THE IDENTITIES OF THE SERICEOUS.LEAVED SPECIES OF

Transcription

Contr.Univ. MichiganHerb. 79 393-413.1993.
THE IDENTITIES OF THE SERICEOUS.LEAVED SPECIES
OF STIGMAPHYLLON (MALPIGHIACEAE) IN THE
AMAZON REGION
ChristianeAnderson
Universityof MichiganHerbarium
North UniversityBuilding
Ann Arbor, MI 48109-1057
INTRODUCTION
Stigmaphyllon, one of the Neotropical wing-fruited genera of the Malpighiaceae,includes nearly 100 species.Most are characterrzedby long-petioled broad
leaves, yellow flowers grouped in umbels or pseudoracemesdisposedin dichasially
an androeciumof 10 heteromorphicstamens,and a gynoebranched inflorescences,
cium whose 3 styles bear apical appendages,the folioles, for which the genus is
named. The samara usually consistsof an ovoid nut with a large flaring dorsal
wing and often also with lateral ornamentation (one to several small lateral winglets, spurs, and/or crests).Both nomenclatural and taxonomic problems have confused the identities of some of the representatives occurring in the Amazon
region in which the leaves are sericeous below, i.e., the trabecula of the hair
straight and sessileto subsessile.This group of plants was found to comprise four
species,S. argenteum C. Anderson (recently described), S. cardiophyllum Adr.
Juss.(a name long ignored), S. convolvulifolium Adr. Juss.(the epithet traditionally attributed to Cavanilles), and S. sinuatum (DC.) Adr. Juss. (a name long
misapplied). For these, the names S. brachiatum Tr. & Pl., S. convolvulifolium
Adr. Juss.,S. fulgens Adr. Juss.,S. hypoleucum Miquel, S. martianum Adt. Juss.,
and S. splendensCuatr. have been most commonly used. These four speciesare
discussedhere in detail. Four sympatric species,S. lacunosum Adr. Juss.,S. maynense Huber, S. paraense C. Anderson, and S. puberun (Rich.) Adr. Juss. are
included in the key presented below; although distinctive, they are occasionally
confused with one of the other four, because they also have abaxially sericeous
leaves.
The very different S. paralias Adr. Juss.,also with the leavessericeousbelow,
is unlikely to be confused with any other speciesof the genus. It is mentioned
here. because it has been recorded from the area of Tucuruf in Par6, Braztl.
Stigmaphyllon paralias is a shrub of dry and sandy areas of eastern Brazil from
MaranhSo to Rio de Janeiro. Its usually elliptical to lanceolate leaves have very
short petioles (up to 1.5 cm long), and the stipules are often fused acrossthe node
into a bifid structure. The inflorescenceis most commonly a solitary umbel of (3-)
4-I5 large flowers or sometimes a dichasium of umbels. The peduncles are very
short, only 0.02-0.2 times as long as the pedicels.The greatly modified "samara"
consists of a nut lacking a carpophore and bearing an apical crest, the rudimentary
dorsal wins.
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C O N T R .U N I V E R S I T Y O F M I C H I G A N H E R B A R I U M
VOLUME 19
TAXONOMY
[Note: Atypical specimens of S. cardiophyllum from E,cuador,considercd putative hybrids,
m a y n o t k e y h e r c ; s e e d i s c u s s i o nb e l o w u n d e r S . c a r d i o p h y l t u m . l
l. Anthers pubescent.
2 . L a m i n a s v e r y s p a r s e l ys e r i c e o u st o g l a b r o u s b e l o w . m a r g i n w i t h i r r e g u l a r l y s p a c e ds e s s i l e
glands and the bases of broken-off filiform glands; anthers of stamens opposite the
p o s t e r i o r - l a t e r a l s e p a l sw i t h t h e c o n n e c t i v e e n l a r g e d a n d b e a r i n g o n l y o n e l o c u l e t d o r s a l
wing of the samara 2.7-4 cm long, the nut 2.5-3.5 mm in diameter, without air chambers:
along river banks in lowland and flood plain forest, at forest edge, and in secondary
g r o w t h a n d c a p o e i r a so f A m a z o n i a n E c u a d o r , p e r u , B r a z i l , a n d B o l i v i a .
S. cardiophvllum Adr. Juss.
2 . L a m i n a s e v e n l y s e r i c e o u sb e l o w ; m a r g i n a l g l a n c l sn a i l - l i k e ,i . e . , a d i s k b o r n e o n a s t a l k u p
to 0.5 mm long (sometimes a few glands sessilc);anthers of stamens oppositc the posteriorlateral scpalsnot modified, the connective and both locules subequally long; dorsal wing
o f t h e s a m a r a r e d u c e d a n d c a . 2 c m h i g h m e a s u r e df r o m b a s e o f n u t , t h e n u t c a . 1 2 . - 5m m
in diameter, the locule surrounded by air chamberst along river banks and in inundated
forest in Amazonian Brazil and Peru.
S . l a c u n o s u r nA d r . J u s s .
l. Anthcrsglahrous.
3. Antcrior stylc and its opposingstamen much larger than the posterior stylesand their
o p p o s i n g s t a m e n s ;p o s t e r i o r s t y l e s f o l i o l a t e .
4 . L a m i n a s s p a r s e l y s e r i c e o u sb e l o w o r r a r e l y t h e h a i r s m o r e a b u n d a n t b u t t h c e p i d e r m i s a l w a y s v i s i b l e ; p e t a l s d i g i t a t e - f i m b r i a t e ; d o r s a l w i n g o f s a m a r a e r e c t a n d t a p e r ed
from the base, lateral winglets and carpophore absent; in rain forests and gallery
f o r e s t s ,a l o n g r i v e r b a n k s , a n d i n m a n g r o v e s w a m p s i n t h e W e s t I n c l i e s ,C e n t r a l A m e r ica. and northcrn South America (not reported from Ecuador): Venczuela (Delta
Amacuro, Monagas), Guyana, French Guiana. Ccllombia (northernmost Choc6 and
Antioquia, Putumayo). Peru (northern Loreto, Huiinuco), Amazonian Bra7ll (Amapri,
P a r 6 , ,A m a z 6 n a s ,A c r e ) .
S. puberum (Rich.) Adr. Juss.
4 . L a m i n a s v e r y d e n s e l y s e r i c e o u sb c l o w , t h e e p i d c r m i s o b s c u r e d t p e t a l s e r o s c t o e r o s e denticulate; dorsal wing of samara flared distally, the nut bearing 3-4 lateral winglets
per side. carpophore present; in wet forests and at water's edge in E,cuador(Napo,
P a s t a z a )a n d P e r u ( A m a z o n a s . H u d n u c c l , L o r e t o . S a n M a r t i n , P a s c o ,M a d r e d e D i o s ) .
o n e c o l l e c t i o n f r o m B r a z l l ( s o u t h w e s t e r nA m a z d n a s ) .
S . m a r - n e n s eH u b e r .
3 . A n t e r i o r s t y l e a n d i t s c l p p o s i n gs t a m e n s h o r t e r t h a n o r a t m o s t s u b e q u a l t o t h e p o s t er i o r
s t y l e sa n d t h e i r o p p o s i n gs t a m c n s ;p o s t e r i o r s t y l e sf o l i o l a t e o r e f o l i o l a t e .
5. Stamens oppositc the posterior-lateral sepals subequal to the stamen opposite the
posterior petal (flag), their anthers unmodified.
6 . L a m i n a s a p p e a r i n g g l a b r o u s b e l o w t o t h e n a k e d c y e . b u t o f t e n v e r y s p a r s e l vs c r i c e o u s ,t h e h a i r s c a . 0 . 1 ( - 0 . 2 ) m m l o n g a n d w i d e l y s p a c e d .n e v e r t o u c h i n g ;i n m o i s t
forcst, along rivers. and also in secondary growth and along roadsidesin Guyana.
Suriname, French Guiana, and northeastern Brazil (Amapii and easternparii). also
recorded from Martinique and Trinidad.
S. convolvulifoliunt Adr. Juss.
6. Laminas sparsely to very dcnsely sericeousbelow, the hairs (0.2-) 0.3-0.5 (-0.7)
mm long. usually touching to overlapping; common in primary and sccondary forest. especiallywet forest, along rivers, in thickets, and at roadsides in Colombia.
Venezuela, the Guianas, northern Brazil, Ecuador. northern Peru, and Amazonian
Bolivia.
S. sinuatum (DC ) Adr. Juss.
5 . S t a m e n s o p p o s i t e t h e p o s t e r i o r - l a t e r a l s e p a l su n l i k e t h e s t a m e n o p p o s i t e t h e p o s t e r i o r
petal (flag), their anthers modified: the connective enlarged and bearing 0-2 reduced
locules.
7. Limb of lateral petals 6-7 mm in diameter, erose; nut of samara without lateral
w i n g l e t s a n d a i r c h a m b e r s ; i n f o r e s t s a n d t h i c k e t s a n d a t r o a d s i d e so f e a s t e r n p e r u
( H u i i n u c o , J u n f n , L o r e t o , P a s c o ,S a n M a r t i n ) .
S. argenteurn C. Anderson.
7 . L i m b o f l a t e r a l p e t a l s 1 0 - l - 5m m i n d i a m e t e r , w i t h f i m b r i a e u p t o 0 . 8 m m l o n g ; n u t
of samara with one or more pairs of lateral winglets, locule surrounded by air
chambers; in woods along rivers, in wet localities in savanna ancl campo, and in
varzeaof Brazil (Goiiis, Maranhdo, western Piauf, Parr{).
S . p a r a e n s eC . A n c l e r s o n .
1993
C . A N D E R S O N :S T I G M A P H Y L L O N I N T H E A M A Z O N R E G I O N
395
Stigmaphyllon argenteum C. Anderson, Novon 2:302.1992.-Tvpe: PEntr.Hudnuco:
Prov. Pachitea,Dtto. Honoria, Bosque Nacional de Iparia, a lo largo del
Rfo Pachitea cerca del campamento Miel de Abejas, 1 km arriba del
pueblo Tournavista o unos 20 km arriba de la confluencia con el Rio
U c a y a l i , 300-400 m,30 May 1967,S chunke V .2018 (hol otype: N Y! ; isoFig.1.
t y p e sC
: OL! F! G! US!).
Vi n e to 1 4 m . Lami nas 2.5-15.3cm l ong, 5.7-14 cm w i de, tri angul ar , ovat e,
elliptical to suborbicular, or sometimes 3-5-lobed, apex acuminate, base truncate
to cordate, sparselysericeousto usually glabrous above, sericeousbelow (trabecula 0.2-0.5 mm long, straight,sessile),margin with irregularly spacedsessileglands
(0.5-0.6 mm in diameter) and with filiform glands (up to 1.5 mm long), with a pair
of prominent but sessileglands at the apex of the petiole, each gland 1.5-3.5 mm
in diameter; petioles 2-10+ cm long; stipules 0.7-1.2 mm long and wide, eglandular. Flowers ca. 15-30 per umbel, these borne in dichasia or compound dichasia.
Peduncles3-7.5 mm long, pedicels4-8.5 mm long; peduncles0.6-1.2 times as long
as the pedicels. Bracts 0.9-1.3 mm long, 0.6-1 mm wide, narrowly triangular;
bracteoles0.7-1.2 mm long, 0.6-1 mm wide, triangular, eglandular. Sepals 1.8-2.3
mm l o n g , 7 .5 _ 2 mm w i de, gl ands 1.6-2.3 mm l ong, 0.6-1.2 mm w i de. L at er al
petals with the limbs orbicular or broadly obovate, glabrous,yellow, margin erose;
anterior-lateral petals: claw 1.8-2.2 mm long, limb ca.J mm long and wide, posterior-lateral petals: claw 0.5-1 mm long, limb 6-6.7 mm long, 4.5-6 mm wide;
posterior petal: claw 2.5-2.8 mm long, apex strongly indented, limb 5-5.6 mm
long, 3.5-4.8 mm wide, elliptical or broadly obovate, margin erose to fimbriatedenticulate, teeth/fimbriae up to 0.5 mm long. Stamens unequal, those opposite
the posterior styles the largest, anthers of those opposite the lateral sepals with
the connective enlarged and the locules reduced; anthers all loculate, glabrous.
those of stamens opposite the anterior-lateral sepalswith 1 or 2 locules, those of
stamensopposite the posterior-lateral sepalswith only 1 locule. Anterior style ca.
2.2 mm long, shorter than the posterior two, glabrous; apex ca. 1.5 mm long, each
foliole ca. 1.4 mm long, ca. 1.2 mm wide, subsquare.Posterior styles 2.6-3 mm
long, glabrous, lyrate; foliole ca.7.4-2 mm long and wide, subsquare.Dorsal wing
o f s a ma ra c a .4 . 5 cm l ong, 1.4-I.7 cm w i de, upper margi n w i th a bl unt t oot h,
lateral winglets absent, nut only prominently ribbed; nut 4-5.5 mm high,3.5-4.5
mm in diameter, areole 3-3.5 mm long,2.5-2.8 mm wide, concave,carpophore up
to 1 .8 m m l o n g . Embryo 5.8-7.3mm l ong, ca.2 ti mes as l ong as w i de, ovoi d, out er
c o ty l e d o n 6 .1 -8 .3mm l ong,2.6-3.9 mm w i de, the di stal 1/6 fol ded over th e inner
cotyledon, inner cotyledon 4-6.6 mm long, 2-3.6 mm wide, straight or the tip
folded back on itself.
Phenology. Collected in flower from April through July; in fruit in May and
from July through September.
Distribution. Lowlands of eastern Peru; in forests and thickets and at roadsides;135-670m.
R E p R p s p N r n r r v eS p e c r v e r u sP. e r u . H u A l u c o : P r o v . P a c h i t e a , r e g i o n o f P u c a l l p a , c a . 2 6 k m S t < l
24 km SSE of Puerto Inca, N of Rio Yuyapichis, 09"37'-09"34'S, 74"56'-74"53'W, Wallndfer Il3l5BB (MICH); vicinity of Tingo Marfa, 3-,5 km from Huiinuco-Tingo Maria rd on Monz6n rd,
M a t h i a s & T a v l o r - 1 6 4 7( F . U C L A ) . - J u x i N : P u e r t o B e r m u d e z , K i l l i p & S m i t h 2 6 6 3 0 ( F . N Y , U S ) ;
P r o v . S a t i p o , E b a n k o f R i o E n e a t m o u t h o f R f o Q u i p a c h i a r t , M a d i s o n 1 0 4 2 7 - 7 0( F ) . - L o n e ' r o :
Quebrada Shanuceabove Yurimaguas,Croat 17999(MICH); Isla de Ushpa-canonear mouth of the
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CONTR. UNIVERSITY OF MICHIGAN HERBARIUM
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,ffi_ffi
7
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itl
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F I G . 1. S t i g m a p h l t l l o n a r g e n t e u m . a . P o r t i o n o f b r a n c h w i t h l a r g e l e a f . b . D e t a i l o f a b a x i a l
surface of lamina. c. Small leaf. d. Flowering branch. e. Posterior petal. f. Androecium, second
stamen from left opposes posterior petal. g. Gynoecium, anterior style to the right. h. Samara.
i.
Two views of an embryo. Scale for a, c. d, h, bar = 1 cm; for b, bar = 0.5 mm; for e-g. bar =
1 mm.
(Based on: a, b, d-g, Schunke V.20tB; c, h, i, Croat 19640.)
R i o l t a y a ' C r o a t 1 9 6 4 0( M I C H ) ; U c a y a l i , B o s q u e N a c i o n a l A l e x a n d e r v o n H u m b o l d t , b e t w e e n K m
9 0 - 1 3 0 o f P u c a l l p a - T i n g oM a r i a r d , 0 8 " 4 8 ' S , 7 5 ' 2 0 ' W , G e n t r y e t a l . 4 t 4 t 3 ( M O ) : w o o d e d b a n k s
on
lower Rio Huallaga, Killip & Smith 29004 (F, GH, NY); Prov. Maynas, vicinity of Iquiros,
Rio
Mom6n, quebrada Momoncillo, McKenna et al. DMK-91 (AMAZ, F, MICH, MO).-pnsgo:
Oxap a m p a , c a . 5 k m u p R f o I s c o z a c i n f r o m v i l l a g e o f I s c o z a c f n , 1 0 " 1 2S , 7 5 ' 1 3 ' W , K n a p p
& Staver
7802A (MICH); PalcazuValley. Rio San Jos6 in the Rio Chuchurras drainage, 10.09'S. 75"20'W.
D.
Smith 4002 (MICH).-SIN
MnnriN: between Tocache Nuevo and Juanjui, 18.7 km S of Rio pulcache, 07"55'S, 76"40 W, Croat 58052 (MICH); vicinity of Aguaytfa, Boquer6n de padre
Abad.
M a t h i a s & T a y l o r - 1 5 9 1 , 6 0 9 2( F , U C L A ) , P r o v . M a r i s c a l C d c e r e s , D t t o . T o c a c h e N u e v o , q u e b r a d a
de Santiago' al E de Puerto Pizana, Schunke V. 6530 (GH, MO); Prov. Mariscal Ciiceres. Dtto,
Tocache Nuevo, quebrada de Cachiyaca, afluente de la quebrada de Huaquista, al E
de puerto
Pizana, Schunke V. B52B (F. MICH. MO).
Stigmaphyllon argenteum is named for the silvery pubescence on the abaxial
leaf surfaces.It is distinguishedby its small flowers, in which the petal limbs are
only up to 7 mm in diameter and the anthers of stamens opposite the posteriorlateral sepalsbear only one locule, and by its samaras,which lack laterai winglets.
Stigmaphyllon cardiophyllum differs in its abundantly pubescent anthers and its
leaves,which may be very sparselysericeousbelow but most commonly areglabrate
to glabrous. In S. sinuatum, the flowers are borne in pseudoracemesinstead of
umbels, the petals are usually larger (up to 13 mm in diameter), the stamens
opposite the posterior-lateral sepalshave unmodified anthers and are subequal to
the stamen opposite the posterior petal, the styles are commonly pubesceni (glubrous in S. argenteum),and the nut of the samara usually bears lateral *lngi"tr.
stigmaphyllon convolvulifolium and S. argenteum are not sympatric
r 993
C ] .A N D E R S O N : S T I G M A P H Y L L O N
IN THE AMAZON
REGION
391
Adr. Juss.,Ann. Sci.Nat. Bot., s6,r.2,13289.1840.cardiophyllum
Stigmaphyllon
Type: "Brasiliaborealis,"collectorunknown (holotype:P!, photos:F!
Fig.Z.
MICH!, fragment:P-JU!).
V i n e to 1 5 m. Lami nasl .3-17 cm l ong,3.1-13.5 cm w i de, ovate to el lipt ical
(the smaller often narrowly elliptical) to suborbicular, sometimes shallowly to
deeply 2-3-lobed, apex acuminate to acuminate-caudate,base truncate to cordate
or sometimes attenuate, especiallyin smaller laminas, very sparsely sericeousto
glabrous above and below (trabecula 0.2-0.9 mm long, straight, sessileto subsessile),margin with irregularly spaced sessileglands (0.2-0.6 mm in diameter)
and the bases of broken-off filiform glands, with a pair of prominent but sessile
glands at the apex of the petiole, each gland 1.1-3 mm in diameter; petioles 2.1-B
cm long; stipules 0.5-1 mm long, 0.5-1.4 mm wide, eglandular. Flowers ca. 15-25
(-30) per umbel, these borne in dichasia or compound dichasia or small thyrses.
P e d u n c l e s(1 .5 -) 2.2-5 mm l ong, pedi cel s4-10 mm l ong; peduncl es0.3-1 ti mes as
long as the pedicels. Bracts 0.7-2 mm long, 0.5-0.8 mm wide, triangular or narrowly so, bracteoles0.4-1 mm long, 0.4-0.8 mm wide, triangular, usually eglandular or sometimes each bracteole with a pair of inconspicuousglands (each ca. 0.1
mm i n d i a m e te r) . S epal s1.2-2 mm l ong and w i de, gl ands 1-1.8 mm l ong, 0. 7- 7
mm wide. All petals with the limbs glabrous, yellow; lateral petals with the limbs
orbicular, margin erose; anterior-lateral petals: claw 1-1.8 (-2.1) mm long, limb
(5 -) 6 -6 .5 mm l o ng and w i de; posteri or-l ateralpetal s:cl aw 0.5-1 (-1.3) mm long,
limb 4-5 mm long and wide; posterior petal: claw 2-2.5 mm long, apex indented,
limb (3.2-) 3.5+ mm long, (2.3-) 3-4 mm wide, broadly obovate to broadly elliptical
to suborbicular, margin irregularly denticulate to denticulate-eroseto sometimes
erose. Stamens unequal, those opposite the posterior styles the largest but their
filaments subequal to those of stamens opposite the anterior-lateral sepals, anthers of those opposite the lateral sepals with the connective enlarged and the
locules reduced, those opposite the posterior-lateral sepals usually with only 1
tiny locule; anthers all loculate, pubescent.Anterior style 2-2.3 mm long, shorter
than the posterior two, glabrous; apex 0.9-1.1 mm long; each foliole 0.5-0.8 mm
long, 0.4-0.8 mm wide, triangular to square. Posterior styles 2.2-2.6 mm long,
glabrous,lyrate; foliole 0.9-1.2 mm long, 0.8-1.4 mm wide, subsquareto subtrapezoidal. Dorsal wing of samara 2.74 cm long, 1.1-1.8 cm wide, upper margin with
a blunt tooth; lateral winglets absent, nut only prominently ribbed; nut 4.I-5.5
mm h i g h ,2 .5 -3 .5 mm i n di ameter, areol e 2.5-3.3 mm l ong,2.1-2.5 mm wide,
convex, carpophore up to 3 mm long. Embryo 4.5-6.4 mm long, ca.2 times as long
a s w i d e , o v o i d , o uter cotyl edon4.2-5.9mm l ong,25-3.1 mm w i de, strai gh t ,inner
cotyledon 3.9-5.5 mm long, 1.8-2.8mm wide, straight.
Phenology. Collected in flower from May through February, in fruit from
June through April.
Distribution. Amazonian lowlands of Ecuador, Peru, Brazll, and Bolivia; along
river banks in lowland and floodplain forest, at forest edge, and in secondary
growth and capoeiras;sea level to 1600m.
: staci6n Biol6gica Jatun Sacha,8 km al este de
. cuador. N,q.poE
R n p R r . s p . N r r R r rS
v np E c l v E N s E
Misahualli,01"04'S, 7J"36'W, Cer6n M. & Iguago 5592 (MICH); R(o Napo, Puerto Napo, Harling
P u e r t o S a r a y a c u ,L u g o 5 . 3 8 9 9 ( G B , M I C H ) .
- f 5 / B ( S ) ; M i s a h u a l l i , S r e i n e r2 7 5 ( } l l c H ) . - P n s r a z n :
Peru. AunznNAS: Rfo Cenepa, vicinity of Humapimi, ca. 5 km E of Chdvez Valdivia, ca. 04'30'S,
7 8 o 3 0 ' W ,A n c u a s h l l 3 4 ( F , M I C H , M O ) ; V a l l e d e S a n t i a g o ,Q u e b r a d a C a t e r p i z a , 0 3 " 5 0 ' S , 7 7 o 4 0 ' W ,
T u n q u i 7 0 S( M I C H , M O ) , A r a m a n g o , W o y t k o w s k i 5 6 2 9 ( G , G H , M O , U S ) . - C u z c o : P r o v . P a u c a r t ambo, Atalaya, near junction of Rio Carb6n and Rfo Alto Madre de Dios, Foster3041 (MICH).-
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CONTR. UNIVERSITY OF MICT{IGAN HERBARIUM
VOLUME 19
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F I G . 2 . S t i g m u p h v l l o n c a r d i o p h y l l u n t . a . F l o w e r i n g b r a n c h ( x 0 . . 5 ) .b . D e t a i l o f a b a x i a l l e a f
s u r f a c c ( x 5 ) . c - f . L e a v e s i l l u s t r a t i n gv a r i a t i o n o f l a m i n a r s h a p e ( x 0 . , 5 ) g
. . P o s t e r i o rp e t a l ( x 2 . 5 ) . h .
A n d r o e c i u m ( x 1 0 ) , s e c o n d s t a m e n f r o m l e f t o p p o s e sp o s t e r i o r p e t a l . i . G y n o e c i u m ( x 7 . - 5 ) a. n t e r i o r
stylc at center. j. Samara (xl). k. Embryo (x2.-5).(Based on: a. b, f-i, Enc:arnaci1n26052: c, Ancuash
I134: d,Wo-v"tkowski5954: c. Nee 36821: i,k. Foster 6506.)
t993
C . A N D E R S O N :S . I I G M A P H Y L L O NI N T H E A M A Z O N R E G I O N
399
HuANrrco: Tingo Marfa, Asplund 12103 (S); Prov. Hu:inuco, Maranjillo, cerca a Tingo Maria. Ferreyra
-l'ingo
Maria on rd to Tocachc, Huallaga Valley, ca. 09"S, 76"W, Gentry
2196 (MICH); 69 km NE, of
e t a t . 3 7 6 3 3( M I C H ) . - J l r N i N : S a n i B e n i , W o y t k o w s k i 5 9 5 4 ( G H , M O . U S ) l M a z a m a r i , W o y t k o w s k i
5 9 7 9 ( M O , L J S ) ;S a n R a m 6 n , W o y t k o w s k i 7 4 1 2 ( G H . M O ) . - L o R E r o : A l t o A m a z o n a s . D t t o . P a s t a z a . R i o P a s t a z a ,A y a l a 2 2 9 5 ( A M A Z , M I C H ) ; C o r o n e l P o r t i l l o , T o u r n a v i s t a . m d r g e n i z q u i e r d a d e l
R i o P a c h i t e a . E n t : a r n a c i 6 n2 6 0 5 2 ( M O . N Y ) ; P r o v . C o r o n e l P o r t i l l o , B o s q u e N a c i o n a l d e v o n H u m boldt, Km tl6, Pucallpa-TingoMaria rd,08"40'S,75'00'W, Gentry & Horner 29181 (AMAZ, MICH);
M i s h u y a c u , n e a r I q u i t o s , K l u g 1 6 0 ( F , N Y , U S ) ; S t r o m g e b i e t d e s M a r a f l c i n v o n I q u i t o s a u f w z i r t sb i s
z u r S a n t i a g o - M i i n d u n g a m P o n g o d e M a n s e r i c h e , 7 7 " 3 0 ' W , T e s s m a n3 9 6 6 ( G , S ) ; P r o v . M a y n a s ,
I q u i t o s , Q u i s t a c o c h a , 0 3 ' 4 8 ' S , 7 3 " 2 5 ' W . V d s q t r e z& J a r a m i l l o 1 2 0 7 2 ( M I C H ) . - M n o n E o E D t o s :
P r o v . ' f a m b o p a t a . L a g o T r c s C h i m b a d a s ,c a . 6 5 r i v e r k m S S W o f P u e r t o M a l d o n a d o . c a . 1 0 - 1 5 a i r
k m N W e f f l u e n c e R i o L a T o r r e ( R i o D ' O r b i g n y ) / R i o T a m b o p a t a , 1 2 " 4 9S . 6 9 " 1 7 ' W , B a r b o u r 5 7 4 4
( M I C H ) l P r o v . M a n u , P a r q u eN a c i o n a l d e l M a n u , R i o M a n u , C o c h a C a s h u S t a t i o n ,1 1 ' 5 0 ' S ,7 1 ' 2 5 ' W ,
F-oste6
r 506 (F).9704 (F, MICH)I small tributary of Rfo Madre de Dios, 1 km below Puerto Maldon a d o . G e n t r l t e t a l . 1 9 6 5 4 ( F , M I C H ) . - S n x M , c n r i N : P r o v . M a r i s c a l C 6 c e r e s ,e n t r e P 6 l v o r a y C h i o t e ,
valle Huallaga. Ferrevra 4464 (MICH, LJS); Prov. San Martin, cerca de Shapaja, Ferreyra 18273
(US); Tarapoto, U/e 61-lS(G, K, MG, NY); Pona to Saposoa, Woytkowski 5442 (F. MO). Brazil.
AcRp: 9 km from Rio Branco on Rio Branco-Porto Acre rd, Lowrie 646 (INPA, MICH); Mpio.
S e r r a M a d u r e i r a . R i o C a e t 6 . a f l u e n t e d o R i o l a c o , R a o s e t a \ . 6 4 3 ( I N P A ) ; R i o J u r u i i - J u r u dM i r i m ,
B6ca do Acre, Purf s, Goeldi 3969 (MG); Tonantins (Solimoes),
Ute 5593(G, Mc).-AunzoxRs:
Jobert 761 (P): Mpio. Sdo Paulo de Olivenqa, near Palmares, Krukoff 8291 (A, BR, F, G. LE,
M I C H , M O , N Y . P . S . U ) . - M R n R N u A o : b e t w e e n V i a n a a n d B a n d e r a n t e ,c a . 0 3 " 0 0 ' S ,4 5 " 1 0 ' W ,D a l y
e t a l . 6 1 8 ( N Y ) ; A l z i l A n d i a , R i o P i n d a r d , 0 3 ' 4 5 ' S , 4 6 " 0 5 ' W ,l a n g o u x & B a h i a 3 2 1 ( M I C H ) . - P n n A :
A l e n q u e r , C o l 6 n i a L a u r o S o d r 6 , K m 1 5 , F r ( t e s2 9 3 7 8 ( l A N ) ; I l h a d e M o s q u e i r o , n e a r P a r 6 , K i l l i p &
Smith 30394 (NY); Rd BR-22, Capanema to Maranh6o, Km 96, Prance & Pennington lB24 (IAN.
Pnz: Prov. Sud
M I C H , N Y ) . B o l i v i a . B e N r : B o p i R i v e r v a l l e y , R u s b y- l S 5 ( K , M I C H , N Y , U S ) . - L I
Yungas, Alto Beni, carretera entre Puente Sapecho y Santa Ana, Seidel & Schulte 2301 (MICH)I
Prov. Nor Yungas, Alto Beni, camino del puente hacia San Antonio, Seidel & Schulte 2320 (MICH).PnNne: Prov. Manuripi. antes de Independencia, Moraes 264 (MICH); Prov. Manuripi. along Rio
Madre de Dios. U0 km (by air) downstreamfrom and W of Chibe, l1'54'S,68"02'W, Nee 31525
(MICH)I Rio Acre, im Walde bei Cobija, Ille 9484 (G, K, MG).-S,+nrn Cnuz: Prov. Andr6s Ibiinez, 12
k m E o f c e n t e r o f S a n t a C r u z o n r d t o C o t o c a , 1 7 " 4 6 ' S , 6 3 ' 0 4 ' W ,N e e 3 6 8 2 1 ( M I C H ) .
Stigmaphytlon cardiophyllum, a common speciesof the Amazonian lowlands,
,
is readily recognized by its very sparsely sericeous to usually glabrous leaves,
small flowers, and samaras.The laminas are mostly ovate to elliptical (the smaller
narrowly so) and only sometimes cordate, as in the type. The limbs of the petals
are only up to 6.5 mm in diameter. The anthers are pubescent,and those opposite
the posterior-lateral sepals bear only one tiny locule (very rarely two). The samara
lacks lateral ornamentation. This speciesmay be confused with forms of S. sinuatum with abaxially very sparsely pubescent leaves, and, in eastern Brazll (Parii),
with S. convolvulifolium, whose leaves are also very sparsely sericeous to glabrous below. Both differ in their larger flowers (limbs of the petals up to 15 mm
in diameter), glabrous anthers (all with two locules), usually pubescent styles
(glabrous in S. cardiophytlum), and samarasthat are usually laterally ornamented
with winglets, crests, and/or spurs. For a separation from S. argenteum,see that
species.
Several collections from Ecuador and one from Peru may represent hybrids
b e t w e e n S . c a r d i o p h y l t u m a n d a n u n k n o w n s p e c i e s ,p o s s i b l y t h e s y m p a t r i c
S. sinuatum.They differ from typical specimensof S. cardiophyllum in their leaves,
anthers,posterior styles,and, sometimes,the samaras.The laminas are persistently sparsely sericeousbelow (the trabecula 0.3-0.5 mm long, sessile).The anthers
are only sparsely pubescent or glabrous, and those of the posterior-lateral stamens may have two unequal locules. The posterior styles are sparsely pubescent
in the proximal 1/3, and the nut of the samara may have a small lateral winglet or
400
CONTR. UNIVERSITY OF MICHIGAN HERBARIUM
VOLUME 19
spur. Pollen of these anomalous specimens is mostly composed of misshapen,
heavy-walled grains. When placed in cotton blue in lactophenol, only 0-20% of
the pollen is stained; the grains that do stain are of unequal size. The following
collectionsare consideredputative hybrids:
E c u a d o r . M o n o r u n - S n N r r A c o :a l r e d e d o r e s d e l p u e n t e s o b r e e l R f o B o m b i o z a , c a r r e t e r a G u a l a quiza-Zamora cerca la Paroquia de Bombioza, M. Baker 6479 (MICH).-Nero:
Estaci6n Bioldgica
Jatun Sacha, 8 km al este de Misahuallf, 01"04'S, 77o36'W, Cer6n M. 2038 (MICH); Payamino,
R e s e r v a F l o r i s t i c a " E l C h u n c h o , " - 5k m a l N W d e C o c a , 0 0 o 3 0 ' S , 7 7 o 0 1 ' W , C e r 6 n M . & N e i l t 2 3 6 7
(MICH); v(a Puerto Napo-Misahuallf, Jaramillo 87 (AAU, NY, QCA); Rfo Napo between Coca
(Puerto Francisco de Orellana) and Armenia Vieja, Harling & Andersson 11977 (GB, MICH);
S a n t a R o s a a t R f o N a p o , L u g o 1 6 8 , 1 9 8 1 ,2 0 0 1 , 2 0 2 7 ( G B , M I C H ) ; M i s a h u a l l i a t R i o N a p o , L u g o
2273 (GB, MICH).-Zevonn-CutNCHIpE: near M6ndez, Camp E-853 (NY, US). Peru. ArrrnzoNAs:
Rfo Cenepa, vicinity of Huampami, ca. 5 km E of Chiivez Valdfvia, ca. 04"30'5, 78'30 W, Ancuash
1 2 6 2( F , M r C H , M O ) .
The name Stigmaphyllon cardiophyllum has long been ignored, because most
later botanists did not see Jussieu'stype and found his description not definitive
enough to assignthis name with confidence.Collections of this specieswere often
left undetermined, assigned to S. convolvulifolium, or labeled with one of the
synonyms of S. sinuatum. Grisebach (1858) did recognize S. cardiophyllum and
correctly listed a Spruce collection with heart-shapedleavesfrom Bentham's herbarium (Spruce 1644, K!); however, Niedenzu (1900) tentatively placed S. cardiophyllum in S. salzmannii Adr. Juss. Niedenzu later (1928) listed S. cardiophyttum
as a separatespeciesbut only quoted Jussieu'sdescription from the Monographie
(1843) and cited Grisebach (1858). Macbride (1949) included S. cardiophyllum at
the species level in his account of the Malpighiaceae of Peru. He noted that
Williams 6883 was distributed as .S.cardiophyllum [correctly identified, F!], but
that it "is probably referable to S. tiliaefolium or allied form." He cited other
collections of S. cardiophyllum under other names, e.g., Tessmann 3412 (NY!)
and 3966 (G! S!) as S. convolvulifolium, and Klug 160 (Fl NY! US!) as S. puberum. Examination of the Amazonian sericeous-leavedspeciesof Stigmaphyllon
and comparison with Jussieu'stype revealed the distinctive speciesto which the
name S. cardiophyllum applies.
Stigmaphyllon convolvulifolium Adr. Juss., Ann. Sci. Nat. Bot., s,i,r.2, 13 289.
1840.-TvpE: FReNcnGutaNn. Cayenne, Martin s.n. (lectotype, designate d b y C . A nderson, 1987:P !).
Stigmaphyllon latifolium Bentham, London J. Bot. 7: 728.1848.-TypE: SuRrN AME .H ostmann 146 (hol otype: K -herb. B entham!, photo: MICH! ; isoty p e s :B M! G! K -herb. H ooker! N Y -fragment! P ! U ! W !).
Vi n e to 1 5 m. Lami nas 5-15 cm l ong, 4.5-11.5cm w i de, ovate to co r dat e or
narrowly so, apex acuminate-mucronate,base cordate, very sparselysericeousto
soon glabrate or glabrous above, very sparselyand minutely sericeous(appearing
glabrous to the naked eye) to glabrate to glabrous below [trabecula 0.1 (-0.2) mm
long, straight, sessile],margin with irregularly spaced sessileglands (0.2-0.5 mm
in diameter) and filiform glands (up to 1.6 mm long), with a pair of prominent but
sessileglands at the base borne halfway on the petiole, each gland l.I-2.6 mm in
d i a m e te r; p e ti ol es 1.5-10 cm l ong; sti pul es 0.3-0.9 mm l ong, 0.5-1.3 m m wide,
eglandular. Flowers ca. 15-40 per pseudoraceme,these borne in dichasia or comp o u n d d i c h a si a or smal l thyrses. P eduncl es4-12.5 mm l ong, pedi cel s 3- 9 m m
1993
C . A N D E R S O N :S T I G M A P H Y L L O NI N T H E A M A Z O N R E G I O N
401
long; peduncles0.T-2 times as long as the pedicels.Bracts 1-7.7 mm long,0.8-1.2
mm wide, triangular; bracteoles1-1.5 mm long, 0.7-7.3 mm wide, oblong to ovate,
eglandular or each bracteole with a pair of inconspicuous glands (each 0.2-0.4
mm in diameter). Sepals7.8-2.5 mm long and wide, glands I.2-\.9 mm long, 0.77.2 mm wide. All petals with the limbs orbicular, glabrous, suffused with red,
margin erose to denticulate-fimbriate,the teeth/fimbriae up to 0.2 mm long; anterior-lateral petals: claw (1 .5-) 2-2.2 mm long, limb 11-12 mm long and wide;
s:cl aw 1-1.5 (-1.7) mm l ong, l i mb 8-10 (-11) mm l ong and
p o s te ri o r-l a te ra petal
l
wide; posterior petal: claw (3.2-) 2.5-3.5 mm long, apex indented, limb 6-8.5 mm
long and wide. Stamens unequal, those opposite the posterior styles the largest,
anthers of those opposite the anterior-lateral sepalswith the connective enlarged
and the locules reduced or rarely with only 1 reduced locule or eloculate; anthers
usually all loculate, glabrous. Anterior style 2.8-3.3 mm long, shorter than the
posterior two, with scatteredhairs in the proximal 113-ll2: apex 1.5-1.9 mm long
sometimes including a spur ca. 0.2 mm long; folioles variable, the larger folioles
0.9-1.5 mm long, 0.7-7.5 mm wide, parabolic to broadly lunate to subrectangular,
sometimes much smaller, ca. 0.6 mm long, ca. 1 mm wide, broadly triangular.
Posterior styles3.14 mm long, with scatteredhairs in the proximal ll4-3l4,lyrate;
foliole 1.5-1.8mm long, 1.6-2 mm wide, suborbicular to subsquareto trapezoidal.
Dorsal wing of samara 3.4-4.2 cm long, I.2-Z cm wide, upper margin with a blunt
tooth; nut bearing a pair of rectangular to semicircular to lunate, entire to grossly
dentate lateral winglets, these 3.5-6.5 mm long, 1.2-2 mm wide, and often also
with a few spurs and crests up to 1.5 mm long and 1 mm wide; nut 4.5-6.7 mm
h i g h , 3 .5 -4 .3mm i n di ameter, areol e 3.3-3.5mm l ong,2' 8-3.5 mm w i de, concave,
carpophore up to 3 mm long. Embryo 6-6.7 mm long, ca.2 times as long as wide,
ovoid, outer cotyledon 9-10.2 mm long, ca.3.3 mm wide, the distal 1/3 folded over
th e i n n e r c o ty l e don,i nner cotyl edon 6.6-1.3 mm l ong,2.8-3.1mm w i de, fo lded at
th e d i s ta l 1 1 4 -1 13.
Phenology.Collected in flower and fruit throughout the year.
Distribution. In moist forest, along rivers, and also in secondary growth and
a l o n g ro a d s i d e s ;Guyana, S uri name, French Gui ana, and northeastern Br azil
(Amapri and eastern Par6), also recorded from Martinique and Trinidad; sea level
to 300 m.
RepRpsEN-rnr.rS
v ep r . c ' r v n N sM. a r t i n i q u e . M a r i g o t , S t e . M a r i e , D u s s 1 4 7 3 ( N Y ) . T e r r a s s o ns . n . i n
1 7 9 6 ( P - J U ) . T r i n i d a d . M a r a c a s . B r o a t l w a y 8 0 5 2( A , B M , M O , S ) ; N E o f P o i n t F o r t i n ' D a v i d s e 2 5 7 8
(F, GH, MO, NY).
Guyana. Northwest distr., Waini River, Marabo Creek, de la Cruz 1267 (NY, US); Pomeroon
dist.. Pomeroon Rivcr, rle la Cruz 3044 (CM, F, GH, MO. NY, US); Kamakusa, upper Mazaruni
R i v e r . c a . , 5 9 " 5 0 ' W ,f l e l a C r u z 4 t 4 9 ( C M , F , G H , N Y , U S ) ; m a r g i n s o f B e r b i c e R i v e r , S o f N e w
D a g e r a a d . c a . 0 6 " N , 5 7 ' 4 3 ' W , M a a s e t a l . 5 5 4 3( M I C H ) 1 E s s e q u i b o , M e y e r s . n . ( G O E T ) . S u r i n a m e .
F l u v . C o p p e n a m e , B o o n 1 0 4 8 ,1 1 0 4 ( U ) ; W i l h e l m i n a G e b e r g t e . L u c i e R i v e r , 0 3 ' 2 0 N , 5 6 ' 4 9 ' W ,
I r w i n e t a l . 5 5 1 0 8( F . K . M I C H , N Y , U , U S ) , I r w i n e t a l . 5 5 4 6 3 ( C , F , K , M I C H , M O , N Y , R B ' U ) ;
Paramaribo, Kramer & Hekking 2-lJB (U); ad ripas fluv. Marowijne, Lanjottw & Lindeman 2962
( N Y . U ) ; J a n d 6 k r e ek , b o v e n S u r i n a m e r i v i e r , 1 l l 2 u u r v a r e n b e n e d e n K a b e l , L i n d e m a n 4 4 5 5 ( l d O ,
U ) ; J o d e n s a v a n n e - M a p a n ek r e e k a r e a , S u r i n a m e R i v e r , L i n d e m a n 5 0 0 1 ( U ) : P e r i c a R i v e r , L i n d e m a n 5 1 4 0 ( C O L , M t C H ) ; S a r a m a c c aR i v e r , r e a r o f v i l l a g e J a c o b k o n d r e . M a g u i r e 2 3 8 4 7 ( F ' G H ' K '
M O , N Y , R B , U , U S ) ; f l u v . G o n i n i , V e r s t e g4 7 ( U ) . F r e n c h G u i a n a . H a u t . R i v . M a n a , a m o n t d c
S a n r A n a n a s , C r e m e r s T 5 3 l ( C A Y , M I C H ) I S d e S t . J e a n d u M a r o n i , C r e m e r s 7 6 7 2( C A Y , M I C H ) ;
rourc de St. Laurent ii Paul Isnard entre les PK 10 ct 40, Cremers 7979 (CAY, MICH); fleuve
I-ampoc, ii 4 km cn amont de son confluent avec I'Ouaqui, Granville 84834 (MICH, P); RN2, d
p r o x i m i t 6 d u p o n t s u r I ' O r a p u . G r a n v i l l e 5 0 3 6 ( C . A Y , M I C H ) ; C r i q u e C a b a r c t - B a s s i nd e I ' O y a p o c k ,
e n t r e l ' e m b o u c h u r ee t l a c r i q u e M d r i g n a n , 0 3 ' 5 5 ' N , 5 1 " 4 8 ' W . G r a n v i l l e 1 0 2 3 5( M I C H ) ; T r o i s S a u t s .
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VOLUME 19
Akattis Alasuka. Ha.raire 566 (CAy); Comt6, entre Rodre Fondd er Belizon, Okleman
tl49 (MICH):
Haut Oyapock, environ 2500 avant I'embouchure de la rividre Eurupoucigne.
oldeman B32Bg
(MICH); entre Cabassou et Degrad des Cannes, ile cle Cayenne, pr|vost 1257 (CAy,
MICH);
rividre Inini' affluent du Moyen-Maroni (Lawa), en amont de Maripasoula, Sas/re
et at.39g6 (CAy,
M I C H , P ) ; S i n n a m a r y ,r o u t e d e S t e . E l i e , K m 1 7 , s a s r r ee t a l . 4 t 9 9 ( M I C H , p ) ; f l e u v c
Approuague,
p r d s d e R 6 g i n a , S u s t r e1 B I 3 ( C A Y , M I C H ) ; m o n t a g n e s d e K a w , a u b e r g e d e B r o u s s e
des Cascades,
0 4 " 3 - 5 ' N ,5 2 1 7 ' w , W e i t z m a n 2 7 2 ( M I C H ) . B r a z i l . P n n A : O b i d o s , R i o P a r u c l e O e s t e ,
Cavalcante B0t
(MG); Gurupii, Rio Amazon. Killip & Smith 30594 (Ny); Jari, cstrada do Munguba,
N. T. Sitva
2 1 - l B ( l A N ' M I C H ) . - A n a n p A : R i o A m a p a r i . S e r r a d o N a v i o , C o w a n 3 8 2 0 6( M I C H .
Ny); Rio Jari,
0 . - 5 - 3 k m S o f S a n t o A n t o n i o d a C a c h o e i r a , 0 0 ' - 5 - 5 ' S5, 2 ' . 5 - 5 ' W ,E g t e r & I r w i n 4 6 0 6 6( M I C H .
Ny.
UB); confluence of Rio Iane with Rio Oiapoque,02'53 N. 52"22'w, Egler & pires1777t (MICH,
NY); Mpio. Mazagdo, BR-156, Bl km wSW of Macapd, ca. 11 km SW of Rio preto.00.0g'S.
- 5 1 ' 4 8 ' w . M o r i & C a r d o s o t 7 4 , 1 2( M I C H ) ; M p i o . C a l g o e n e . B R - 1 5 6 , 5 3 - 7 2 k m W S W
of Calqocne.
c a . 0 2 " 3 3 - 3 u ' N , 5 1 ' 1 6 ' WR
, a b e l oe t a t . 2 9 6 4 ( M I C H ) ; R i o A r a g u a r i , b e r w e e n0 1 . 0 2 N ,
- 5 1 . 4 5 ' Wa n d
0 0 ' 5 7 ' N . - 5 1 " 2 9w , P i r e s e t a l . 5 0 9 0 7 ( I A N , M G , M I C H , N y ) ; b e t w e e n R i o s c u i u b i m
and Flecha.
0 1 ' 4 - 5 ' N ,- 5 0 ' 5 8 ' W .P i r e s & C a v a l c a n t e5 2 4 3 5 ( l A N , M G . M I C H , N y ) .
Stigmaphyllon convolvulifolum is distinguishedby its ovate to cordate leaves.
whose lower surfaces are glabrate to glabrous but often very sparselyand minutely
sericeousbelow (appearing glabrous to the naked eye); the sessilehairs are onl;
0 .1 (-0 .2 ) mm l ong. Onl y the stamensopposi te the anteri or-l ateralsepalshau!
enlarged connectives and reduced locules, and the styles are pubescent in the
basal f 14-314.The petals. especially the flag, are usually suffused with red. This
species is most likely to be confused with the widely distributed and extremely
variable S. sinuatum and in eastern Brazil (Parrl) with S. cardiophyltum (see thit
species); S. convolvulifolium and S. argenteum are not sympatric. Stigmaphytton
convolvulifolium and S. sinuatum are similar in many aspectsbut can always be
separated by the nature of the pubescenceon the abaxial leaf surface. In the
range of S. convolvulifolium, the common form of S. sinuatum has the leaves
densely silver-sericeousbelow. A variant in French Guiana and in Amazonian
Brazll has the leaves lessabundantly pubescentabaxially though never as sparsely
as in s. convolvulifolium; the hairs are (0.2-) 0.3-0.5 (-0.7) mm long.
The epithet "convolvulifolium" is often ascribedto Cavanilles,who published
Banisteria convolvulifolia in 1790; however, Cavanilles cited Linnaeus's earlier
Banisteria dichotoma f=5s;fmaphyllon dichotomum (L.) Griseb.] in synonymy.
Jussieu (1840) intended to make the combination in Stigmaphyllon, but because
Cavanilles'sname was superfluous, the name Stigmaphyllon convolvulifolium dates
from Jussieu's publication. The lectotype is a Martin collection, annotated bv
J u s s i e u ,th a t matchesJussi eu' sdescri pti on.
stigmaphyllon sinuatum (DC.) Adr. Juss., Ann. Sci. Nat. Bot., s6,r.2, l3: 2gg.
1840. Banisteria sinuata DC., Prodr. 1: 588. 1824.Stigmaphyllon hastatum
v a r. B si nuatum (D C .) N i ed., Ind. Lect. Lyc. B runsberg.p. aest. lg00: 24.
1900.Stigmaphyllon sagittatum var. B sinuatum (DC.) Nied., Pflanzenreich
IV . l a I Q):506. I928.-TvpE : FnE N cHGuraN n.P errottets.n. (hol o t ype:G D C !, mi crofi che:MIC H !, photos: F! GH ! MIC H ! N y!).
Fig. 3.
Banisteria heterophylla willdenow, Sp. pr. 2: 742. 1799, non stigmaphylb,
heterophyllum Hooker, 1843. Banisteria splenderzsDC., Prodr. 1: 588. 1824.
nom. superfl. Stigmaphyllonfulgens Adr. Juss.,Ann. Sci. Nat. Bot.. sdr.2.
13: 289- 1840, nom. superfl. Stigmaphyllon splenrlens Cuatr., Webbia 13:
5 3 1 . 1 9 58,nom. superfl .-TypE : vE N E zueLA". ad ori nocum,,, B redem ever
s .n . (h o l otype:B -W 8855,mi crofi che:MIC H !).
l 993
C. ANDERSON: STIGMAPHYLLON
IN THE AMAZON
REGION
403
martianumAdr.Juss.,Ann. Sci.Nat. Bot., s6r.2,13:.289.1840.Stigmaphyllon
Dec, Martiw s.n.(holoTypE:Bnnztl. Amazdnas:"in sylvisJapurensibus,"
type: M!, photos:F! GH! MICH! NY!).
Stigmaphytton richardianum Adr. Juss., Ann. Sci. Nat. Bot., s6r.2, 13:289.
1840.-TvpE: FRE,NcuGulnNn. Cayenne, July' Richards.n.(holotype:P!,
fra g me n t:P -JU !, photos: F! MIC H !).
Stigmaphyllon hypoleucum Mrquel, Linnaea 18: 51. 1844.-TvpE: SuRtNnrulE.
"ad fluv. Boven Cottica, in sylva," Oct 1842,Focke 683 (holotype:U!).
Stigmaphyllon purpureum Bentham, London J. Bot.7: 128. 1848.-TYpE:GuvnNn. Picarara,Robt. Schomburgk, lst coll.737 (holotype: K!, photo:MICH!;
i s o t y p e sG
: ! P! W!).
Stigmaphylton brachiatum'friana & Planchon, Fl. Novo-Gran. in Ann. Sci.
Soc. Nat. Bot., s6r. 4, 18: 316. 1842.-TvpE: ColoMBIA. Meta: Villavicenc i o ,4 5 0 m . Tri ana s.n. (hol otype: P !, photos: F! GH ! MIC H ! MO ! ; isot y p e s :C O L ! G ! ) .
Stigmaphyllon monancistrum Niedenzu, Ind. Lect. Lyc. Brunsberg. p. hiem.
1899-1900:9. 1899.-Typp: "Colombia" [VeNEzuEr-n].Aragua: Maracay,
ad rivulis in crepidis, Moritz 779 (lectotype, here designated:LE!; fragme n t o f B dupl i cate:N Y !).
Vine to 30 m. Laminas 6-2I cm long, 4.510 cm wide, triangular to ovate to
cordate to elliptical to broadly so to orbicular to oblate to reniform, apex mucronate
to emarginate-mucronateto short-acuminate,base acute to truncate to cordate to
deeply auriculate, very sparselysericeousto glabrous above, sparselysericeousto
deniely silver-sericeousbelow [trabecula (0.2-) 0.3-0.5 (-0.7) mm long, straight,
mostly sessilebut sometimes with a tiny stalk up to 0.1 mm long, especially if
vesture is very dense],margin grossly and shallowly crenate to subentire and with
irregularly spaced sessile glands (0.4-1.5 mm in diameter) at the sinuses and
sometimesalso with filiform glands (up to 1.5 mm long), with a pair of prominent
but sessileglands at the apex of the petiole, each gland 1-3.5 mm in diameter;
p e ti o l e s 1 .8 -1 3 c m l ong; sti pul es0.5-1.3 mm l ong,0.5-1.5 mm w i de, egl a ndular .
Flowers ca. 15-35 (-40) per pseudoraceme,these sometimesinterrupted, borne in
compound dichasia or small thyrses. Peduncles 1.5-11 mm long, pedicels 3-9.5
mm long; peduncles 0.4-1.5 times as long as the pedicels. Bracts 0.7-2 mm long,
0 .5 -1 .4 mm w i d e , tri angul ar or narrow l y so; bracteol es0.9-1.6 mm l ong,0. 1- 1. 3
mm wide, triangular to parabolic to ovate, eglandular or more commonly each
bracteole with a pair of inconspicuousglands (each 0.1-0.5 mm in diameter) or
sometimesonly with a glandular area in the basal ll3-112. Sepals7.5-2.5 (-3) mm
long, 1.5_2.5(-3) mm wide, glands (1-) I.2-2.2 mm lon$, 0.8-1.3 mm wide. All
petals glabrous or sometimes pubescent abaxially, yellow or suffused with red,
margin erose to erose-denticulateto denticulate, teeth up to 0.3 mm long, lateral
petals with the limbs orbicular; anterior-lateral petals: claw (1-) 1.5-2.5 mm long,
limb (8-) 10-15 mm long and wide, posterior-lateral petals: claw (0.5-) 0.8-2 mm
long, limb (7-) g-13 mm long and wide, posterior petal: claw 2.6-3.5 (-4) mm
long, apex indented, limb (5-) 7-11 mm long and wide, broadly elliptical to broadly
obovate to orbicular. Stamens unequal, those opposite the anterior style and/or
th e p o s te ri o r s t yl es the l argest, anthers of those opposi te the anteri or - lat er al
sepals with the connective enlarged and the locules reduced or sometimes with
only 1 locule or rarely eloculate; anthers glabrous.Anterior style (2.G) 3.21(-4.8)
404
VOLUME 19
-1
.*r_
"
FIG' 3. Stigmaphyllon sinuatunr. a. Fruiting branch (x0.5). b. Flower (x2.-5).
c. Flower with
p e t a l s r e m o v e d ( x - 5 ) ;a n t e r i o r s e p a l a t r i g h t . d . P o r t i o n o f a n d r o e c i u m ( x - 5 ) ;
stamen at left opposite
posterior petal. e. Flower with petals and androecium removed (x5); anterior
style in center. f. Fruit
w i t h t w o s a m a r a s( x 0 . 7 5 ) . ( B a s e d o n W u r d . a c k3 4 4 3 4 . )
1993
IN THE AMAZON
REGION
405
mm long, shorter than or subequal to the posterior two. glabrous or commonly
w i th a ro w o f h a i rs adaxi al l yi n the proxi mal 115-213:
apex (1.1-) 1.4-2.6mm long,
often including a spur 0.1-0.5 mm long, each foliole 0.1-1.9mm long, 0.G1 .9 (-2.1)
mm wide, triangular to parabolic to subrectangular or folioles absent and the
a p e x e x te n d e d i n to a cl aw 0.7-1.3 mm l ong. P osteri or styl es (2.8-) 3-4.5 ( - 5. 2)
mm long, glabrous or commonly with a row of hairs adaxially in the proximal 1/4T 1 2 ,l y ra tefo
; l i o l e 1-2.2 mm l ong, (0.8-) L2-2.3 mm w i de, subsquareto subo r bicular or sometimesparabolic or rectangular.Dorsal wing of samara 3.5-5.5 cm long,
1 -1 .8 c m w i d e , upper margi n w i th a bl unt tooth; nut smooth or beari ng 1- 3
s u b c n ti re to c o a rsel ytoothed to l aceratel ateral w i ngl ets,these up to 6 mm long
a n d 3 .5 m m w i d e . and/or spurs and crestsup to ca.2 mm l ong and w i de; nut 4. 5- 7
m m h i g h ,2 .8 * 4 .4 mm i n di ameter, areol e 2.3-4 mm l ong,2.5-4.5 mm w i de, conc a v e , c a rp o p h o re up to 5 mm l ong. E mbryo 5.1-6.8 mm l ong, ca. 1.5-2 ti mes as
l o n g a s w i d e , o v o i d, outer cotyl edon (4.1-) 5.4-8.5 mm l ong,2.5* 4.2 mm wide,
s tra i g h t o r th c d i stal l l 5-315 fol ded over the i nner cotyl edon,i nner cotyl edon 46 .3 mm l o n g ,2 .2 -4 mm w i de, strai ghtor fol ded at the di stal l l 4-213.
Phenology.Collected in flower and fruit throughout the year.
D i s tri b u ti o n . C ommon i n the l ow l andsof C ol ombi a, V enezuel a,the Gu ianas,
northern Brazil, Ecuador, northern Peru, and Amazonian Bolivia; in primary and
s e c o n d a ryfo re s t, especi al l yw et forest but al so i n w hi te sand vegetati on,along
ri v e rs .a t ro a d s i d es.i n thi ckets:sea l evel to 1000m.
Rr,pnr,sr.NlR'lrvS
n p r . r ' r v r - . x(sa n t e r i o r s t y l e f o l i o l a t e ) . C o l o m b i a : A v n z o r n s : c a . 6 k m W o f
Leticia at Santa Isabclla,04'10'S,69"58'W. Gillett & Dickenson 16521(COL, MO); Araracuara,
n e a r R i o C a q u e t i { .M a g u i r e e t o 1 . 4 4 1 4 2( M I C H . N Y ) l L e t i c i a . c a . 1 k m N E o f t o w n . P l o w m a n e t a l .
: ept. Rio
2 2 9 1 ( E C O N . F . G H . N Y ) ; L e t i c i a , S c h u l t e s8 2 2 2 ( C O L , G H ) V e n e z u e l a . A H r n z - o x n s D
Negro, along Rio Marawinuma, vicinity of Ccrro de la Neblina base camp, ca. 00'50 N, 66"09'W,
L i e s n e r 1 5 9 6 6( M I C H ) . N e e - 1 0 8 8 0( M I C H ) : T a m a t a m a , L l . W i l l i a n t s 1 5 2 1 7( F , N Y , U S ) . W u r d a c k &
Adtlerlev 43641 (F, MICH, NY. U).-AxzoAn,<;ut: along Rio Querecual. SW of Bergantin. Steyerm a r k 6 1 5 0 6 ( F . N Y . V E N ) . - A n A ( ; u A : R a n c h o G r a n d c , B a d i l l o 1 7 9 - l( M Y ) . - B , q n r r a s : A l l a m i r a .
C L r r r t t nM - 6 1 9 ( N Y ) . - B o r . i v R n : N E d e C a n a i m a . S d e C e r r o V e n a d o . c a . 0 6 ' 1 6 ' N , 6 2 ' 4 6 ' W . A g o s t i n i 2 9 9 ( U S ) ; C i u d a d B o l f v a r , M a q u a n t a R i v e r , c a . 0 8 ' 1 0 ' N . B a i l e l -t & B a i l e l t l 4 l l ( 4 . N Y ) : C a l z e t a
en la Botclla, Rfo Cuyuni. Bernardi 6497 (MER, NY); a 48 km NE del caserio Los Rosos, este
r i l t i m o a 1 7 k m d e U p a t a , s o b r e l a c a r r e t e r a n u e v a U p a t a - S a n F 6 l i x . B l a n c : o4 - 1 4( M E R . M O , N Y ,
US); dist. Ccdcflo. 6 km from Maniapure toward Caicara, 06"55'N, 66"30'W. Boont & Grilb 6487
( M I C H ) ; M p i o . R a f l L c 6 n , R f o P a r a g u a , 0 4 o l B ' N , 6 2 " 0 - 5 ' WD, e l g u d o 1 0 1 ( M I C H ) ; K m 1 0 5 - 1 1 2d e
la carretera El Dorado-Sta. Elena, Morillo et al. 2932 (MICH); Mpio. Piar, camino desde El Plomo a
S t a . B a r b a r a , 0 6 ' 4 . 53 0 " N . 6 2 ' 4 f J ' W , P i c 6 n N a v a l 5 B B ( M I C H ) ; U r u y d n , A u y a n t e p u i . S c h n e e1 4 3 7
'l'umcremo,
(MY); alrcdcdoresdc
c a m i n o T u m e r c m o - B o c h i n c h e ,c n t r e P u e s t o ( G N ) C o r u m o y
C a f l o M a t u c o d c l R i o N c g r o , S t e r g i o se t a l . 3 5 6 - 1 ( M I C H ) ; S i e r r a I m a t a c a . a l o n g R f o R c f o r m a . 1 k m
a b o v e j u n c t i o n w i t h R f o T o r o , S t e y e r m u r k 8 7 9 1 7 ( M I C H , N Y . L l . U S ) l a c e r c a n d o s ea l a s c a b e c e r a s
del Rfo Nichare (afflucnte dcl Rio Caura) en la direcci5n de la Sierra Maignalida y Sierra Cervat a n a , a r r i b a d e l a d e s e m b o c a d u r ac o n e l R i o C i c u t a ( I c u t a ) , 0 6 " 1 - 5N , 6 5 ' 0 5 W , S t e y e r m a r k & G i b s o n
9 5 7 3 6 ( N Y , U S ) l L a P r i s i 6 n , M e d i o C a u r a , L l . W i l l i a m s l l 5 3 9 a ( F ) ; E l P a l m a r . L l . W i l l i a m . s1 2 9 1 9
(A, F, *, S)' along Fundaci6n rd, Wurdack 34431 (NY).-DElra
AvRc'uRO:near border (=Rfo
Grande o Toro) between Estado Bolfvar and Terr. Delta Amacuro, ca.08'04'N,6l'44'W, Breteler
-1757(MER, VEN, NY, U, US)l Depto. Tucupita,ca. 13 km by rd ESE of the town of SierraImata,
08'32 N. 62"23'W, Davidse & Gonzdlez 16623(MIC.H, MO)ldownstream from San Victor, Rfo Amacuro, Sierra Imataca, Steyermark 87299 (G, NY, P).-MrnaNtx: dist. Paez, Cerro Riber6n between Rfo
G u a p o a n d R i o C h i q u i t o , 4 4 . 5 k m ( s t r . l i n e ) S E o f C a u c a g u a ,1 0 ' 0 5 ' N , 6 6 " 0 1 ' W , D a v i d s e & G o n z d l e z
13604 (VEN)l between La Cortada and Turumo Bridge, Pittier 11184 (G, K,NY, US, VEN); Los
Mariches, Pittier 12991 (F, G, M, MO, NY, US, VEN); Cerros del Bachilla, between Quebradas
C o r o z a l a n d S a n t a C r u z , S o f S a n t a C r u z , 1 0 k m b y a i r W o f C f p i r a , S t e l t e r n t a r k& D a v i d s e 1 1 6 4 4 3
(MICH, MO).-MoNAciAs: ca. B km ESE of Jusepfn. Pursell et al. 9098 (NY, US, VEN).-Sucnr:
Rio Tatracual, 25 km outside Cumanii on Cumanacoa rd, Sobel & Strudwick 2274 (MICH). Guyana.
406
VOLUME 19
Near Mazaruni Forest Station, Archer 2463 (GH, K, US); Pomeroon Dist., Waramuri Mission,
Moruka Rivcr, de la Cruz 2578 (GH, NY, US); Waini River, NW Dist., 08"20 N. 59"40'W, de la
Cruz 3619 (F, GH, MO, NY, US)l upper Demerara-Berbice region. ca. 27 km from Ituni along
Ituni-Kwakwanni rd, 05'22 N, 58'07'W, Gillespie 2991 (MICH); Potaro-Siparuni region. trail from
K a t o t o P a r a m a k a t o i , 0 4 ' 4 1 ' N , , 5 9 ' 5 0 ' W ,H a h n 5 6 2 2 ( M [ C . H ) ; A k y m a . D c m e r a r a R i v e r a b o v e W i s m a r , { ) 5 ' 0 9 ' N , H i t c h c o c k 1 7 4 1 4( G H , N Y . U S , S ) ; K a n u k u M t s , R u p u n u n i R i v e r , n e a r " t h e f a r m " o f
t h e C a p t a i n o f S a n d c r e e k , 0 3 " 0 7 ' N , 5 9 ' 2 6 ' W , J a n s e n - J a c o b se t a l . 2 0 6 ( M I C H ) : G u n n ' s . E s s e q u i b o
R i v e r , 0 1 " 3 9 N , 5 6 " 3 8 ' W , J a n s e n - J a c o b se t a l . 1 5 1 6 ( M I C H ) ; P o t a r o - S i p a r u n i r e g i o n , C h e n a p o u , 5 0
k m u p s t r e a mf r o m K a i e t e u r F a l l s ,0 5 ' 0 0 ' N , 5 9 ' 3 4 ' W , K v i s tJ 2 5 ( M I C H ) : C u y u n i - M a z a r u n ir e g i o n ,
NW of Conoch Tipu,05'48'N,61o03'W, McDowell2629 (MICH); Cuyuni River. by portagerd near
lower Camaria Landing, Sandwith 664 (K, NY, U); Essequibo River, near mouth of Orono Creek,
c a . 0 1 ' 3 - 5 ' N ,A . C . S m i t h 2 B 2 l ( A , F , G , K , M O , N Y , P , S , U ) ; W e x t r e m i t y o f K a n u k u M t s , d r a i n a g e
of Takutu River. A. C. Smith 3166 (F, G, K, MO, NY, P, S, U, W); ca. 5 km SW of Mabura Hill
t o w a r d s E s s e q u i b o R i v e r , 0 5 ' 1 9 ' N , 5 8 " 3 8 ' W , S t o f f e r s e t a l . - 1 5( M I C H ) ; M a r u d i M t s , M a z o a H i l l ,
n e a r N o r M a n M i n e s c a m p , 0 2 ' 1 5 ' N , 5 9 ' 1 0 ' W , S t o f f e r se t a \ . 2 0 7 ( M I C H ) . S u r i n a m e . S c o t e l w e g ,
Archer 265B (US); Zandrlj. Archer 2761 (US): Republieck, van Doesburg,Jr., 70 (U): dist. Saramacca.
Experimental Farm Coebiti, Everaarts519 (MICH); in montibus Bakhuis inter flum. Kabalebo et
Coppename sinistrum, around Kabalebo airstrip, Florschiitz & Maas 2502 (F, U); dist. Nickcrie.
area of Kabalebo Dam project. ca. 22 km SW of Avanavero damsite, Heyde & Lindeman l0-l
(MICH, U); without locality, Ilostmann f029 (BM, G, GH, P, U, W); Wilhelmina Gebergte. Zuid
R i v e r , 4 5 k m a b o v e c o n f l u e n c ew i t h L u c i e R i v e r , 0 3 ' 1 0 - 2 0 ' N . 5 6 ' 2 9 - 4 9 ' W . l r w i n e t a l . 5 7 6 3 1 ( K ,
M G , M I C H . N Y , S , U , U S ) ; S u r i n a m e R i v e r n e a r K a b e l s t a t i o n ,L a n j o u w I I I 8 5 ( U ) ; N i c k e r i e D i s t . .
area of Kabalebo Dam project, 04-05'N, 57"30 -58"W, Lindeman et al. a7 (U); Lely Mts. SW
plateaus,Lindeman et al. 266 (NY, U); Waijombo River, Linder B9 (GH, NY); vicinity of Sectie O,
K M 6 8 , M a g u i r e & S t a h e l2 5 0 0 0 ( B R , F , G , G H , K , M O , N Y , P . R B , U ) ; B r o w n s b e r g N a t u r e P a r k ,
90 km S of Paramaribo, Mazaroni plateau, trail to Witti-creek. Mori & Bolten 8397 (MICH. NY);
surroundings of Blakawatra, camp 8, 60 km SE of Paramaribo, den Outer 872 (U): fluv. Saramacca
i n f . p r o p e M i n d r i n e t t i , P u l l e 3 a ( U ) ; J o d e n s a v a n n e - M a p a n e k r e e ka r e a ( S u r i n a m R . ) , S c h u l z 7 - 1 1 - l
(AAU, COL. MICH, NY, [J, US). French Guiana. Sinnamary, piste de Ste. Elic, Km 15, Billiet &
J a d i n 1 l 0 l ( B M , B R , C A Y , N Y ) ; p i s t e S a i n t L a u r e n t v e r s P a u l I s n a r d ,e n t r e K m 3 0 e t 4 0 , B i l l i e t &
Jadin 1577 (tlAY, MICH); piste forestidre allant de la route N2 vers Nancibo, Billiet & Jadin 1845
(CAY); le long de la piste de la route de Cayenne ii R6gina, ca. 30 km dc R6gina, Cremers 5991
( M I C H ) ; B o u r g d ' A p a t o u , B a s s i n d u M a r o n i , 0 5 ' 0 9 ' N , 5 4 ' 2 0 ' W . F l e u r y - l - 1 4( M I C H ) ; H a u t O y a p o c k .
T r o i s S a u t s ,G a r n i e r 1 0 - l ( C A Y ) ; H a u t O y a p o c k , 6 2 k m e n v . e n a m o n t d e S a u t B o k o , G r a n v i l l e
2465 (NY); Haute Camopi, Mont Belv6ddre, Granville 7010 (CAY, MICH); rochc plate Roche
K o o u t o n - B a s s i nH a u t - M a r o u n i n i , I k m W d e l a R o c h e , 0 2 ' 5 3 ' N , 5 4 " 0 4 ' W , G r a n v i l l e 9 5 1 6 ( M I C H ) ;
S t . G e o r g e s d e l ' O y a p o c k , p i s t e d e M a r i p a , G r e n a n d 2 1 3 6 ( C A Y . M I C H ) ; i l c d e C a y e n n e .M o n t
B a c h r e l , 0 4 ' 5 5 N , 5 2 ' 1 9 ' W , H o f f 5 2 2 a ( M I C H , P ) ; v i l l a g e d e Z i d o c k v i l l e . T r o i s S a u t s ,J a c q u e m i n
I 6 1 0 ( C A Y . M I C H ) ; S a i . i l , 0 3 ' 3 7 ' N , 5 3 ' 1 2W . M a r s h a l l & R o m b o l d 1 6 9 ( C A Y , M I C H ) ; M a r i p a s o u la, Oldeman 1647 (MICH)i rividre Tonigrande, prds de port lnint, Oldeman 1656 (MICH. P)l Saiil,
Oldeman I9B2 (CAY, MICH, P); fleuve Approuague, rivibre Arataye, Sauts Parare, Ponc1,22l
(CAY); Acarouani, Sagot 91 (BM, G. P, S, W); St. Laurent region, ca. 5 km from Rte D9 at
C h a r v e i n , 0 5 " - 5 1N , 5 3 ' 5 1 ' W , S k o g & F e u i l l e t 7 4 8 1 ( C A Y , N Y , P ) ; m a i n r d t h r o u g h M o n t a g n e d e
K a w , 0 4 " 3 5 N , . 5 2 ' l - 5W , W e i t z m a n - 1 0 8( M I C H ) . B r a z i l . A c ' n E :C r u z c i r o d o S u l , P r o j e c t o R A D A M /
BRASIL, acroporto, Monteiro & Damido 209 (INPA, MG). Ramos 116 (INPA), Ramos & Mota 208
( I N P A ) ; n e a r S t a . L u c i a , K m 4 0 o n T r a n s a m a z 6 n i c aH w y , E o f C r u z e i r o d o S u I , 0 7 " 0 8 ' N , 1 2 " 3 3 ' W ,
Pruski 3498 (MICH).-Aunen:2-10
km N de Ferreira Gomes, BR-156, Austin 7257 (MICH); Rio
Amapari, rd to Porto Terezinha, Cowan 38166 (K, NY, RB), Cowan -184738(NY); Rio Amapd,
S e r r a d o N a v i o , l o w e r s l o p e s o f F r i t z A k e r m a n O r e B o d y , C o w a n & M a g u i r e 3 8 0 8 6( C O L . G , G H ,
M I C H , M O , N Y , P , S , U , W ) : R i o O i a p o q u e , 6 k m S E o f C l e v e l a n d i a , 0 3 ' 4 8 ' N ,5 1 ' 5 3 ' W , I r w i n e t a l .
1 7 3 8 2( l A N , M I C H , N Y ) ; M p i o . O i a p o q u e ,B R - 1 5 6 , 6 0 k m S S E o f O i a p o q u e , 0 3 o 1 8 ' N , 5 1 ' 3 9 ' W ,
Mori & Cardoso 17134 (MICH), Macapii, Igarapd do Lago, Rabelo & Non 771 (MG): Matapi,
R i b e i r o 1 . 5 9 7( I N P A , M I C H , M O , N Y , R B ) . - A v n z o N A S : M p i o . T c f d . R i o S o l i m d e s ,v i l a N o g u e i r a ,
Amaral et al. 95 (INPA, NY); Manaus-P6rto Velho hwy, Km 124, Campbell et al. P20920 (INPA,
MICH, MO, NY, S, U), Manaus, lgarapd do Parque 10. Chagas INPA 3679 (INPA, SP); Rio
U a t u m a . M p i o . I t a p i r a n g a ,C i d e t a l . 5 9 2 ( I N P A , M I C H ) ; B R - 1 7 2 . M a n a u s - C a r a c a r a iK, m 9 7 , C i d e t
al. 945 (INPA, MICH); Mpio. Marad, Rio Japurii, affluente do Rio Solim6es, Cid & Linta 3134
(INPA, MG. MICH); BR-307, Mpio. Cruzeiro do Sul,7-tl"S,72-73"W, Cid Ferreira et ul.52lB
(lNPA, MICH); Mpio. S5o Paulo de Olivenqa, 6 km S of town center, 03'30'S. 68"57 W. Daly et al.
4442 (MICH)l Barcelos, Duarte 7160 (lNPA, RB, SP)l Manaus, estrada do Mindf, Ducke 856 (F,
IAN. MG, MO, NY, R. RB, US); Mpio. Mau6s,ca.20 km E of Mau6s,Antarctica Guarand Planta-
I 993
407
tion, Hill et al. l-i|52 (JBSD, MICH, TEX): Manaus. Rua Duque de Caxias, Maas & Maas 362
(lNPA, U); caatinga do Porto Camanaus, Madison et al. PFE 414 (INPA); basin of Rio Demcni,
v i c i n i t y o f T o t o t o b i , P r a n c e e t a l . 1 0 3 5 5( l N P A , M G , M I C H , N Y , U ) ; M a n a u s . I N P A . e s t r a d a d o
A f e i x a . K m 3 . P r a n c e & R a m o s 2 0 9 2 2 ( I N P A , M I C H , M O , N Y , L J ) ;v i c . o f P i c o R o n d o n , P e r i m e n t a l
N o r r c . K m 2 1 1 - 2 2 0 . 0 1 " 3 2N , 6 2 ' 4 t l ' W , P r a n c e e t a l . 2 B B 2 1( M I C H ) , R o d r i g u e se t a l . 1 0 5 8 4( M I C H ) ;
Santo Antonio do lqa. p. Vila Militar, M. Silva2ll2 (MG); Fonte Boa, M. Silva2l83 (MG); Barra
[ = M a n a u s l , S p r u c e l 8 B 0 ( G . G H , L E , M , M G ) l M p i o . H u m a i t d , e s t r a d a H u m a i t i i - L i i b r e a , K m - 5 9 ,a 3
k m a o N , T e i r e i r a e t a l . 9 8 0 ( l N P A , M I C H ) ; M p i o . H u m a i t d , e s t r a d a H u m a i t i i - J a c a r e c a n g a .K m 1 5 0 .
a 63 km ao S, Teixeira et al. 1340 (INPA, MICH).-MannNuAo:
Mpio. Monq5o, basin of the Rio
Turiaqu, Kiiapor Indian Reserve, Balie 886 (NY); ca. 50 km from Santa Luzia on Hwy to AqailAnd i a , 0 4 " 0 5 ' S .4 5 ' - 5 7W , D a l v e t a l . D 7 - 1 6( M I C H ) ; R i o A l t o T u r i a q u , B a r r a n q u i n h a . 0 3 ' 0 0 ' S , 4 - 5 ' 4 _ 5 ' W ,
Jangoux & Bahia l6l (MG. NY); margen do Rio Cururupu, Lisboa 47 (P.B, SP)l Km 375-380 da
r o d o v i a B e l d m - B r a s i l i a . O l i v e i r a1 0 7 2 ( f A N , U B ) . - P n n A : B e l t e r r a . B l a c k 4 T - 1 6 6 0( l A N ) : A l t a m i r a .
Km 74 da estrada Transamazdnica ltaituba, Cavalcante& M. Silva 2780 (MG); Mpio. Oriximinii.
R i o T r o m b e t a s , L a g o d e M a t e n s , 1 9 k m S d e P 6 r t o T r o m b e t a s, C i d e t a l . 1 7 9 3 ( I N P A , M G , M I C H ) i
Jacaracd Island. tla Costa Ia9 $), ca.70 km from Tucuruf, ca. 04'll'S. 49"04'W, Daly et al. 1435
(INPA, MICH); boca do Lago de Faro, Ducke 88677 (MG); 4-5 km W of Sao Francisco do Pard
t o w a r d C a s t a n h a l . G e n t r l t I - 1 1 6 6( I N P A , M I C H , M O , N Y ) ; d i s t . A c a r i i , T h o m 6 A s s r i , P a u V e r m e l ho, Mexia 5926 (A. F. G. GB, GH, MICH. MO, NY. S. U. WIS); Bel6m, horta do IAN. Mrrrga Pires
& B l a c k a 0 - 1( G H ) : M p i o . A l m e i r i m , M o n t e D o u r a d o , e s t r a d a M T D , W e m d i r e g d o d m i n a d e
b a u x i t a , M u r s a P i r e s e t a \ . 6 2 0 ( M I C H ) ; B R - 1 6 3 , K m 1 1 3 1 ,C i u a b i i - S a n t a r 6 mh i g h w a y , v i c i n i t y o f
Igarap6 Natal, Prance P25427 (MG, MICH); Bel6m, terreno da EMBRAPA, Ramos & Rosdrio I4
( M G ) ; O u r 6 m . R o d r i g u e s 1 0 2 1 ( . | l 4 G ) ;T u c u r u f , m a r g e m d i r e i t a d o R i o T o c a n t i n s , R o s d r i o 9 J ( M G .
N Y ) : M p i o . A l m e i r i m . M o n t e D o u r a d o , S a n t o s a - 1 7( N Y ) ; S a n t a r d m , K m 7 0 d a e s t r a d a d o P a l h d o ,
arredorcs do Acampamento do Igarapd Guaranii, M. Silva & R. Souza 2522 (MICH, NY. U). M.
Silva & R. Souza 2522 (CAS, CM, MG, MICH,Ny)'
7-11 km NW of AMZA camp 3-Alfa on rd to
camp 4-Alia,05'47'S,,50'34'W, Sperling 6057 (}/ICH); Santar6m, Spruce 767 (G, GH, GOET, M,
NY. W)l Macau airstrip. I l/2 hrs upstream from Lageira airstrip,05'5.5'S, -54'26'W.Strudwick &
Sobel 3474 {MICH); Taperinha bei Santardm, Zernv 589 (W).-RoruooNrn: Pdrto Velho-Cuiabd
h w y . 2 - 5k m S o f N o v a V i d a , F o r e r o & W r i g l e y 7 0 B 4 ( I N P A , M G , M I C H , N Y ) ; b a s i n o f t h e R i o
Madeira. cerrado bctween Jaciparanii and Rio Madeira, Prance et al. 5180 (INPA, MG, MICH,
NY); Km 166-169,Madeira-Mamor6 railroad near Mutumparand, Prance et al. 5690 (INPA, MG,
M I C H , N Y ) l f o o t h i l l s o f S c r r a d o s P a c a : i sN o v o s , 1 2 k m N N E o f G u a j a r i i - M i r i m , P r a n c e e t a l . 6 6 5 8
(F. INPA. MICH. NY); P6rto Velho-Cuiabiihwy, vicinity of Santa Barbara, 15 km E of Km 117,
R a m o s & P r a n c e6 9 0 5 ( I N P A , M I C H , N Y ) : P 6 r t o V e l h o . R e p r e s aS a m u e l , 0 8 ' 5 5 ' S . 6 3 " 1 6 ' W , T h o m a s e t a \ . 4 9 4 9 , 5 0 J 4( M I C H ) . - R o n n r v a , : A r i t u m 5 r e g i o n .o n a n a z i m u t h o f 0 1 l o f r o m B o a V i s t a a t
a distanceof 210 km, Coradin & Cordeiro 943 (INPA, MICH, NY); SEMA Ecological Reserve,Ilha
de Maracii,03'21'N. 61"2J'W, Milliken M792 (MICH); Canto Galo, Rio Mucajaf, betwccn Pratinha
and Rio Apiari, Prance et al. 3964 (INPA, MG, MICH, NY, U): Serra Tepequem, Prance et al. 41-17
( I N P A , M G . M I C H , N Y . U ) ; v i c i n i t y o f U a i c i l a i r s t r i p , R i o U r a r i c o e i r a ,0 3 ' 3 3 ' N , 6 3 " 1 1 ' W , P r a n c ee t
a l . 1 0 9 0 9 ( I N P A , M G . M I C H . N Y . U ) ; M p i o . C a r a c a r a f , e s t r a d a M a n a u s - C a r a c a r a f ,K m 5 2 9 - 5 5 0 .
d o s S a n t o s & C o | h o 7 0 0 ( I N P A , M I C H ) ; e s t r a d a M a n a u s - C a r a c a r a i .B R - 1 7 4 , K m 3 2 9 , a r m y p o s t N
o f W a i m a r i - A t o a r i I n d i a n R e s e r v e ,S t e w a r d e t a l . 9 ( M I C H ) ; R i o S u r u m u , a n e i n e m B a c h c d e r S e r r a
d o M e l , U l e B l 8 5 ( M G ) B o l i v i a . B e r u r :V a c a D i e z , 3 k m E o f R i b e r a l t a o n r d t o G u a y a r a m e r i n ,
1 1 " 0 0 ' S . 6 6 ' 0 5 ' W . S o l o m o n 7 6 8 2 ( l d I C H ) . P e r u . H u A N u c r o :P r o v . L e o n c i o P r a d o , M o e n a . c e r c a a
-fereza,
Tingo Maria, Woytkowski ll87 (MICH); Santa
valle del Huallaga, Woytkowski 1232
( E C O N ) . - L < t n e r o : P r o v . C o r o n e l P o r t i l l o , P a m p a d e S a c r a m e n t o ,c e r c a P u c a l l p a , F e r r e y r a H . l l 8 4
(GH, MICH, US); Prov. Alto Amazonas, entre Yurimaguas y Chambira, Ferreyra H.4897 (MICH,
U S ) ; P r o v . C o r o n e l P o r t i l l o , o n R i o A g u a y t i a , 0 8 ' 5 0 ' S , 7 5 " 2 0 ' W , F o s b e r g 2 8 8 7 5( M O ) ; B a l s a p u e r t o .
Klug 3075 (A, BM, F, G, K. MO, NY, S, US); Prov. Ucayali, Canchahuayo (Rio Ucayali), 07"05'S,
7 5 ' 1 0 ' W , V d s q u e z e t a l . 6 9 7 0 ( M O ) ; M a r i s c a l C a s t i l l a , C a b a l l o c o c h a , 0 3 ' - 5 5 ' S ,7 0 ' 3 0 ' W , V d s q u e z &
Jaramillo 9Jl9 (MICH).-SnN ManriN: ca. 10 km NE of Tarapoto. Gentrv et al. 37915 (MICH); San
M a r t f n , - 5 - 1 - 5k m E o f S h a p a j a o n r d t o C h a z u t a , 0 6 " 3 6 ' S , 7 6 ' 1 0 ' W , K n a p p & M a l l e t 7 0 2 6 ( M I C H ) ;
Prov. Mariscal Caceres, Dtto. Uchizo, en la carretera a Rio Uchizo 2 km del caserio Nuevo Progreso,
Schunke V. 3219 (COL, F, G, NY, US); Prov. Mariscal Caceres,Dtto. Tocache Nuevo. Quebrada
Luis Siilas (5 km NE de Puerto Pizana), Schunke V. 6578 (GH, MO).
R e p n e s p N r R r t v pS
. p E c r r v r e N( as n t e r i o r s t y l e c l a w e d ) . C o l o m b i a . A n a a z o N n s :L e t i c i a , O c t 1 9 4 6 ,
B l a c k . r . r z .( l A N ) ; R f o l a g a r a - P a r a n d ( a f f l . R f o P u t u m a y o ) , L a C h o r r e r a , G a s c h e & D e s p l a t s 5 9 ( K ,
MICH).-ANrroetrA:
M p i o . C a u c a s i a , a l o n g r d t o N e c h f , 2 4 k m f r o m C a u c a s i a - P l a n e t aR i c a r d ,
0 8 ' 0 4 ' N , 7 5 " 0 - 5 ' W ,Z a r u c c h i e t a l . 4 9 0 3 ( M I C H ) . - C , q o u e r A : 6 k m S E o f M o r e l i a a l o n g r d t o R f o
Pescado (SW of Florencia), Davidse 5644 (COL, MICH); Rio Arteguaza, 9 km S of Florencia,
408
VOLUME 19
Plowman & Kennedy 2282 (F. GH, M, NY, P, S, US).-ME'rn: Villavicencio, Killip 34346 (COL, S);
Sierra de la Macarena, Rio Guapaya, Philipson et al.2124 (COL).-VRupes: rd from Mitu to Monf o r t , D a v i s 1 0 7 ( C O L , G H , M I C H ) . V e n e z u e l a . A u n z o N e s : a l r e d e d o r e sd e S a n J u a n d e M a n a p i a r e ,
0 - 5 ' 1 8 ' N , 6 6 ' 0 3 ' W ,A g o s t i n i 1 5 0 4 ( M I C H ) ; R i o O r i n o c o , I s l a d e l R a t 6 n , 0 5 ' 0 2 ' N , 6 7 ' 4 5 ' W , B r e t e l e r
4721 (F, K, MO, NY, US, WAG)l Depto. Atures, 26 km SE de Puerto Ayacucho por la carretera
P u e r t o A y a c u c h o - E l G a v i l i i n , 0 - 5 ' 3 2 ' N ,6 7 o 2 4 ' W , C u e l l o & F e r n d n d e z 5 0 8 ( M I C H ) : E l G a v i l i i n , 3 0
k m a l E d e P u e r t o A y a c u c h o , F e r n d n d e z2 q 5 0 ( M Y ) I R f o V e n t u a r i , L a C e i b a , b a j o d e l S a l t o T e n c u a ,
F . o l d a t s1 4 7 4 ( N Y , V E , N ) ; D e p t o . R f o N e g r o , c e r c a d e S h a b o n o Y a n o m a m i u b i c a d o a l a i z q u i e r d a
d e l R f o O r i n o c o y a l a d e r e c h a d e l a d e s e m b o c a d u r ad e l R i o M a v a c a , 0 2 ' 3 0 ' N , 6 5 ' 1 0 ' W , G u a n c h e z
656 (MICH); Cerro Camani, Maguire 31802 (NY)l ad flumina Casiquiari, Vasiva, et Pacimoni,
Spruce 3277 (BP.. G, K. NY, W); Depto. Atabapo, Rfo Cunucunuma, entre el Cerro Duida y
Pozo del Diablo,
H u a c h a m a c a r i , 0 3 " 4 0 ' N , 6 - 5 o 4 5 ' W ,S t e y e r m a r k e t a l . I 2 6 l 7 B ( M I C H ) . - A n n c u n s
caflada del Rio Yuare, Maracay, Baldillo 3782 (F, MY); Rancho Grande, Maracay, F.errari 745
( M Y ) , V o g e l s a n gE I 0 ( M Y ) . - B n R r N n s : T i c o p o r o F o r c s t R e s c r v e , 0 8 ' 1 5 ' N , 7 0 " 4 5 ' W , B r e t e l e r 3 6 7 8
( G , M E R . N Y , S , U , U S , V E N ) ; e n t r e K m 4 6 9 - 4 7 0 ,c a r r e t e r a B a r i n a s - S a n C r i s t 6 b a l , C d r d e n a s d e
C l u e v a r ae t a I . 2 5 6 5 ( M Y ) l 1 - 2 k m N E o f B u m b u m , c a . 6 8 k m S W o f B a r i n a s , G e n t r y e t a l . l l l 4 2
( M I C I H , M O . V E N ) . - B o r i v n n : P i c a C a i c a r ad e l O r i n o c o - S a nJ u a n d e M a n a p i a r e , R f o S u a p u r e . 2 0 2
km al S de Caicara. ca. 07'N, 67'W, Delascio & L6pez 2766 (VEN); Santa Marfa de Erebato, Rio
Erebato, 05'0-5'N, 64'40 W, Steyermark et al. 109817 (K, NY).-CnnABOBO: Dtto. Valencia, carretera Valencia-El Palotal-El Paito-Los Naranjos,Bunting 4597 (NY); Guataparo. Valencia, Saer
8 6 5 ( V E N ) . - D r s ' r ' n r r o F r , o n R R r -C: a r a c a s ,B r e d e n t e y e r2 0 6 ( W ) : D e p t o . V a r g a s , c a m i n o e n t r e O s m a
y Oritapo, Benltez de Rojas 5BB(MY).-FnlccrN: carretera Yaracal-Araurima, 8 km de la carretera
Y a r a c a l - ' f u c a c a s ,C a r d o z o e t a \ . 6 5 ( M I C H ) ; E l G u a n i i b a n o , 2 T k m S d e P u e r t o C u m a r e b o , F / o r a
F a l c 6 n . l . / J ( M I C H , M O , U ) ; D t t o . S i l v a , c a . 2 1 k m W o f T u c a c a s ,W i n g f i e l d 1 2 6 9 6( M I C H ) . - L n n n :
Dtto. Palavccino, carretcra cntre Manzanito y El Altar, Burandt, Jr., & Smith V0075 (MICH).M E R r o n :a b o v c d a m s i t e o n R i o C a p a r o , 3 l k m E S E o f S a n t a B a r b a r a , c a . 0 7 " 4 7 ' N , 7 1 " 2 8 ' W ,L i e s n e r
& G o n z d l e z .9 2 6 7 ( M I C H , V E N ) . - P o R l l r c i L i E S A : D t t o . O s p i n o , c a r r e t e r a L a A p a r i c i 6 n d e O s p i n o 'N.
Moroturo. 09'31
69'26 W, Aymard & Cuello 6577 (ll{IC.H); Dtto. Araure, carretera Hoja BlancaGuayabal-El Rechazo, Cuello & Cuello 59 (MICH).-TAcutnn: E of San Crist6bal and 40 km W of
Bocon6, L6pezSanta Barbara, rd W or NW of Abajales, Sobel & Strudwick 2108 (NY).-Tnulr-lo:
P a l a c i o s - 1 9 . 5( M E R . V E N ) . - Z u r - r n : D t t o . C o l c i n , h a c i e n d a E l R o s a r i o , 1 8 k m E d e l a c a r r e t e r a
Machiques-La Frfa. 12 km N de Rio Catatumbo, Bunting 6475 (MICH, VEN); Dtto. Col6n, entre
C a s i g u a E l C u b o y K m 8 d e l a v i a r u m b o a l P a l m i r a , B L t n t i n g7 3 2 8 ( M I C H ) ; D t t o . P c r ( j a , c n t r e K m
16 de la carretcra Machiques-La Fria y Calle Larga y San Jos6,Bunting l0Bl6 (MICH); Dtto. Perfja,
between Rio Yasa and Rio Tucuco along the Machiques and Los Angeles de Tucuco rd. 09'50- 5 6 ' N . 7 2 ' 4 0 - 4 4 W . D u v i d s e e t a l . 1 8 3 9 0 ( M I C H ) . E c u a d o r . M o n o u e - S n N r r A G O :T u n a n t z a , J i b a r o
s e t t l e m e n l ,n e a r M a c u m a , c a . 5 0 k m N E o f M a c a s , L u g o S . 3 7 l B ( G B , M I C H ) ; E l C e n t r o S h u a r
K a n k a i m ( C a n g a i m e ) ,R f o K a n k a i m ( C a n g a i m e ) ,0 2 ' 2 0 ' S , 7 7 " 4 1 ' W , S h i k i R B A E 2 1 9 ( N Y ) . - N n p o :
c a r r e t e r a H o l l f n - L o r e t o - C o c a , e n t r e A v i l a y R f o P o c u n o , 0 0 ' 3 9 ' S, 7 7 o 2 2 ' W, C e r 6 n e t a l . 2 8 7 9 ( M I C H ) ;
P a r q u c N a c i o n a l Y a s u n i . P o z o P e t r o l e r o " C o w i " d e C o n o c o , 0 0 " 5 - 5 ' S ,J 6 " 2 0 ' W , C o e l l o i 7 / ( M I C H ) ;
P u e r t o F r a n c i s c o d c O r e l l a n a ( C o c a ) , c a . 4 0 k m S E o f t o w n , A u c a o i l f i e l d , 0 0 " 4 2 ' S ,7 6 ' 5 2 ' W ,
B a l s l e v & M a d s e n 1 0 5 9 5( A A U , M B M , M O , N Y ) ; n e a r e n d o f t h e A u c a o i l f i e l d r d , 0 0 ' 4 4 ' 5 ,
7 6 " 5 4 ' W , B r a n t l b y g e & A s a n z . a- 1 0 1 0 9( A A U . M I C H ) ; r d C o c a ( P u e r t o F r a n c i s c o d e O r e l l a n a ) t o
C u r a r a y , c a . 4 0 k m S E o f C o c a , H a r l i n g e t a l . 1 4 7 5 7( G B , M I C H ) ; D u r e n o o n R i o A g u a r i c o , H a r l i n g
& A n t l e r s s o n1 6 6 0 5 ( G B , M I C H ) I R ( o A u y a b e n o , n e a r P u e r t o M o n t f f a r , 0 0 o 0 6 ' 5 , 7 6 o 0 1 ' W , H o l m Nielsenet ul.2l2lJl (AAU)l Rfo Aguarico, Monte Cristi,00'18'S,76'll7'W, Holm-Nielsenet al.
, eill et a\.6216
2 1 6 6 6 ( A A U ) l E s t a c i 6 nE x p e r i m e n t a ld e I N I A P , S a n C a r l o s , 6 k m S E d e l o s S a c h a s N
( M I C H ) ; a 2 k m d e J a t u n S a c h a .e n v f a a T e n a , 5 k m d e M i s a h u a l l f ,0 1 " 0 8 ' S ,7 7 " 3 0 ' W , P t t l a c i o s2 7 8 9
( M I C I H ) . - P n s r n z n : 3 l k m N o f P u y o o n r d t o T e n a . s i d e r d E o f C a j a b a m b a ,t ) 0 ' 1 5 ' S , 7 7 ' 5 0 ' W ,
B o o m & B e a r d s l e y8 4 . 1 6( M I C H ) ; C u r a r a y ( J e s r i sP i t i s h k a ) , H a r l i n g & A n d e r s s o n 1 7 4 5 0( G B , M I C H ) ;
R i o C h u l l a n a ,c a . 1 , 5k m N o f P u e r t o S a r a y a c u ,L u g o S . 4 l 8 4 ( G B . M I C H ) ; R f o C u r i a c u , c a . t Jk m W
o f P u e r t o S a r a y a c u ,L u g o S . 1 2 3 8 ( G B , M I C H ) ; v i c i n i t y o f E , l P o r v e n i r , c a . 5 k m W o f P u y o p u n g a ,
Lugo S. 4942 (GB. MICH): trail to Copataza,l0 km S of Sarayact. Lugo S. 5519 (GB, MICH);
c a r r e t e r a d e P c t r o - C a n a d a ,v f a A u c a , 1 1 5 k m S d e C o c a . 6 k m S d c l R f o T i g u i n o . Z a k & R u b i o 4 - 1 3 9
( M O ) B r a z i l . P n n A : T u c u r u i , R a m o s 1 1 1 5( I N P A ) . P e r u . A n a ' r z o x n s v: a l l e d e l R f o S a n t i a g o ,c a . 6 5
km N dc Pinglo. Quebrada Caterpiza. fIuashikat 1ll0 (MICH, MO)l Prov. Bagua, B km E of
Montenegro at Km 286 E ol Olmos on the Mcsones-Muro hwy, Hutchison & Wright 3781 (F, GH,
K , M . M I C H , M O . N Y , P , U S ) l M i r a n a , W o y t k o w s k i5 6 4 9 ( G , G H , M O . U S ) l P r o v . B a g u a .v a l l e y o f
t h e R i o M a r a f r 6 n a b o v e C a s c a d a sd e M a y a s i , K m 2 7 6 - 2 8 0 o f M a r a f l 6 n r d . W u r d a c k I B 4 l ( F , N Y . S ,
U S ) . - L o r < r r o : M a y n a s . S h u s u n a ,c a r r e t e r aa Z u n g a r o c o c h a ,A y a l a 4 3 7 ( A M A Z , M O ) : R f o Y u v i n c t o ,
l 993
IN THE AMAZON
REGION
409
afflucnt du Rfo Putumayo, Rio Putumayo, Barrier 441 (AMAZ, MICH)I cerca de Zungaro. Cocha.
1 - 5k m S W d e l q u i t o s , D o d s o n 2 8 0 9 ( M O , S E L , U S ) ; N a u t a , R i o M a r a f r 6 n a b o v e m o u t h o f R i o
U c a y a l i , 0 4 " 3 0 ' S ,7 3 ' 3 0 ' W , G e n t r l t e t a l . 2 9 9 6 8 ( A M A Z , M I C H ) ; M a y n a s . Q u e b r a d a S u c u r s a r i ,R i o
N a p o , 0 3 " 1 5 ' S .7 2 ' 5 5 ' W , G e n t r v e t a l . 1 2 6 7 3 ( M I C H ) ; M i s h u y a c u . n e a r I q u i t o s , K l u g l 1 . l ( F , N Y ,
US); Prov. Loreto. Pampa Hermosa and vicinity, Rio Corrientes, 1 km S of junction with Rfo
M a c u s a r f , 0 3 ' 1 5 ' S . 7 2 " - 5 0 ' W ,L e w i s e t a l . 1 0 6 5 1 ( M O ) ; M a y n a s , I q u i t o s , Q u i s t a C o c h a , M c D a n i e l
t0907 (F. MO); Rcqucna, Rfo Tepiche, Santa Elena, McDaniel & Marcos 11263(F, MO)l Santa
Ana on the Rfo Nanav. Ll. Williams 1225(F. US).
Stigmaphyllon sinuatum is a common and polymorphic species of the lowl a n d s o f n o rth e rn S outh A meri ca to w hi ch severalnames have been app lied. I t s
flowers are borne in pseudoracemesarranged in compound dichasia and thyrses
with axes to the sixth order. The androecium differs from that of most speciesin
th a t th e p o s te r i or-l ateralstamens bear unmodi fi ed anthers. Onl y the a nt er ior l a te ra l s ta me n shave the connecti veenl argedand the l ocul esreduced;som et im es
one or rarely both locules are absent. All anthers are glabrous. The styles are
usually bearded adaxially but sometimes glabrous, and the posterior ones are
always foliolate; the anterior style either bears two folioles or lacks them, or
ra re l y i s i rre g u l a rl y ornamented.The samara,l i ke that of most speci es,has a lar ge
flaring dorsal wing, and the nut commonly bears on each side 1-3 lateral winglets
and/or spurs and crests.
Stigmaphyllon sinuatum is highly variable in the shape and abaxial pubescence of the leaves, the color of the petals, and the presence of folioles on the
anterior style. Representativescombining the various expressionsof these characters have been segregatedas species.Although the extremes are striking and may
show some geographic correlation, they are linked by intermediates occurring
throughout the range and thus are not accorded taxonomic recognition here. In
Fig. 4. the various combinations of leaf pubescence and ornamentation of the
anterior style are mapped; however, this map reflects more the itineraries of
c o l l e c to rsth a n the range of the speci es.
The laminas vary from triangular to ovate to cordate to elliptical to orbicular
to oblate to reniform. and are cordate to auriculate at the base to sometimes
truncate or acute in smaller ones. In general, plants from the eastern and western
p a rt o f th e ra n ge have rather broad l ami nas (i .e.,broadl y el l i pti cal to orbicular t o
oblate to reniform), and those from the central part are more likely to have
triangular to ovate to elliptical ones. The appressedabaxial pubescenceimparts a
silvery metallic sheen.The abundance of the hairs varies greatly from sparse(the
hairs barely if at all touching) to moderate (the hairs touching and/or overlapping
b u t th e e p i d e rm i svi si bl e) to so densethat the epi dermi si s hi dden by a si lver m at
of hairs. The very dense pubescenceis most common in plants from the eastern
p a rt o f th e ra n g e (Fi g. a).
The petals range from entirely yellow to yellow with varying amounts of red.
If the petals in bud are yellow, they may remain yellow at maturity, or the flag
and sometimes also the lateral petals may show red streaks. If the petals are red
in bud, the expanded petals may be mostly red with only a yellow spot on the
adaxial surface,may be mostly yellow but with a broad red band along the margin, or only the flag may be red and the lateral petals mostly yellow with a little
red, either in streaks or along the margin. Sometimes the red pigment is evident
only in bud and the expanding petals but is absent from the mature petals. Mostly, pure yellow limbs are slightly larger than those suffused with red. Specimens
from Colombia, Ecuador, and Peru nearly always have yellow petals lacking red,
410
VOLUME I9
S t i g m a p h y l l o ns i n u a t u m
leaf pubescence
dense moderale sparse
=eff::' o
o
o
ictu*
tr
tr
A
A
I
; unequal
Eornamen-A
tation
l./
o
8--
\itrr
\ _--*\\
+-
H
I
F I G . 4 . D i s t r i b u t i o n o f t h e v a r i a n t s i n l e a f p u b e s c e n c ea n d o r n a m e n t a t i o no f t h e a n t c r i o r s t v l e
of Stigmaphvllon .sinuatum.
but yellow petals with minor red markings have also been reported (e.g., Ecuador Harling et al. 14757;Colombia: Davis 107;Peru: McDaniel 10907).The petals
are abaxially sericeousin nearly all collections from Amapd, Brazll. and rarely in
specimens from other parts of the range (e.9., Teixeira et al. 1340, Amaz6nas,
Brazll, Murga Pires et al. 620, Par6, Brazll; Wurdack 34434, Bolfvar, Venezuela).
Pubescent petals are otherwise known only in S. sirzgulareC. Anderson, a very
different speciesof Venezuela (Tiichira) and Colombia (Norte de Santander).
The names Banisteria heterophylla, B. splendens,Stigmaphyllon splendens,S.
fulgens, S. hypoleucum, and S. purpureum are based on collections from eastern
Venezuela and the Guianas. The laminas of plants from the northeastern part of
the range are usually orbicular to oblate or reniform and densely silver-sericeous
below (Fig. 4, closed symbols). The petals have broad red-orange margins or may
be entirely red with only a central yellow spot abaxially. The anterior style bears
folioles.
In plants from the western part of the range as well as from the area of Bel6m
(Parii), the Rio Tocantins and its tributaries, and MaranhSo, all in Brazil. the
laminas are most often broadly cordate to broadly ovate or sometimes narrower,
though the orbicular to oblate/reniform shape is also infrequently encountered;
the abaxial hairs are sparsely to moderately abundant (Fig. 4, open and partly
closed symbols, respectively). Most commonly, the petals are yellow. The form
with abaxially sparsely to moderately sericeous laminas and foliolate anterior
styles is commonly assignedto S. martianLtm.a name based on a collection from
Manaus (Amazdnas, Brazil). The majority of specimensfrom the vicinity of Manaus, including the type of S. martianLtm,have triangular to ovate laminas that are
acute to truncate to cordate at the base and sparsely sericeousbelow; the petals
are yellow.
Much overlap occurs in the general rangesof the leaf variants.The laminas of
plants of Guyan a are often not quite as densely pubescent as those from French
Guiana and Suriname. In a few collections from French Guiana, including the
type of S. richardianum, the elliptical to broadly so to cordate (but not orbicular)
1993
411
laminas are sparselyto moderately sericeousbelow, and the petals often lack red
pigments (e.g., Grenand 2136, Oldemqn 1982,Jacquemin l610). A few collections
of these variants are also known from Guyana (e.g., de la Cruz 2578, Sandwith
664), Suriname (e.g., Lindeman et al. 47'), easternVenezuela (e.g.,Sobel & Strudwick 2274, Sucre), eastern Brazil (".g., Cowan 38473B,lrwin et al. 47382,Amapi{).
Some collections from Manaus have yellow-red petals and very broad (suborbicular) though abaxially sparsely pubescent laminas (..9., Campbell et al. P20920,
Maas & Maas 362, Chagas INPA 3679). Most collections from the vicinity of
Santardm have broad, abaxially densely sericeouslaminas and red-yellow petals;
yet, plants with sparselyto moderately pubescentlaminas and yellow petals occur
there as well (e.g., Silva & Souza 2522). In Roraima, the entire spectrum of
variation in laminar shape and pubescenceand in petal color is encountered. In
some specimensfrom this region and elsewhere (e.9., Prance 3964, 10909,Roraim4 Thomas et al. 5034, Rond6nia), the abaxial surface of the young leaves and
smaller ones near the inflorescenceis densely sericeous,whereas that of the larger leaves is only moderately pubescent.The broad laminar shape and yellow-red
petals typical of the eastern variant is also found in plants collected along the Rio
Madeira (e.g., Teixeira et al. 980. Prance et al. 5IB0) and in northern Rond6nia
(..g., Prance et al. 6658,Thomas et al. 4949) as well as along the Rio Mamor6 and
adjacent Bolivia (e.g., Solomon 7682): however, the laminas are only sparsely to
moderately pubescent below. In most collections from Ecuador and Peru, the
laminas also are broadly elliptical or cordate to orbicular.
In a few speciesof Stigmaphyllon characterized by foliolate anterior styles,
individuals are occasionallyencountered in which the folioles of the anterior style
are greatly reduced or one or both are absent. Such atypical forms are known in
S. adenodon Adr. Juss. (Peru), S. lindenianum Adr. Juss. (Yucatan Peninsula,
Mexico). and S. sagraeanum Adr. Juss. (Cuba). In S. sinuatum, plants from the
greater part of the range have foliolate anterior styles (Fig. 4, circles),but in many
from western Venezuela (and also Amazonas and western Bolivar), Colombia,
Ecuador, and northern Peru (Amazonas and Amazonian Loreto) the apex of the
anterior style is efoliolate but extended into a claw 0.7-1.3 mm long (Fig. 4,
squares).Exceptionsto this geographicalpattern are rare but not unknown; for example, in Ramos I I l5 from Tucuruf, Par6, Brazil, the anterior stylesare efoliolate, and in
Maguire et al. 44142 from Araracuara, Amazonas, Colombia, they are foliolate.
The names Stigmaphyllon brachiatum and S. monancistrum are based on the
efoliolate form. In his treatment of Stigmaphyllon in the Guayana Highlands, W.
R. Anderson (1981) commented on the morphological diversity of this species
and questioned the validity of taxonomic recognition of the efoliolate variant as
S. brachiatum.
Intermediate style forms occur, infrequently, throughout the range of S. sinaatttm (Fig. 4, triangles). In these individuals, the anterior style may bear only one
or two greatly reduced folioles, only one large foliole, or one reduced and one
large foliole. The size and/or presenceof such folioles may vary even within the
same inflorescenceor even the same umbel. Such intermediates otherwise share
the characters of "normal" plants of their part of the geographic range. The
following collections serve as examples of plants with variable anterior styles:
Guyana, Jenman 6875, two large to tiny folioles; Suriname, BBS 254,259, one
foliole or two unequal folioles; Venezuela, Trujillo 4127 (Aragua), Pittier 5699,
12105, Diederichs 8B (Yaracuy), Steyermark 61019 (Azo6tegul), Stergios 6224,
8902, Bernardi 7226 (Bolivar), all with one foliole or two unequal folioles; Colombia, Garcfa B. lB24B (Norte de Santander), one foliole or two unequal folioles;
412
VOLUME 19
Brazll, Maas PI28l5 (Acre), Vieira et a\.546 (Ronddnia), all with one foliole or
two unequal folioles; Peru, Huashikqt 1375 (Amazonas), two folioles or one folio
a n d a n a rro w l i p. In one V enezuel ancol l ecti on (R omero 545,Y aracuy), the ant erior styles varies from hooked, i.e., without folioles, to bearing I-2large folioles.
The absenceor variability of folioles of the anterior style might indicate irregularities causedby hybridization; however, a preliminary survey of pollen showed no
correlation between absenceof folioles and presenceof malformed pollen grains.
In most samples of all three variants of ornamentation of the anterior style only
few misshapengrains were found; the pollen was mostly 95-99% normal (at least
9 1 % n o rm a l ). The few excepti onsare Ol deman l 9B 2 (French Gui ana,86 % nor mal), Maguire 44142 (Colombia, Amazonas, 85% normal), Prance et al. P25427
(Brazil, Par6,,58-72% normal), all three collections with foliolate anterior styles;
and Breteler 4721 (Venezuela, Amazonas, 83-84"/" normal), anterior styles efoliolate. Only in two collections did the pollen consist entirely of misshapen,thickwalled grains: Broom & Beardsley 8436 (Ecuador, Pastaza,anterior styles efoliolate) and Bernardi 7226 (Yenezuela, Bolivar, anterior styleswith variable folioles).
Because S. sinnatum is so variable and widespread, sympatric species with
abaxially sericeouslaminas have often been confused with it; for separation from
these species,see the key presented above and the discussionsof S. argenteum,S.
cardiophy llum, and S. convolvulifolium.
In addition to taxonomic difficulties. S. sinr.ratumhas also been a source of
nomenclatural problems. The specieswas first described by Willdenow (1799) as
Banisteriaheterophylla.De Candolle (1824) considered this name unsuitable and
in his Prodromrzs substituted the name B. splendens. Cuatrecasas(195U) published the name Stigmaphyllon splendens;however, that name is also superfluous,
becausehe cited in synonymy two legitimate available names,S. hypoleucum and
S. purpureLtm.The epithet "heterophyllum" is no longer available in Stigmaphytlon,because it has already been used for another species[5. heterophyllum Hook€r = S. littorale Adr. Juss.].Matters were also confused by Lamarck (1783), who
misapplied a Linnaean name, B. fulgens, to the specieshere discussed[8. futgens
L. = S. emarginatum (Cav.) Adr. Juss.].This error was noted by Cavanilles(1790),
who had correspondedwith J. E. Smith, then the owner of the Linnaean herbarium. Cavanilles treated the speciesunder the name " B. fulgens Lam." When Jussieu (1840, 1843) published S. fulgens, he noted that he was basing the name on
Banisteria.fulgensin the senseof Lamarck, not Linnaeus, but also listed B. heterophylla Willd. and B. splendensDC. in synonymy. S. fulgens Adr. Juss. is thus a
superfluous name. Niedenzu (1900, 1928) followed Jussieuand cited this species
under the name "5. fttlgens (Lam.) Juss."
W. R. Anderson (1981) also discussedthe nomenclatural difficulties. He noted that the names S. fulgens and S. splendensare illegitimate and that the name S.
heterophyllum had already been used for another species.He adopted S. hypoteucum Miquel, the earliest legitimate name for the variant with abaxially densely sericeous leaves,but also indicated that this name would be displacedby S. richardianum
Adr. Juss.and S. martianum Adr. Juss.,if their types are included in this species.
De Candolle's B. splendens (i.e.. B. heterophylla Willd.) was based on the
variant in which the lower leaf surfacesare very densely pubescent.De Candolle
also had a specimenof the form in which the leavesare only moderately sericeous
below, a Perrottet collection from French Guiana; this specimen is the type of his
name Banisteria sinuata, also published in the Prodromus (7824). Jussieu (1840,
1843) made the combination in Stigmaphyllon, but expressedsome doubts that
I 993
C ] .A N D E R S O N :S T I G M A P H Y L L O N I N ] . H E A M A Z O N R E G I O N
413
his concept of the speciesand de Candolle's B. sinuata were the same. lJnfclrtun a te l y , i t p ro v e d that the P errottet speci mensthat Jussi cusaw at P and annot at ed
as S. slnuatum belong to S. palmatum (Cav.) Adr. Juss.,and were not duplicates
o f d e C a n d o l l c ' s type of B . si nuata,as Jussi eubel i eved. In Jussi eu' sdef ense,it
should be noted that S. palntatum is also greatly variable in leaf shape and that its
type. crroneously, was said to be from Hispaniola; the type of S. palmatum has
d e e p l y p a l m a te l ami nas,w hereas the P errottet speci mensJussi cu saw h ave unl o b e d l e a v e s .N i edcnzu confounded the matter further by recogni zi ngde Cand o l l e ' s B . s i n u a ta as a vari ety, fi rst (1900) of S . hastatumGri seb. [a super f luous
name based on B. ungulata Vell. (=5. auriurlatum (Cav.) Adr. Juss.),which Grisebach misapplied to S. littoralef and later (192tt) of S. sagittatum [sensu Niedenzu.
including S. hostatum: S. sagittatum (Cav.) Adr. Juss. = S. palmatumf. Jussieu's
a n d N i e d e n z u ' s mi si nterpretati onsof B . si nttatal ed l ater botani ststo apply t he
name S. sinuatunt to specimensof either S. littorale or S. palmatum. My recent
c x a mi n a ti o no f the type of B . si nuataccl nfi rmedthat i t bel ongsto the speciesher e
(DC.) Adr. Juss.
discussed.and that in Stigmaphyllonthe correct name is S. sirzurztunt
ACKNOWLEDGMENTS
I t h a n k W i l l i a m R . A n d e r s o n I ' o r h i s c o m m e n t s .I a m g r a t e f u l t o R . V i c k e r y ( t s M ) . A . C h a r p i n
( G ) . M J . S a n d s ( K ) . a n d A . L o u r t c i g ( P ) f o r t h e i r k i n d a s s i s t a n c cd u r i n g m y v i s i t t o t h c i r i n s t i t u t i c r n s .I t h a n k t h c c u r a t o r s o f t h e f o l l o w i n g h e r b a r i a f o r m a k i n g t h e i r c o l l e c t i o n s o f S t i g n t u p l r v l l o t t
a v a i l a b l et o m e : A . A A U . A M A Z . B M , B R , C , C A S , C A Y . C M , C O L . E C O N . F . G . G B . G H .
G O E T . I A N . I N P A . J B S D . K , L E . M . M B M . M E , R .M G . M I C H . M O . M Y , N Y . P . R . R B , S , S E L .
SP. UCLA. [J. UI]. US. VEN, W. WAG, WIS. Figurcs l-3 wcrc drawn by Karin Douthit. Fig. 3 is
r e p r i n t e d b v p c r m i s s i o n f r o m M e m o i r s o . f t h e N a w Y o r k B o t a n i c a l G a r t l e n 3 2 : 1 6 9 . 1 i g . - 5 .c o p y r i g h t
1 9 8 1 t. h e N e r v Y o r k B o t a n i c a lG a r d e n .
LITE,RATURECITED
A n d e r s o n . C . 1 9 8 7 .S t i g m a p h y l l o n( M a l p i g h i a c e a e )i n M e x i c o .C e n t r a l A m e r i c a .a n d t h c W c s t I n d i c s .
Contr. Univ. MichiganHerb. 16:l-4tJ.
A n d c r s o n . C . 1 9 9 2 .T w o s p e c i e so f S t i g m a p h y l l o n( M a l p i g h i a c e a e )f r o m P e r u . N o v o n 2 : 3 0 2 - 3 0 - 5 .
A n d e r s o n . W . R . 1 9 8 1 .M a l p i g h i a c e a e l. n T h e B o t a n v o . ft h e G u u r u r t u H i g h l t t n d - P u r t X I b v B a s s e t t
M a g u i r c a n d c o l l a b o r a t o r s .M c m . N e w Y o r k B o t . G a r d . 3 2 : 2 1 - 3 0 5 .
vol. l. Paris:Trcuttelet Wiirtz.
naturalis reg,nivegetabilis,
Candolle. A. P. de 1824.Protlronlus systentatis
C a v a n i l l e s .A . J . 1 1 9 0 .N o n a t l i s s e r t a t i ob o t a n i c t t .M a d r i d : T y p o g r a p h i a r e g i a .
. cbbia l3: 343-66'1.
C u a t r c c a s a sJ. . 1 9 5 t i .P r i m a F l o r a C o l o m b i a n a . 2 . M a l p i g h i a c e a e W
G r i s c h a c h . A . H . R . 1 t 3 - 5 8M. a l p i g h i a c e a e .l n F l o r u b r a s i l i e n s i se. d . C . F . P . v o n M a r t i u s , l 2 ( 1 ) : 1 - 1 0 t 3 .
. n n . S c i . N a t . t s o t . ,s i r . 2 , 1 3 : 2 4 1 - 2 9 1 . 3 2 1 3 3 8 .
. a l p i g h i a c e a r u ms y n o p s i s A
J u s s i c u .A d r . d c . 1 1 i 4 0M
J u s s i e u .A c l r . d c . 1 8 4 3 .M o n o g r a p h i e d c l a l a m i l l e d e s m a l p i g h i a c i e s .A r c h . M u s . H i s t . N a t . P a r i s 3 :
. s - 1 5 1 .2 5 5 - 6 1 6 . t . . - 2 . 1 .
L a m a r c k , J . B . A . P . M . d e 1 7 8 3[ 1 7 8 5 ] .B a n i s t e r i a . l n E n c l ; c l o p t d i em t t h o d i q u e . B o t a n i q u e ,e d . J . B .
A . P . M . d e L a m a r c k . l : 3 6 6 - 3 6 9 .P a r i s :P a n c k o u c k e .
M a c b r i d e . J . F . 1 9 4 9 .M a l p i g h i a c c a e .I n F l o r a o f P e r u , P u b l . F i c l d M u s . N a t . H i s t . . B o t . S e r . l 3 ( 3 ) :
7 8 I - r 3 7 1.
N i e d c n z u . F . 1 9 0 0 .D c g c n c r e S t i g m a t o p h y l l o ( p a r s p o s t e r i o r ) . I n d . L e c t . L y c . B r u n s b e r g .p . a e s t .
1 9 0 0 :I 3 2 .
N i e d e n z u , F . 1 9 2 8 . M a l p i g h i a c e a e .I n D a s P f l u n z e n r e i c h ,e d . A . E n g l c r . I V . l 4 l : l - t 3 7 0 . L e i p z i g :
Wilhelm Engelmann.
s l a n t a r u m .v o l . 2 . B e r l i n : N a u k .
W i l l d e n o w , C . L . 1 7 9 9 .S p e t ' i e p

THE IDENTITIES OF THE SERICEOUS.LEAVED SPECIES OF

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