal fide Niedenzu.
B. allophila Sieber ex Adr. Jussieu =Stigmaphyllon ovatum (Cavanilles) Niedenzu fide Niedenzu.
B. angulata Vellozo =Stigmaphyllon affine Adr. Jussieu fide Niedenzu.
B. angulosa L. =Stigmaphyllon angulosum (L.) Adr. Jussieu fide Niedenzu.
B. anoptera (Adr. Jussieu) Morton =Heteropterys anoptera Adr. Jussieu.
B. antifebrilis Grisebach =Mascagnia psilophylla (Adr. Jussieu) Grisebach var.
antifebrilis (Grisebach) Niedenzu fide Niedenzu.
B. apiculata (Miquel) Robinson in Small =Heteropterys macrostachya Adr. Jussieu fide Niedenzu.
B. arborescens (Brandegee) Small =Heteropterys arborescens Brandegee.
B. argyrophaea (Adr. Jussieu) Morton =Heteropterys argyrophaea Adr. Jussieu.
B. aurea Vellozo = unidentifiable; cf. Anderson & Gates (1975, p. 54).
B. auriculata Cavanilles =Stigmaphyllon auriculatum (Cavanilles) Adr. Jussieu.
B. beecheyana (Adr. Jussieu) Robinson in Small =Heteropterys beecheyana Adr.
Jussieu.
B. benghalensis L. =pp Hiptage benghalensis (L.) Kurz, pp Heteropterys laurifolia (L.) Adr. Jussieu, fide Niedenzu.
B. billbergiana Beurling =Stigmaphyllon ellipticum (H.B.K.) Adr. Jussieu fide
Niedenzu.
B. bonariensis Hooker & Arnott =Stigmaphyllon littorale Adr. Jussieu fide Niedenzu.
B. brachiata L. =Heteropterys brachiata (L.) Humboldt, Bonpland & Kunth.
B. brachiata Sprengel =Heteropterys longifolia (Swartz) Niedenzu fide Niedenzu.
B. brachiata Willdenow =Stigmaphyllon ovatum (Cavanilles) Niedenzu fide Niedenzu.

B. brachyptera DC. =Stigmaphyllon ovatum (Cavanilles) Niedenzu fide Niedenzu.
B. bracteata DC. =Tetrapterys citrifolia (Swartz) Persoon fide Niedenzu.
B. brevipes DC. =Gaudichaudia sp., probably G. mollis Chodat.
B. caerulea Lamarck =Heteropterys laurifolia (L.) Adr. Jussieu fide Niedenzu.
B. calcitrapa Desvaux ex Hamilton =Stigmaphyllon ovatum (Cavanilles) Niedenzu fide Niedenzu.
B. canaminensis Rusby =Heteropterys canaminensis (Rusby) Niedenzu.
B. chrysophylla Lamarck =Heteropterys chrysophylla (Lamarck) H.B.K.
B. chrysophylla Bello =Heteropterys wydlerana Adr. Jussieu fide Niedenzu.
B. ciliata Lamarck =Stigmaphyllon ciliatum (Lamarck) Adr. Jussieu.
205
B. coccinea Vellozo =non Malpighiaceaefide Adr. Jussieu.

B. convolvulifolia Cavanilles =Stigmaphyllon convolvulifolium (Cavanilles) Adr.
Jussieu.
B. cordata Vellozo =Tetrapterys phlomoides (Sprengel) Niedenzu fide Niedenzu.

B. corymbosa Larranaga =Stigmaphyllon sp.
B. cotinifolia (Adr. Jussieu) Robinson in Small =Heteropterys cotinifolia Adr.
Jussieu.
B. cuiabensis Grisebach =Hiraea cuiabensis (Grisebach) Grisebach.

B. deformis Desvaux ex Hamilton =Stigmaphyllon angulosum (L.) Adr. Jussieu
fide Niedenzu.
B. dichotoma L. =Stigmaphyllon lalandianum Adr. Jussieu or S. convolvulifo-
lium (Cavanilles)Adr. Jussieu fide Niedenzu.
B. discolor (J. D. Smith) Niedenzu =Mascagnia stannea (Grisebach) Niedenzu
vel aff. fide Anderson.

B. dispar (Grisebach) Niedenzu =Barnebeya dispar (Grisebach) Anderson &
Gates.
B. diversifolia H.B.K. =Stigmaphyllon diversifolium (H.B.K.) Adr. Jussieu.
B. dumetorum (Grisebach) Morton =Heteropterys dumetorum (Grisebach) Nie-
denzu.
B. eglandulata Vellozo =Tetrapterys crebriflora Adr. Jussieu fide Niedenzu.

B. eglandulosa DC. =Heteropterys suberosa (Willdenow) Grisebach var. can-
dolleana (Adr. Jussieu) Niedenzu fide Niedenzu.
B. elliptica H.B.K. =Stigmaphyllon ellipticum (H.B.K.) Adr. Jussieu.

B. emarginata Cavanilles =pp Stigmaphyllon tomentosum (Desfontaines) Nie-
denzu, pp S. emarginatum(Cavanilles)Adr. Jussieu fide Niedenzu.
B. eneaptera Sess6 & Mocinio ex DC. =non Banisteriopsis, possibly Triopterys.

B. fagifolia DC. =Hiraea fagifolia (DC.) Adr. Jussieu.
B. ferruginea Seemann =pp Tetrapterys seemannii Triana & Planchon.
B. fimbriata Vellozo =non Banisteriopsis fide Adr. Jussieu.
B. fisheriana Regel & Koernicke =Heteropterys ternstroemiifolia Adr. Jussieu.
B. floribunda DC. =Stigmaphyllon sp. fide Adr. Jussieu.
B. fruticosa Vellozo =Heteropterys umbellata Adr. Jussieu fide Niedenzu.
B. fulgens Lamarck =Stigmaphyllon hypoleucum Miquel.
B. fulgens L. =Stigmaphyllon emarginatum (Cavanilles) Adr. Jussieu fide Nie-
denzu.
B. fulgens G. F. W. Meyer =Stigmaphyllon puberum (Richard) Adr. Jussieu fide
Niedenzu.
B. gayana (Adr. Jussieu) Robinson in Small =Heteropterys gayana Adr. Jussieu.

B. glauca Desfontaines =Stigmaphyllon ciliatum (Lamarck) Adr. Jussieu.
B. glaucophylla Martius in Mohl =Verrucularia glaucophylla (Martius) Adr. Jussieu.
B. helicina (Grisebach) Morton =Heteropterys orinocensis (H.B.K.) Adr. Jus-
sieu.
B. heterocarpa Standley =Heteropterys heterocarpa (Standley) Standley.

B. heterophylla Willdenow =Stigmaphyllon hypoleucum Miquel Adr. Jussieu fide
Niedenzu.
B. hexandra Vellozo =Schwannia elegans (Adr. Jussieu) Adr. Jussieu fide Nie-
denzu.
B. hispida Mociiio et Sesse ex Ramirez =Gaudichaudia podocarpa Adr. Jussieu.

B. humboldtiana DC. =Stigmaphyllon humboldtianum (DC.) Adr. Jussieu.
B. hyposericea Standley & L. O. Williams =Heteropterys obovata (Small)
Cuatrecasas& Croat.
B. kraussiana Hochstetter =Acridocarpus natalitus Adr. Jussieu fide Niedenzu.
206
Flora Neotropica
B. kuntzei Small =Heteropterys obovata (Small) Cuatrecasas & Croat.

B. lancifolia West & Schlechtendal =Heteropterys laurifolia (L.) Adr. Jussieu
fide Niedenzu.
B. laurifolia L. =Heteropterys laurifolia (L.) Adr. Jussieu.

B. laurifolia Richard ex Adr. Jussieu =Stigmaphyllon periplocifolium (Desfon-
taines) Adr. Jussieu fide Niedenzu.
B. ledifolia H.B.K. =Stigmaphyllon diversifolium (H.B.K.) Adr. Jussieu fide
Niedenzu.
B.
B.
B.
B.
B.
leona Cavanilles =Heteropterys leona (Cavanilles) Exell.

longifolia Swartz =Heteropterys longifolia (Swartz) Niedenzu.
lupuloides L. =Rhamnaceae.
macradena DC. =Heteropterys macradena (DC.) W. R. Anderson.
macrocarpa Persoon =Heteropterys longifolia (Swartz) Niedenzu fide Nie-
denzu.
B. macrophylla Colla = Hiraea sp.

B. macrophylla Adr. Jussieu =non Banisteriopsis.
B. macrostachya Vellozo =?Heteropterys byrsonimifolia Adr. Jussieu fide Nie-
denzu.
B. magnoliaefolia Desvaux ex Hamilton =Heteropterys longifolia (Swartz) Nie-
denzu fide Niedenzu.
B. maritima Richard =Stigmaphyllon ovatum (Cavanilles) Niedenzu fide Nie-
denzu.
B. mathewsana (Adr. Jussieu) Gleason ex Rusby =Heteropterys mathewsana
Adr. Jussieu.
B.
B.
B.
B.
B.
mediterranea Vellozo =Schwannia muricata Adr. Jussieu fide Niedenzu.

megacarpos Vellozo =Stigmaphyllon megacarpon (Vellozo) Grisebach.
michoacanensis Sesse & Mocino =non Banisteriopsis.
microphylla Jacquin =Heteropterys purpurea (L.) H.B.K. fide Niedenzu.
microphylla Hamilton =Stigmaphyllon periplocifolium (Desfontaines) Adr.
Jussieu fide Niedenzu.
B. microphylla Sesse & Mociiio =Heteropterys purpurea (L.) H.B.K.

B. monoptera Vellozo =Heteropterys chrysophylla (Lamarck) H.B.K. fide Nie-
denzu.
B. mossii Morton =Heteropterys mossii (Morton) Cuatrecasas.

B. mucronata DC. =pp Stigmaphyllon ellipticum (H.B.K.) Adr. Jussieu, pp S.
diversifolium(H.B.K.) Adr. Jussieu fide Niedenzu.
B. multialata Vellozo =Tetrapterys glabra (Sprengel) Grisebach.

B. multiflora Adr. Jussieu =Sphedamnocarpus multiflorus (Adr. Jussieu) Nie-
denzu.
B.
B.
B.
B.
B.
B.
multiflora DC. =Heteropterys multiflora (DC.) Hochreutiner.

mutabilis Vellozo =Thryallis brachystachys Lindley fide Kew Index 1895.
nemorum Brandegee =Heteropterys sp.
nervosa (Adr. Jussieu) R. O. Williams =Heteropterys nervosa Adr. Jussieu.
nitida Lamarck =Heteropterys nitida (Lamarck) H.B.K.
nitida Vellozo =Stigmaphyllon ciliatum (Lamarck) Adr. Jussieu fide Nie-
B.
B.
B.
B.
B.
B.
obovata Small =Heteropterys obovata (Small) Cuatrecasas & Croat.

orinocensis H.B.K. =Heteropterys orinocensis (H.B.K.) Adr. Jussieu.
ovata Cavanilles =Stigmaphyllon ovatum (Cavanilles) Niedenzu.
pallida (Brandegee) Standley =Heteropterys pallida Brandegee.
palmata Cavanilles =Stigmaphyllon palmatum (Cavanilles) Adr. Jussieu.
palmeri (Rose) Robinson in Small =Heteropterys gayana Adr. Jussieu.
denzu.

B.
B.
B.
B.
B.
207
paniculata Mocifio & Sesse ex DC. =Heteropterys laurifolia fide Niedenzu.

parvifolia Ventenat =Heteropterys purpurea (L.) H.B.K.
patens Grisebach =Heteropterys patens (Grisebach) Adr. Jussieu.
pentandra Sesse & Mocifio =Gaudichaudia sp.
periplocifolia Desfontaines =Stigmaphyllon periplocifolium (Desfontaines)
Adr. Jussieu.
B. picta H.B.K. =Stigmaphyllon ovatum (Cavanilles) Niedenzu.
B. portillana (Watson) Robinson in Small =Heteropterys portillana Watson.
B. pragua Vellozo = probably Heteropterys syringifolia Grisebach fide Niedenzu.
B. procumbens Sesse & Mocifio =non Banisteriopsis, probably Gaudichaudia
sp.
B. pruriens E. Meyer =Sphedamnocarpus pruriens (E. Meyer) Szyszylowicz.
B. pubera Richard =Stigmaphyllon puberum (Richard) Adr. Jussieu.
B. pubiflora DC. =Heteropterys laurifolia (L.) Adr. Jussieu fide Niedenzu.
B. purpurea L. =Heteropterys purpurea (L.) H.B.K..
B. quapara Aublet =Hiraea quapara (Aublet) Sprague.
B. quapara var. mucronulata DC. =Heteropterys berteroana Adr. Jussieu fide
Niedenzu.
B. reticulata (Poiret) Robinson in Small =Heteropterys multiflora (DC.) Hochreutiner.
B. retusa (J. D. Smith) Robinson in Small =Heteropterys retusa J. D. Smith.
B. riedeliana Regel =Mascagnia riedeliana (Regel) W. R. Anderson.
B. rosea Standley =Heteropterys sp.
B. rotundifolia Buc'hoz =Stigmaphyllon sp.
B. rotundifolia Sesse & Mocifio =non Banisteriopsis.
B. sagittata Cavanilles =Stigmaphyllon sagittatum (Cavanilles) Adr. Jussieu.
B. sanguinea Rusby =Heteropterys beecheyana Adr. Jussieu fide Niedenzu.
B. sericea Cavanilles =Heteropterys sericea (Cavanilles) Adr. Jussieu.
B. simulans Small =Heteropterys simulans (Small) Niedenzu.
B. sinemariensis Aublet =Mascagnia sinemariensis (Aublet) Grisebach.
B. sinemariensis DC. =Stigmaphyllon puberum (Richard) Adr. Jussieu fide Niedenzu.
B. sinuata DC. =Stigmaphyllon sagittatum (Cavanilles) Adr. Jussieu fide Niedenzu.
B. solitaria Vellozo =Hirea bahiensis Moricand fide Niedenzu.
B. splendens DC. =Stigmaphyllon hypoleucum Miquel.
B. suberosa (Grisebach) Robinson in Small =Heteropterys suberosa Grisebach.
B. tenuis Lindley =Heteropterys glabra Hooker & Arnott fide Niedenzu.
B. terminalis Vellozo =unidentifiable.
B. terminariifolia Mohl =Byrsonima vacciniifolia Adr. Jussieu.
B. ternata Mocifio & Sesse =Stigmaphyllon ellipticum (H.B.K.) Adr. Jussieu.
B. ternata Vellozo =Sapindaceae.
B. ternstroemiifolia (Adr. Jussieu) Niedenzu =Heteropterys ternstroemiifolia
Adr. Juss.
B. tetraptera Sonnerat =Hiptage benghalensis (L.) Kurz fide Niedenzu.
B. thyrsoidea Grisebach =Heteropterys cordifolia Moricand fide Niedenzu.
B. tiliifolia H.B.K. =Stigmaphyllon humboldtianum (DC.) Adr. Jussieu.
B. tiliifolia Ventenat =Ryssopterys tiliifolia (Ventenat) Adr. Jussieu fide Jussieu.
B. timoriensis DC. =Ryssopterys timoriensis (DC.) Blume.
B. tomentosa Desfontaines =Stigmaphyllon tomentosum (Desfontaines) Niedenzu.
208
Flora Neotropica
B. tomentosa Schlechtendal =Heteropterys beecheyana Adr. Jussieu fide Jussieu.

B. trifora Vellozo =unidentifiable.
B. trifoliolata Steudel =Sapindaceae.
B. umbellata Sesse & Mocino =unidentifiable.
B. umbellulata DC. =Stigmaphyllon periplocifolium (Desfontaines) Adr. Jussieu.

B. unicapsularis Lamarck =Hiptage benghalensis (L.) Kurz.
B. varia Sprengel =Stigmaphyllon tiliifolium (H.B.K.) Niedenzu fide Niedenzu
[=S. humboldtianum (DC.) Adr. Jussieu].
B. variifolia DC. =Stigmaphyllon tiliifolium (H.B.K.) Niedenzu fide Niedenzu
[=S. humboldtianum (DC.) Adr. Jussieu].
B. velutina Martiusex Colla =unidentifiable.

B. verruculosa Swartz =Heteropterys purpurea (L.) H.B.K. fide Niedenzu.
B. virgata Sesse & Mocifno =probably Gaudichaudia.
B. vitifolia Sesse & Mocinio ex DC. =probably Stigmaphyllon tiliifolium (H.B.K.)
Niedenzu fide Niedenzu [=S. humboldtianum(DC.) Adr. Jussieu]
B. volubilis Sesse & Mocifio =Gaudichaudia sp.

B. whitei (Rusby) Niedenzu =Heteropterys sp.
B. wydlerana (Adr. Jussieu) Robinson in Small =Heteropterys wydlerana Adr.
Jussieu.
Banisteriopsis cabrerana Cuatrecasas =Diplopterys cabrerana (Cuatrecasas)
Gates.
B. discolor (J. D. Smith) Small =Mascagnia stannea (Grisebach) Niedenzu vel.
aff. fide Anderson.

B. whitei Rusby =Heteropterys sp.
II. DIPLOPTERYSAdr. Jussieu in Delessert, Icon. sel. 3: 20, tab. 33. 1837
[1838].
Lianas, stipules interpetiolar,small to minute. Leaves simple, opposite, the
petiole appressed-sericeous,eglandularor apicallybiglandular,the laminaentire,
bearingnumerousminute glands along the margin,the gland on each side of the
midribnearest the base often enlarged,acuminateto long-acuminateat the apex,
adaxiallyglabrous,abaxiallysparselyappressed-sericeous.Inflorescenceaxillary,
of 4-floweredumbels, borne singly or in racemes of up to 7 umbels. Bracts and
bracteoles eglandular,lingulate,spreading,persistent, appressed-sericeousabaxially, glabrous adaxially. Pedicels sessile, appressed-sericeous. Flowers hermaphroditic. Calyx of 5 sepals, usually the anterior sepal eglandular, rarely
1-glandular,the 4 lateral sepals biglandularabaxially. Petals 5, yellow, clawed,
the limb orbicularto broadlyelliptic, externally sparsely sericeous in the middle,
long-fimbriate,eglandular,the posterior petal with the claw longer and thicker,
the limb obovate. Stamens 10, all fertile, the filamentsglabrous, those opposite
the sepals longer than those opposite the petals, those of the 3 posterior stamens
inflexed between the posterior styles, anthers with the locules sparsely hairy or
glabrous, the marginof the pollen sacs along the line of dehiscence membranaceous, the connectives papillose, those opposite the sepals larger than those
opposite the petals. Gynoecium of 3 carpels, each uniovulate, free, adnate to a
pyramidaltorus, densely hairy, embryo adaxial, pendent, anatropous.Styles 3,
free, hairy at the base or for up to half their length, stigmas capitate. Fruit of 3
mericarps without carpophore, sometimes only 1 or 2 developing. Nut of the
mericarpwith hardwoody pericarp,bearinga dorsalcrest and 2 to several lateral
winglets or crests parallelto the areole and interconnectedby ridges, appressedsericeous. Locule of the mericarpmore or less spherical, quite smooth.
209
Type species. Diplopterys pauciflora (G. F. W. Meyer) Niedenzu.
Distribution.Two of the four species are Amazonianin distribution,one species

occurs in northeasternSouth America (Para, French Guiana)and the remaining
species is from Veracruzin Mexico.
The genus Diplopterys was first described by Adrien de Jussieu for the single
species D. paralias, which is a synonym for D. pauciflora. The genus has traditionallybeen allied with the mascagnioidgenera because of its fruit without a
dorsal wing and with well-developedlateralwinglets, and Niedenzu includedthe
mascagniolidgenera Jubelina and Mezia in Diplopterys. However, the flowers
of Diplopterys are very similar to those of Banisteriopsis, and specimens of
Diplopterys in flower are often misidentifiedas Banisteriopsis. Species of Diplopterys closely resemble species within the subgenus Pleiopterys section Anisopterys of Banisteriopsisin the marginalglands on the lamina, sericeous petals,
and the form of the androeciumand gynoecium. The lingulate, spreadingbracts
and bracteoles of Diplopterys distinguishspecimens of this genus from all but
two species in section Anisopterys;the long-fimbriate,sparsely sericeous petals
of Diplopterys distinguish it from these two species, Banisteriopsis patula and
B. krukoffii,which have dentate to lacerate, densely sericeous petals.

In fruit, Diplopterysappearsvery distinctfrom Banisteriopsiswith its absence
of a dorsal wing, but the reductionof the dorsal wing to a dorsal crest is found
in three species within Banisteriopsis, B. cipoensis, B. paraguariensis and B.
sepium; if the fruit of Diplopterys is regardedas a banisterioidfruit with welldeveloped lateral winglets and the dorsal wing reduced to a crest, the derivation
of Diplopterysfrom a banisterioidancestor is readily postulated.
The four species of Diplopterys form a cohesive group with the members of
the group more similarto each other than to any Banisteriopsis species. I have
chosen to maintainthis small group as a separate genus because of the unusual
nature of the fruits; I feel that inclusion of Diplopterys within Banisteriopsis
would distort the definition of the genus unnecessarily. The interconnectionof
the lateral winglets by ridges to give a more or less reticulatesurface to the nut
is not found within Banisteriopsis. Perhapsthis reticulationtraps air to provide
buoyancy and the fruits are water-dispersed;the naturalhabitat of Diplopterys
appearsto be riverine. Large nut size and reductionof the dorsal wing to a crest
are represented in Banisteriopsis. Large nut size is usually found in species in
which the whole samara, includingthe dorsal wing, is greatly enlarged;only in
B. sepium is the nut enlargementassociated with a reductionof the dorsal wing.
But this species and the two other species of Banisteriopsis with reduced fruit
wings are each very similarto one or more other species within the genus with
well developed fruit wings, and differprincipallyfrom these species in the degree
of development of the fruit wing; the three species in Banisteriopsis with the
dorsalwing reducedto a crest are not at all similarto each other in other respects.
Key to the Species of Diplopterys
(See key to Banisteriopsis for separation of Diplopterys and Banisteriopsis,
and for explanationof terms used in the keys.)

Key to Flowering Specimens
1. Bracts and bracteoles 1.0-1.5 mm long; leaves up to 10.2 cm long and 4.7 cm wide.
3. D. cururuensis.
1. Bracts and bracteoles 2.0-3.5 mm long; leaves up to 21.0(-25.9) cm long and 9.0 cm wide.
2. Posterior styles longer than anterior style, or all 3 styles subequal.
210
Flora Neotropica
3. Posterior styles 2.4-3.0 mm long, anteriorstyle 1.8-2.0 mm long; petiole 4-11 mm
1. D. pauciflora.
long.
3. Styles subequalor the posteriorstyles slightlylonger than anteriorstyle, styles 1.42. D. cabrerana.
1.6 mm long; petiole (4-)8-15(-22) mm long.
4. D. mexicana.
2. Posteriorstyles shorterthan anteriorstyle.
Key to Fruiting Specimens

1. Anteriorstyle longer than posteriorstyles (fruitunknown).
4. D. mexicana.
1. Anteriorstyle shorterthan posteriorstyles or styles subequal.
2. Bracts and bracteoles 1.0-1.5 mm long; nut up to 7 mm tall and long, bearing2 welldeveloped winglets on each side, the anteriorwinglet longerthan the posteriorwinglet,
3. D. cururuensis.
up to 8 mm high, the nut surface smooth.
2. Bractsand bracteoles2.0-3.5 mmlong; nut up to 15 mm tall and long, bearing2-4 crestlike winglets on each side of the same height, often interconnectedby ridges to form a
reticulum.
3. Petiole (4-)8-15(-22) mm long; nut bearingca. 4 ridges or wingletson each side, the
surface between the areole and the anteriorwinglet smooth, between the winglets
and between the posteriorwingletand the dorsalcrest irregularlyfoveolate.
2. D. cabrerana.
3. Petiole 4-11 mm long; nut bearing2 winglets on each side, the surfacebetween the
areole and the anteriorwingletand the posteriorwingletand dorsalcrest smooth, the
1. D. pauciflora.
2 winglets more or less interconnectedby ridges.
1. Diplopterys pauciflora (G. F. W. Meyer) Niedenzu in A. Engler, Das Pflanzenreich IV, 141: 230. 1928.
TriopteryspaucifloraG. F. W. Meyer, Primit.Fl. essequib.: 183. 1818.
Tetrapterys pauciflora (G. F. W. Meyer) DC., Prodr. 1: 587. 1824.
Diplopterysparalias Adr. Jussieuin Delessert, Icon. sel. 3: 20. 1837.Type. Richards.n., French
Guiana,fl fr (holotype, P n.v.).
?Diplopteryspaucifloravar. latifolia Niedenzu,Arb. Bot. Inst. Lyc. 4: 22. 1912.Type. ?, French
Guiana,Cayenne, fl (holotype, B destroyed).
Liana, young branches appressed-sericeous, old branches glabrate, with the

secondary xylem sometimes developing very asymmetrically as a flange on 2
sides of the stem, giving the stem a flattened appearance. Stipules minute, triangular, usually connected by an interpetiolar line. Leaves with the petiole 4-11
mm long, appressed-sericeous to glabrate, channelled adaxially, eglandular, the
lamina 7.0-17.4 cm long and 2.4-7.9 cm wide, ovate to lanceolate, truncate at
the base, acuminate at the apex, with the margin plane to slightly revolute and
bearing minute glands, with the gland nearest the base on each side of the midrib
enlarged, glabrous adaxially, sparsely appressed-sericeous abaxially, the hairs
with the trabecula 0.2-0.3 mm long, eventually glabrate, with the reticulation
prominent adaxially and the 8-10 pairs of lateral veins prominent abaxially. Inflorescence axillary, of 4-flowered umbels, borne singly or in racemes of 3-7
umbels, the branches up to 5 cm long, appressed-sericeous; bracts and bracteoles
2.0-3.5 mm long, lingulate, rounded at the apex, sparsely sericeous abaxially,
spreading, persistent; pedicels sessile, 7-12 mm long, 0.4-0.5 mm wide, goldenappressed-sericeous. Sepals 1.8-2.8 mm long, 1.2-1.6 mm wide at the base, 0.61.0 mm wide at the apex, deltate, inpressed and reflexed apically, appressedsericeous abaxially, glabrous adaxially, the 4 lateral sepals each biglandular, projecting 0.6-1.8 mm beyond the glands, the glands 1.6-2.6 mm long, 0.8-1.2 mm
wide. Petals yellow, sparsely sericeous externally, long-fimbriate, eglandular, the
4 lateral petals reflexed between the sepals, with the claw 1.0-2.0 mm long, the
limb 5-8 mm long and 4-8 mm wide, broadly elliptic, the antero-lateral petals
slightly larger and more concave than the postero-lateral petals, the posterior
211
petal with the claw erect, 2.6-3.0 mmlong, 0.5-0.7 mm wide, apically constricted,
the limb 4-6 mm long and 4.5-5.0 mm wide, obovate, plane. Stamens with the
filamentsconnate proximallyfor up to half their length, those opposite the sepals
2.6-3.4 mm long, those opposite the petals 1.8-2.6 mm long, those opposite the
postero-lateralpetals almost as long as those opposite the antero-lateralsepals,
those of the 3 anteriorstamens erect, those opposite the antero-lateralsepals and
postero-lateralpetals directed posteriorly, those of the 3 posterior stamens inflexed between the posteriorstyles, the antherswith the locules 0.8-1.0 mm long,
sparsely hairy to glabrate, the connectives of the stamens opposite the sepals
0.8-1.0 mm long, 0.5-0.8 mm wide, 0.4-0.6 mm deep, broadly obovate, those
opposite the petals 0.5-0.8 mm long, 0.4-0.5 mm wide, 0.2-0.3 mm deep, oblong.
Ovarydensely hairy, 1.0-1.2 mm tall, the anteriorstyle 1.8-2.0 mm long, straight
and erect, the posterior styles 2.4-3.0 mm long, diverging and directed posteriorly, hairy for up to half their length, stigmas capitate. Fruit of 3 mericarps
without carpophore,the nut orbicularup to 15 mm tall and long, bearinga semicirculardorsal crest 2-6 mm high and usually bearing2 lateral wings parallelto
the areole on each side up to 4 mm high, the lateralwings + interconnectedwith
ridges, with the surface of the nut between the proximal lateral wing and the
areole and the distal lateralwing and the dorsalcrest smooth, sparselyappressedsericeous, the hairs with the trabecula0.1-0.2 mm.
Type. Herb. Meyer332, Guyana, Essequibo, fl (holotype, GOET n.v., MICH
neg. #3488 and 3489).
Distribution.(Fig. 44). A riverinevine in Guyana,FrenchGuiana,and northern
Brazil.
Collected in flower in November to January,and March,June and August, and
in fruit in Januaryand March.
GUYANA. Hb. Meyer 332 fl (GOET). FRENCH GUIANA. Black 54-17488 fl (IAN); 54-17585
fl (IAN); Martin s.n. fl (F); Oldeman 1368 fl (CAY); B-1027 fr (CAY); B-2696 fr (CAY); B-2722 fl
(CAY); B-2723fl (CAY). BRAZIL.Para:Ducke s.n. fl (MG);Fr6es 34110 fl (IAN); 34609 fl (IAN);
Sampaio 5501 fl (R); 5512 fl (R); Spruce 249 fl (NY).
This species is most closely related to D. cabrerana from which it can be

distinguished by its narrower leaves with shorter petiole, enlarged glands on
the leaf marginnear the base, not on the petiole, by its posteriorstyles which are
much larger than the anterior style, and by its fruit which has only two welldeveloped lateral crests on each side, with the area between the distal lateral
crest and the dorsal crest smooth. The two Froes collections from Para, Brazil,
have leaves which are smaller and narrowerthan the leaves of other collections
of this species.
2. Diplopteryscabrerana(Cuatrecasas)B. Gates, Brittonia31: 109. 1979.
Fig. 45.
Banisteriopsis cabrerana Cuatrecasas, Webbia 13: 493. 1958.
Banisteriopsisrusbyanasensu ethnobotanists,non (Niedenzu)Morton.
Liana, young branchesgolden-appressed-sericeous,old branchesglabrate,with

xylem sometimes developing very asymmetricallyas two lobes, giving the stem
a flattened appearance. Stipules minute, triangular, sparsely sericeous, often
joined by an interpetiolarline. Leaves with the petiole (4-)8-15(-22) mm long, appressed-sericeous to glabrate, channelled adaxially, apically biglandular, the
glands convex and prominent, the lamina coriaceous, falcate, (8.5-)10.0-21.0
(-25.9) cm long and (2.9-)4.1-9.0 cm wide, elliptic to broadly elliptic, truncate
Flora Neotropica
212
X
':",; ,
FIG. 44.
...
'.
}'.
Distribution of Diplopterys. Closed square, D. pauciflora; open square, D. cabrerana;
mexicana.
closed circle,
circle, D.
circle, D.
cururuensis; open
closed
D. cururuensis;
D. mexicana.
open circle,
at the base, long-acuminate at the apex, the acumen up to 3 cm long, with the
margin bearing minute glands, plane to slightly revolute, glabrous adaxially, abaxially sparsely appressed-sericeous, the hairs with the trabecula 0.2-0.3(-0.6) mm,
with the reticulation prominulous to prominent adaxially and the 6-8(-10) pairs
of lateral veins prominent abaxially. Inflorescence axillary, of 4-flowered umbels,
the umbels borne singly or in short racemes or condensed panicles, appressedsericeous; bracts and bracteoles (1.5-)2.0-3.0 mm long, lngulate, sparsely sericeous abaxiastriy,glabrous adaxially, spreading, persistent; pedicels sessile, 5-12
in and
to wide
mm
in fruit,
0.4-0.5
1.0 mm
1.4 mm
at themm
base
0.7-0.8upmm
relong,wide
diameter,
sparsely
appresseddeltate,
apically,
inpressed,
sericeous to glabrate. Sepals 1.5-2.2 mm long, appressed-sericeous abaxially,
the anterior
0.7-0.8 inmm
with
a few
hairsmm
mmappressed
mid1.0-1.8
wide.adaxially,
Petals yellow,
sericeous
thewide,
sparsely
sepal
apically
long, 0.5-1.0
narrowly elliptic, eglandular or rarely bearing 1 gland, the 4 lateral sepals up to
flexed apically, biglandular, projecting 1.0-1.3 mm beyond the glands, the glands
adaxally,
dle of the limb exteglabrous
long-fimbriate, eglandular, the 4 lateral petals reflexed
between the sepals, the antero-lateral petals with the claw 1.0-1.5 mm long, the
limb 7.0-8.0 mm long and wide, concave, the postero-lateral petals with the claw
0.5-1.0 mm long, the limb 5.0-7.0 mm long, 4.0-5.5 mm wide, broadly elliptic,
plane, the posterior petal with the claw erect, 2.5-3.5 mm long, up to 0.6 mm
wide, apically constricted, the limb 4.5-5.5 mm long and 3.0-4.5 mm wide, obovate. Stamens with the filaments connate basally for 0.4-1.0 mm, those opposite
the sepals
sepals 2.0-2.8
the petals
mm long,
long, glabrous,
2.0-2.8 mm
mm long,
1.6-1.8 mm
those opposite
the
glabrous,
long, those
opposite the
petals 1.'1.8
those of the 3 anterior stamens erect, those opposite the antero-lateral sepals and
213
I
.......
?Jr
":,,
~:~,:_~
"~.:?
..~..
;'i
"f:
":~":
':,'',' - .... '."'
.......
. .2..
...........,
,, , "
'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-?
~
~
z?
?'??~~~~~~ ~i
.i.i
c?: ,.. ....:....

~?
:-!
anerorstlei
cntr,x
0 g mriar
let..:axa
wng,abxil
ie
vie ...de
?.mntolf
.nero
""'~?:"
.post pot.io
x
pea,~
.A.se.~lopst
.st.e it sorgh
cros-secion
righ), h . eia wt
o
wns
bxa
i
(et
aai
vw
rg) cs.
section(below),
x1~~~~~~~~.
?Daw..rnDuht
-, cuts&Cbea129;g
rkf87;h
',i:/ms
1496.?]:.
.,?
FIG. 45.
Diplopterys cabrerana. a, habit x 0.5; b, leaf base with glands x 1.5; c, umbel of 4
lower a posterior (flag), x 4; e, part of androecium,

buds x 2; d, petals, upper apostero-lateral,
(flag),
postero-lateral,
stamento left opposite anteriorsepal, stamen to right opposite posteriorpetal, x 10; f, gynoecium,
anteriorstyle in center, x 10;g, mericarpwith short wings, abaxialview (left), adaxialview (middle),
cross-section(right), x 1; h, mericarpwith long wings, abaxialview (left), adaxialview (right),crosssection (below), x 1. (Drawn by Karin Douthit; a-f, Schultes & Cabrera 17297; g, Krukoff 8971; h,
Williams 14965.)
214
Flora Neotropica
postero-lateralpetals directedtowardsthe posteriorpetal, those of the 3 posterior

stamens inflexed between the posterior styles; anthers with the locules 0.8-1.0
mm long, sparsely hairy to glabrate,the connectives of the stamens opposite the
sepals 0.8-0.9 mm long, 0.7-1.0 mm wide, 0.4-0.6 mm deep, broadly obovate,
papillose, those opposite the petals 0.5-0.6 mm long, 0.4 mm wide and 0.2-0.3
mm deep, oblong, smooth to papillose. Ovary densely hairy, 1.0 mm tall, the
styles 1.4-1.6 mm long, the posteriorstyles slightlylongerthan the anteriorstyle,
straightand parallel or the posterior styles diverging,bearing stiff straighthairs
at the base, stigmas strongly capitate. Fruit of 3 mericarps,without carpophore,
the nut orbicular,up to 15 mm long and wide, bearinga crest-like dorsal wing 15 mm high, and usually 4 ridges or winglets on each side 1.0-10.0 mm high,
irregularor dissected along their margin,interconnectedwith ridges such that the
surface of the nut between the winglets, and between the winglets and the dorsal
crest, is irregularlyfoveolate, the surface of the nut between the areole and the
proximalwinglet smooth, appressed-sericeousthroughout,the hairs with the trabecula 0.1-0.2 mm long.
Type. Schultes & Cabrera17297, Colombia,Vaupes, rio Piraparana,tributary
of the rio Apaporis, CanioTeemefna,fl (holotype, US).
Distribution. (Fig. 44). Flowering and fruiting collections of this species are
from river marginsin AmazonianBraziland Venezuela, and Vaupes in Colombia,
with two collections from near Belem in Para, Brazil. Sterile collections of the
drugplant are from Peru, Ecuador, and Colombia.
Collected in flower in Septemberand fruit in April and October to December.
COLOMBIA. Vaupes: Hugh-Jones 2 st (ECON); Schultes & Cabrera 13134 st (ECON); 17297 fl
(US); 17459 bud (NY, US). Putumayo: Chindoy B. 281 st (US); Cuatrecasas 10597 st (F, US);
Garcia-Barriga et al. 18684 st (ECON); Klug 1971 st (BR, F, G, GH, M, MICH, MO, NY, P, S, SP,
U, US, W); Naranjo 9 bud (ECON); Pinkley 449 st (S); Plowman 2034 st (US); 2159 st (US); 2177
st (GH, S). VENEZUELA. Amazonas: LI. Williams 14965 fr (GH, US, VEN). ECUADOR. von
Hagen 141 st (NY); Mathias 5260 st (UCLA). PERU. Loreto: Mathias 3883 st (UCLA); San Martin:
Plowman 6040 st (MICH). BRAZIL. Amaz6nas: Krukoff8971 fr (GH, MICH, MO, NY, US). Para:
Archer 7917 fr (IAN, NY); Pires 4747 fr (IAN, US).
Common names. Colombia: "Mene kahi ma"; "nyoko-buku guda hubea ma"
(barasana); "yaje oko" (kofan); "ka-hee-ko" (Karapana); "chagropanga" (Inga);
"yageuco"; "oco yage"; "chagropanga azul pisco." Ecuador: "yaji'" "yaje.
Peru: "yaco-ayahuasco"; "yage."
This species is used as an ingredientin the preparationof the hallucinogenic

drink variously called "yage," "ayahuasca," "natem." Its leaves contain the
hallucinogendimethyl tryptamine(Der Marderosianet al., 1968; Agurell et al.,
1968).It has been wrongly called Banisteriopsisrusbyanaby ethnobotanistsand
botanists since Morton first misappliedthis name to a sterile collection (Klug
1971) of the drug plant (Morton, 1931). Many of the collections of this species
have been made by ethnobotanistsas voucher specimens of the drug plant and
are sterile. The drug plant apparentlyrarely flowers. It is often planted by the
native peoples who use it in secluded places in the forest, and perhaps the few
fertile collections may representescape from such cultivation.Perhapsthe natural
habitatof this species is riverine;its fruits are well suited to water dispersalwith
the large air-filledlocule and complex lateralwinglets which could act as flotation
devices. Because of its wide use as a drugplant, it is commonlytransplantedby
means of cuttings. Perhapsthe two collections from Belem are the result of such
a transplant.
215
3. DiplopteryscururuensisB. Gates, sp. nov.

Liana, rami juniores appresso-sericei, vetustiores glabrati. Stipulae minutae,
triangulares,jugo interpetiolarijunctae. Folia coriacea, petiolo 5-8 mm longo,
appresso-sericeo,canaliculatoadaxialiteret biglandulosoad apicem, lamina5.710.2 cm longa, 2.8-4.7 cm lata, ovata, basi obtusa, apice acuminata, margine
parumrevoluta et glandulisminutisinstructa,adaxialiterglabra,abaxialitersparsim appresso-sericea, pilis sessilibus, trabecula recta 0.2-0.3 mm longa, venis
venulisque adaxialiterprominulis,et 7-10 paribusvenorumlateraliumabaxialiter
prominentibus.Inflorescentiaaxillaris, in umbellis 4-floriferiscompositis, usque
5 umbellis in racemis dispositis, appresso-sericea;bracteis bracteolisque 1.0-1.5
mm longis, lingulatis,apice rotundatis,abaxialitersparsimsericeis, divergentibus,
persistentibus;pedicellis sessilibus 10-12 mm longis, 0.5 mm diametro, aureoappresso-sericeis.Sepala 2.0-2.4 mm longa, usque 1.0 mm lata, incurvata,apice
rotundataet reflexa, appresso-sericeaabaxialiter,glabraadaxialiter,sepalumanticum angusteoblongum,4 lateraliadeltataet biglandulosa,glandibus1.6-2.4 mm
longis, 0.8-1.2 mm latis. Petala non suppetunt. Staminafilamentisconnatis basi
usque 1.0 mm, illis sepalis oppositis 2.2-2.6 mm longis, illis petalis oppositis 1.82.2 mm longis, 3 anticis rectis erectisque, illis sepalis antero-lateralibuset petalis
postero-lateralibusoppositis postice spectantibus,3 posticis inter stylos posticos
inflexis; antheraeloculis 0.6-0.8 mm longis, glabris, connectivis petalis oppositis
0.4-0.6 mm longis, 0.3-0.5 mm latis, 0.2-0.3 mm profundis,oblongis vel anguste
ellipticis, illis sepalis postero-lateralibusoppositis 0.7-0.8 mm longis, 0.6 mm latis
et 0.4 mm profundis,late obovatis, ceteris sepalis oppositis non suppetunt.Ovarium dense sericeum, 1.0 mm altum; stylus anticus rectus erectusque, 2.0 mm
longus, basi pubescens, styli postici 2.2-2.6 mm longi, plus minusve divergentes,
basi pubescentes, stigmata capitata. Fructus ex 3 mericarpiis(vel abortu 1-2)
constans, sine carpophoro, nuce usque 7 mm alta et longa, orbiculari, crista
dorsali semicirculariusque 4 mm alta et 2(-3) alulis utrinque instructa, alulis
proximalibususque 8 mm altis, et marginedissectis, appresso-sericea,pilis sessilibus, trabecula0.1-0.2 mm longa.
Type. Anderson 10803, Brazil, Para, Rio Cururu, 1-10 km upriver(SE) from
village of Pratati,Alto Tapajos, 8?S, 57?5'W,fr (holotype, IAN; isotype, MICH,
NY).
Distribution.(Fig. 44). Known only from two collections in inundatedforest
along the Rio Cururu.
Collected in fruit in February.
BRAZIL. Para: Anderson 10803 fr (IAN, MICH, NY); 10973 fr (IAN, MICH, NY).
This species is distinguishedby its small leaves, short bracts and bracteoles,
and its relatively small fruit with well-developed "hiraeoid" lateral wings.
4. DiplopterysmexicanaB. Gates, sp. nov.
Liana, rami juniores appresso-sericei, vetustiores glabrati. Stipulae minutae,
triangulares,saepe jugo inter-petiolarijunctae. Folia chartacea, petiolo 7-8 mm
longo, appresso-sericeovel glabrato,canaliculataadaxialiter,eglanduloso,lamina
8.2-15.2 cm longa, 3.6-6.7 cm lata, anguste elliptica vel elliptica, basi cuneata,
apice acuminata,margineplanaet glandulisminutisinstructa,glandulisbasi prope
costam majoribus,adaxialiterglabra, abaxialitersparsim appresso-sericea,pilis
rectis sessilibus trabecula0.2-0.3 mm longa, venis venulisque prominulisadaxi-
216
Flora Neotropica
aliter, et 7-8 paribusvenorumlateraliumprominentibusabaxialiter.Inflorescentia

axillaris, in umbellis 4-floriferiscomposita, appresso-sericea,usque 7 umbellisin
racemis dispositis; bracteis bracteolisque2.0-2.5 mm longis, lanceolatis, rotundatis apice, abaxialitersparsim sericeis, divergentibus,persistentibus;pedicellis
sessilibus 12-15 mm longis, 0.5 mm diametro, sparsimappresso-sericeisvel glabratis. Sepala 2.8-3.0 mm longa, incurvata et reflexa apice, appresso-sericea
abaxialiter,glabra adaxialiter,apice rotundata,sepalum anticum anguste oblongum usque 1.4 mm latum, 4 lateraliadeltata,0.8 mm lata ad apicem, biglandulosa,
glandes 1.0-1.2 mm superantia,glandibus 2.0-2.4 mm longis et usque 1.0 mm
latis. Petala lutea, extus sparsimsericea, longe fimbriata,eglandulosa,4 lateralia
inter sepala patentia, ungue 1.5-2.0 mm longo, 0.4-0.6 mm lato, limbo 7-10 mm
longo, 6-9 mm lato, late elliptica, antero-lateralibusquampostero-lateralibusparum majoribuset concavioribus, petalum posticum ungue erecto 3.0 mm longo
et 0.6 mm lato, crasso, limbo 6.0-6.5 mm longo et 5.0-6.0 mm lato, obovato,
piano. Staminafilamentisconnatis basi usque 1.4 mm, illis sepalis oppositis 3.43.8 mm longis, illis petalis oppositis 2.4-2.8 mm longis, 3 anticis rectis, erectis,
illis sepalis antero-lateralibuset petalis postero-lateralibusoppositis postice spectantibus, 3 posticis inter stylos posticos inflexis; antherae loculis 1.0-1.4 mm
longis, glabris, connectivis sepalis oppositis 0.8-1.0 mm longis, 0.6-0.8 mm latis,
0.4-0.6 mm profundis,obovatis, illis petalis oppositis 0.7-1.0 mm longis, 0.4-0.5
mm latis, 0.2-0.5 mm profundis.Ovarium1.5 mm altum, dense sericeum; stylus
anticus 2.6-2.8 mm longus, rectus erectusque, usque medium pubescens, styli
postici 2.3-2.4 mm longi, basi divergenteset pubescentes, pilis V-formibus,sessilibus, brachiismultuminaqeualibus,brachiolongo usque 1.5 mm, alterominuto,
stigmatacapitata. Fructus non suppetit.
Type. M. Vdzquezet al. V-951, Mexico, Veracruz, Hermanos Cedillo-LaEscuadra por el Rio Soloxuichil,Mun. Hidalgotitlan,fl (holotype, JALAPA; isotypes, F, MICH).
Distribution.(Fig. 44). Known only from the type from Veracruz, Mexico, in
tall evergreenforest.
Collected in flower in August.
MEXICO.Veracruz:Vdzquezet al. V-951fl (F, JALAPA, MICH).
This species is distinguishedby its chartaceousleaves, largerflowers, and the

anteriorstyle longer than the posterior styles.
Excluded Species
Diplopterys bracteosa (Grisebach) Niedenzu =Jubelina bracteosa (Grisebach)
Cuatrecasas.
D. includens (Bentham) Niedenzu =Mezia includens (Bentham) Cuatrecasas.
D. involuta (Turczaninow)Niedenzu =Mezia includens (Bentham)Cuatrecasas.

D.
D.
D.
D.
D.
D.
marshalliana Riley =Banisteriopsis lucida (Richard) Small.

microcarpa Sandwith =Mascagnia microcarpa (Sandwith) Anderson.
riparia (Adr. Jussieu) Niedenzu =Jubelina riparia Adr. Jussieu.
rosea (Miquel) Niedenzu =Jubelina rosea (Miquel) Niedenzu.
spruceana Niedenzu =Mascagnia heterocarpa Anderson.
uleana Niedenzu =Jubelina uleana (Niedenzu) Cuatrecasas.
ACKNOWLEDGMENTS
I would like to express my thanks to the many people who have contributed
to the completionof this monograph:to my doctoralcommitteefor their time and
217
help, to MarthaM. Jones and Betsy Strauchfor their excellent typing, to Karin
Douthit for her help with the plates, and especially to Dr. WilliamAndersonfor
his unfailinghelp and encouragement,not only as my doctoral committee chairman, but also continuingto the present, and to my husband, George Estabrook,
without whose supportthis monographwould never have been completed.
Field work for the researchwas supportedby a grantfrom the National Science
Foundation, GB-37314,to W. R. Anderson at the New York Botanical Garden
for research in systematics and evolutionary biology, and a Doctoral Dissertation ImprovementGrant, DEB 76-01757,to W. R. Anderson at The University
of Michigan.
I wish to thank those in charge of the following herbariafor makingherbarium
materialavailablefor this study: BotanicalMuseumand Herbarium,Copenhagen;
Botanische Staatssammlung,Miinchen;British Museum(NaturalHistory), London; Conservatoireet JardinBotanique, Geneva; Field Museum of NaturalHistory, Chicago; Gray Herbariumand Arnold Arboretumof HarvardUniversity,
Cambridge;HerbariumBradeanum,Rio de Janeiro;Herbariumof CEPLAC-CEPEC, Itabuna,Bahia;Instituto de Botanica, Sao Paulo; Institutode Conservagao
da Natureza, Rio de Janeiro; Instituut voor Systematische Plantkunde,Rijksuniversiteit, Utrecht; Jardim Botanico do Rio de Janeiro; Jardin Botanique de
l'Etat, Brussels; Missouri Botanical Garden, Saint Louis; Museu Botanico Municipal, Curitiba;Museu Nacional, Rio de Janeiro;Museu ParaenseEmilio Goeldi, Belem; MuseumNational d'HistoireNaturelle, Laboratoirede Phanerogamie,
Paris; Naturhistorisches Museum, Wien; NaturhistoriskaRiksmuseet, Stockholm; Segao da Botanica, IPEAN-EMBRAPA,Belem; Systematisch-Geobotanisches Institut, Universitat Gittingen, Gottingen;The Herbariumand Library,
Kew; The New York BotanicalGarden,Bronx, New York;UniversidadNacional
del Nordeste, Departamentode Botanica y Ecologia, Corrientes;Universidade
de Brasilia,Departamentode Botanica, Brasilia;UniversityHerbarium,The University of Michigan,Ann Arbor;U.S. National Museum(Departmentof Botany),
Washington.
The distributionmaps in Figs. 14, 16 and 25 were based on Goode Base Map
203, those in Figs. 27, 33 and 37 on Goode Base Map 103, and that in Fig. 35 on
Goode Base Map 203 NE, all Copyright,The University of Chicago.
LITERATURE CITED
Agurell, S., B. Holmstedt & J. E. Lindren. 1968. Alkaloid content of Banisteriopsis rusbyana. Amer.
J. Pharm. 40: 148-151.
Anderson,W. R. 1967. A proposal to conserve the generic name Banisteria Adr. Jussieu against
Banisteria L. Taxon 16: 472-474.
1977. Byrsonimoideae,a new subfamilyof the Malpighiaceae.Leandra7: 5-18.

& B. Gates. 1975. Notes on Banisteriopsisfrom South-CentralBrazil. Contr. Univ. Mich.
Herb. 11: 51-55.
Baldwin,J. T. 1946. Banisteriacaapi Spruce:its chromosomes.Bull. TorreyBot. Club73: 282-285.
Cuatrecasas,J. 1958. PrimaFlora Colombiana.Malpighiaceae.Webbia13(2):343-664.
Der Marderosian,A. H., H. V. Pinkley& M. F. DobbinsIV. 1968. Native use and occurrenceof
in leaves of Banisteriopsisrusbyana.Amer. J. Pharm.140: 137-147.
N,N-Dimethyltryptamine
Eiten, G. 1972. The cerradovegetationof Brazil. Bot. Rev. 38: 201-341.
Ferri, M. G. 1963. Historicados trabalhosbotanicos s6bre o cerrado. Simp6sio s6bre o Cerrado,
15-50. Universidadede Sao Paulo.
Fouet,M. 1966. Contributiona l'etude cytotaxonomiquedes Malpighiacees.Adansonia6: 457-505.
Gates, B. 1977. A Monographof the CentralBrazilianspecies of Banisteriopsis(Malpighiaceae).
Thesis, The Universityof Michigan,1-336.
. 1977a. Proposalto conserve the genericname BanisteriopsisRobinsonin Smallwith a new
type. Taxon 26: 593.
Flora Neotropica
218
. 1979. New names in Banisteriopsisand Diplopterys(Malpighiaceae)of the GuayanaHighland. Brittonia31: 108-109.

Grisebach,A. H. R. 1858. Malpighiaceae.In: Martius,C. F. P. Fl. bras. 12(1): 1-108.
Haffer,J. 1969. Speciationin Amazonianforest birds. Science 165: 131-137.
Humboldt,F. H. A. von, A. J. A. Bonpland& C. S. Kunth. 1821[1822]. Malpighiaceae.Nova Genera
et Species Plantarum 5: 145-174 (4? ed.).
Jussieu,Adriende. 1832. Malpighiaceae.In: Saint Hilaire,A. F. C. P. Fl. bras. merid.3: 1-86.

. 1840. Malpighiacearum
synopsis. Ann. Sci. Nat. Bot., Ser. 2, 13: 247-291; 321-338.
.1843. Monographiede la famille des Malpighiac6es.Arch. Mus. Hist. Nat. 3: 5-151; 255616.
Kostermans,A. J. G. H. 1938. Studies in South AmericanMalpighiaceae,Lauraceaeand Hernandiaceae, especially of Surinam.1-356.
Macbride,J. F. 1950. Malpighiaceae.Flora of Peru. Field Mus. Publ. Bot. 13(3):781-999.
McVaugh,R. 1968. Reportof the Committeefor Spermatophyta.Conservationof genericnames, X.
Taxon 17: 462-466.
Moraes,V. & K. Arens. 1971. Potassioe sua lavagemem folhas de plantasde cerradoem comparagao
com plantascultivadas.III Simp6sio s6bre o Cerrado,199-200. Universidadede Sao Paulo.
Morton,C. V. 1931. Notes on yage, a drugplant of Colombia.J. Wash. Acad. Sci. 21: 485-488.
.1967. A proposalto conserve the generic name Banisteria H.B.K.. Taxon 16: 74-76.
.1968. A typificationof some subfamily,sectional and subsectionalnames in the Malpighiaceae. Taxon 17: 314-324.
Niedenzu,F. 1900-1901. De genre Banisteria, parts 1 & 2. Ind. Lect. Lyc. Brunsb.
.1912-1913. MalpighiaceaeAmericanae.Verz. Vorles. Konig. Ak. Braunsb.
.1928. Malpighiaceae.In: Engler,A. Das PflanzenreichIV, 141: 1-870.
O'Donell,C. A. & A. Lourteig. 1943. MalpighiaceaeArgentinae.Lilloa 9: 221-316.
Pal, M. 1964. Chromosomenumbersin some IndianAngiospermsI. Proc. IndianAcad. Sci. 60B:
347-351.
Ratter, J. A., P. W. Richards, G. Argent & D. R. Gifford. 1973. Observations on the vegetation of
northeasternMatoGrossoI. The woody vegetationtypes of the Xavantina-Cachimbo

Expedition
area. Phil. Trans. Roy. Soc. London B 266: 449-492.
Rawitscher,F. 1948. The water economy of the vegetation of the 'campos cerrados' in southern
Brazil. J. Ecol. 36: 237-268.
Simpson,B. 1971. Pleistocene changes in the fauna and flora of South America. Science 173: 771780.
Small,J. K. 1910. Malpighiaceae.North AmericanFlora 25: 117-171.
Standley,P. C. & J. A. Steyermark.1946. Flora of GuatemalaVol. V. Fieldiana,Botany 24: 1-502.
Vanzolini,P. E. 1970. Zoologia, sistematica,geographiae a origemdas especies. Univ. Sao Paulo,
Inst. Geogr., Ser. Monografiase teses, 3.
Vogel, S. 1974. Olblumenund olsammelndeBienen. Tropischeund subtropischePflanzenwelt,Nr.
7. F. Steiner, Wiesbaden.267 pp.
NUMERICAL LIST OF TAXA

1. Banisteriopsis cornifolia (H.B.K.)
Robinsonin Small
a. var. cornifolia
b. var. elliptica (Niedenzu)Gates

c.
2. B.
a.
b.
var. standleyi Gates

acapulcensis (Rose) Small
var. acapulcensis
var. Ilanensis Gates
3. B. ferruginea (Cavanilles)Gates
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
megaptera Gates
brevipedicellata Gates
wurdackii Gates
cinerascens (Bentham) Gates
longipilifera Gates
laevifolia (Jussieu) Gates
argyrophylla (Jussieu) Gates
byssacea Gates
paraguariensis Gates
harleyi Gates
14. B. vernoniifolia (Jussieu) Gates

15. B. schizoptera (Jussieu) Gates
16. B. stellaris (Grisebach)Gates

17. B. acerosa (Niedenzu)Gates
18. B. malifolia(Nees et Martius)Gates
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
a.
b.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
var. malifolia
var. appressa Gates
confusa Gates
latifolia (Jussieu) Gates
variabilis Gates
hirsuta Gates
hatschbachii Gates
megaphylla (Jussieu) Gates
irwinii Gates
campestris (Jussieu) Little
cipoensis Gates
angustifolia (Jussieu) Gates
andersonii Gates
arborea Gates
219

31. B. calcicola Gates
32. B. membranifolia (Jussieu) Gates
33. B. pulchra Gates
a. var. pulchra
b. var. glabrata Gates
34. B. adenopoda (Jussieu) Gates
35. B. caapi (Spruceex Grisebach)Morton

36.
37.
38.
39.
B.
B.
B.
B.
schwannioides (Grisebach) Gates

prancei Gates
multifoliolata (Jussieu) Gates
quadriglandula Gates
40. B. muricata(Cavanilles)Cuatrecasas
41. B. oxyclada (Jussieu)Gates
42. B. pauciflora (H.B.K.) Robinson in Small
43. B. goiana Gates
44. B. martiniana (Jussieu) Cuatrecasas
a. var. martiniana
b. var. subenerviaCuatrecasas
45. B. velutinissima Gates
46. B. elegans (Triana& Planchon)Sandwith

47. B. polygama (Niedenzu) Gates
48. B. padifolia (Niedenzu)Gates

49.
50.
51.
52.
53.
54.
55.
56.
57.
B.
B.
B.
B.
B.
B.
B.
B.
B.
wilburii Gates
pubescens (Niedenzu) Cuatrecasas
grandifolia (Niedenzu) Gates
alternifolia (Steyermark) Gates
maguirei Gates
pulcherrima (Sandwith) Gates
nummifera (Jussieu) Gates
sellowiana (Jussieu) Gates
anisandra (Jussieu) Gates
58. B. gardneriana(Jussieu)Andersonet Gates

59. B. lyrata Gates
60. B. parviflora (Jussieu) Gates
61. B. salicifolia (DC.) Gates
62. B. scutellata (Grisebach)Gates
.B.
o4. B.
65. B.
66. B.
67. B.
basifixa Gates
parviglandula Gates
pseudojanusia (Niedenzu) Gates
magdalenensis Gates
valvata Anderson et Gates
68. B. heterostyla(Jussieu)Cuatrecasas
69. B. lutea (Grisebach)Cuatrecasas
70. B. cristata (Grisebach)Cuatrecasas
71.
72.
73.
74.
75.
76.
77.
78.
79.
B.
B.
B.
B.
B.
B.
B.
B.
B.
leiocarpa (Jussieu) Gates

parvifolia (Niedenzu) Gates
populifolia (Niedenzu) Gates
lucida (Richard) Small
nutans (Niedenzu) Gates
peruviana (Niedenzu) Gates
woytkowskii Gates
nigrescens (Jussieu) Gates
cachimbensis Gates
80. B. hypericifolia(Jussieu)Andersonet Gates

81. B. virgultosa(Jussieu)Andersonet Gates
82. B. amplectens Gates
83. B. rondoniensis Gates
84. B. pubipetala(Jussieu)Cuatrecasas
85.
86.
87.
88.
89.
90.
91.
B.
B.
B.
B.
B.
B.
B.
platyptera (Grisebach) Cuatrecasas

longialata (Niedenzu) Gates
erianthera (Jussieu) Gates
schunkei Gates
caduciflora (Niedenzu) Gates
krukoffii Gates
patula Gates
92. B. sepium (Jussieu)Gates
D1. Diplopterys pauciflora (G. F. W. Meyer)
Niedenzu
D2. D. cabrerana(Cuatrecasas)Gates
D3. D. cururuensis Gates
D4. D. mexicana Gates
220
Flora Neotropica
LIST OF EXSICCATAE
Ackermann,s.n.(9); s.n.(26).
Ab., A., 605(60).
Agostini, G., 655(40).
Alain, Bro., 294(42);1608(42);4987(42).
Alexander,E. J., 187(2a).
Allemao,Fr. et al., 213(31);214(40);215pp(4and
34); 216(16);221(69)s.n.(4); s.n.(40).
Allen, C., 125(2a);701(40);787(2a).
Allen, P. H., 972(40);1030(40);1116(40);1129(6);
2018(40);2091(6);2776(40).
Almeda,F., 2943(49);2978(49);3453(49);3983(2a);
4207(2a).
Alston, A. H. G., 5941(lb); 8625(46).
Alston, H. G., 108(40).
Amaral,A., 1696(40).
Ancuash, E., 284(74).
Anderson,A., s.n.(74).
Anderson, W. R., 6341(57);6350(10);6393(16);
6446(58); 6504(9); 6572(84); 6658(22);
6671(14); 6686(18b); 6738(26); 6776(9);
6835(24); 6930(10); 6931(41); 6981(24);
7240(14);7273(9);7320(9);7357(24);7360(41);
7442(14); 7508(84); 7512(9); 7567(10);
7578(20); 7638(24); 7647(24); 7694(14);
7820(41); 7905(26); 7954(18b); 7977(21);
7980(26); 7997(16); 8088(16); 8105(18a);
8267(24); 8352(18a); 8383(29); 8445(28);
8500(29); 8503(15); 8522(28); 8528(18a);
8532(80); 8663(10); 8739(10); 8759(28);
8909(84); 8971(30); 9183(41); 9198(31);
9382(21); 9394(16); 9435(41); 9442(9);
9460(33a); 9461(9); 9472(41); 9485(10);
9680(10); 9814(16); 9910(37); 10000(21);
10000A(21); 10005(14); 10075(24); 10077(9);
10161(10); 10178(41); 10226(15); 10291(21);
10294(20); 10803(D3); 10973(D3); 11142(10);
11143(9); 11144(41); 11148(40); 11160(21);
11177(17); 11201(63); 11204(26); 11206(84);
36124A(29);36159(27);36291(41);36325(18a);
36421(15);36424(26);36484(84);36584(15);
36590(18b);
36742(81);36761(18b);
36762(18b);
36833(16); 36905(18b); 36906(26); 36981(67);
37079(16);37133(31);37150(84).
Andrade,A., 342(58).
Andrade,N., SP25008(9).
Andre, E., K22(40);K23(40);K24(40);1651(40);
1652(40); 1874(la).
Anisits, J. D., 2040(19); 2709(40).
Antonio, C. H., 100(40); 100b(40).
Araujo,F., s.n.(32).
Araujo,S. et al., 73(34); 109(40).
Archer, W. A., 2396(44a); 2951(40); 3005(16);
3117(40); 7917(40); (D2).
Archer, W. A. & A. Gehrt, 127(55).
Arenas, P., 295(19).

Argent, G. C. et al., 6342(84).
Argent, G. C., in Richards, 6448(84);6478(58);
6646(58); 6663(55).
Argumosa,J. A. de, 126(35).

Ariste-Joseph,Bro., 1906(40);A883(68);s.n.(68).
Aristeguieta, L., 4097(2b); 4894(2b); 6104(70);
6146(lb); 6414(lb).
Armond,N., 76(40);80(84);82(84).
Asplund, E., 10323(40).
Assis, V., 19(26);28(26);32(84);143(26);166(26);
204(9); 217(41); 5728(26).
Atala, F., 59(24); 64(21); 65(26).
Austeri, P. A., 11988(26).

Avalone, C. L. et al., 5(21).
Avery, G. N., 906(69).
Azevedo, L. G., 15(42).
Badcock, W. J., 90(40).

Badillo, V. M. & J. Holmquist,6042(44a).
Bailey, L. H. & E. Z., 425(85).
Baker, R. E. D., s.n.(74).
11234(40); 11235(9); 11249(24); 11252(21); Balansa,2395a(40);2402(40);2402a(40);4581(19).
11255(26); 11260(19); 11264(16); 11277(19); Balopure,K. M., 504(9).
11305(16); 11308(84); 11318(16); 11328(9); Bang, A. M., 1365(40);2061(40);2457(84and 69);
2489(55);2811(40and 84); 2813(40).
11341(9); 11376(16); 11387(17); 11389(16);
11396(26); 11399(9); 11415(10); 11417(9);

11424(26); 11426(24); 11427(16); 11459(10);
11469(26); 11470(14); 11471(25); 11480(22);
11481(26); 11484(16); 11488(16); 11490(14);
11493(23); 11504(16); 11506(24); 11518(18a);
11525(9); 11529(26); 11533(80); 11548(80);
11550(80); 11551(15); 11558(26); 11560(15);
11562(28); 11564(30); 11579(15); 11583(16);
11585(30); 11588(18a); 11592(28); 11594(29);
11603(80);
11601(30);
11595(28x29);
11605(16); 11606(40); 11607(41); 11618(34);
11712(40); 11713(32); 11727(64); 11730(32);
11732(32); 11733(40); 11734(4); 11735(4);
Banis, R., 5(9).
Barbierherb., s.n.(74).
Barclay, A. S. et al., 3488(40).
Barnes, V., 50(53).
Barreto, M., 49(9); 720(24); 1938(3);7510(28);
7562(10); 7563(10); 7600(26); 7601(26);
7602(24); 7604(26); 7606(24); 7609(27);
7610(26); 7612(21); 7614(21); 7615(21);
7616(18a); 7617(18a); 7619(18a); 7695(57);
7696(57); 7698(9); 7699(9); 7700(9); 7701(9);
7703(9); 7704(57); 7705(41); 7706(41);
7709(34); 7710(34); 7724(84); 7725(84);
8552(26); 9321(15); 10298(26); 10301(9);
10332(26); 10512(26); 10569(9); 12034(15);
12066(15); s.n.(57).
11743(18b); 11750(15); 11751(84); 11784(21);

11785(21); 11787(33a); 11788(33a); 11789(33a);
Barroso, G. et al., 608(23); 613(26); 773(14);
11793(40).
RB141453(9);RB141454(43);RB141455(26);
Anderson, W. R. et al., 35156(15); 35190(80);
RB143049(16).
35191(30); 35192(18a); 35203(29); 35204(28);
35304(26);35399(28);35506(30);36124(28); Belem, R. P. et al., 375(28); 420(9); 1219(92);
1521(91); 2678(3); 3655(18a); 3747(28);
4013(18a);4021(16);4022(21).
Belshaw, C. M., 3156(40).
Benoist, R., s.n.(69).
Bento, D. & J. Pickel, 794(40);1134(16);2958(9).
Bernardi,A. L., s.n.(74).
Bernardi,L., 1627(44a);7955(40).
Bernoulli& Cario, 3004(40).
Bertero, 2201(68).
Berthoud-Coulon,288(74);295(74).
Bertoni, 2548(40);2747(40);3852(40).
Beyrich, s.n.(3).
Bicalho, H. D., 27(57).
Black, G. A. et al., 47-1170(36);47-1815(84);483406(74); 51-11022(16); 51-13610(84); 5113716(40); 51-13781(40);52-14593(19);5214641(40); 52-14867(59);52-15188(69);5215189(19); 52-15550(84);54-16659(84);5417488(D1);54-17585(D1);55-18696(84).
Blanchet, J. S., 1632(91); 2576(55); 2737(40);
2845(16);2860(15);3174(55).
Blanco, C., 676(59);715(59).
Bockermann,W., 101(26);213(55).
Boerboom,J., 8793(44a);9113(44a).
Boeke, J. D., 1400(40);1443(7).
Bokermann,W., 213(55).
Bond, F. E. et al., 76(44a).
Bonpland,s.n.(40); s.n.(42).
Bordo, A., 49(16).
Bowie, J. & Cunningham,232(40);522(3).
Brade, A. C., 5312(26); 5950(34); 5951(34);
6018(26); 6029(16); 7312(10); 11282(62);

11879(18a);11880(18a);12324(26);12328(78);
13012(26); 13435(9); 13770(15); 13772(29);
13778(15);13779(15);13780(28);13783(18a);
13784(18a);13787(26);15428(26);17809(26);
17811(16);17812(34);17813(41);17814(10);
17815(10).
Braga,P. J. S., 492(40);502(32).
Braga, R. & R. Lange, 12(84);127(84);128(84);
129(84).
Bredemeyer,92(lb); 204(40);s.n.(lb).
Breedlove, D. E. et al., 13487(40).
Brenes, A. M., 3899(46); 4163(46); 4242(46);
4621(40); 5106(40); 5123(46); 5247(46);
5316(40); 5369(46); 5418(46); 6372(40);
6372b(40); 6809(46); 13229(40); 15145(46);
16714(40);17001(40);17307(40);18949(40);
19414(40);s.n.(40).
Brewer, C. H., 134(74).
Breteler, F. J., 4254(40).
Bristol, M. L., 759(35).
Brito, S., 96(26).
Britton, N. L. et al., 603(74);5868(92);6287(42);
10323(42);12602(42);12841(42).
Broadway,W. E., 4032(74);6910(74);7172(44a).
Brooke, W. M. A., 5568(69);5703(84).
Bruijn,J. de, 1265(lb); 1364(84).
B. T., Herb. LehmannianiumColombianum,
403(47);445(47); 557(47);660(46); 1093(la);
1094(46).
Buchtien,0., 669(55);912(19);913(40);1794(33b);
1801(40); 1804(19); 1809(48); 1821(75);
221
1822(75); 1824(75); 3797(55); 3798(84);
4656(84).
Bunbury,C., 1(91).
Bunting,G. S., 2964(54);3042(54).
Burchell, W. J., 560A(84); 2808(91); 3105(3);
4602(26); 4689(34); 5475(16); 5859(84);
6068(84); 6103(43); 6665(9); 6666(24);
6913(40); 6946(10); 6950-2(9); 7026(14);
7292(41); 7325(69); 7339(84); 8671(16);
8682(18b); 8729-2(16 and 78); 8746-6(16);
8752-4(40);8759(40);9244(40).
Cabrera, A. L. & H. A. Fabris, 15977(69);
15992(69);20987(69).
Calder6n,S., 68(40);224(2a);894(40);931(40).
Calmon,M., 430(26).
Camargo,J. C., 37(41).
Camp,W. H., E6(40);E3773(40).
Campos,C. J. et al., 118-25572(10).
Campos,J. M. de F., 35(69);47(57);48(84).
Campos, S. M., 116(26).
Carauta,J. P. P., 759(16);773(16).
Carcerelli,C., 16(40);30(34).
Cardenas, M., 4365(84); 5634(69); 5683(84);
5692(55);5891(7).
Cardona, F., 1178(74); 1422(74); 1428(74);
1820(44a);1903(44a);2009(44a);2044(44a);
2518(74).
Carrasquilla,L., 266(40).
Carvalho,A., 164(41).
Carvallo,J. C. de M., s.n.(9).
Casaretto,1408(3);2817(28).
Castellanos, A., 520(16); 571(69); 22632(26);
25010(15);25037(18a);25121(40);25309(15).
Calvalcante, P. B., 2108(18b); 2110(18b);
2137(18b); 2168(16); 2637(16); 2638(18b);
2691(16).
Cezio, 132(34).
Chagas,J., 1251(84);1437(84);s.n.(55); s.n.(84).
Chardon,C. E., 167(40).
Chase, A., 7687(69).
Chaves, C. M., 20(9).
ChindoyB., P. J., 256a(35);281(D2).
Chodat, 135(19);3779(69).
Claussen,P., 15(26);19(57);35(26);46(9);60A(9);
61A(9); 62A(9); 63A(9); 64A(9); 65A(9);
66A(21);69A(26);70A(21);71A(26);72A(18a);
73A(26);74A(26);77A(41);78A(24);79A(18a);
80A(18a); 86A(57); 106a(28); 108a(40);
121(57); 163(26); 189(9); 402(40); 415(26);
538(41); 542(24); 543(9); 544(10); 545(40);
546(26); 547(18a);548(41); 549(84); 615(9);
616(57);618(26);1018(26);1189(9);1521(18a);
1538(57); 1771(9); 18.1536(28); s.n.(9);
s.n.(10);s.n.(16);s.n.(18a);s.n.(21);s.n.(24);
s.n.(26); s.n.(28); s.n.(40); s.n.(41); s.n.(57);
s.n.(84).
Clayton, W. D., 4889(15).
Clemente, Bro., 258(42); 2258(42); 2838(42);
2900(42); 2903(42); 4253(42); 5088(42);
5120(42).
Cobra,L. Q. & J. Oliveira,247(84).
Cochran,D., s.n.(9); s.n.(26).
Flora Neotropica
222
Coelho, L., 5809(55); s.n.(44a); s.n.(55).
Coleman, J. R., 555(26).
Collenette, C. L., 124(84).
Combs, R., 628(42).
Commerson, s.n.(3).
Commissao Geogr. e Geol. de Minas, 804(34);
1123(32); 1190(64).
Commissao Geogr. e Geol. de Sao Paulo, 36(9);
90(26); 478(26); 554(16); 569(10); 862(57);
1049(16); 1204(16); 1233(9); 4384(41);
SP12009(16);
6166(41); SP12007(16);
SP12010(16).
Commissao Rondon, 169(69); 170(69); 1892(58);
2236(84).
Commissao Rondon Inspegao de Fronteiras,
22037(84); 22128(44a); 22822(84).
Constantino, D., 3(34); 7(60).
Conzatti, C., 3725(2a).
Cook, O. F. & G. B. Gilbert, 982(75).
Coradin, L., 547(7).
Cordeiro, M. R., 556(6); 902(83); 903(83);
917(19).
Costa, B. da, 104(9).
Cousin, A., s.n.(40).
Cowan, R. S. et al., 1182(74); 1815(54); 2030(54);
2093(74).
Cox, D. K. & Geezman, SCG 1147(2a).
Cox herbarium, 1092(2a).
Croat, T. B., 12536(40); 20065(44b); 33246(46);
33790(49).
Cuatrecasas, J. et al., 8985(86); 10597(D2);
10598(35); 10599(35); 11061(35); 12094(40);
13301(84); 13446(40); 14372(35); 15043(84);
17359(46); 19990(46); 24980(40); 25359(74);
25395(74).
Curran, H. M., 38M(40); 46M(40); 79M(40);
149(70); 170M(lb).
Cutler, H., 8079(69); 8405(18b); 8407(18b);
8410(69).
Dahlgren, B. E. et al., 122(40); 814(69); 885(84);
916(84); 959(84).
Damazio, L., 145(28); 328(32); 864(34); 872(26);
885(9); 890(18a); 892(84); 899(57); 1204(28);
1210(84); 1682(9); 1714(24).
Daubenmire, R., 203(31).
Davidse, G., 1484(46); 3738(lb); 4232(40);
4276(2b); 10840(11); 10884A(24).
Davidson, C., 3565(44b); 4795(84); 4995(84).
Davis, P. et al., 2355(58); 2419(84).
Davis, P. H., 60119(26); 60175(10).
Dawe, T., 37(40); 356(40).
Dawson, E. Y., 14198(10); 14203(24).
De La Cruz, J. S., 1043(44a); 1044(44a);
1262(44a); 1386(74); 1388(74); 1439(74);
1718(44a); 1803(44a); 2431(44a); 2684(44a);
3081(74); 3182(74); 4075(44a); 4160(44a);
4251(44a); 4271(44a); 4433(74); 4494(44a).
Dedecca, D., 9224(40); 9266(41).
Delascio, F. et al., 3400(2b); 4688(2b); 5087(2b).
Delgado, E., 74(74).
Devereux, C. P., 15(84).
Diaz, C. et al., 788(77).
Diehl, 2434a(69).
Dionisio, E. & Otavio, 270(60).

Dobermir, 45(69).
Dombey, s.n.(40).
Don, G., 88(84).
Dorsett et al., 201B(26); 202B(26, pp).
Drake, E., s.n.(42).
Dravassos, L., s.n.(27).
Drouet, F., 2522(69); 2523(84).
Dryander, E., 2674(35).
Duarte, A. P., 447(26); 1257(69); 1293(18b);
1300(16); 1428(16); 1958(27); 2050(28);
2441(24); 2442(21); 2637(18a); 2757(28);
2800(16); 4256(32); 5584(41); 7879(29);
10657(26).
Duarte, L. et al., 104(84); 105(57); 113(84);
286(15); 313(18a); 396(40); 483(69).
Duchassaing, s.n.(40).
Ducke, A., 153(35); 390(58); 793(69); 806(58);
2262(40); 2373(69); 2484(84); 2855(84);
2873B(40); 2905(36); 2978(84); 7573(45);
8519(36); 8587(84); 11018(36); RB25258(35);
RB25259(35); RB25260(35); RB35599(35);
s.n.(84); s.n.(D1).
Dudley, T. R., 9085(40).
Dugand G., A., 74(2a); 339/767(68); 547(68);
751(68); 1056(40); 1060(68); 1138(40);
2320(68); 2773(2a); 3335(2a); 3443(2a);
5025(68); 5102(68); 5330(40); 5331(2a);
5784(2a); 6704(68).
Dupre, s.n.(9); s.n.(26).

Dusen, P., 256a(26); 7915(60); 8062(34); 8326(60);
8330(34); 9079(65); 9585(34); 9637(9);
9705(26); 9970(34); 11010(26); 11645(26);
11671(26); 11673(26); 14860(26); 14991(26);
16518(9); 16622(40); s.n.(34); s.n.(65).
Dwyer, J. D. et al., 5040(49).
Edwall, G., 3376(84).
Eggers, 1145(44a); 13547(lb); 15485(35); 15494(35).
Egler, W. A., 827(84); 986(84); RB59841(30);
s.n.(29).
Eiten, G. et al., 1604(26); 1626(26); 1629(9);
1717(9); 2532(9); 2573(16); 2651(9); 2784(21);
2987(16); 3073(10); 3113(21); 3588(15);
4220(16); 4457(67); 4619(16); 4627B(69);
4875(69); 5402(84); 5403(84); 5425(84);
5486(84); 5541(84); 5580(69); 5918(40);
5938(84); 6018(16); 8391(84); 8452(84);
9970(26); 10114(16); 10512(15); 10600(15);
10663(15); 10666(15); 10804(15); 10987(27);
11070(26); 11086(27).
Ekman, E. L., 1153(42); 1422(42); 1530(40);
6521(42); 7639(42); 8027(42); 8972(42);
14924(42).
Elias, Bro., 387(68); 442(40); 443(4); 492(68);
697(68); 1198(68); 1288(68); 1442(68).
Emmerich, M., 97(21); 334(58); 3288(26); 3385(26).
Emygdio, L., 103(10); 138(26); 139(16); 146(10);
193(41); 2704(26); 2795(26).
Erwan, 70(41).
Estabrook, G. F., 1(26); 4(25); 6(14); 7(25); 9(25);
10(14); 11(14); 12(16); 13(23); 14(21); 17(58);
18(23).
Eugenio, J., 731(40); 741(69).

Eupunino, A., 255(32); 264(55); 283(55).
Ewan, J. A., 16717(84).
Eyerdam, W. J. et al., 22274(69).
Falcao, J. I. A. et al., 1059(69).
Fanshawe, D. B., 2400/5136(44a).
Fassett, N. C., 25992(40).
Felippe, G. M., 6(26); 111(84); 122(16); 149(16).
Fendler, A., 182(40).
Fennell, J. L., 282(35).
Fernandez, A., 1120(54).
Ferrari, G., 1021(44a).
Ferreira, A. et al., R1181(84); R1843(84).
Ferreira, A. G. et al., 96(21).
Ferreira, A. R., s.n.(19).
Ferreira, M. B., 28(18a); 34(24).
Ferreyra, R., 7133(7); 14196(73).
Fiebrig, K., 56(69); 689(19); 826(40); 1016(40);
2677(40); 4193(69); 5114(40); 5198(40).
Filho, A. F. C., 11(60).
Focke, s.n.(74).
Fonseca, S. G. da, 266(84); 306(84); 1064(84);
1209(84).
Forero, E. et al., 1189(46).
Forest Dept. British Guiana, 2929(44a); 5136(44a);
6967(74); 7031(74); 7849(54).
Forestry Bureau Surinam, 5127(74); 5315(74);
6663(74).
Forest Service French Guiana, 6180(74).
Fosberg, F. R., 19458(6); 21625(49); 27700(72).
Foster, R., 1478(6).
Frazao, A., s.n.(26); s.n.(32); s.n.(38); s.n.(40);

s.n.(55).
Freire, C. V., 677(40); 747(40); 784(40).
Fries, R. E., 520(69).
Fr6es, R. L., 3002(41); 20238(28); 20638(84);
21228(6); 21427(35); 22331(90); 23731(87);
23754(87); 25130(36); 25491(36); 25770(74);
27088(40); 28491(35); 29698(16); 30024(6);
30061(6); 31248(35); 33300(57); 33317(57);
33332(84); 34033(26); 34110(D1); 34609(D1).
Fromm, E., 1325(4); 1674(32); 1678(10); 1892(18a);
1894(84).
Fryxell, P., 1960(80); 1961(16).
Funck, N., 308(68); 377(40); 497(40).
Gaillard, 245(35).
12037(40);
H., 11834(la);
Garcia-Barriga,
12332(la); 13706(44b); 14250(35); 14415(35);
14570(35); 18060(40); 18604(35); 18684(D2);
20055(35).
Gardner, G., 18(56); 338(32); 844(40); 1481(69);
1483(16); 1484(18b); 2071(31); 2074(16);
2075(16); 2499(84); 2500(84); 2502(58);
3061(9); 3068(24); 3624(40); 3625(9); 4100(21);
4472(29); 4473(28); 4475(84).
Garganta, 927(40).
Gates, B. E., 303(26); 307(2b); 323(84); 324(18a);
325(15); 326(10); 327(15); 328(18a); 329(20);
330(15); 331(58); 333(26); 334(18a); 335(28);
336(15); 337(18a); 338(57); 339(84); 340(15);
341(28); 342(15); 343(26); 344(26); 345(16);
346(28); 347(28); 348(26); 349(80); 350(15);
351(29); 352(26); 353(26); 354(26); 355(26);
223
356(15); 357(26); 358(26); 359(30); 360(18a);
361(30); 362(15); 363(15); 364(28); 366(80);
367(28); 368(26); 369(26); 370(26); 371(26);
372(26); 373(57); 374(26); 375(26); 376(15);
377(15); 378(30); 379(16); 380(32); 381(32);
382(32); 383(32); 385(26); 386(27); 387(24);
388(9); 389(26); 390(21); 391(24); 392(15);
393(21); 394(16); 395(18a); 396(26); 397(9);
398(21); 399(10); 400(41); 401(40); 402(58);
403(21); 404(24); 405(26); 406(16); 407(16);
409(25); 410(20); 411(25); 412(24); 413(21);
414(18a); 415(18a); 416(21x18a); 417(20);
418(21); s.n.(62).
Gates, B. E. & G. F. Estabrook, 37(22); 224(22).
Gaudichaud, C., 86(9); 586(26); s.n.(60); s.n.(78).
Gay, C., 971(74); s.n.(74).
Geay, F., 858(74).
Gehrt, A., SP3216(24); SP3326(41); SP5483(34);
SP43679(55); SP45861(69).
Gemtchfjuicov, I., s.n.(58).
Gentry, A. et al., 3348(6); 3356(6); 3544(40);
6159(75); 10294(2b); 13826(85); 14142(46);
17242(46); 20031(6); 20375(44b); 26586(50).
Gillis, W. T., 11237(35).
Gines, H., 4843(74); 5018(74); 5720(44a).
Giuletti, N. et al., 700(26).
Glaziou, A. F. M., 1059(38); 1325(3); 2512(62 and
63); 3884(78); 3888(40); 3894(63); 6489(3);
8293(26); 8582(34); 8583(62); 8586(38);
8587(56); 8588(3); 9363(38); 9364(34);
9676(55); 9679(84); 9682(16); 10357(32);
10365(60); 10370(62); 10381(16); 10383(84);
11811(63); 11812(62); 11816(40); 12475(34);
12475a(16); 12483(78 and 84); 12484(55 and
58); 12485(15); 12486(18a); 12489(58);
12490(57); 12492(4); 13598(18a); 13602(26);
13606(28); 13607(4); 13615(4); 14567(28);
16362(62); 16362a(40); 17488(34); 18162(10);
18162a(9); 18164(28 and 29); 18927(18a);
18928(18a); 18932(15); 18935(80); 18936(80);
18938(14 and 28 and 41 and Heteropterys);
18939(15); 18942(57); 18945(41); 18947(34);
18948(32); 18950(32); 20231(84); 20231a(40);
20750(20); 20751(18a and 20); 20752(18ax
21); 20754(57); 20755(41); 20756(57);
20758(84); s.n.(41); s.n.(84).
Goeldi, A., s.n.(84).
Gollmer, s.n.(40).
Goes, O. C. et al., 368(63); 378(63); 662(63).
Gomes, J. C., 240(10); 356(41); 424(9); 1106(21);
1111(21).
Gomes, J. F., SP1732(9); SP1735(40).
Gomes, J. G., 376(40); 459(40); 460(40).
Gomes, M., 2632(61).
Gomes, V., 1215(64).
Goodland, R., 61(26); 179(26); 830(57).
Gottsberger, I. S., 38(9); 157(21); 219(16);
336(84); 6571(21); 13-26970(69); 25-23771(69);
47R-1020(16); 49R-4573(21); 114R-11272(26);
142-25771(81); 271R-301172(21).
Goudot, s.n.(68).
Gounelle, 9(28); 34(26); s.n.(26); s.n.(31); s.n.(84).
Granville, J. J. de, 1265(55).
224
Flora Neotropica
Griffith,R. E., 9(42).

Grosourdy,R. de, s.n.(40).
Guedes, T. N., 13(84);377(18b);444(16).
Guillemin,353(26);s.n.(9); s.n.(21); s.n.(41).
Giinther(leg Buchtien), 1813(40).
Haas, J. H. & R. P. Belem, 370(58).
Hagen, W. von, 141(D2).
Halle, F., 640(74).
Hammar,A., 36(9);52(84).
Handro,0., 65(26);s.n.(35); s.n.(60).
Handro,W., 4(16);27(16).
Harley, R. M. et al., 10034(55); 10913(84);
11312(14); 11434(26); 15028(16); 15100(28);
15227(15); 15523(28); 15640(13); 15855(28);
15856(28); 15875A(13); 16108(16); 16789(28);
16855(18b); 18078(55); 18854(28); 18926(15);
19035(18b); 19329(16); 19619(28); 19963(18b).
Hartman,T., 33(16).
Harvey, D. R., 5188(40);5189(41).
Hassler, E., 812(69); 812a(69); 1292(40); 1919(19);
3115(40); 3274(69); 3774(40); 3817(40);
3986(19); 4028(19); 4089(40); 4451(69);
5118(69); 6452(19); 6682(19); 6726(19);
6726a(19); 7172(69); 7278a(40); 7355(69);
8218(33a); 8317(33a and 40); 9012(40);
9832(12); 9832a(12); 9856(12); 10189(41);
10252(21);
10252a(21);
10303(33a);
10303a(33a); 10403(10); 10481(17); 10537(69);
10562(69); 10562a(69); 10831(21); 10832(10);
10833(84); 11676(40); 12107(40); s.n.(40).
Hatschbach, G., 666(34); 1916(34); 2760(34);
3055(26); 3118(34); 3788(26); 5376(26);
5541(34); 5543(60); 6034(60); 6777(34);
6851(26); 6852(9); 7347(21); 8573(26);
8842(21); 14247(21); 16987(84); 18871(26);
21060(40); 21584(84); 21586(21); 21853(84);
23705(10); 23706(17); 23759(19); 24267(16);
24295(19); 24328(33a); 24531(84); 28066(84);
29016(28); 29232(9); 31199(26); 31208(9);
31231(9); 31319(10); 31488(9); 31519(26);
31540(27); 31546(28); 31629(18a); 31631(29);
31633(15); 31634(28); 31657(15); 31867(21);
31894(16); 31940(17); 31945(21); 32393(84);
Heithaus, E. R., 423(40).

Helmreichen,72(34).
Hemmendorff, E., 86(11); 128(84); 134(84);
154(84).
Herbier Imperial du Br6sil, 580(60); 583(78);

855(78).
Heriberto,Bro., 182(40);302(68);306(68).
Heringer, E. P., 74(18b); 103(57); 349(18b);
401(9); 476(10); 554(40); 1215(64); 3333(9);
3345(9); 3382(18a); 3439(21); 3477(21);
3558(57); 3713(26); 4029(84); 4049(57);
5691(69); 7185(26); 7483(41); 7930(84);
8009(18a and 21); 8873/1067(15); 8928(14);
9096/1290(20);
9450(57);
10102(18a);
10206(20); 10492(84); 11475(26); 11511(20);
11868(57); 11868A(10); 12171(43); 13190(16);
13193(18a and 21); 13197(10); 13198(26);
13253(18a); 13258(26); 13260(26); 13262(21);
14068(43); 14164(43); 14386(26); 14400(40);
14441(26); 14521(18a); 14580(10); 14606(21);
14654(58); 15417(26); 15443(40); 15451(26);
15789(18b); 15805(20); 15885(58); 15890(58);
16771(40); 21960(26); 22163(29).
Hermann, F. J., 11115(84).
Herzog, 1182(69).
Heyde, E. T. & Lux, 241(40); 3703(40); 4121(2a);
6309(40).
Heyligers, P. C., 797(44a).

Hioram,Bro., 2030(42).
Hipolito, s.n.(19).
Hirsh, R., P901(40).
Hoehne, F. C., 169(69); 170(69); 1581(33a);
1629(19); 1659(19); SP1828(60); 1891(37);
1892(58); 2236(84); 3098(34); 3923(34);
SP3965(60); 4758(40); 4803(9); 4895(19);

4896(19); SP4925(26); 5801(26); 5803(26);
5140(9); 6704(28); 6706(28); 17711(63);
19113(34); SP20330(9);
SP20430(16);
SP28521(78); SP53744(34); s.n.(40).
Hoffman, 228(40).
Hollister, G., s.n.(54).

Holt, E. G. et al., 155(70);689(74).
Holton, I. F., 800(la); s.n.(40).
Hostmann, 1027(74);2801(74).
33946(19); 33981(24); 34022(9); 34693(82);
Houllet, 1044(40); 1045(10); 1046(9); 1047(26);
34723(17); 34814(84); 34958(21); 35026(82);
1048(24);1049(21);s.n.(41).
36672(10); 36745(23); 36793(25); 36882(14);
Howard,R. A., 5238(42);5810(42).
36898(21); 36956(41); 36973(20); 36976(20);
J.,
36985(24); 36989(21); 36999(21); 37273(57);

38124(9); 38234(9); 38237(43); 38243(24);
38316(18b); 38336(33a); 38470(40); 38476(41);
38485(18b); 38612(21); 38613(8); 38646(33a);
38669(12); 38771(55); 39386(9); 39395(41);
39492(15); 39496(16); 39498(28); 39499(80);
39504(18b); 39935(60); 40806(9); 40844(29);
40891(15); 41220(18a); 41350(18a); 41414(28);
42115(67).
Haught, O., 2247(2a); 2368(40); 3733(40); 3807(40);
3996(68);6258(1a);6348(40).
Havard,V., 54(42).
Hayes, S., 229(40); 284(40); 770(6).
Hazlett, D. L., 2025(40).

Heiner, A., 193(84);438(10); 439(34); s.n.(26);
s.n.(34); s.n.(41).
Huber,
164(55); 250(40).
Hugh-Jones,S., 2(D2);3(35).
Huidobro,R., 2210(40).
Humboldt& Bonpland,s.n.(la).
Hunnewell,F. W., 16039(75).
Hunt, D. R. 5442(16);6227A(21).
Hunter,A. A. et al., 539(46).
Ibarrola,T., 4039(40).
Idrobo,J. M. et al., 1329(84);1724(46).
Imaguire,N., 1833(40).
Inst. MiguelLillo, 3078(69).
Irwin, H. S. et al., 2681(32),5157(18a);5347(57);
5485(69); 5574(16); 5702(26); 5772(16);
5975(10); 6035(57); 6058(57); 6167(84);
6347(84); 6362(84); 6591(84); 7241(16);
7404(21); 7838(58); 7843(16); 7961(18a);
225
Johnson, S. C. & Son CarnaubaExpedition,

885(84);916(84);959(84).
10303(18a);10719(26);10838(84);10901(15); Joly, A. B., 1035(27);1122(28);1165(28);1237(28);
10952(26); 11121(18a); 11184(24); 11242(24);
3532(26);4638(27).
Jonsson, G., 256a(26).
11265(15); 11398(9); 11429(15); 11524(26);
11553(18a); 11634(15); 11721(14); 12162(16);
Jorgensen,P., 4219(40);4478(40).
Jussieu, J. de, s.n.(40).
12303(26); 12419(27); 12453(16); 12464(16);
13369(18a); 13440(18a); 13555(21); 13558(26);
Kappler,A., 417(74);1027(74);1900(74).
13849(24); 14020(40); 14027(40); 14099(20);
Karsten,H., (4)55(85);s.n.(lb).
14302(10); 14318(24); 14341(18b); 14457(16);
Keel, S. & J. Guedes, 280(84).
14459(18b); 14491(18b); 14547(10); 14615(18b);
Kellerman,W. A., 7847(2a).
14628(26); 14976(41); 15007(9 and 24);
Kiegel, 1182(74).
15446(20); 15811(10); 16004(10); 16169(16);
Killip, E. P. et al., 14506(2a); 21087(68);
16550(16); 16669(9); 16673(10); 16951(10);
8294(84); 8377(58); 9660(16); 9808(26);

9909(21); 9909a(26); 9934(21); 10297(26);
17140(10); 17805(33a); 17806(10); 18091(21);

18707(15); 18980(9); 18992(24); 19006(24);
19020(9); 19046(9); 19049(24); 19303(15);
19522(26); 19731(28); 20002(27); 20101(28);
20354(26); 20778(28); 20827(30); 20904(28);
20904A(29); 20948(16); 21185(84); 21235(16);
21429(10); 21490(16); 21530(18b); 21665(16);
21666(9); 21824(80); 21834(15); 21837(18a);
21916(29); 22059(15); 22237(28); 22238(26);
22288(80); 22507(28); 22600(28); 22678(29);
22747(30); 22752(28); 22977(28); 23114(23);
23262(18a); 23401(18a); 23421(18a); 23625(28);
23715(18a); 23746(9); 23918(16); 24109(14);
24159(24); 24242(9); 24276(24); 24487(14);
24705(25); 24752(23); 24960(14); 25028(9);
25040(26); 25329(26); 25384(24); 25508(16);
25538(26); 25608(18a); 25765(9); 25904(26);
25974(24); 25975(9); 26081(10); 26086(26);
26294(16); 26408(9); 26477(24); 26562(26);
26563(10); 26567(24); 26670(18a); 26736(9);
26768(26); 26808(9); 26846(24); 26894(16);
26967(9); 27034(26); 27065(9); 27066B(16);
27088(30); 27361(41); 27455(28); 27488(18a);
27506(26); 27507(15); 27631A(16); 27631B(15);
27928(18a); 27954(30); 28319(80); 29080(28);
29266(28); 29562(28); 30504(34); 30762(16);
30773(84); 30879(13); 30909(28); 30909a(13);
31078(28); 31081(28); 31371(84); 31436(16);
31730(24); 31856(84); 32056(14); 32335(28);
32401(16); 32882(16); 32896(14); 33086(26);
33139(14); 34044(15); 34098(15); 34107(20);
34359(57); 34382(10); 34428(15); 34504(21);
34517(58); 34717(24); 34742(20); 34745(58);
34795(84); 34954(9); 34961(9); 47437(74);
55415(44a).
Jack, J. G., 6851(42); 7687(42); 8055(42); 8132(42).
Jamain, A., s.n.(42).
Jardim Botanico, Belo Horizonte, 430(84);
863(57); 864(57); 1756(84); 12328D(26);
12333(28); 12568(9); 12596(24).
Jelski, C. de, 343(76).
Jenman, E., 243(54); 4846(44a).
Jesus, J. A. de, 393(39); 1992(3).
Jim6nez-M., A., 344(40); 1284(40); 3517(40);
3788(6).
Jim6nez, O., s.n.(40).
Jobert et al., 450(55); 958(84); 1079(16); 1185(84).
Johns, W., 6201(34).
26082(89); 29922(44b); 38193(la); 38987(46).

Kirkbride, J. H., 1979(74); 2677(40); 2886(79);
2957(79).
Klug, G., 136(44b);167(44b);706(6); 1175(44b);

1934(35); 1964(35); 1971(D2); 3032(40);
3329(47); 3506(40); 3573(40); 4373(40).
Koczicki, 295(84).
K0ie, M., 4980(la).
Kramer,K. U. & W. H. A. Hekking,2753(74).
Krapovickas, A. et al., 12476(19); 19255(40);
31497(40);31579(40);31650(40);31697(19);
31763(7); 31935(9); 31936(41); 32254(41);
32895(8); 32897(8); 32916(17); 32930(21);
33053(16); 33097(26); 33105(9); 33140(9),
33268(9); 33296(43); 33299(26); 33300(9).
Krieger, L., 1589(9).
Krukoff, B. A., 6444(87); 6791(90); 7669(35);
8511(32); 8550(32); 8758(5); 8779(5); 8873(5);
8971(D2); 10022(86); 10048(84); 10590(86);
10857(55); 11184(55); 11234(84).
Kuhlmann, J. G., 148(84); 150(55); 441(6);

732(40); 2058(37); 2059(83); 2300(64);
2306A(40); RB35801(27); RB41444(91);
RB60455(35).
Kuhlmann,M., 383(40);399(41);624(40);701(34);
986(55);2819(40);3062(57);3508(26);3510(9);
3791(21);4143(16);4544(21).
Kummrow,455(26).
KunhardtJr., H. R., 2(54);21(54).
Kuntze, O., s.n.(19); s.n.(40); s.n.(41); s.n.(84).
Labouriau,L. & Valio, 1246(58).

Labouriau,M. S., 1(9);42(16);757(18a).
Labroy, 119(84).
Lam, s.n.(42).
Lambert,s.n.(74).
Lane, F., s.n.(55).
Lang, H. & A. C. Persaud,291(44a).
Langdon, 30(35).
Langsdorff, s.n.(26); s.n.(34).
Lanjouw, J. et al., 287(74);440(44a); 1913(74);

3465(74).
Lankester,C. H., 612(40).

Lanstyack,L., s.n.(26).
Laroche, R. et al., 153(34).
Lasser, T., 819(40).
Lawrance, A. E., 508(40); 639(49).
Leblond, 49(74);s.n.(74).
Lehmann, F. C., 2628(40); 5159(50);6102(40);
7558(la); 8491(47).
226
Lehmann Herbarium, 403(47); 445(47); 557(47);
660(46); 1093(la); 1094(46).
Le6n, Bro., 5613(42); 6251(42); 6380(42); 7352(42);
10204(42); 13357(42).
Levy, P., 386(40); s.n.(40).
Liebmann, s.n.(3).
Liesner, R. L., 7375(90).
Lillo, 18096(40).
Lima, A., 53-1372(40); 61-3935(92).
Lima, Alcina et al., 3(18a).
Lima, A. S., 7379(26).
Lima, D. A., 25(9).
Lima, J. A., s.n.(42).
Lima, J. I. de, RB52477(34); RB55459(26);
RB56851(26); RB56852(9); RB60721(41);
RB69472(40); s.n.(26).
Lima, M., s.n.(3).
Lima, N., 9(26); 128(18a); 255(20).
Lima, S., 350(66); 13281(66); 14239(66).
Lindeman, J. C. et al., 973(21); 986(84); 3171(26);
4409(40); 6010(18a); 6046(21).
Linden, J., 284(40); 698(40); 2075(42).
Linder, D. H., 164(74).
Lindman, C. A. Me., 2851(9); 2867(33a);
A2175(69).
Lisb6a, A., 2443(41); 2484(69); 2490(18b);
2517(84).
Lizot, J., 77(74).
Lofgren, A., 36(9); 62(26); 90(26); 101(55);
270(84); 478(26); 481(21); 504(84); 537(21);
554(16); 563(18b); 569(10); 601(16); 670(18b);
862(57); 876(84); 898(84); 1049(16); 1204(16);
1233(9); 1351(55); SP12007(16); SP12009(16).
Lorentz & Hieronymus, 486(69); 615(40).
Liidervaldt, H., 153(26).
Ludze, B. et al., s.n.(26).
Lugo, M., 179(47).
Luiz, 16(34).
Lund, W., 1058(56); s.n.(3); s.n.(9); s.n.(10);
s.n.(16); s.n.(26); s.n.(28); s.n.(57).
Luschnath, s.n.(55).
Liitzelburg, P. von, 151(58); 336(84); 394(16);
446(15); 460(15); 1582(58); 1824(58); 5801(15);
7165(29); 11216(69); 12019(69); 12493(18b);
12510(18b); 20658(40); 20660(7); 20686(40);
22037(84); 22128(44a); 22822(84); 26000(16);
26080(16); 26139(18b); 26211(18b); 26267(18b).
Lutz, B., 1723(3).
Macedo, A., 37(16); 233(9); 415(69); 492(84);
1043(9); 1045(10); 1046(41); 1139(84);
1140(69); 2260(41); 2500(69); 2505(84);
2512(55); 3226(17); 3530(43); 3673(20);
3692(58); 3899(58); 3901(84); 4014(84);
4163(9); 4178(26); 4198(9); 4287(16); 4419(14);
5119(26); 5277(26); 5382(58).
Machado, A., 27(26); 47(28).
Machado, 0., 268(6).
Machris Brazilian Expedition,
14198(10);
14203(24).
Madison, M. T. et al., 5138(50).
Magalhaes, H., 1190(64).
Magalhaes, M., 6(1); 167(18a); 240(16); 1219(28);
1441(24 and 26); 1444(9); 1445(84); 1721(18a);
Flora Neotropica
2140(26); 2212(26); 2213(18a); 4688(58);
9800(15); 15217(28); 15304(4); 15434(4);
15497(18a); 17018(34); 17173(26); 17505(28);
17559(9); 18088(15); 18390(15); 18391(15);
18955(84);
18908(26 and Heteropterys);
19127(18a).
Maguire, B. et al., 23125(54); 23354(74);
24615(44a); 24632(44a); 24715(44a); 27302(40);
28323(53); 29807(53); 30010(53); 30096(53);
30244(53); 31699(53); 31805(35); 32499(44a);
32850(53); 33888(44a); 34721(74); 34748(74);
35163(53); 35285(53); 35495(53); 35965(40);
40235(40); 41638(44b); 43889(54); 44168(26);
44524(26); 44536(26); 44618(26); 46235(74);
49165(80); 53524(44a); 53561(54); 53661(44a);
53669(54); 53704(44a); 56181(84); 56246(84);
56430(58); 56489(58).
Malme, G. O. A., 3328(10); 3386(37); 3414(41);
s.n.(10); s.n.(40); s.n.(41); s.n.(84).
Mandon, G., s.n.(35).
Manso, P. da S., 119(84); 325(40).
Marcos, 23129(35); 23132(35); 23134(35);
23135(35).
Marin, 1651(69).
Marinis, G., 200(9).
Markgraf, F. et al., 3240(15); 3242(15); 3522(15);
3588(26).
Martin, 92(44a); s.n.(D1).
Martin, R., 467(41).
Martin, R. T. et al., 1089(35); 1805(35).
Martinet, 1397(84).
Martins, H. F., 121(62); 122(41); 123(40).
Martius, K. F. P. von, 325(40); 517(78); 608(34);
658(84); 722(10); 1127(28); 1162(21); 1164(9);
1165(26); 1272(15); 1273(18a); 1342(18a);
1352(18a); 1360(15); 1423(28); 1428(28);
1761(81); s.n.(9); s.n.(10); s.n.(14); s.n.(15);
s.n.(16); s.n.(18a); s.n.(24); s.n.(28); s.n.(40);
s.n.(55); s.n.(69); s.n.(74); s.n.(78); s.n.(84);
s.n.(87); s.n.(92).
Mason, H. L., 13838(la); 13881(la).
Mathews, A., 682(40); 2024(40); 2126(71).
Mathias, M. E. et al., 3510(40); 3614(40);
3883(D2); 3925(35); 3975(35); 5260(D2);
5262(35); 6067(35).
Matos, M. E. R., 91(16).
Mattos, J. R. et al., 8159(10); 8987(10); 9537(26);
9613(26); 11636(26); 11637(9); 11646(10);
14416(40); s.n.(16).
Matuda, E., 598(40); 1744(40); 17064(40);
17266(40).
McCarty, G. S., 126(55).
Meader, R. E., 10(37).
Medina, E., 384(74).
Melin, D., 96(35).
Ml1inon, 245(74); 285(74); 913(74); s.n.(44a);
s.n.(74).
Mello, F., s.n.(55); s.n.(84).
Mendes, O. T., 245(40).
Mendonga, 749(28).
Mexia, Y., 4396(32); 4419(40); 4589(40); 4630(64);
4641(32); 5545(26); 5546(9); 5557a(41); 5605(15
and 16); 5666(9); 5679(24); 5711(15); 5732(28
and 29); 5804(80);
5818(15); 5818a(15);
5895(18a); 6636(35); 6837(35); 7291(35);

8027(40); 8029(40).
Mikan, s.n.(56).
Mimura, I., 17(26); 45(84); 147(26); 234(26);
566(84).
Molina R., A., 86(40); 694(2a); 931(40); 3909(2a);
3910(2a); 5135(40); 7205(46); 12233(46);
13004(40); 13217(2a); 13312(40); 18168(6).
Molina, J. A. et al., 18N.S.0.48(40).
Mocquerys, 6(75); s.n.(40); s.n.(75).
Montes, J. E., 15562(40).
Moore, S., 15(84); 104(84).
Moraes, J. C. de, 1977(69).
Moreira, C. et al., s.n.(9); s.n.(21); s.n.(40).
Moreira, J. I., 5779(26).
Morello, J., s.n.(26).
Moretti, 995(D1).
Mori, S. et al., 1836(75); 3604(49); 4599(46);
4940(46); 6084(49); 7111(49); 8722(6);
9198(44b); 9965(39); 10046(38).
Morillo, G. & B. Manara, 1083(70); 1141(70);
1165(70).
Morin, C. V. P., R9789(69).
Moritz, 1193(40); 1888(40).
Morong, T., 672(40).
Morton, C. V., 4075(42).
Mosen, H., 365(84); 827(26); 1148(9); 1150(40);
1151(34 and 40); 1152(40); 1153(26); 1154(26);
1159(16); 1795(41); 1796(40); 1797(40);
1798(9); 1960(34); 1961(34); 3841(16);
3842(34); 3844(10); 3967(55); 4047(9); 4048(9);
4049(26).
Moura, C., 64(40).
Mulford Biol. Explor. of the Amazon Basin,
110(84); 126(55); 663(86); 970(35).
Muller, 93(26).
Muller, F. von, 14(3).
Mutis, J. C., 2062(40); 2365(84); 2368(84);
2396(84); 2411(84); 2418(84); 2420(la);
2525(84); 4200(40); 4722(40).
Nadeaud, s.n.(3).
Naranjo, C. et al., 5(35); 9(D2).
Nee, M., 10517(49).
Nees von Esenbeck, 1(18a).
Nelson, E. W., 3278(40).
Netto, L., 71(18a); 79(26); 81(21); 82(28); 83(41);
84(41); 90(3); 92(9); s.n.(34); s.n.(55).
Neuwied, Prinz von, s.n.(18a); s.n.(31).
Newman, s.n.(84).
Niederlein, 165(2a).
Novaes, J. de C., 627(16); 628(84); 631(34);
672(40); 674(9); 675(26); s.n.(40); s.n.(41);
s.n.(84).
Nuiez, 0. V., 6577(35); 6585(35); 6587(35);
6589(35).
Occhioni, P., 1215(34); 4203(35); 4836(60);
4891(60); 5044(18a); 5632(16); 5699(21);
5767(16); 5792(84); 5809(84); 7298(82);
7573(16); 7576(16); s.n.(34).
Occhioni, T., s.n.(26).
Octacilio, P. & 0., 4216(9); 4285(9).
227
Oldeman, B-1027(D1); B-1144(74); B-1303(44a);
B-1362(74); 1368(D1); B-2696(D1); B2722(D1); B-2723(D1).
Oliveira, E. de, 3298(74); 3501(59).
Oliveira, J. E. de, 1145(57); 1201(57); 1566(40);
s.n.(10); s.n.(28).
Onishi, E. et al., 68(20); 295(84); 430(84).
Opler, P. A., 467(40).
d'Orbigny, 302(28); 581(40); 700(69); 755(69);
804(41); 909(84).
Orozco, J. M., 439(40).
0rsted, 7(lb); 31(2a); 5361(40); 5362(lb); 5363(2a);
s.n.(40).
Osten, C., 8524(69); 9043(69).
Pabst, G. F. J., 3279(26); 3376(28); 4507(3);
2942(18a); 6828(9).
Padilla, S. A., 86(40); 87(40); 629(40).
Palacios et al., 3192(26); 3258(26); 3458(26);
3657(26); 3747(9).
Palmer, E., 219(2a).
Paul, Bro., 189(40).
Paula, J. E. de, 58(24); 101(16); 154(26).
Pearce, R., s.n.(35); s.n.(40).
Peckolt, 235(91).
Peckolt, T., 180(40).
Peckolt & Freire, 482(62).
Pedersen, T. M., 3883(40).
Pena, B. S., 474(6).
Pennell, F. W., 2820(40); 2838(la); 2929(la);
4734(68); 6344(47).
Peredo, T., 355(40); s.n.(40).
Pereira, E., 1245(40); 1400(15); 1530(28); 1720(29);
2443(26); 2540(28); 4266(62); 4363(56);
4616(18a); 5415(34); 5866(92); 10756(34);
s.n.(9); s.n.(28).
Perez-Arbelaez, E., 639(35); 2106(la); 6057(47).
Perez-Medina, L. et al., Leticia 9(35).
Perottet, s.n.(74).
Persaud, A. C., 130(44a); 132(74).
Philcox, D., 3716(26); 3862(24); 3978(9); 4199(84);
4257(26); 4365(16); 4446(9); 4497(9); 4697(19);
4723(16); 4724(19); 4825(20); 4841(18a).
Philipson, W. R. et al., 1719(84).
Pickel, 2906(85).
Pickel, D. B., 133(69); 275(69).
Pierotti, S. A., 1205(40).
Pilger, 355(16).
Pinheiro, G. S. et al., 677(41).
Pinkley, H. V., 445(35); 449(D2).
Pires, J. M. et al., 19(35); 1640a(69); 2297(16);
2338(16); 2576(6); 2588(40); 2726(28);
2743(28); 2773(26); 2785(26); 4747(D2);
9014(9); 9023(10); 9025(26); 9051(20);
9072(26); 9093(18a); 9428(20); 9526(24);
9560(10); 9660(20); 9963(6); 12524(36);
13191(18b); 13640(35); 57945(41); 57965(9);
57994(16); 58019(69); 58020(9); 58037(18a);
58167(21).
Pittier, E., 13398(40); 13452(1b); 14137(74).
Pittier, H., 757(40); 3079(40); 4949(40); 6117(51);
7539(40); 8161(40); 8299(51); 8867(70);
9080(40); 10307(70); 10417(70); 10654(40);
10731(lb); 10779(lb); 10797(40); 11346(70);
228
11470(2b); 11762(70); 11798(70); 12234(lb);
12493(40); 12528(2b); 12591(70); 12592(40);
13036(lb); 13477(40); 13880(2a); 14741(2b);
14943(2b); 15119(lb); 15180(70); 15719(74).
Plee, s.n.(lb).
Plowman, T., 2034(D2); 2159(D2); 2177(D2);
2575(35); 3521(68); 5558(71); 5966(40);
6040(D2); 6041(35); 6052(40); 6488(6).
Poeppig, E., 99.1752(40); 1214(40); 1288(48);
1497(75); 1879(40); 2433(40); 2474(89);
s.n.(40).
Pohl, J. E., 362.1520d(26);
489(11); 717860(84); 982.1532d(26);
1257.1523d(9);
1514.1528d(10);
1328(14);
1496(16);
1516d.2330(84); 1519d.5670(57); 1529(18a
and 21); 1563(16); 1677(20); 1695.1533d(40);
2524.3273(16); 2684(24); 2691.1531d(43);
2858(18a and 21); 2994.1521(18a); 3056(28);
3094(58); 5719(28); s.n.(9); s.n.(10); s.n.(14);
s.n.(38); s.n.(43); s.n.(84).
Poiret herbarium, s.n.(3).
Porter, D. M. et al., 4318(46).
Porto, P. C. et al., 517(26); 1232(26); 2146(26);
2166(9); 2201(10); 2434(40); 2513(40);
2881(26); s.n.(40); s.n.(56).
Prance, G. T. et al., 1628(84); 5152(69); 5177(40);
5422(84); 5707(6); 5883(6); 5891(55); 6273(6);
6355(6); 6483(6); 6960(59); 7498(35); 8872(87);
10740(74); 11056(74); 11203(74); 14357(55);
14568(37); 14615(84); 19358(84); 20998(35);
20999(35); 23552(44a); 25108(84); P25282(79);
58167(21); 58533(81); 58607(84); 59071(57);
59176(84); 59504(84).
Prater, s.n.(26); s.n.(41).
Prescott, G. W., 1538(35).
Primack, R. & J. Utley, 716(46).
Puga, B., 22(40).
Puttemans, A., SP12010(16); s.n.(21).
Rabello, C., 1133(32).
Raben, F. C. C., 29(9); 141(84); 802(26); 892(26);
930(26); s.n.(26).
Raimundo, SP1122(32).
Ratter, J. A. et al., R11(18a); R32(58); R37(37);
R161(84); R280(84); R453(84); R896(24);
R1065(9); R1156(37); R1243(10); R1338(24);
R1470(37); R2751(15); R2764(26); R2809(16);
R2826(10); R2886(24); R2993(20); R3179(15);
R3226(20); R3418(58); R3444(17); R3452(21);
R3509(58); R3759(57); R3992(26).
Regnell, A. F., 1.29(26); 11.27(34); 11.29(41);
11.30(9); 11.1915(84); 11.2435(84); 111.149(69);
111.314(10);
111.313(16);
111.312(84);
III.315(18a, 19 and 34); 111.316(19);111.317(34
and 40).
Reichardt, 91(84); 92(18a); 93(21).
Reinhardt, s.n.(9).
Reitz, R. & R. M. Klein, 12424(65).
Revilla, J., 364(44b); 650(45); 712(44b); 2242(6).
Ribeiro, B. G. S., 1077(16).
Richard, L. C., s.n.(74).
Richardson, W. D., 747(44a).
Riedel, L. 26(26); 29(34); 46(9); 62(57); 134(9);
152(26); 355(63); 489(17); 578(34); 586(26);
Flora Neotropica
611(57); 801(19); 837(26); 908(30); 1004(9);
1009(41); 1013(40); 1016(40); 1017(40);
1022(40); 1023(40); 1036(3); 1037(56);
1527(26); 1898(16); 1964(9); 2210(21);
2469(18a); 11.1827(34); s.n.(9); s.n.(10);
s.n.(16); s.n.(19); s.n.(21); s.n.(26); s.n.(30);
s.n.(39); s.n.(41); s.n.(57); s.n.(62); s.n.(84).
Riley, L. A. M., 94(74).
Rivier, L., 1(35); 3(35).
Rizzo, A., 3976(14); 4065(14); 4068(14); 4639(26);
4682(14); 4684(26).
Robert, A., 523(55); 591(16).
Rodrigues, W. et al., 1251(84); 3110(84); 3292(84);
4312(84); 4576(44b).
Rodriguez, D., 1112(69).
Rodriguez, J. V., 79(31); 893(2a); 1363(31);
1630(31); 3249(2a).
Rodriguez, R. S., 3345(74).
R6hl, 34(51).
Roig, J. L. et al., 2087(42).
Rojas, T., 4159(12); 11467(40); 12917A(69);
13285(40); 13552(69); 13629(69).
Romariz, D., 0101(28); 0119(28); 0313(26);
0403(15).
Rombouts, J. E., 2675(84).
Romero-Castaneda, R., 10091(74).
Rond6n Commission, 169(69); 170(69); 1581(33a);
1629(19); 1659(19); 1891(37); 1892(58);
2236(84); 2438(16); 4758(40); 4803(9);
4895(19); 4896(19); 6387(26); 6704(28);
6706(28); 22037(84); 22128(44a); 22822(84).
Rose, J. N., 19952(18a); 21764(40).
Roth, L., 1588(9); 1592(40).
Rouhier, A., s.n.(35).
Ruano, J. M., 1341(2a).
Rugel, 848(42).
Ruiz, H. & Pavon, 33/64(69); 33/82(86); s.n.(40).
Rusby, H. H. et al., 110(84); 319(la); 354(la);
511(40); 514(40); 515(84); 663(86); 732(69);
852(55); 1033(55); 1109(40); 2171(86);
2418(40).
Rutkis, E., 209(35).
Saco, J. C., 1866(41).
Saer, J., 755(la).
Sagastegui-A., A., 5859(69); 5879(40).
Sagot, P., 96(74); s.n.(44a).
Sagra, R. de la, 29(42); 765(42); 882(42); s.n.(42).
Saint Hilaire, A. de, B1.125(26); B1.428(28);
B1.1104(18a);
B1.649(40);
B1.860(10);
B1.1117(28); B1.1132(15); B1.1142(28);
B1.1154(18a); B1.1365(15); B1.1617(84);
C1.642(20);
C1.117(26);
B1.1691(80);
C5.164(26);
C2.1186(78);
C1.762(84);
D.382(26); D.383(26); D.716(36); 705(32);
s.n.(9); s.n.(15); s.n.(20); s.n.(26); s.n.(28);
s.n.(57); s.n.(84).
Saint John, H., 20729(40).
Salzmann, 92(55).
Sampaio, A. J. de, 240(26); 240a(26); 5501(D1);
5512(D1); 6081(40); 6396(26); 6558(26);
6853(26); 6855(28); 7178(41); 7366(10);
7468(24); s.n.(28).

Sandwith, N. Y., 143(44a); 292(44a); 1430(54);
1441(44a).
Santoro, J., 527(10); 597(10); 849(84).
Santos, E., 1347(4); 1633(41).
Santos, N., R40549(40); R40550(9); R40551(9);
R40552(10);
R48595(9);
R48597(9);
R72015(16); R72031(40); s.n.(26).
Santos, R. S., s.n.(18a); s.n.(57).
Santos, R. S. & A. Castellanos, 24058(84);
24059(57); 24136(84); 24137(57); 24157(84);
24183(84); 24197(58); 24204(80).
Santos, T. S. dos, 442(39); 1318(4).
Sastre, C., 1105(8); 1218(91); 2173(44b).
Schenck, 3315(34).
Schinini, A., 4206(40); 4346(19); 4399(19);
4400(19); 4417(19); 4441(40); 7922(19);
9030(33a).
Schlim, L., 108(40).
Schmidt, L., s.n.(74).
Schneider, M., 1063(40).
Schomburgk, Rich. or Rob., 111(40); 844(40).
Schomburgk, Rich., 594(70); 773(7); 874(74);
999(44a); 1505(74).
Schomburgk, Rob., 488(7); 651(44a).
Schott, s.n.(38).
Schreiner, s.n.(40).
Schreiter, 11110(40); 11253(40); 11429(69).
Schiich, 5682(9); s.n.(40); s.n.(55).
Schultes, R. E. et al., 3037(35); 3346(35);
3836(44b); 13133(35); 13134(D2); 13156a(35);
13595(44b);
13615(44b);
14569(44b);
16387(44b); 17209(35); 17297(D2); 17459(D2);
s.n.(35).
Schulz, A. G., 5496(69).
Schulz, J. P., 8651(44a).
Schunke, C., 306(77).
Schunke, J. M., 63(44b).
Schunke-Vigo, J., 1952(40); 2037(40); 3863(40);
4635(88); 4709(40); 5882(40); 6344(35).
Schwacke, C. A., 21(16); 11.14(3); 11.93(26);
111.510(84); 5594(3); 6719(40); 7194(10);
7358(32); 7673(32); 8355(41); 8371(15);
8377(80); 8892(34); 10397(32); 10433(61);
10864(91); 12810(57); 13910(18a); s.n.(3);
s.n.(9); s.n.(10); s.n.(26); s.n.(34); s.n.(40).
Schwarz, G. J., 7193(40); 7330(40).
Seemann, s.n.(40).
Seler, 2419(40); 2428(2a).
Seifriz, W., 249(lb).
Sellow, F., 1288(16); 11.234(56); III.B1838c.
1304(9); III.B1839c.1305(32); III.B1841c.
1307(26); III.1843(18a); IV.304(78); IV.
5733(34); IV.5932(34); IV.5933(60); VI.
9561(57); s.n.(3); s.n.(10); s.n.(18a); s.n.(26);
s.n.(28).
Semir, J. et al., 496(27); 546(27); 4373(57);
4934(28 and 29); 5027(18a).
Sendulsky, T., 666(34).
Senege, M. V., s.n.(40).

Serre, s.n.(40).
Sesmero, E., s.n.(40).
Shank, P. J. et al., 4551(40).
Shattuck, 0., 645(6).
229
Shimoya, C., 3640(9).
Sick, H., s.n.(79).
Silva, A., 133(74).
Silva, H. D. C. da, s.n.(18).
Silva, J. F., 67(84).
Silva, M., 1746(84); 1871(84).
Silva, N. T. da et al., 729(10); 2726(84); 2859(59);
3656(86); 59727(35).
Silveira, A., 804(34).
Siqueira, O. P., 5(16).
Skutch, A. F., 4737(lb).
Smith, Au., 1829(46); P2415(40).
Smith, A. C., 3152(70); 3176(40); 10076(74).
Smith, C. E. et al., 3257(40).
Smith, F. D., 180(70); 253(70); 265(lb).
Smith, G. Dorrien, 202(21); 237(16).
Smith, H. H., 29(9); 35(26); 1509(46); 1524(68);
R72381(9).
Smith, J. D., 1698(46); 3703(40); 4121(2a);
5167(40); 6309(40); 8981(2a).
Smith, L. B., 6717(84); 6955(21); 6988(26);
14443(34); 14683(26); 14850(9).
Smith, S. G., 1135(la).
Sneidern, K. von, 5563(47).
Snethlage, E., 2552(40).
Snethlage, H., 615(84); 621(84); 701(84).
Sobrinho, J. S., 261(18a).
Sobrinho, L., 208(58).
Soejarto, D. D. & H. Cardozo, 674(44b).
Sota, E. de la, 636(40).
Spruce, R., 156(84); 240(84); 249(D1); 1012(36);
1727(84); 2712(35); 3276(74); 3885(40);
4199(69); 4272(84); 4530(40); s.n.(36); s.n.(84).
Standley, P. C. et al., 1868(40); 2417(40);
2688(40); 3471(2a); 4498(2a); 13888(2a);
19447(2a); 19691(2a); 19908(2a); 20949(2a);
21339(2a); 23240(40); 25377(40); 26876(40);
28725(2a); 36046(40); 62143(40); 75778(2a);
76178(2a); 77122(2a); 77454(2a); 87352(2a);
87795(lc); 88835(lc); 89073(2a); 91573(46);
91589(46); 92581(46); 92666(46).
Steinbach, J., 1075(7); 1189(40); 1601(69);
1712(69); 1764(40); 2819(84); 3018(84);
3242(40); 3260(7); 5602(40); 6386(69);
6438(69); 6495(55); 6530(55); 6817(74);
7043(40); 7087(40); 7305(84); 7384(7);
7603(69); 8186(69).
Stephan, M. D., s.n.(9); s.n.(28); s.n.(40);
s.n.(57).
Steward, W. C. et al., P12817(40); P20184(84).
Steyermark, J. A. et al., 83(54); 835(54);
1110(44a); 16776(6); 30745(40); 31646(2a);
31993(2a); 32938(2a); 37627(40); 38077(lb);
51164(40); 55102(51); 58559(44a); 58689(54);
59180(44a); 59710(44a); 60807(44a); 61914(52);
87415(44a); 89510(74); 90937(40); 91740(85);
92964(54); 93047(44a); 97422(40); 97497(51);
99089(51); 100303(51); 100363(40); 102333(lb);
104132(44a); 104601(44a); 105613(74);
105918(53); 109672(53); 111290(54).
Stiibel, 22(68).
Sucre, D. et al., 283(21); 667(41); 2651(40);
2661(32); 9231(16); 9599(3); 10268(84).
230
Tamayo, F. T., 253(75); 538(40); 616(40);
2160(40).
Tamberlik, s.n.(9); s.n.(10); s.n.(33a); s.n.(41).
Tate, G. H. H., 1120(40).
Tavares, S., 558(69).
Teixeira, A. de S., SP18834(9).
Tejera, E., 8(40).
Tessman, G., 4184(44b); 6049(40).
Thieme, C., 63(40).
Thouin, L., s.n.(3).
Tillett, S. S. et al., 44905(54); 45387(44a);
45467(44a); 45593(44a); 45597(54).
Toledo, F. T. de, 664(34); s.n.(34).
Tonduz, A., 1575(40);3079(40); 4581(40); 4719(40);
4746(40); 9080(40); 13477(40).
Toro, R. A., 585(49); 606(49).
Torres, J., 110(86).
Tovar, G., P2(40).
Tovar, 0., 4651(40).
Triana, J., 3394(la); s.n.(40); s.n.(46).
Trinta, Z. A., 251(4); 598(34); 602(10); 816(18a);
818(84); s.n.(34); s.n.(40).
Troll, C., 30(69); 957(7); 1783(40); 2693(55).
Trujillo, B., 3541(54); 5965(40).
Tryon, R. M. & A. F., 6791(27); 6813(26).
Tschudi, s.n.(84).
Tiirckheim, H. von, II.1785(46).
Tutin, T. G., 694(54).
Ule, E., 127(62); 143(84); 144-2787(57); 145(41);
421(26); 423(14); 2458(28); 2460(28);
2461(18a); 2462(18a); 2464(10); 5044(84);
6350(55); 6439(84); 6705(87); 6862(40);
7071(4); 7192(16); 7394(15); 7445(31);
7534(18b); 8892(84); 9477(6); 9480(40).
Uribe-Uribe, L., 2878(la).
Usteri, P. A., 28a(34); s.n.(41).
Valio, I. M., 247(21); 265(21); 267(26).
Van Herman, H. A., 189a(42).
Van Landsberge, 263(40); s.n.(40).
Van der Veen, R., 21(26).
Vargas, C., 1144(69).
Vargas C., C., 1842(40); 2044(69); 4571(48);
C7196(40); 12970(48); 17232(40).
Vargas, 302(40).
Vauthier, 4(3); 8(3); 91(38); 448(80); 455(15);
458(26); s.n.(21); s.n.(56); s.n.(80).
Vazquez, M. et al., V-951(D4).
Velloso, H., 432(60).
Venturi, S., 1426(69); 5190(69).
Vidal, J., 1-323(16); 1-324(16); 1-378(16); 1-390(40);
I-392(pp 10, pp 11); 1-446(10); I-467A(16); I484(24); 1-492(21); 1-497(34); 1-507(16); I513(16); 1-573(16); 1-590(16); 1-609(34); I620(26); 1-707(10); 1-708(40); 1-730(16); I778(11); 1-894(21); 1-1958(9); 11-304(62); II66123(28); 11-6202(26); 11-6268(26); II6343(26); 11-6373(24 and 26); 11-6383(26); II6385(26); 11-6391(26);11-6393(26);11-6777(32);
III-321(26); IV-1140(26); V-10(26); V-52(26);
V-156(9); V-163(9); V-175(9); V-179(9); V184(26); V-214(18a); V-221(26); V-227(18a);
V-238(26); s.n.(18a); s.n.(34); s.n.(40).
Flora Neotropica
Vidal, N., s.n.(9).
Viegas, A. P. et al., 3203(26); 3698(41); 3769(40);
3918(16); 4508(41); 5392(16).
Vieira, G. et al., 392(89).
Vogel, P. C., 471(70); 471a(70); 1120(40);
1121(40); 1123(40); 1124(40).
Wachsmund, A., s.n.(26).
Wacket, s.n.(16); s.n.(26).
Warming, E., 26(26); 29(24); 31(69); 286(84);
307(40); 353(57); 432(84); 1022(84); s.n.(9);
s.n.(10); s.n.(18a); s.n.(21); s.n.(24); s.n.(26);
s.n.(34); s.n.(40); s.n.(41); s.n.(57); s.n.(84).
Wawra, 39(60).
Weberbauer, A., 1918(74); 4794(73); 6063(72);
6160(40); 6230(73); 6466(72); 7935(40).
Weddell, A., 931(18a and 26); 1179(18a);
1592(18a); 1604(26); 1607(9); 1679(18a);
1954(9); 1959(24); 1996(43); 2073(20);
2074(14); 2094(41); 2465(84); 2574(16);
2608(16); 3278(33a); s.n.(21); s.n.(26).
Weddell, H. A., 4207(48).
Wedel, H. von, 2776(46); 2791(46).
Weir, J., 93(34); 112(26); 140(16); s.n.(34).
Werckle, 2065(40).
Werdermann, 2138(84); E.2375(69).
Wessels Boer, J. G., 977(74).
West, J., 7191(69).
Wetmore, R. H. & E. C. Abbe, 63(6).
White, G. E., 970(35).
White, O. E., 255(69).
Widgren, 179(26); 389(40); 390(9); 395(34); 753(26);
760(26); 990(38); 1006(34); s.n.(1); s.n.(18);
s.n.(26); s.n.(38).
Wilbur, R. L. et al., 11371(49); 13615(49);
19468(49).
Williams, LI., 330(35); 971(44b); 3348(35);
3523(35); 3741(35); 5426(84); 6157(84);
7274(74); 7401(40); 7776(74); 8111(36);
8224(35); 10304(lb); 10362(40); 10453(40);
10585(40); 10712(40); 10918(51); 11627(59);
13600(40); 14965(D2); 15719(74).
Williams, L. O. et al., 5079(26); 5099(9); 5189(10);
5470(26); 5505(18a); 5547(40); 5569(26);
5694(26); 5703(18a); 5728(26); 5771(18a);
5779(26); 5831(9); 5974(26); 6058(40);
6256(18a); 6268(18a); 6290(18a); 6304(21);
6399(9); 6528(41); 6586(34); 6686(41);
6793(34); 6929(21); 7335(84); 8139(26);
8196(58); 10572(40); 10878(2a); 11901(2a);
18334(40); 26705(40); 41306(40); 42403(40);
42559(lb).
Williams, R. 0., 10121(52).
Williams, R. S., 62(40); 102(7); 165(40); 192(40);
796(69); 809(40); 1511(84).
Wohon, A., 7(19).
Woolston, A. L., 790(40); 827(40); 955(40);
999(40); 1380(40); 1609(40).
Woronow, G., 7608(40).
Woytkowski, F., 19(44b); 37(40); 5045(35);
5072(40); 5074(35); 5076(35); 5119(35);
5332(40); 5581(84); 5588(40); 5640(40);
5674(40); 5719(40); 5762(40); 5879(84);
5909(84); 5917(69); 6499(69); 6801(69);
231
7197(40);7230(40);7258(40);7259(40and84);
7265(6);
7328(77);
7392(84);
7878(40);
8066(86);8135(47).
Wright, C., 95(42); 152(42); 347(42); 1172(42);
1584(42); 2152(42); 2158(42).
Wullschlaegel,
181(40); 182(40); 54(74);
54.196(74); 54.1357(74); 181(40); 182(40);

183(70);1387(74);1388(74);s.n.(74).
Wurdack, J. J. et al., B-156(84); 1051(47);
40976(6);41145(6);43130(74);43650(74).
Zarucchi,J. L. et al., 1817(84).
INDEX TO SCIENTIFICAND COMMONNAMES

Accepted taxa are in Roman type, with the new or newly combined names in
boldface; synonyms and commonnames are italicized. Page numbersin boldface
indicate detailed treatmentsof taxa. The letter f. before a page numberindicates
a figure.
Abejucado 55
Ala de cucaracha 46
Ala de zompopo 123
Almizclillo 42
Auca ayahuasca 198
Ayahuasca 113, 123, 214
Ayahuasca amarilla 113
Ayahuasca de los brujos 123
Ayahuasca negra 113, 123
Ayahuasca rosada 123
Banisteria L.
SubgenusEubanisteria(Grisebach)Niedenzu 3, 57
SubgenusHemiramma(Grisebach)Niedenzu 3, 57
SubgenusPleiopterysNiedenzu 3, 164
SubgenusSciurostylisSkottsberg164
Section Actinoctenia Niedenzu 176
Section AnisopterysGrisebach176
Section Camptostylis Niedenzu 57
Section Eubanisteria Grisebach 57
Section HemirammaGrisebach57
Section LeiantheraGrisebach57
Section Leiococca Niedenzu 3, 57
Section MonocteniaNiedenzu 3, 51
Section Orthostylis Niedenzu 57
Section Pseudobyrsonima Niedenzu 3
Section Trichanthera Niedenzu 57
SubsectionAnadenia Niedenzu 176

Subsection Cosmiothamnus Niedenzu 57
Subsection Microzeugma Niedenzu 57
SubsectionNarcothamnusNiedenzu 57
SubsectionOctadeniaNiedenzu 176
Subsection Pachyzeugma Niedenzu 57
Subsection Psilopetalum Niedenzu 64
SubsectionSericopetalumNiedenzu 176
Series CyrtantheleNiedenzu 176
Series OrthantheleNiedenzu 176
Series Psilothece Niedenzu 57
Series Trichothece Niedenzu 57
acanthocarpaAdr. Jussieu 120, 126

var. glandulifera Niedenzu 121
acerosa Niedenzu 74
aculeata Miller 204
adamantiumMartiusex Adr. Jussieu69

adenopodaAdr. Jussieu 110
var. subrotundaNiedenzu 110
albicans Grisebach126
albida Willdenowex Steudel 204
allophila Sieber ex Adr. Jussieu 204
alternifolia Steyermark 142
angulata Vellozo 204
angulosa L. 204
angustifolia Adr. Jussieu 98
anisandra Adr. Jussieu 57, 150
anoptera(Adr. Jussieu)Morton204
antifebrilis Grisebach 204
apiculata (Miquel)Robinsonin Small 204

appendiculata(DC.) Adr. Jussieu 177
arborescens(Brandegee)Small 204
argentea (H.B.K.) Sprengel 18, 120, 126
var. acuminataNiedenzu 121
var. obtusiuscula Niedenzu 121
var. transiensNiedenzu 121

argyrophaea(Adr. Jussieu)Morton204
argyrophylla Adr. Jussieu 57, 61
atrosanguineaAdr. Jussieu 121, 125

var. benthamiana(Adr. Jussieu) Macbride 121
aurea Vellozo 204
auriculata Cavanilles 204
beecheyana (Adr. Jussieu) Robinson in

Small 204
benghalensis L. 204
benthamianaAdr. Jussieu 121, 125

billbergiana Beurling 204
blanchetiana Adr. Jussieu 147
bonariensisHooker & Arnott 204

bopiana Rusby 196
brachiata L. 2, 204
brachiata Sprengel 204
brachiata Willdenow 204
brachyptera DC. 204
bracteata DC. 204
brevipes DC. 204
232
Flora Neotropica
caapi Spruceex Grisebach112, 217

caduciflora Niedenzu 198
caerulea Lamarck 204
calcitrapaDesvaux ex Hamilton204
calocarpa Miquel 177
campestris Adr. Jussieu 57, 93
var. glaucescens Grisebach 93
var. lancifolia Niedenzu 98
var. montana Adr. Jussieu ex Grisebach 98

var. ovata Grisebach93
var. rotundataGrisebach93
canaminensis Rusby 204

chrysophylla Bello 204
chrysophylla Lamarck 204
ciliata Lamarck 204
cinerascens (Bentham) Grisebach 54
var. glabrescens Niedenzu 52
var. platyphylla Niedenzu 54
cinerea Rusby 147
clausseniana Adr. Jussieu 57, 150
coccinea Vellozo 205
comans Martiusex Adr. Jussieu71

var. latifolia Adr. Jussieu71
var. pubifolia Adr. Jussieu71
var. stellaris Adr. Jussieu71
constricta Grisebach 191
convolvulifolia Cavanilles 205
cordata Vellozo 205
cornifolia(H.B.K.) Sprengel41
var. elliptica (Niedenzu)Niedenzu 43
var. leiocarpa (Niedenzu)Niedenzu44
corymbosa Larrafiaga 205
cotinifolia(Adr. Jussieu)Robinsonin Small

205
cristata Grisebach 171

crotonifolia Adr. Jussieu 76
var. angustata Grisebach76
cuiabensis Grisebach 205
deformisDesvaux ex Hamilton205
dichotoma L. 205
discolor (J. D. Smith)Niedenzu 205

dispar (Grisebach)Niedenzu 205
divaricata Adr. Jussieu 177
diversifolia H.B.K. 205
dumetorum(Grisebach)Morton205
eglandulata Vellozo 205
eglandulosa DC. 205
eglandulosaAdr. Jussieu 176, 201

elegans Triana& Planchon 134
ssp. cordata Niedenzu 134
var. ciliata Niedenzu 134
var. pubescens Niedenzu 140
var. grandifolia Niedenzu 141
ssp. ovata Niedenzu 134
var. guatemalensis Niedenzu 134

elliptica H.B.K. 205
emarginata Cavanilles 205
eneaptera Sess6 & Mociiio ex DC. 205

erianthera Adr. Jussieu 197
fagifolia DC. 205
ferruginea Cavanilles 48
ferruginea Seemann 205
fimbriata Grisebach 183
fimbriata Vellozo 205

fischeriana Regel & Koernicke 205
floribunda DC. 205
fruticosa Vellozo 205
fulgens Lamarck 205
fulgens L. 205
fulgens G. F. W. Meyer 205
gardneriana Adr. Jussieu 153
var. nummifera (Adr. Jussieu) Niedenzu 147
f. blanchetiana Niedenzu 147
f. luschnathiana Niedenzu 147
gayana (Adr. Jussieu) Robinson in Small 205
glauca Desfontaines 205
glaucophylla Martius in Mohl 205
grata Grisebach 57
guatemalensis Niedenzu 44
hassleriana Chodat 164, 168
helicina (Grisebach) Morton 205
heterocarpa Standley 205
heterophylla Willdenow 205
heterostyla Adr. Jussieu 167
hexandra Vellozo 205
hispida Mociiio & Sess6 ex Ramirez 205
hispida Niedenzu 168
humboldtiana DC. 205
hypericifolia Adr. Jussieu 186
hyposericea Standley & L. O. Williams 205
illustris Rusby 121
intermedia Adr. Jussieu 69, 70
jasminellum Adr. Jussieu 167
kraussiana Hochstetter 205
kuntzei Small 206
laevifolia Adr. Jussieu 57
var. grata (Grisebach) Niedenzu 57, 60
lanceolata Niedenzu 98
lancifolia West & Schlechtendal 206
latifolia Adr. Jussieu 82
var. paraguariensis Niedenzu 83
laurifolia L. 206
laurifolia Richard ex Adr. Jussieu 206
ledifolia H.B.K. 206
leiocarpa Adr. Jussieu 172
var. parvifolia Niedenzu 174
leona Cavanilles 206
leptocarpa Bentham 130, 132
lobulata E. Meyer 177
longialata Niedenzu 195
longifolia Swartz 206
lucida Richard 176, 177
lupuloides L. 206
lutea Grisebach 168
macradena DC. 206
macrocarpa Persoon 206
macrophylla Colla 206
macrophylla Adr. Jussieu 206
macrostachya Vellozo 206
magnoliaefolia Desvaux ex Hamilton 206
maracaybensis Adr. Jussieu 41, 42
var. angustifolia Grisebach 44
var. cornifolia (H.B.K.) Niedenzu 41
var. elliptica Niedenzu 43
var. leiocarpa Niedenzu 44
maritima Richard 206
Index
233
martiniana Adr. Jussieu 130
mathewsana(Adr. Jussieu)Gleasonex Rusby 206

mediterranea Vellozo 206
megacarpos Vellozo 206
megaphylla Adr. Jussieu 89
membranifolia Adr. Jussieu 57, 106
f. subglabrataNiedenzu 106, 107
metallicolor Adr. Jussieu 57, 120, 126
var. aurea Niedenzu 121

var. falcata Niedenzu 121
var. pruinosa (Martiusex Adr. Jussieu)

Niedenzu 121
var. sericea Niedenzu 121
var. subrotundaNiedenzu 121
var. subsalicina Niedenzu 121
michoacanensis Sess6 & Mocino 206
microphylla Hamilton 206
microphylla Jacquin 206
microphyllaSess6 & Mociiio 206

minarumMartiusex Adr. Jussieu71
monoptera Vellozo 206
monostyla Niedenzu 152, 154
montanaAdr. Jussieu 98, 99
mossii Morton 206

mucronata DC. 206
multialata Vellozo 206
multiflora DC. 206
multiflora Adr. Jussieu 206
multifoliolata Adr. Jussieu 118
muricataCavanilles120
var. atrosanguinea(Adr. Jussieu)Macbride 121
mutabilis Vellozo 206
nemorumBrandegee206
nervosa (Adr. Jussieu)R. O. Williams206
nigrescens Adr. Jussieu 183
nitens Grisebach61
nitida Lamarck206
nitida Vellozo 206
nitrosiodora Grisebach 164, 168
nummiferaAdr. Jussieu 147

nutans Niedenzu 179
obovata Small 206
orbicularis Niedenzu 171, 172
orinocensis H.B.K. 206
ovata Cavanilles206
ovata Niedenzu 49
pentandraSess6 & Mociiio 207

periplocifolia Desfontaines 207
peruviana Niedenzu 180
picta H.B.K. 207
platyptera Grisebach 194
f. grandifolia Niedenzu 197
podadena Turczaninow127
polita Miquel 177

polygama Niedenzu 136
populifolia Niedenzu 175
portillana(Watson)Robinsonin Small 207

praecox Grisebach168, 171
pragua Vellozo 207
procumbensSess6 & Mocifio207
pruinosa Martiusex Adr. Jussieu 121
pruriens E. Meyer 207
pseudo-janusia Niedenzu 162
pubera Richard207
pubiflora DC. 207
pubipetalaAdr. Jussieu 164, 176, 191

var. constricta (Grisebach) Macbride
191
var. oblongata Grisebach191
var. scandens Grisebach191
var. virgultosaAdr. Jussieu 188
f. constricta(Grisebach)Niedenzu 191
f. cordata Niedenzu 191, 194
f. spruceana(Grisebach)Niedenzu 191
purpurea L. 207
quapara Aublet 207
var. mucronulata DC. 207
quitensis Niedenzu 112
reticulata(Poiret)Robinsonin Small 207

retusa (J. D. Smith)Robinsonin Small 207
riedeliana Regel 207
rigida Adr. Jussieu 186, 187

rosea Standley207
rotundifolia Buc'hoz 207
rotundifolia Sess6 & Mocino 207
rusbyana Niedenzu 195
sagittata Cavanilles 207
salicifolia DC. 157
salicifolia sensu Grisebach128
sanguinea Rusby 207

schizoptera Adr. Jussieu 57, 69
var. minarum(Martiusex Adr. Jussieu)

Grisebach71
schlimiiTurczaninow121
oxyclada Adr. Jussieu 126

var. rhombifoliaSkottsberg126
schomburgkiana Bentham 121

schwannioides Grisebach 80, 114
scutellata Grisebach 158
f. brevialata Niedenzu 126

padifolia Niedenzu 137
sellowiana Adr. Jussieu 149
var. subleiocarpa Niedenzu 49
pallida (Brandegee)Standley206
palmata Cavanilles206
palmeri (Rose) Robinsonin Small 206
paniculata Mocino & Sess6 ex DC. 207
parviflora Adr. Jussieu 156
var. boliviensis Niedenzu 137
var. tomentosa Niedenzu 158

parvifolia Ventenat 207
patens Grisebach207
pauciflora H.B.K. 127

pearcei Rusby 191
f. obtusa Niedenzu 158

var. blanchetiana (Adr. Jussieu) Niedenzu 147
f. grandifolia Niedenzu 147
sepium Adr. Jussieu 203

sericea Cavanilles207
simulans Small 207

sinemariensis Aublet 207
sinemariensis DC. 207
sinuata DC. 207
solitaria Vellozo 207
sphaerandra Rusby 147
234
splendens DC. 207
spruceana Grisebach 191, 194
stellaris Grisebach 71
var. intermedia Adr. Jussieu 69
subcordata Gardner 110
suberosa (Grisebach) Robinson in Small 207
tenuis Lindley 207
terminalis Vellozo 207
terminariifolia Mohl 207
ternata Mociiio & Sesse 207
ternata Vellozo 207
ternstroemiifolia (Adr. Jussieu) Niedenzu
207
tetraptera Sonnerat 207
thyrsoidea Grisebach 207
tiliifolia H.B.K. 207
tiliifolia Ventenat 207
timoriensis DC. 207
tomentosa Desfontaines 207
tomentosa Schlechtendal 208
triflora Vellozo 208
trifoliolata Steudel 208
umbellata Sesse & Mocino 208
umbellulata DC. 208
unialata Vellozo 48
unicapsularis Lamarck 208
varia Sprengel 208
variifolia DC. 208
velutina Adr. Jussieu 80
velutina Martius ex Colla 208
velutina sensu Niedenzu 83
vernoniifolia Adr. Jussieu 67, 68
verruculosa Swartz 208
virgata Sesse & Mocifio 208
virgultosa Adr. Jussieu 188
viridis Niedenzu 156, 157
vitifolia Sesse & Mocifio ex DC. 208
volubilis Sess6 & Mocifio 208
wettsteinii Kralik 93
whitei Rusby 208
williamsii (Rusby) Niedenzu 121
wydlerana (Adr. Jussieu) Robinson in Small
208
Banisterieae 2
Banisteriopsis Robinson in Small
Subgenus Banisteriopsis 3, 4, 6, 8, 11, 12,
14, 16, 17, 18, 27, 39, 41, 50, f51, 56, 61
Subgenus Hemiramma (Grisebach) B. Gates
3, 4, 6, 8, 11, 12, 14, 16, 17, 18, 57, 200
Subgenus Pleiopterys (Niedenzu) B. Gates 3,
4, 5, 6, 8, 11, 12, 14, 16, 17, 18, 164, 200,
209
Section Anisopterys (Grisebach) B. Gates 5,
6, 11, 12, 14, 16, 18, 166, 176, 209
Section Sciurostylis (Skottsberg) B. Gates 5,
6, 11, 12, 14, 16, 17, 18, 164, f170
acapulcensis (Rose) Small 27, f43, 44
var. acapulcensis f43, 45
var. lanensis B. Gates 19, f43, 45, 46,
f47
acerosa (Niedenzu) B. Gates 19, 30, 31, 35,
f72, f74, 74
adenopoda (Adr. Jussieu) B. Gates 16, 26,
30, 35, 82, 105, 110, fl10
Flora Neotropica
alternifolia (Steyermark) B. Gates 10, 25, 38,
130, f139, 142
amplectens B. Gates 23, 40, f182, f187, 189
andersonii B. Gates 29, 37, f100, 101, f102,
104
angustifolia (Adr. Jussieu) B. Gates 29, 38,
68, f96, 97, 98, 102, 103
anisandra (Adr. Jussieu) B. Gates 8, 27, 39,
57, 150, fl51, 154
arborea B. Gates 28, 37, flOO, f102, 103
argentea (H.B.K.) Robinson in Small 121
argyrophylla group 6, 10, 17, 29, 36, 61, f62
argyrophylla (Adr. Jussieu) B. Gates 19, 29,
36, 37, 38, 57, f59, 61, f62, 64, f65, 66, 67
basifixa B. Gates 8, 26, 32, 33, 159, f160, 160,
162
brevipedicellata B. Gates 27, 39, f47, 50, f51
byssacea B. Gates 30, 37, f62, 63, f65, 66
caapi (Spruce ex Grisebach) Morton 6, 10,
14, 18, 19, 29, 30, 34, 108, 112, 116, fll17,
119
cabrerana Cuatrecasas 211
cachimbensis B. Gates 24, 40, 185, f187, 190
caduciflora (Niedenzu) B. Gates 5, 6, 23, 26,
27, 39, 167, 198, f199
calcicola B. Gates 6, 28, 35, 37, f96, 104
campestris group 6, 28, 37, 93, f94, f96, f102,
105
campestris (Adr. Jussieu) Little 10, 16, 29,
37, 38, 57, 93, f94, f96, 98, 104, 105
cinerascens (Bentham) B. Gates 27, 39, f51,
54
cipoensis B. Gates 17, 19, 29, 37, f94, 97,
f102, 209
clausseniana (Adr. Jussieu) Anderson &
Gates 150
confusa B. Gates 31, 35, 76, f79, 80, 112
cornifolia (H.B.K.) Robinson in Small 2, 6,
20, 28, 39, 41, f43, 49, 50, 52, 54, 55
var. cornifolia 42, f43
var. elliptica (Niedenzu) B. Gates 42,
f43, 46
var. leiocarpa (Niedenzu) Cuatrecasas
44
var. standleyi B. Gates f43, 44
cristata (Grisebach) Cuatrecasas 24, 26, 32,
168, f170, 171, f173
discolor (J. D. Smith) Small 208
dugandii Cuatrecasas 44, 46
elegans (Triana & Planchon) Sandwith 10,
25, 32, 33, 130, 134, f139, 140, 141
ssp. cordata (Niedenzu) Sandwith 135
var. ciliata (Niedenzu) Sandwith
135
var. pulcherrima Sandwith 145
erianthera (Adr. Jussieu) B. Gates 24, 40,
197, f199, 201
ferruginea (Cavanilles) B. Gates 27, 39, 48,
50, f51
gardneriana (Adr. Jussieu) Anderson &
Gates 8, 16, 17, 27, 39, 149, fl51, 152
goiana B. Gates 28, 36, f17, 120, f122, 128
grandifolia (Niedenzu) B. Gates 8, 25, 38,
130, f139, 141
Index235
guatemalensis (Niedenzu) Robinson in Small
44
harleyi B. Gates 29, 36, 61, f62, f65, 66
hatschbachii B. Gates 31, 34, 76, f85, 87
heterostyla (Adr. Jussieu) Cuatrecasas 5, 6,
8, 9, 16, 23, 26, 33, 164, 166, 167, f170,
f173
hirsuta B. Gates 31, 34, 76, f85, 86
hypericifolia (Adr. Jussieu) Anderson &
Gates f9, 16, 19, 23, 40, 166, 176, f182,
186, f187, 189
illustris Rusby 121
inebrians Morton 112, 113
irwinii B. Gates 28, 30, 35, 88, f90, f91, 92
krukoffii B. Gates 6, 19, 23, 39, f199, 200,
203, 209
laevifolia (Adr. Jussieu) B. Gates 6, 8, 14, 16,
17, 19, 26, 36, 57, f59, f60
latifolia (Adr. Jussieu) B. Gates 8, 9, 27, 30,
31, 34, 76, 82, f85
leiocarpa group 6, 14, 21, 25, 32, f170, 172
leiocarpa (Adr. Jussieu) B. Gates 25, 32,
f170, 172, f173, 175, 176
leptocarpa (Bentham) R. O. Williams 130
longialata (Niedenzu) B. Gates 24, 40, 194,
195, f199
longipilifera B. Gates 6, 28, 39, f47, f51, 56
lucida group 6, 18, 177, f184, 185, f187, 195
lucida (Richard) Small 24, 40, 176, 177, 180,
f182, f184
lutea group 6, 12, 24, 26, 32, 168, f170
lutea (Grisebach) Cuatrecasas 8, 9, 19, 24,
32, 164, 168, f170, 172, f173
lyrata B. Gates 26, 27, 39, f151, 154, f155
macedoana L. B. Smith 74
magdalenensis B. Gates 6, 26, 33, 34, f160,
163
maguirei B. Gates 25, 33, 38, 130, f133, 143,
f144
malifolia group 6, 12, 18, 31, 34, 75, 118, 119
malifolia (Nees & Martius) B. Gates 30, 31,
34, 35, 70, 76, f77, f79, 81, 82, 86
var. appressa B. Gates f79, 79, 116
var. malifolia f77, 78, f79, 80
martiniana group 6, 11, 12, 14, 16, 18, 24, 33,
129, f133, f139
martiniana (Adr. Jussieu) Cuatrecasas 17,
25, 32, 33, 130, f133, 134, 150
var. elegans (Triana & Planchon) Cuatrecasas 135
var. grandifolia (Niedenzu) Cuatrecasas 141
var. laevis Cuatrecasas 132, 133
var. martiniana 131, f133
var. popayanensis Cuatrecasas 136
var. subenervia Cuatrecasas 132, f133
megaphylla group 6, 18, 28, 30, 35, 88, f90,
f91
megaphylla (Adr. Jussieu) B. Gates 11, 28,
30, 35, 89, f90, f91, 93
megaptera B. Gates 28, 39, f47, 49, f51
membranifolia group 6, 12, 18, 30, 35, 105,
fl10
membranifolia (Adr. Jussieu) B. Gates 16,
26, 31, 34, 36, 57, 106, f107, 109, fll10, 113,
119
metallicolor (Adr. Jussieu) O'Donell &
Lourteig 121
var. sericea (Niedenzu) O'Donell &
Lourteig 121
var. subrotunda (Niedenzu) O'Donell &
Lourteig 121
multifoliolata (Adr. Jussieu) B. Gates 6, 18,
30, 34, 118, 120
muricata group 6, 12, 14, 17, 28, 36, 105,
f117, 120
muricata (Cavanilles) Cuatrecasas 19, 28, 32,
36, 37, 57, 109, 112, 118, 120, f122, 127,
128, 129, 130
nigrescens (Adr. Jussieu) B. Gates 24, 40,
180, 183, f187
nitrosiodora (Grisebach) O'Donell & Lourteig 168
nummifera group 6, 11, 12, 27, 38, 147, 150,
fl51
nummifera (Adr. Jussieu) B. Gates 11, 16,
27, 39, 147, f151
nutans (Niedenzu) B. Gates 24, 41, 179, 181,
f184, 185, 195
oxyclada (Adr. Jussieu) B. Gates 10, 19, 28,
36, f 17, 120, 126
padifolia (Niedenzu) B. Gates 25, 32, 38, 130,
137, f139
panamensis Cuatrecasas & Croat 179
paraguariensis B. Gates 17, 19, 29, 36, f62,
64, f65, 209
parviflora group 6, 12, 14, 16, 17, 33, 156,
f160
parviflora (Adr. Jussieu) B. Gates 10, 26, 33,
156, 158, f160, 163
parvifolia (Niedenzu) B. Gates 25, 32, f170,
174, 176
parviglandula B. Gates 8, 27, 33, f160, 161,
163
patula B. Gates 6, 19, 23, 33, 176, f199, 201,
209
pauciflora (H.B.K.) Robinson in Small 28,
36, 120, 127
peruviana (Niedenzu) B. Gates 24, 40, 179,
180, f184
platyptera (Grisebach) Cuatrecasas 24, 40,
180, 194, f199
polygama (Niedenzu) B. Gates 25, 32, 38,
130, 136, 138, f139, 143
populifolia (Niedenzu) B. Gates 25, 32, f170,
174, 175
prancei B. Gates 6, 29, 35, 38, f115, 116, f117
pseudo-janusia (Niedenzu) B. Gates 6, 16,
27, 33, f160, 162
pubescens (Niedenzu) Cuatrecasas 25, 32,
33, f139, 140
pubipetala group 6, 39, 190, f199, 200
pubipetala (Adr. Jussieu) Cuatrecasas 18, 23,
24, 40, 164, 176, 178, 179, 191, f192, 195,
196, 201
pulcherrima (Sandwith) B. Gates 16, 25, 33,
130, f133, 145, f146
pulchra B. Gates 19, 29, 30, 35, 108
Flora Neotropica
236
var. glabrata B. Gates 109, f 10
var. pulchra 109, fllO, 110
quadriglandula B. Gates 6, 29, 36, 119
quitensis (Niedenzu) Morton 112
rondoniensis B. Gates 24, 40, f182, f187, 189
rusbyana (Niedenzu) Morton 196
rusbyana sensu ethnobotanists 211, 214, 217
salicifolia (DC.) B. Gates 26, 32, 157, f160
scalariformis Schery 135
schizoptera group 6, 17, 30, 31, 35, 61, 69,
f72, f74
schizoptera (Adr. Jussieu) B. Gates 30, 31,
32, 35, 57, 69, 71, f72, f74
schomburgkiana (Bentham) Robinson in
Small 121
schunkei B. Gates 24, 40, f192, 198, f199, 200
schwannioides (Grisebach) B. Gates 6, 16,
17, 28, 35, 114, fll5, fl17, 154
scutellata (Grisebach) B. Gates 10, 27, 33,
158, f160, 161, 162
sellowiana (Adr. Jussieu) B. Gates 27, 39,
149, fl51
sepium (Adr. Jussieu) B. Gates 6, 17, 19, 23,
33, f192, f199, 203, 209
speciosa Small 134
stellaris (Grisebach) B. Gates 16, 30, 31, 35,
71, f72, f74, 75
sublucida Rusby 54
valvata Anderson & Gates 5, 6, 14, 16, 23,
25, 33, 164, f165, f170
variabilis group 76, f85
variabilis B. Gates 8, 11, 31, 34, 76, 79, 83,
f85, 88
velutinissima B. Gates 25, 32, 33, 130, f133,
133
vernoniifolia (Adr. Jussieu) B. Gates 19, 29,
36, f62, 67
virgultosa (Adr. Jussieu) Anderson & Gates
23, 40, f187, 188
whitei Rusby 208
wilburii B. Gates 25, 33, 130, 138, f139
williamsii Rusby 121
woytkowskii B. Gates 24, 40, 181, f182, f184
wurdackii B. Gates 27, 39, f51, 52, f53, 55,
56
Barboleta do campo 60
Bejuco colorado 168
Bejuco de casa 123
Bejuco golandrina 168
Bejuco hoja de plata 123
Cip6 prata 60, 63, 127, 153

Cip6 preto 194
Coloradito 168
Crista de gallo 194
Crista de galo 153
Cuchi-ayahuasca 113
Diplopterys Adr. Jussieu
bracteosa (Grisebach) Niedenzu 216
cabrerana (Cuatrecasas) B. Gates 20, 211,
f212, f213
cururuensis B. Gates 209, f212, 215
includens (Bentham) Niedenzu 216
involuta (Turczaninow) Niedenzu 216
marshalliana Riley 177
mexicana B. Gates 210, f212, 215
microcarpa Sandwith 216
paralias Adr. Jussieu 209, 210
pauciflora (G. F. W. Meyer) Niedenzu 209,
210, f212
var. latifolia Niedenzu 210
riparia (Adr. Jussieu) Niedenzu 216
rosea (Miquel) Niedenzu 216
spruceana Niedenzu 216
uleana Niedenzu 216
E-pe-pee-yoo-wee 133
Flor do dia 79
Gaudichaudioideae 2
Guarand-rana macho 178
Heteropterys H.B.K.
acapulcensis Rose 44
appendiculata DC. 177
argentea H.B.K. 120
bopiana (Rusby) Niedenzu 196
cinerascens Bentham 54
cornifolia H.B.K. 20, 40
micans Skottsberg 57
pirayuensis Morong 121
Hiraeeae 2
Huillca bejuco 170
Jubistylis Rusby
mollis Rusby 18, 168
Ka-hee-ko 214
Lucu'-tu'-654
Caapi 113
Carasucio 46
Cauupuri mariri 113
Chagropanga 214
Chagropanga azul pisco 214
Chinak 132
Cielo ayahuasca 113
Cipo de prata 152
Cipo de rego 194
Cipo de Sdo Jodo 170
Cipo folha de prata 127
Mabude hi 132
Malpighia L.
malifolia Nees & Martius 76
Mao de onca 113
Margadin 187
Marmelinha da flor branca 73
Mene kahi ma 214
Mune-yek 178
Natema 113, 214
Index
Nepe 113
Nucna-wasca 133
Nyoko-buku guda hubea ma 214
Oco yage 214
Pastora 123
Patillo 42
Planitorae 2
Pragua 112
Purga-huasca 113
Purga-huasca de los perros 113
Pyramidotorae 2
Rama de mogo 80
Sarcello 123
Shuri-fisopa 113
Sombra de tora 123
Stigmaphyllon Adr. Jussieu
nigrescens 0. Kuntze 177
237
Tetrapterys Cavanilles
acutifolia Cavanilles
var. ferruginea Grisebach 48
crebriflora Martius
var. lundiana Grisebach 48
pauciflora (G. F. W. Meyer) DC. 210
Tiwaco-mariri 113
Totorinmo 178
Triopterys L.
brasiliensis Poiret 48
pauciflora G. F. W. Meyer 210
Yaco-ayahuasco 214
Yage 113, 214
Yage del monte 113
Yage sembrado 113
Yageu'co 214
Yaje 214
Yaje oko 214
Yaji 214
Yukuna 132

Banisteriopsis, Diplopterys (Malpighiaceae) - CNCFlora

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Banisteriopsis, Diplopterys (Malpighiaceae) - CNCFlora

Uploaded by

Copyright:

Available Formats

Organization for Flora Neotropica

FLORA NEOTROPICA MONOGRAPH

The New York Botanical Garden

The New York BotanicalGarden

The New York Botanical Garden

Library of Congress Cataloging in Publication Data

Gates, Bronwen, 1945Banisteriopsis, Diplopterys(Malpighiaceae)

A MONOGRAPH OF BANISTERIOPSIS AND

University of Michigan Herbarium, Ann Arbor, Michigan 48109, U.S.A.

denzu (1900, 1912), who monographedthe Malpighiaceaefor Das Pflanzenreich

belong to the genus Heteropterys),and Robinson (in Small, 1910)introducedthe

The Malpighiaceaeare characterizedby opposite leaves, stipules, a five-parted

wing and have well-developedlateralwings. However I considerthat Diplopterys

of subgenus Hemirammaforms the basis for subgenus Banisteriopsis, and his

banisteria form the amended subgenus Hemiramma. The subgenus Pleiopterys

is maintained,based on some of the charactersused by Niedenzu, such as the

Paniculate, flowers borne alternately

Short-pedunculate, often thickened

Cymes or dichasia of four-flowe

Calyx glandular, glands free apically

Calyx glandular, rarely eglandu

Eglandular or glandular, the gla

Ovary velutinous or sericeous,

Short, up to 1 mm, broad, nonfunctional

2-4 mm long, functional

Spherical; surface smooth laterally or

Laterally compressed, rarely

Basal appendage on fruit

None or rarely a shallow appen

One to two pairs on the petiole or

Curved,either sigmoidor incurved

Section Anisopterys(spp. 74-92)

Banisteriopsis and Diplopterys

base, is the most common habit of Banisteriopsis species in savannavegetation,

gynoecium of species in the subgenus Pleiopterys. Most of the hairs in both

_' - *-".., '

ovary hairs of B. hypericifolia. Pubescence composed of hairs of types A or B is described as

up to 2 cm fromthe stem. This thickcork layerdevelopsonly on old stems of

plopterys and subgenus Banisteriopsis, section Anisopterys, and the B. nummi-

The flowers of Diplopterys and Banisteriopsis are borne on pedicels which in

some species in the subgenus Banisteriopsis and section Anisopterys of Banis-

is denoted by a distinctjoint; the peduncle bears a pair of bracteoles which are

B. membranifolia group), but are persistent in most species of Banisteriopsis,

tiniana groupof Banisteriopsisthe ultimatebranchesbear many flowers arranged

plopterys and most species of subgenus Hemiramma and subgenus Pleiopterys,

Banisteriopsis and Diplopterys

FIG. 2. Types of inflorescencein Banisteriopsis, and arrangementof bracts and bracteoles. A,

lateral and postero-lateralsepals, and an unpairedsepal, the anteriorsepal (Fig.

Banisteriopsis and Diplopterys

The petals of Diplopterys and of subgenera Banisteriopsis and Pleiopterys of

martiniana and B. nummiferagroups have yellow petals; the other species in

terys have externallysericeous petals. In addition, B. heterostyla and B. valvata

enlarged. The glandulartissue in Diplopterys and in the subgenus Pleiopterys

There are no recorded chromosomecounts for Diplopterys.

Source of the count

Fouit, 1966, mitotic root

leaf cell count

2. Numbersobtainedin the currentstudy

Lianas, shrubs, vining or not, treelets, or shrublets with woody xylopodium.

a short peduncle 0.5-4.0(-7.0) mm long. Flowers hermaphroditic.Calyx of 5

the trabeculae more or less parallel, the pubescence is described as sericeous

' ' ' '

? . ? ?-., .~? ',.