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BANISTERIOPSIS, DIPLOPTERYS
(Malpighiaceae)
BRONWEN GATES
^.\
C
f?,
T
TROPIC
...
Of
CANCER
FLORA!
NEOTROPICA,
TROPIC
Of
CAP R ICORN
Copyright ? 1982
All material subject to this copyright may be photocopied for the non-commercial purpose of
scientific or educational advancement.
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
Flora Neotropica
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
Banisteriopsisand Diplopterys
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
Pedicels
Calyx glands
Flower color
Stamen connectives
Eglandular
Gynoecium
Ovary short-appressed-sericeous,
styles glabrous or basally
appressed-pubescent
Carpophore
Nut
Shallow, up to 2 mm high
Banisteriopsisand Diplopterys
Table II
Comparisonof the Sections of Subgenus Pleiopterys
Section Sciurostylis(spp. 67-73)
Leaf glands
Inflorescence
Calyx glands
Absent
Petals
Glabrous,except B. heterostyla
Styles
Fruit
Carpophoreup to 2 mm long,
apparentlyfunctional;hairs on
fruit appressed,stiff and irritating
to the skin
Slender, not thickenedand woody in
fruit
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
--
--^~~~~~~~~~~~
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
stem often becomes very thick and forms a series of prominentridges projecting
Banisteriopsisand Diplopterys
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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
Banisteriopsisand Diplopterys
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-
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
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
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-
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-
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
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14
Flora Neotropica
..
15
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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
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-
Banisteriopsisand Diplopterys
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.
Banisteriopsisand Diplopterys
19
Table IV
ChromosomeNumbers in Banisteriopsis
1. Numbersfrom the literature
Species
B. muricata (as Banisteria
argentea)
"B. laevifolia"
B. caapi
Count
2n = 20
n = 20
2n = 20
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.
20
Flora Neotropica
Banisteriopsisand Diplopterys
21
22
FloraNeotropica
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wing..................;
Banisteriopsisand Diplopterys
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
86. B. longialata.
long.
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
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.
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.
Flora Neotropica
26
68. B. heterostyla.
63. B. basifixa.
61. B. salicifolia.
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.
66. B. magdalenensis.
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.
63. B. basifixa.
27
62. Pedicels sparselyvelutinous;stem velutinous.
62. B. scutellata.
65. B. pseudojanusia.
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.
89. B. caduciflora.
Subgenus Banisteriopsis.
6. B. wurdackii.
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.
3. B. ferruginea.
Flora Neotropica
28
4. B. megaptera.
1. B. cornifolia.
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.
25. B. irwinii.
24. B. megaphylla.
Banisteriopsisand Diplopterys
29
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.
92. Laminahairyabaxially.
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.
14. B. vernoniifolia.
12. B. paraguariensis.
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.
long.
20. B. latifolia.
18. B. malifolia.
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.
Banisteriopsisand Diplopterys
31
21. B. variabilis.
21. B. variabilis.
B. schizoptera group.
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.
70. B. cristata.
Banisteriopsisand Diplopterys
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.
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.
64. B. parviglandula.
abaxially.
63. B. basifixa.
30. Sepals tomento-sericeousabaxially;base of lamina stronglycordate, almost claspingthe stem, petiole 2-4(-7) mm long.
50. B. pubescens.
44. B. martiniana.
54. B. pulcherrima.
34
Flora Neotropica
36. Petiole up to 3 mm long; laminaovate to rotund.
66. B. magdalenensis.
35. B. caapi.
38. B. multifoliolata.
32. B. membranifolia.
late.
B. malifolia group.
21. B. variabilis.
35
B. megaphylla group.
25. B. irwinii.
reduced.
mm long.
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.
36
Flora Neotropica
appressed-sericeous to glabrate, the hairs
sessile, the trabecula0.1-0.25 mm long.
32. B. membranifolia.
B. muricata group.
42. B. pauciflora.
cm long, ovate to
41. B. oxyclada.
14. B. vernoniifolia.
13. B. harleyi.
68. Lamina chartaceous, with the reticulation impressed and the areoles
betweenthe veins shallowlyinflated.
10. B. argyrophylla.
12. B. paraguariensis.
13. B. harleyi.
37
11. B. bvssacea.
10. B. argyrophylla.
4 mm high.
27. B. cipoensis.
30. B. arborea.
26. B. campestris.
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.
77. Laminahairyabaxially.
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.
26. B. campestris.
long-ciliate.
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.
10. B. argyrophylla.
85. Leaves with a single pair of glands at the apex of the petiole.
B. nummifera group.
Banisteriopsisand Diplopterys
39
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.
nut.
5. B. brevipedicellata.
7. B. cinerascens.
101. Laminaabaxiallytomentose.
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.
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.
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. Laminaabaxiallytomentose.
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.
81. B. virgultosa.
80. B. hypericifolia.
77. B. woytkowskii.
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.
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].
Banisteria maracaybensis Adr. Jussieu var. cornifolia (H.B.K.) Niedenzu, Ind. Lect. Lyc.
Brunsb.:5. 1900.
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.
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
,+
'
..-'*.
...'
, f,.^,7 ,d^4
tf-?
'.
V.
+,,
,s
;-?
..
--
+"
-.'
:.:
4-
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).
Banisteria guatemalensis Niedenzu, Ind. Lect. Lyc. Brunsb.: 6. 1900. Type. Seler 2428, Gua-
Banisteria maracaybensis var. leiocarpa Niedenzu, Ind. Lect. Lyc. Brunsb.: 5. 1900. Types.
totypes, MO, NY). J. D. Smith 4121 (leg Heyde & Lux), Guatemala, Santa Rosa, Naranjo,
Banisteriopsisand Diplopterys
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.
46
Flora Neotropica
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
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
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,
Tetrapterys acutifolia var. ferruginea Grisebach, Linnaea 13: 236. 1839. Type. Sellow s.n., Bra-
Banisteriopsisand Diplopterys
49
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).
50
Flora Neotropica
Banisteriopsisand Diplopterys
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.
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.
fO(GH,
NY); 8779
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).
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
54
Flora Neotropica
Vine or spreadingshrub,branchesdensely appressed-sericeous,eventuallyglabrate, light to dark brown, with prominentpale lenticels. Stipules up to 1.5 mm
Banisteriopsisand Diplopterys
55
Flora Neotropica
56
(MICH);32897 fl fr (MICH).
Banisteriopsisand Diplopterys
57
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.
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
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-
'-
'
?'.
Banisteriopsisand Diplopterys
61
Figs. 9, 12.
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-
Banisteriopsisand Diplopterys
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
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).
64
Flora Neotropica
florescentia floribus in umbellis 4-floriferisvel racemis usque 12-floriferisfasciculatis, umbellis in cymis dispositis, semel vel bis composita, laxa densave, cinereo- vel 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).
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 ~;..
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
Banisteriopsisand Diplopterys
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
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).
Banisteriopsisand Diplopterys
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)
Fig. 13.
70
Flora Neotropica
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
Banisteriopsisand Diplopterys
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.
Jussieu, fragment).
Banisteria minarum Martius ex Adr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 283. 1840. Type.
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.
Flora Neotropica
72
'Is:.
A
:
FIG. 13. Banisteriopsis schizoptera group. A-E, B. stellaris. A, flower bud x 3.2; B, flower x 5;
Banisteriopsisand Diplopterys
73
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.
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).
Banisteriopsisand Diplopterys
75
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
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
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).
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
79
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).
80
Flora Neotropica
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.
Banisteriopsisand Diplopterys
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).
82
Flora Neotropica
Banisteriopsisand Diplopterys
83
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
85
Banisteriopsisand Diplopterys
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
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
Banisteriopsisand Diplopterys
87
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
Banisteriopsisand Diplopterys
89
Fig. 18.
BanisteriamegaphyllaAdr. Jussieu, Ann. Sci. Nat. Bot., Ser. 2, 13: 282. 1840.
Flora Neotropica
90
r9?
i
i'
,:
'
''
?ij
"
..
,..
*:
.-
..
FIG. 18. Banisteriopsis megaphylla group. A-F, B. megaphylla. A, flower bud x 4.2; B, flow-
?..- ..
91
. .
..
. . .
. ...
*1
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
Banisteriopsisand Diplopterys
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
Banisteria campestris var. ovata Grisebach, Linnaea 13: 196. 1839. Type. Sellow s.n., Brazil,
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,
Banisteriopsisand Diplopterys
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).
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
Banisteriopsisand Diplopterys
97
Flora Neotropica
98
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)
Banisteriopsisand Diplopterys
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
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.)
Banisteriopsisand Diplopterys
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.
102
Flora Neotropica
t
-44
-0Q
'
A.
..
40" .
i0
23.
FIG.
Distribution
cipoesis;closd
FIG
of
of part
cirle,
Banisteriopsis
anerson\
group,
campestris
cicle,
opn
arore
27,
species
29,
30.
B.~~~~~~~~
Triangle,
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;
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).
Banisteriopsisand Diplopterys
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).
Banisteriopsisand Diplopterys
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
capitatis. Fructus ex 3 samaris (vel abortu 1-2) constans; samara carpophoro
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).
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
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).
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
Banisteriopsisand Diplopterys
109
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,
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-
Banisteriopsisand Diplopterys
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
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.
113
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.
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
Fig. 26.
Banisteriopsisand Diplopterys
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.
118
Flora Neotropica
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.
Banisteriopsisand Diplopterys
119
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.
120
Flora Neotropica
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.
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,
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,
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).
Banisteriopsis williamsii Rusby, Bull. New York Bot. Gard. 8: 97. 1912. Type. R. S. Williams
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.
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.,
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
x 1.4. J-L,
B. goiana.
gynoecium x 7. (A-F, Anderson 11712; G, Gates 401; H, Woytkowski7615; J-L, Hatschbach &
38237.)
Kummrow 38237.)
Banisteriopsisand Diplopterys
123
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);
Banisteriopsisand Diplopterys
125
Montes 15562 fr (G, W); Schwarz 7193 fl (P, S). Corrientes: Huidobro 2210 fl (S, W); Ibarrola 4030
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
126
Flora Neotropica
Banisteriopsisand Diplopterys
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.
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
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).
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-
Banisteriopsisand Diplopterys
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
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)
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.
Banisteriopsisand Diplopterys
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.
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).
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 &
133
+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
Williams 971 f
19 closed
open (F,
square
US). San Martinan var. subenervia
,US)
114569t(;
circle,
ft (A,
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).
Banisteriopsisand Diplopterys
135
223. 1943.
Banisteriopsis scalariformis Schery, Ann. Missouri Bot. Gard. 7: 92. 1943. Type. H. von Wedel
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
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
Banisteriopsisand Diplopterys
137
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).
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).
139
Banisteriopsisand Diplopterys
+.
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.
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).
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).
Banisteriopsisand Diplopterys
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).
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.
Banisteriopsisand Diplopterys
143
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;
Banisteriopsisand Diplopterys
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.
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.)
Banisteriopsisand Diplopterys
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
Banisteriopsisand Diplopterys
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.
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
(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).
Banisteriopsisand Diplopterys
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
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
FIG.. 34.
155
Banisteriopsis lyrata. a, leafy shoot and inflorescence on older leafless stem x 0.5; b,
156
Flora Neotropica
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.
Banisteriopsisand Diplopterys
157
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).
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
Banisteriopsisand Diplopterys
159
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.
Banisteriopsisand Diplopterys
161
162
Flora Neotropica
Banisteriopsisand Diplopterys
163
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.
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.
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
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
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.
Banisteriopsisand Diplopterys
167
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).
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
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).
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
Banisteriopsisand Diplopterys
169
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
Banisteriopsisand Diplopterys
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.
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
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.
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,
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).
Banisteriopsisand Diplopterys
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
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).
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.
Banisteriopsisand Diplopterys
177
Diplopterys marshalliana Riley, Kew Bull.: 135. 1925. Type. L. A. M. Riley 94, Trinidad, Arena
178
Flora Neotropica
Banisteriopsisand Diplopterys
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.
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).
Banisteriopsisand Diplopterys
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.
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;
Banisteriopsisand Diplopterys
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).
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
Banisteriopsisand Diplopterys
185
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,
Fig. 39.
187
*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.
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.
Banisteriopsisand Diplopterys
189
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
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)
Banisteriopsisand Diplopterys
191
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,
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FIG.
loer,sd
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42. Ban~~~~~~~~iseipipuieaaB.shneanB.spu.AEB.pbpta.,ifor-
yocu
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ri
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tmna
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ri
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.(-,Gts32;F
eer
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hc n lsy .-. mln,aial
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ptas
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)
Banisteriopsisand Diplopterys
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).
Banisteriopsisand Diplopterys
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,
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).
Banisteriopsisand Diplopterys
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).
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).
199
Banisteriopsisand Diplopterys
?.TROPICA...E..CA
TI
"
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n ' rr;~
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. ?
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amp no!1_,
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
Banisteriopsisand Diplopterys
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).
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
have chartaceous leaves, but the abaxial leaf pubescence is more dense, the
Banisteriopsisand Diplopterys
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
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
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.
Banisteriopsisand Diplopterys
205
Jussieu.
Jussieu.
fide Niedenzu.
denzu.
denzu.
Niedenzu.
sieu.
Niedenzu.
B. hexandra Vellozo =Schwannia elegans (Adr. Jussieu) Adr. Jussieu fide Nie-
denzu.
Cuatrecasas& Croat.
206
Flora Neotropica
fide Niedenzu.
Niedenzu.
B.
B.
B.
B.
B.
denzu.
denzu.
denzu.
Adr. Jussieu.
B.
B.
B.
B.
B.
denzu.
denzu.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
B.
denzu.
207
208
Flora Neotropica
Gates.
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.
Banisteriopsisand Diplopterys
209
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,
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.
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.
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).
Banisteriopsisand Diplopterys
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).
Flora Neotropica
212
X
':",; ,
FIG. 44.
...
'.
}'.
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
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
214
Flora Neotropica
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."
Banisteriopsisand Diplopterys
215
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
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
Banisteriopsisand Diplopterys
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.
Flora Neotropica
218
Robinsonin Small
a. var. cornifolia
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
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
B.
B.
B.
B.
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
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
.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.
84. B. pubipetala(Jussieu)Cuatrecasas
85.
86.
87.
88.
89.
90.
91.
B.
B.
B.
B.
B.
B.
B.
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).
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).
Banisteriopsisand Diplopterys
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);
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).
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
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);
3996(68);6258(1a);6348(40).
Havard,V., 54(42).
Hayes, S., 229(40); 284(40); 770(6).
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);
Banisteriopsisand Diplopterys
225
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).
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
Banisteriopsisand Diplopterys
and 29); 5804(80);
5818(15); 5818a(15);
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).
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
Banisteriopsisand Diplopterys
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,
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
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
acerosa Niedenzu 74
aculeata Miller 204
anoptera(Adr. Jussieu)Morton204
antifebrilis Grisebach 204
232
Flora Neotropica
calcitrapaDesvaux ex Hamilton204
calocarpa Miquel 177
campestris Adr. Jussieu 57, 93
var. glaucescens Grisebach 93
var. lancifolia Niedenzu 98
cornifolia(H.B.K.) Sprengel41
var. elliptica (Niedenzu)Niedenzu 43
var. leiocarpa (Niedenzu)Niedenzu44
corymbosa Larrafiaga 205
deformisDesvaux ex Hamilton205
dichotoma L. 205
dumetorum(Grisebach)Morton205
eglandulata Vellozo 205
eglandulosa DC. 205
Index
233
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
ovata Cavanilles206
ovata Niedenzu 49
podadena Turczaninow127
pubera Richard207
pubiflora DC. 207
pallida (Brandegee)Standley206
palmata Cavanilles206
palmeri (Rose) Robinsonin Small 206
paniculata Mocino & Sess6 ex DC. 207
parviflora Adr. Jussieu 156
var. boliviensis Niedenzu 137
patens Grisebach207
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
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