Download A Synopsis of Neotropical Plukenetia (Euphorbiaceae) Including

A Synopsis of Neotropical Plukenetia (Euphorbiaceae) Including Two New Species
Author(s): Lynn J. Gillespie
Source: Systematic Botany, Vol. 18, No. 4 (Oct. - Dec., 1993), pp. 575-592
Published by: American Society of Plant Taxonomists
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Systematic
Botany(1993), 18(4): pp. 575-592
? Copyright1993 by the American Society of Plant Taxonomists
A Synopsis of Neotropical Plukenetia (Euphorbiaceae)
including Two New Species
LYNN J. GILLESPIE
Botany Department NHB#166, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560
A synopsis of the eleven neotropical species of Plukenetia(Euphorbiaceae) is given.
ABSTRACT.
The genus Plukenetiais redefinedto include the neotropical monotypicgenera Eleutherostigma
and
Vigia (previously known as Fragariopsis).As treated here Plukenetiamay be distinguished fromall
othermembersof the tribePlukenetieae by itsfour-carpellateovary.Two new species, P. supraglandulosa fromFrench Guiana and the Brazilian state of Amapa and P. stipellata fromMexico and
Central America, are described and illustrated.Plukenetiasupraglandulosais closely allied with P.
penninerviaand P. multiglandulosa
and differsin the presence of scattered laminar glands on the
upper leaf surfaceand longer inflorescences.The three species share a very similar androecium of
dimorphic stamens,which is correctlydescribed for the firsttime. Plukenetiastipellatadiffersfrom
the closely related South American and West Indian species, P. volubilis,
by its stipellate leaf blade
base, shorterstylesand longer slender stamens.Two new combinations,P. serrataand P. lehmanniana, are proposed. A key to the 11 species in the Neotropics is provided.
PlukenetiaL. is a pantropical genus of 16 species-of twining vines and lianas belonging to
the Euphorbiaceae. The genus is notable forits
four-carpellateovary,stylesthatare partiallyto
entirelyfused and oftenmassive, and scandent
habit. Other diagnostic charactersinclude one
or more pairs of adaxial basilaminarglands, numerousstamens,and pistillateflowerswith four
sepals. Two new species ofPlukenetiawere identifiedwhile working on preliminaryrevisional
studies of the genus and the treatmentof the
tribe Plukenetieae for the Flora of the Guianas.
Since manyofthe neotropicalspecies are poorly
known and specimens are frequentlymisidentified,itseems appropriateto provide a synopsis
of the 11 neotropicalspecies. This will also serve
to introduce several nomenclatural and taxonomic changes,and will make available the new
names and combinations for floristicstudies
priorto a complete revision of the genus. While
species of Plukenetiaare forthe most part well
defined,some collections appear to be morphologically intermediateor to combine characters
of several species and cannot be easily placed.
Field studies and more collections are needed
to determinethe statusof these anomalous collections beforea monograph can be completed.
Plukenetiabelongs to the tribe Plukenetieae
of the uniovulate subfamily Acalyphoideae
(sensu Webster 1975). Members of Acalyphoideae characteristicallylack latex, and have
eglandular inflorescencebracts,apetalous flowers and staminate flowerswith valvate sepals.
The tribe Plukenetieae is distinguished by the
charactercombination of entire styles that are
partlyto completelyconnate and oftenmassive,
and bisexual racemose or spicate (or rarelypaniculate) inflorescencesbearing cymoseclusters
of flowerswith pistillate flower(s)at the basalmostnode(s) (unisexual inflorescencesare found
in a few taxa including AcidotonSw. and a small
percentageof species of TragiaL.). Members are
frequentlyscandent,a veryunusual habitin the
family. Webster (1975) divided the tribe into
two subtribes,the Tragiinae characterized by
the presence of stinging hairs and the Plukenetiinae, which lack stinging hairs. The tribe
was monographed by Pax (1890) and Pax and
Hoffmann(1919, 1931; their subtribePlukenetiinae is equivalent to Webster's tribe Plukenetieae), and morerecently,pollen morphology
was surveyed (Gillespie, in press).
The circumscriptionof genera belonging to
subtribe Plukenetiinae (sensu Webster) has
changed considerably over the last 150 years
[although the subtribe was not formallyrecognized prior to Webster(1975), changes in its
constituent genera can be traced]. The three
genera,Angostyles
Benth.,Astrococcus
Benth.,and
Haematostemon
(Muell. Arg.) Pax & K. Hoffm.,
distinguished by their shrub or tree habit and
tricarpellateovary, are well defined and have
been consistentlyrecognized by most authors;
they appear to be more closely related to each
otherthan to Plukenetia(Gillespie, in press). The
remaining genera may be recognized by their
575
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576
SYSTEMATIC BOTANY
[Volume 18
twining vine or liana habit and, with one ex- ence in degree of style fusion and represents
ception, by a four-carpellate ovary. Baillon only one end of a continuum(Fig. 1). Likewise,
(1858), Bentham(1880), Mueller (1866), and Pax the single unique featurecharacterizingVigia,
(1890) adopted a broad generic concept forspe- an unusually enlarged globose staminaterecepcies with a scandent habit. Mueller and Baillon tacle, is simply a differencein size, since all
recognized only two genera, Plukenetia(called species have globose or convex receptacles [re(Pax
Sajorumby Baillon) and FragariopsisA. St. Hil. ferto the discussions under P. lehmanniana
(=Vigia Vell. Conc.), while Bentham and Pax & K. Hoffm.)Huft & L. J.Gillespie and P. serrata
recognized only Plukenetia;all these authors (Vellozo) L. J. Gillespie in the section "Enusubdivided Plukenetiainto three to six sec- meration of Taxa" below for a more complete
and Vigiaare
tions.In contrast,Pax and Hoffmann(1919, 1931) discussion]. When Eleutherostigma
recognized seven genera. In addition to Pluke- excluded, Plukenetiais a paraphyletic genus
theytreatedAnabaenaAdr. lacking any uniquely derived defining charnetiaand Fragariopsis,
Juss. (which they renamed Anabaenella,but is acters.
Pax and Hoffmann (1919, 1931) recognized
now correctlyknown as RomanoaTrev. St. Leon;
see Gillespie, in press; Radcliffe-Smith1980), two sections in their narrowly defined, neo(Muell. Arg.) Pax & K. Hoffm. tropical genus Plukenetia,sects. Euplukenetia
Angostylidium
[also conHassk. as (=sect. Plukenetia)and Cylindrophora
Pax), and Pterococcus
(=Tetracarpidium
genera rather than sections of Plukenetia,cre- sidered sections by Mueller (1866) in his more
ated a new genus, Apodandra,fortwo species of broadly defined circumscriptionof the genus].
Plukenetia,and described a new monotypicge- In this treatmentof neotropical species, two inMore recently, Webster formalspecies groups are outlined, which cornus, Eleutherostigma.
(1975) recognized four genera among species respond to more broadly circumscribed verhaving the vine or liana habit, Plukenetia,Eleu- sions of the two sections.However, recognition
Fragariopsis(=Vigia) and Anabaena of a formal sectional classificationwill await
therostigma,
revision of the paleotropical species and a phy(=Romanoa).
In the present treatmentonly two genera, logenetic study of the genus.
Neotropical species of Plukenetiamay be diPlukenetiaand Romanoa,are recognized among
vided
into two informalspecies groups based
scandent members of subtribe Plukenetiinae;
and Vigia are included on styleshape, androecium morphology,pollen
both Eleutherostigma
within Plukenetia.The principal defining syn- exine structure,fruitsize, and leaf architecture.
apomorphies of Plukenetiaare the four-carpel- The firstgroup (species group 1, "Cylindrolate ovary and the associated characterof four phora") is characterizedby stylesthat are only
pistillate sepals. Romanoa is considered a dis- partly fused into a cylindrical stylar column
tinct monotypic genus on the basis of its pis- (mostly fused in P. volubilisL. and mostly free
Figs. lA-C, 9C), androecia in
tillateflowerswith a tricarpellateovaryand five in P. lehmanniana;
have distinctfilaments(Fig.
all
stamens
which
the
on
sepals, and the sistertaxon of Plukenetia
basis of numerous shared characters of floral 9D-E), pollen with a foveolate tectum(Figs. 4and pollen morphology(Gillespie, in press). An 6; Gillespie, in press), large fruit,and palmately
alternative and phylogenetically equivalent veined or triplinerved, cordiform, ovate or
(Adr. broadly elliptic leaves (Figs. 3, 9A). The followwould be to considerR. tamnoides
treatment
Juss.)R.-Smithas the most plesiomorphic spe- ing four species comprise this group: P. lehas circumscribedhere manniana,P. polyadeniaMuell. Arg., P. stipellata
Plukenetia
cies ofPlukenetia.
includes 11 neotropicalspecies,one species from L. J.Gillespie, and P. volubilis.
The second species group (species group 2,
southeastAsia, threefromAfrica,and one from
"Euplukenetia") is characterized by entirely
Madagascar.
The neotropicalmonotypicgenera,Eleuthero- fused styles(Figs. 1D, 1OH), all or at least most
stigmaand Vigia,are included within Plukenetia anthers sessile (Fig. lOG), pollen with a reticsince neitherare defined by unique characters ulate tectum(Figs. 7-8; Gillespie,in press),small
considered significantat the generic level. The drycapsules (exceptP. serrata),and elliptic,pinshortly natelyveined leaves (Figs. 2, 1OA;exceptP. verdefining character of Eleutherostigma,
connate ratherthan mostlyto entirelyconnate rucosaSmith). Three species belonging to this
Muell. Arg., P. loretensis
styles,appears to be only a quantitativediffer- group, P. brachybotrya
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GILLESPIE: PLUKENETIA
19931
1mm
577
"mu
lmm
(Acevedo
FIG. 1. Pistillate flowersof Plukenetiashowing variation in style morphology. A. P. lehmanniana
& Daly 1691,US). B. P. polyadenia(Black52-14173,P). C. P. volubilis(Zarucchi2378,US). D. P. verrucosa(Irwin
48796,US); note the pubescent and indistinctlybilobed ovary lobes between the sepals. Note the cyme axis
and bracteole(s) subtending the pedicel in A-C (not shown in D).
Ule and P. serrata,have androecia in which all
anthers are sessile on a globose receptacle. P.
Muell. Arg.,
Jabl.,P. penninervia
multiglandulosa
and P. supraglandulosaL. J. Gillespie form a
closely relatedspecies complex (referredto here
as the P. penninerviaspecies complex) characterized by androecia consisting of two distinct
typesofstamensand by a shortstout-cylindrical
stylarcolumn (Fig. 10H). This unusual type of
androecium, correctlydescribed here for the
firsttime,consistsof an outer whorl of four(or
rarely five) stamens having filamentsand an
inner clusterof sessile anthersdensely crowded
on a small globose receptacle (Fig. lOG). Previous authors have overlooked the dimorphic
nature of the stamens and have usually describedthe stamenssimplyas having veryshort
filaments.Plukenetiaverrucosaalso has dimorphic stamensbut differsin having a subglobose
stylarcolumn and narrowlycordiformor ovate
triplinervedleaves.
Using the hypothesisof Romanoaas the sister
taxon of Plukenetia, many of the above characters can be tentativelypolarized. Romanoais
characterized by palmate-veined cordiform
leaves, partlyfusedstyles,stamenswith distinct
filaments,and a fossulate-foveolatepollen tectum. This would indicate that entirely fused
styles,sessile anthers,and reticulatepollen tectum are synapomorphiesof the second species
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578
[Volume 18
SYSTEMATIC BOTANY
sd
3
, \
N~~~~~~~~~~~~~~~~
sb~~~
Ab
b
f~~~~~~~
l1cm
FIGS. 2-3. Leaf Architectureof Plukenetia.b = basilaminar gland pair; sd = scatteredlaminar glands on
adaxial surface;sb = scatteredlaminarglands on abaxial surface. 2. P. supraglandulosa
(Granville3626,CAY). 3.
P. stipellata(Gillespie 413, US).
group, and that pinnately veined leaves are a
synapomorphywithin that group.
The presence of stipels or a glandular knob
at the petiole apex is a featureunique to Plukenetiaamong members of the tribe Plukenetieae. The following three species have paired
stipels: P. serrata,P. stipellata,and P. verrucosa;
while the following have a single glandular
P. polyadenia,and P. volknob: P. lehmanniana,
ubilis.Since a small knob is occasionally present
between the pair of stipels in P. stipellata,these
two structuresappear to be distinctand probably not directlyhomologous. The stipels are
most likely comparable to the similarlyplaced
stipels of DalechampiaL., a genus related to the
Plukenetieae (in which it is sometimesincluded
as a separate subtribe; Webster,in press), and
may have originated frompaired acropetiolar
glands that are commonly found in species of
Euphorbiaceae. They are presumably not homologous with morphologically similar structuresin otherfamilies."Stipels" is used here in
a broad sense to indicate a paired appendage
located at the leaf blade base and not in the
strictfunctionalsense of secondary stipules at
the base of leafletsof a compound leaf.
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579
GILLESPIE: PLUKENETIA
1993]
*UI
-IM7~~~~~
s1-~~~~s
V^
4I4.'
7~~~~~~~J
@
U
44
-
b~~~
FIGs. 4-8. Pollen morphology of Plukenetia. 4-6. P. stipellata(Gillespie413, US). 7-8. P. supraglandulosa
(Cowan38204,US). 4, 7. Polar view. Scale bars = 10 ,um. 5. Equatorial view. Scale bar = 10 ,um. 6, 8. Exine
sculpture.Scale bars = 2 Mm.
While the majorityof species are relatively
well defined,a number of collections appear to
be intermediate,to combine charactersof several species, or to be somewhat distinct.These
anomalous collectionsare noted and brieflydescribed. Although at least some may represent
new species, our knowledge of them is inadequate and additional collections are needed to
determinetheir status. Furthercollections are
also necessaryto determinemore precisely the
range of variation in known species and the
existence of possible hybrids.It is hoped that
by drawing attentionto these problematiccollections, field botanists will be encouraged to
collect populations of Plukenetiamore extensively, particularlyin poorly known and undercollectedareas of Amazonian Brazil and the
eastern slopes of the Andes.
One explanation for the above problematic
collectionsis the existenceof species complexes
consisting of populations that are geographically or altitudinallyisolated but morphologically little differentiated.An example is the P.
penninerviaspecies complex consisting of one
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580
SYSTEMATIC BOTANY
[Volume 18
lcm
u1
ln'm
lcm~~~~~~~~~~~~~~c
lcm
H
AS
lrnm
lmm~~~~~~~~~~~
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All use subject to JSTOR Terms and Conditions
n~~~~~~~~~~~~~
1993]
GILLESPIE: PLUKENETIA
species widespread in Mesoamerica and Venezuela and a numberof isolated populations scattered throughoutSouth America (P. multiglandulosa, P. supraglandulosaand the anomalous
Colombian population of P. penninervia).Plukenetiavolubilisand P. stipellatacomprise a second species complex. While considered as disKEY TO THE NEOTROPICAL
581
tinctbut closely relatedspecies here,theycould
alternativelybe treatedas subspecies of a single
variable widespread species. Plukenetiabrachybotryaand the numerous problematic collections cited under that species may representa
thirdspecies complex.
SPECIES OF PLUKENETIA
1. Leaf blade pinnately veined, acute, obtuse, or rounded at base; glandular knob lacking at midrib base,
stipels presentonly in P. serrata;scatteredlaminar glands usually presenton lower leaf surface(except
in P. penninervia
and P. serrata);styles entirelyconnate, 1-4 mm long; all anthers sessile or stamens
dimorphic with inner anthers sessile and outer with short filaments(as in Fig. lOG).
2. Fruit ca. 4 cm or more in diam., fleshy;pistillate flowers 1-10 per inflorescence;all anthers sessile
and scattered on globose receptacle, receptacle 2 mm or more in diam., clearly visible between
anthers; stylarcolumn massive-obovoid; pair of stipels present adaxially at midrib base ........
7. P. serrata
................................................................................
2. Fruit 1-2 cm in diam., capsular; pistillate flower1 per inflorescence;all or only some antherssessile
and densely crowded on globose receptacle,receptacle less than 1 mm in diam., not visible between
anthers;stylarcolumn various; stipels absent.
3. Stamens of one type: 15-50 sessile anthers on globose receptacle; stylarcolumn massive-globose
or slender-cylindrical,(1) 2-4 mm long.
4. Stylarcolumn slender-cylindrical;ovaryand capsule with pointed horn on each carpel; stamens
3. P. loretensis
15-25; basilaminar glands in 1 to several pairs ..........
....................
4. Stylarcolumn massive-globose; ovary and capsule with rounded tubercle on each carpel; stamens (24) 30-50; basilaminar glands in a single pair .......
.............. 1. P. brachybotrya
3. Stamens of two types: outer whorl of 4 (5) stamens with filaments,inner cluster of 6-12 sessile
anthers on globose receptacle; stylarcolumn stout-cylindrical,1-2 (3) mm long.
5. Basilaminarglands in 3-5 pairs; young shoots and petioles densely hirsute;blade hirsutebelow,
4. P. multiglandulosa
hairs persistent.
..............................................
5. Basilaminar glands in 1-2 (3) pairs; young shoots and petioles puberulent; blade sparsely
puberulent below, oftenbecoming glabrous.
6. Scattered laminar glands absent or rarely 1-8 near margin on lower leaf surface only; leaf
blade chartaceous to coriaceous, mostly subcoriaceous, margin distinctlyserrulate and
oftendistinctlyglandular; inflorescence1-3 cm long (to 6 cm only in Colombia) ......
.
5. P . penninervia
...................................................................
6. Scattered laminar glands numerous, present on both upper and lower leaf surfaces; leaf
blade chartaceous,margin minutelyserrulate,appearing undulate, never distinctlyglan9. P. supraglandulosa
dular; inflorescence2-8 cm long .............
....................
1. Leaf blade palmatelyveined or triplinerved,cordate,rounded, or broadly obtuse at base; glandular knob
or stipels presentadaxially at midribbase; scatteredlaminar glands absent on lower leaf surface;styles
partlyto entirelyconnate, 1.5-30 mm long; all stamens with distinctfilamentsor stamens dimorphic
with inner anthers sessile (only in P. verrucosa).
7. Stylarcolumn globose, 1.5-2 mm long, styles entirelyconnate; outer stamens with filaments,inner
10. P. verrucosa
anthers sessile; fruitless than 1.5 cm in diam., 4-lobed, capsular ..................
7. Stylarcolumn cylindrical,2-30 mm long, styles only partlyconnate (sometimes free only at tip); all
stamens with filaments;fruitgreater than 2.5 cm in diameter, 4-lobed or subglobose, fleshy or
capsular.
FIG. 9. Plukenetiastipellata,with staminate flower of P. volubilisfor comparison. A. Habit (Leisner2068,
US). B. Adaxial leaf base showing pair of stipels between pair of basilaminar glands (Gillespie413, fromFAA
preserved material). C. Pistillate flower(Gillespie413, US). D. Staminate flower(Gillespie413, US). E. Staminate flowerof P. volubilis(Vargas13994,US). F. Capsule (Pittier324, US, dried specimen). G. Seed, ventral
view (Smith1651,US). H. Seed, lateral view (Smith1651, US).
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582
[Volume 18
SYSTEMATIC BOTANY
1m
j2mm~~~~~~~~m
(I
ZZNK
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]2mm
~~~~~~~~~~~~~~~~~~~~~~~~~~
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~
i
~
~
2m
1993]
GILLESPIE: PLUKENETIA
583
8. Styles free for one half or more of length, column 3-6 mm long, arms 3-6 mm long; staminate
sepals 5; ligulate (strap-shaped)disc segmentspresentbetween stamens;fruitsubglobose, squarish in cross-section ..........................
2. P. lehmanniana
8. Styles free for less than one half of length, column 2-30 mm long, arms 1-2.5 (3) mm long;
staminatesepals 4 or 5; staminate disc segments absent, or if present minute and not ligulate;
fruitvarious.
9. Stylarcolumn 3-7 mm long; fruitsubglobose, squarish in cross-section,greaterthan 6 cm in
diam., indehiscent; staminatebud narrowly oblong-ellipsoid; filaments2-2.5 mm long; inflorescencesmostlyunisexual, dimorphic,pistillate inflorescencewith 3-4 (10) flowers;leaf
base rounded or broadly obtuse ........................
6. P. polyadenia
9. Stylar column 6-30 mm long; fruit4-lobed, 2.5-4 (6) cm in diam., dehiscent; staminate bud
subglobose; filaments0.4-1.2 mm long; inflorescencebisexual with one or two pistillate
flowersat basal nodes; leaf base cordate or truncate.
10. Pair of stipels presentadaxially at petiole apex, glandular knob minute or lacking between
stipels; stamens 25-40, filamentsslender, 1-1.2 mm long; stylarcolumn 6-11 mm long;
8. P. stipellata
staminatesepals 5, rarely4 ....................
10. Pair of stipels absent, single glandular knob present adaxially at petiole apex; stamens 1630, filamentsstout-conical,0.4-0.5 mm long; stylar column 15-30 mm long; staminate
11. P. volubilis
sepals 4 .........................
ENUMERATIONOF TAXA
1. PLUKENETIA BRACHYBOTRYA Muell. Arg., Lin-
67?45'W, Croat51620 (F, MO); Tuiri, on left bank of
naea 34: 158. 1865. Apodandrabrachybotrya Rio Mapiri, Krukoff10753 (G, F, K, MO, NY,
(Muell. Arg.) J.F. MacBr., Field Mus. Nat. US). Pando: cerca de Porveniren direcci6n a Chive,
Casas & Casas 8149 (NY).
Hist., Bot. Ser. 13(3a): 117. 1951.-TYPE:
BRAZIL.
Acre: 35 km fromRio Branco on Rio BranPeru, Herb.Pavon (holotype: G-BOIS!; isoco-Santa Rosa road, Lowrieet al. 315 (GH, MO, NY,
types: G!, G-DC!)
US).
Plukenetiabuchtienii
Pax, Feddes Repert. 7: 110.
ECUADOR.
Napo: Estacion Biologica JatunSacha,
1909. Apodandrabuchtienii(Pax) Pax & K. 8 km E of Misahualli, 1?4'S, 77?36'W, Ceron& Iguago
Hoffm.,Pflanzenr. IV.147.IX.(Heft 68): 21. 5474 (DAV, MO).
1919.-TYPE: Bolivia, Charopampa near
PERU.
Huanuco: Bosque Nacional de Iparia, road
Mapiri, Buchtien1962 (holotype: presum- to Ayamira along Rio Pachitea, Schunke1314 (NY,
ably B, destroyed; lectotype, here desig- US). Loreto:12.5kmSWofIquitos,Croat20058(GH,
MO, NY, P, US). Madre do Dios: ManuiPark,Cocha
nated: US!; isolectotype:NY!).
Distribution
andhabitat. Found in thewestern
part of the Amazon Basin in Brazil (Acre and
possibly Amazonas, Mato Grosso,and Para, see
discussion below), Ecuador, Peru, and Bolivia.
A twining vine or liana found in wet lowland
and lower montane forest,often in disturbed
areas, sea level to 900 m.
Representative
specimensexamined. BOLIVIA. Beni:
Serrania del Pilon Lajas, prov. Ballivian, Smith13222
(MO). La Paz: 27.8 km N of Caranavi, 15?33'S,
Cashu uplands, 11?45'S, 71?0'W, Nuinez 6087 (F, MO,
NY).
The species may be distinguished from the
closely related and vegetativelysimilarspecies,
P. loretensis,
by its globose stylarcolumn (typically 2-3 mm in diam.), capsule with rounded
tubercle on each carpel lobe, and anthers that
are typically30-50 in number.
Plukenetiabuchtienii
is treated here as a synPlukenetiabrachybotrya
onym of P. brachybotrya.
FIG. 10.
Plukenetiasupraglandulosa. A. Habit. B. Adaxial leaf base showing pair ofbasilaminarglands. C.
Abaxial leaf surfaceshowing scatteredlaminar glands. D. Leaf axil with young inflorescenceshowing basal
positionof pistillateflower. E. Terminalsectionof inflorescenceshowing staminateflowerbuds in condensed
cymes. F. Staminateflowerbud beginning to open. G. Staminateflower. H. Pistillateflower. I. Immature
capsule. J.Seed, showing dorsal, ventral,and lateral views, respectively. (A-C, F, G, I based on Granville
3626, CAY, US; D, E, H based on Cowan 38204,US; Jbased on Granvilleet al. 10783,CAY).
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584
SYSTEMATIC BOTANY
[Volume 18
has been considered a very poorly known spe- both present on specimens of one collection
cies with descriptions based primarily on (Gentry43598), suggestingthatthe fleshyfruits
Mueller's (1866) partlyincorrectdescriptionin are abnormal.
Flora Brasiliensis.Although Mueller originally
described the species as having about 30 an- 2. Plukenetia lehmanniana (Pax & K. Hoffm.)
thers,as distinctfromP. penninervia
with 12-15
Huft & L. J.Gillespie, comb. nov.-Eleutheanthers,the following year Mueller (1866) rerostigmalehmannianumPax & K. Hoffm.,
versed the numbers and described P. brachyboPflanzenr. IV.147.IX.(Heft 68): 11. 1919.tryaas having 12-15, and the latterwith 20-30
TYPE: Colombia, Narifio, Ricaurte, 1000anthers.This mistakepersisteduntilthe present
1400 m, Lehmann5158 (holotype: B, dewith anther number always given incorrectly
stroyed;lectotype,here designated: K!; isoas 12-15 forP. brachybotrya
(e.g., MacBride 1951;
lectotype:K!). Fig. 1A.
Pax and Hoffmann1919). The only other char- PlukenetiachapoensisCroizat, Caldesia 2: 431.
actersgiven by MacBride to distinguishthe spe1944.-TYPE: Colombia, Boyaca, region of
cies fromP. buchtienii
were differencesin leaf
Mt. Chapon, 3600 ft,Lawrance276 (holoshape and number of secondary veins, neither
type: A!; isotypes: BM! K! MO! NY! US!).
of which are valid when the full range of variDistribution
and habitat. Known only from
ation is examined.
Colombia and Ecuador, where it appears to be
Both P. buchtieniiand P. loretensis(Pax and
restrictedto the area west of the Andean divide
Hoffmann1919), and, subsequently,P. brachyin Ecuador and to the Pacific coast region and
botrya(MacBride 1951) were segregatedas a dismountain valleys of southern and central Cotinctgenus, Apodandra,based on the androecilombia. A robust, somewhat woody, twining
um of sessile anthers on a globose receptacle.
vine found in seasonal to wet, lowland to monHowever, both of these character states are
tane forest,150-2100 m.
shared with other species of Plukenetia.Plukenetiaserrataalso has all antherssessile on a gloRepresentative
specimens
examined. COLOMBIA. Calbose receptacle,while the remaining members das: Pacora,Daniel4943(US). El Valle: hoyadel Rio
of species group 2 (P. penninervia
species com- Cali, vertienteizquierdadel Rio Pichinde,El Cairo,
plex and P. verrucosa)have most antherssessile Cuatrecasas21971 (US).
ECUADOR. Carchi:vicinity of Chical, W of Malon a globose receptacle.
& Shupp26396
donadoon trailtoPenasBlancas,Gentry
Several collections, particularly the few
(DAV, MO). Esmeraldas:roadfromLitato San Loknown from central and eastern Amazonian renzo,19 km N of Lita,Acevedo& Daly 1658 (NY),
Brazil, appear to diverge fromthose of typical 1691(US). Guayas:Teresita,
3 kmW ofBucay,HitchP. brachybotrya;
these aberrant collections are cock 20490 (NY, US). Los Rios: Rio Palenque Bioincluded here, but additional collections may logicalStation,km56 Quevedo-Santo
Domingo,Dodshow that they representdistinctspecies. The son5771(MO,US); RioPalenqueFieldStation,
halfway
single collection (Lowrieet al. 30) seen fromthe betweenQuevedo and SantoDomingode los Colo9701(DAV, MO). Pichincha:Reserva
Brazilian stateof Amazonas has an androecium rados, Gentry
Quitoroad,10 kmN
of ca. 20 anthers,a depressed globose stylarcol- ForestalENDESA,Quito-Puerto
of
main
km
road,
113,
Acevedo
etal.
00?05'N,
79?02'W,
umn distinctlywider than long (ca. 1 x 1.8
1658(US), 1694(US); 20 kmW ofSantoDomingode
mm), and a narrowersubcoriaceous leaf blade.
los Colorados,Cazalet& Pennington
5089 (DAV,K,NY,
Collections fromPar'a and Mato Grosso, Brazil US).
(Plowmanet al. 8741,Sperlinet al. 5837, and Thomas et al. 4732) have distinctlylarger obovate
Pax and Hoffmanndistinguishedtheirmonostylarcolumns (3.5-4.5 mm long). Several col- typic genus Eleutherostigma
fromPlukenetiaand
lections fromPeru (e.g., Croat20058 and David- related genera based on the stylesbeing shortly
son 5310) have smaller stylarcolumns (1-2 mm connate ratherthan entirelyor mostlyconnate.
long) and longer inflorescencesand in some Styles are connate for one-thirdto one-half of
cases fewer anthers (ca. 25). A number of oth- theirlength into a cylindricalcolumn with the
erwise typical collections fromMadre de Dios, four style arms slender-cylindrical and rePeru appear to have fleshyfruitca. 2.5 cm in curved when mature (Fig. 1A). Among species
diameter,much largerthan the normal drycap- with stylespartlyfused into a cylindricalstylar
sules. Large fleshyfruitand typicalcapsules are column (species group 1), P. polyadeniahas styles
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1993]
GILLESPIE: PLUKENETIA
connate for approximatelytwo-thirdsof their
length (Fig. 1B), while P. stipellata(Fig. 9C) and
P. volubilis(Fig. 1C) have stylesconnate forseven-eighthsor more. The characterof stylemorphology used to defineEleutherostigma
is merely
a quantitativedifferencein degree of style fusion. Since there are no significantfloral,vegetative or palynological qualitative characters
distinguishing the genus from Plukenetia,
Eleutherostigma
is not worthyof generic recognition [Huft (unpubl. manuscript) previously
came to this conclusion]. Indeed, P. lehmanniana
fitseasily within the boundary of species group
1 and appears to be closely related to the other
three members. Croizat (1944) previously recognized thisalliance by placing his new species
P. chapoensis(=P. lehmanniana)in sect. Cylindrophora.
The species maybe distinguishedby the charactercombinationof mostlyfreestyles,ligulate
staminate disc segments, large fleshy subglobose fruit,and single conspicuous knob at the
petiole apex. Plukenetialehmannianais the only
species of Plukenetiafound west of the Andes
in Ecuador and southwestColombia; recordsof
P. volubilis
in thisarea (Dodson and Gentry,1978;
Dodson et al., 1985) appear to be based on misidentifiedspecimens of P. lehmanniana.
585
tryetal. 15598(DAV,F,MO); RioPutumayo,
atmouth
of Rio Zubineta,Klug 2178 (A, GH, K, MO, NY,
US). San Martin:SW of Aeropuertode Tocache
3686(GH, G, MO, NY, US).
Nuevo,Schunke
VENEZUELA. Amazonas: La Neblina Base Camp,
0050'N,66010'W,Leisner& Funk16526(DAV, MO);
CerroSipapo,BaseCamp,Maquire& Politi27371(NY,
US). Bolivar:Ikabaru,nearCampoDiamentifero
de
6649(NY).
Uaipare,Bernardi
Plukenetialoretensis
is unique among neotropical species in having stylesentirelyfused into
a slenderelongate-cylindrical
stylarcolumn,and
a distinctlyfour-hornedovary and maturecapsule. These characters,along with an anther
number of 15-25 and basilaminar glands frequently in multiple pairs on either side of the
midrib, distinguish it from the vegetatively
similar species, P. brachybotrya.
4.
MULTIGLANDULOSA Jabl., Mem.
New York Bot.Gard. 17: 143. fig.23. 1967.TYPE: Venezuela, Amazonas, Cerro Paru,
1800 m, Cowan& Wurdack31400 (holotype:
NY!; isotypes: NY!, US!).
PLUKENETIA
Distribution
and habitat. Known only froma
single collectionon CerroPariuwhere it is found
in montaneforeston talus at base of escarpment
at 1800 m.
The species may be distinguished from the
3. PLUKENETIA
LORETENSIS
Ule, Verh. Bot. Verelated species, P. penninervia
closely
and P. sureins. Prov. Brandenburg 81. 1980.-Apoits
praglandulosa,
by
persistent
yellow-hirsute
dandra loretensis(Ule) Pax & K. Hoffm.,
indumentumand multiglandularleafbase comPflanzenr.IV.147.IX.(Heft 68): 21. 1919.ofthreeto fivepairsofbasilaminarglands.
posed
TYPE: Peru, Loreto, Iquitos, Apr 1903, Ule
The
androecium,
previously described inaccu6837 (isotype: G!, photo F 24572!).
rately,consists of an outer whorl of four staDistribution
and habitat. Found in the upper mens with shortfilamentsand an inner cluster
and middle Amazon Basin in Peru, Bolivia, and of eight to ten sessile anthers on a globose reBrazil (Amazonas, Mato Grosso,and Rondonia), ceptacle, very similar to the androecium of P.
and in southernVenezuela and southeastGuy- supraglandulosa
(Fig. lOG).
ana. A twining vine or slender liana in moist
to wet, lowland forest,sea level to 700 m.
5. PLUKENETIA PENNINERVIA Muell. Arg., LinRepresentative
specimensexamined. BOLIVIA Beni:
naea 34: 158. 1865.-TYPE: Venezuela, near
3 kmE ofPiberaltaon roadtoGuagaramerin,
11?00'S,
Biscaina, Fendler 2412 (holotype: G-DC!,
66?5'W,Solomon7972 (MO, NY)
photo F 7110!; isotypes: K!, GH!, MO!).
BRAZIL. Amazonas: Rod. Transamazonicaa 400 km
Plukenetiaangustifolia
Standley, Publ. Field Code Humaita, 7?15'S, 60?00'W,Cid 5983, INPA 127.449
lumbian Mus., Bot. Ser. 4: 314. 1864.-TYPE:
(F, NY); Mun. Sao Paulo de Olivenqa,BelemCreek,
Honduras, Atlantida Dept., Lancetilla Val8791(A,G,K,MO, NY,P, US). Mato Grosso:
Krukoff
ley, near Tela, Standley56708 (holotype: F!;
BR-174, Aripuana, Silva & Rosdrio 4795 (MO,
isotypes:A!, K!, US!).
PortoVelho-Cuiaba,
NY). Rond8nia:BR-364,
12?13'S,
60?61'W,Cid et al. 4359, INPA 120.732 (MO, NY).
GUYANA. Kanasenay, Guppy203, FD 7179 (NY).
PERU. Loreto: Rio Nanay, Purto Almendrez, Gen-
Distributionand habitat. Widespread from
southernMexico to Venezuela. A twining vine
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All use subject to JSTOR Terms and Conditions
586
SYSTEMATIC BOTANY
[Volume 18
of four (or sometimes five) stamens on short
filaments,ca. 0.3 mm long, and an inner cluster
of 10 to 12 sessile anthers on a globose recepexamined. BELIZE. Big
specimens
Representative
tacle. An annular disc, sometimes segmented,
Creek,Schipp156(BM,G, K, NY, UC, US).
Puntarenas:Rinc6nde Osa, Utley& is usually evident between the outer stamens
COSTA RICA.
and inner anthers.
Utley1215(F).
Collections from the departmentsof Choco
GUATEMALA. Izabal: vicinityof Quirigua, Steyermark23921(GH, US). Peten:Dos Lagunos,5 kmon and El Valle in Colombia (Cuatracasas 14006,
17472; Espina & Garcia 15333; Gentry& Renteria
IxcanrioRoad,Elias8361(MO, US).
Martinez- 23761, 24367; Gentryet al. 47799) are included
MEXICO. Oaxaca: Chiltepec and vicinity,
Calder6n
291(A,UC, US). Tabasco:La Palma,Balan- here but may representyet another species in
can, Matuda3288 (A, K, NY). Yucatan: Tuxpefia, the P. penninervia
species complex. They differ
Campeche,Lundell1368(US).
in havfromtypicalcollections of P. penninervia
NICARAGUA. Rio San Juan:3.5 kmNE ofBoca de
ing largerseeds, longer inflorescences,and lamSabalo, 11?3'N, 84?26'W,Moreno25463 (DAV, F,
inar glands usually present on the lower leaf
MO). Zelaya: road betweenWani and Siuna near
surface.
Leaves often have a somewhat orange
Pipoly4693(DAV, MO).
Rio Matis,13043'N,84049'W,
79045'W, or reddish brown undersurfaceat least when
PANAMA. Colon:SantaRitaRidge,9020'N,
dry,appearing quite distinctfromthe greyish
8461(MO).
McPherson
and
VENEZUELA. Bolivar:RioToro,betweenRioLa Re- or yellowish green leaves of P. penninervia
formaand PuertoRico,N of El Palmar,SteyermarkP. supraglandulosa.Collections from Pastaza
88148(F, GH, K, NY, P, U, US). Delta Amacuro:ca. province,Ecuador (Lugo4150,4219,4399),which
13 kmby road ESE of townof SierraImata,8023'N, superficiallyresemble P. loretensis,appear to
62023'W,Davidse & Gonzdlez16501 (DAV, MO, represent another taxon in the P. penninervia
1NY). DistritoFederal:along Rio Chichiriviche,
species complex.
Stey67014'30"W,
10032'30"N,
2 kmS ofChichiriviche,
Sierra
& Espinosa
112709(F,MO, NY). Falcon:
ermark
Muell. Arg. in Mart.,
POLYADENIA
3 kmE ofCerro 6. PLUKENETIA
de San Luis,Montanade Paraguariba,
334.
1874.-Elaeophora
polyFl.
11(2):
bras.
Los
Miranda:
MO).
617
Galicia,FloraFalc6n (DAV,
adenia(Muell. Arg.) Ducke, Arch. Jard.Bot.
Caracas-Cabo Codera, Aristeguieta4852 (NY,
Rio de Janeiro 5: 146. 1930.-TYPE: Peru,
US). Portuguesa:betweenLa Estacionand La Laguna,15-18kmNNW of Ospino,9025-27'N,69030"Maynas", Poeppig2385 (holotype: W, fragetal. 127019(F,MO,U). Zulia:entre
31'W,Steyermark
ment at F!, photo F32544!; isotype: G!; poslas Tres Marias(10025'N,70055'W)y Rio Chiquito,
sible isotype: NY!, labeled Poeppig 2585).
& Stoddart
8913(DAV, NY).
Bunting
Fig. 1B.
ElaeophoraabutifoliaDucke, Arch. Jard. Bot.
is the mostcommon and
Plukenetiapenninervia
Rio de Janeiro4: 112. fig. 9. 1925.-Plukewidespread species in the P. penninerviaspecies
netiaabutifolia
(Ducke) Pax & K. Hoffm.,Nat.
complex, which is characterizedby dimorphic
Pflanzenfam. ed. 2. 19c: 141. 1931.stamensand a shortstout-cylindricalstylarcolSYNTYPES:Brazil, Para, near Belem, Ducke,
umn. The species may be distinguished fromP.
HJBR17892(not seen); Xingu R., Kuhlmann,
multiglandulosaby its glabrescentleaves and sinHJBR 17895 (isosyntype: U!); Tajaparu R.,
basilaminar
glands
circular
of
usually
gle pair
Ducke,HJBR17893 (isosyntype:P!).
(rarelyseveral pairs in collectionsnorthofHon-
in disturbed areas of dry to wet, lowland to
montane forest,sea level to 1500 m.
by the
duras only), and fromP. supraglandulosa
absence of scatteredlaminar glands (occasionally presenton the lower leaf surfacebut never
on the upper surfaceapartfromthe basilaminar
gland pair), distinctlyserrulateand oftenglandular leaf blade margin, and shorterinflorescences. The androecium has been incompletely
or incorrectlydescribed in the past, mostlikely
because of its minute size (less than 1 mm in
diameter). Very similar to that of P. supraglandulosa (Fig. lOG), it consists of an outer whorl
and habitat. Widespread in the
Distribution
Guianas and easternVenezuela, and in the Amazon Basin of Ecuador, northernPeru, and Brazil (Amapa, Amazonas, Par'a). A canopy liana
found in moist to wet, lowland or lower montane forest,sea level to 1000 m.
examined.BRAZIL.Amaspecimens
Representative
pi: Igarape do Navio-Mazagao,Rabeloet al. 2140
(NY). Amazonas:Mun. Sao Paulo de Olivenqa,ba8821(F, G, K, MO, NY, P,
sin ofcreekBelem,Krukoff
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All use subject to JSTOR Terms and Conditions
1993]
GILLESPIE: PLUKENETIA
US); Mun. Eirunepe,Igarape Folguedo Assahtuba,Froes
12098 (A, F, MO, NY, US). Para: Macapa, Rio Matapy,Ducke 20619 (K, P, US).
ECUADOR.
Napo: Rio Aguarico,upriverfrombridge
at Aguarico, Gentry9717 (F, MO, NY). Pastaza: Comuna Shuar Amuntay, 2031'S, 76048'W, Ceron et al.
4319 (DAV, MO).
FRENCH GUIANA.
Sauil,Mori& Gracie18668(K, US).
Camaria Falls, Cuyuni R., Fanshawe3383
GUYANA.
= FD 6947 (FDG, K, NY); Mt. Ayanganna, NE side,
below talus along base, Tilletet al. 44950 (MO, NY,
US).
PERU.
Amazonas: 1 km de La Poza, Rio Santiago,
Huashikat172 (MO). Loreto: Rio Napo near Mazan,
Mexia 6470 (F, G, GH, K, NY, US).
587
cies, up to ten in P. serrata)or occasionally staminate if the pistillate flower(s) is lacking.
A single geographically distant collection
(Ventura522) fromPuebla, Mexico superficially
resemblesP. polyadenia
in leaf shape, single knob
at the petiole apex, and apparently unisexual
pistillate inflorescences, but has a pistillate
flowervery similar to that of P. stipellata.
7. Plukenetia serrata (Vell. Conc.) L. J.Gillespie, comb. nov.-Vigia serrataVell. Conc.,
Fl. Flum. 9: t. 127. 1832.-TYPE: Illustration
t. 127 in Vell. Conc., Fl. Flum. 9. 1832.
Fragariopsis
scandensSt. Hil., Leqons bot. 426.
SURINAM.
Jodensavanne-Mapanekreek area,
1840.-Plukenetiascandens(St. Hil.) Pax, Nat.
Schultz7284 (MO, U, US).
Pflanzenfam.III. 3(5): 67. 1890.-TYPE: BraBolivar: Mun.Raul Leoni,85 kmSSE
VENEZUELA.
zil, St. Hilaire72 (holotype: P!; isotype: P!).
of EntreRios, ca. 5018'N,63059'W,Aymard& Fernandez
7272 (F, MO). Delta Amacuro: Cerro Moco Hierro,
[No collection number was cited by St. HiE de Rio Grande, ENE de El Palmar, Steyermark
93082
laire. Of his 2 collections, 72 and 95, at P,
(F, G, K, NY; US).
a specimen of collection 72 has an original
label quoting the protologue.]
The species may be distinguishedvegetative- Accia scandensSt. Hil., Leqons bot. 499. 1840.ly by its broadly elliptic,distinctlytriplinerved
TYPE: not designated, probably based on
leaf blade thatis rounded or obtuse at the base.
one of St. Hilaire's collections at P [or perFruits are large, subglobose, fleshy,and indehaps a nomenclatural synonymof F. scanhiscent (only P. lehmanniana has similar fruit).
dens].
Ducke (1925, 1930) considered the species to Botryanthe
discolorKlotzsch, Arch. Naturgesch.
7: 191, 204. tab. 9b. 1841.-Fragariopsisdisbe dioecious with dimorphic unisexual inflocolor(Klotzsch) Baill., Etude Euphorb. 498.
rescences,and noted thatstaminateindividuals
1858.-TYPE: Brazil,Sellows.n.(holotype: B,
are much more common than pistillateindividuals. Pistillateinflorescencesare 1-3 (7) cm long,
destroyed; lectotype,here designated: P!,
fragment at F!). [Klotzsch's name "Bowith flowers solitary at each node, while statryantheconcolor" given in the same pubminate inflorescencesare much longer, 10-33
lication was not validly published.]
cm long, with flowers in one-sided (or somepolyandrus
Baill.,Etude Euphorb.498.
times dichasial) cymes. Of the 35 fertilecollec- Fragariopsis
P1. 13, figs. 29-36. 1858.-TYPE: not desigtions examined, about two-thirdsare staminate
nated, possibly Guillemin798 from Rio de
and one-fifthare in fruit.Only two have pisJaneiro,Brazil (holotype: P!; isotypes: G!,
tillate inflorescences as described by Ducke
P!).
(Black 52-14137; Ducke, HJBR 17893), while three
Muell. Arg. in Martius,Fl.
have bisexual inflorescencesthat are morpho- Fragariopsis
warmingii
bras. 11: 338. tab. 52. 1866.-Plukenetia warlogically similarto staminateinflorescencesbut
mingii(Muell. Arg.) Pax, Nat. Pflanzenfam.
include eitherpistillateflowers(Ce'ronet al. 4319,
III. 3(5): 67. 1890.-TYPE: Brazil, Minas GeNeill et al. 8370) or immaturefruit(Schultz 7284)
at the basalmost nodes. This evidence suggests
rais, near Lagos Santa, Warmings.n. (holothat individuals may be either dioecious or
type: P!, fragmentat F!; probable isotype:
monoecious,or perhaps thatall are monoecious,
F!).
but are frequentlytemporally dioecious. The
dioecious condition and unisexual pistillateinDistribution
and habitat. Restrictedto southflorescencesare unknown elsewhere in the ge- east Brazil (Bahia, EspiritoSanto, Minas Gerais,
nus. Inflorescencesof all other species are typ- Rio de Janeiro,and Sao Paulo). A vine or liana
ically bisexual with pistillateflowerssolitaryat of wet forest,oftenfound at the forestedge, sea
the basalmost nodes (usually one in most spe- level to 1000 m.
This content downloaded from 210.72.93.209 on Tue, 10 Feb 2015 22:33:30 PM
All use subject to JSTOR Terms and Conditions
588
SYSTEMATIC BOTANY
Representative
specimens
examined.BRAZIL. Bahia:
RodaviaBA-265,ca. 25 kmNW de Caatiba,Morietal.
9409(DAV,NY). EspiritoSanto/Riode Janeiro:
Itabapoana,Samp1001,HJBR25979(US). Rio de Janeiro:NouvelleFribourg(=Nova Friburgo),
Claussen
83 (G). Sao Paulo: Sao Paulo,groundsof theInstitutoBotanico,Davidse10480(NY); Pirajussara,
Gehrt
12559 (NY).
Although generally treated as a monotypic
genus, Fragariopsis(=Vigia), based on androecial
morphology and fleshy fruit, there appears to
be no reason to exclude the species from Plukenetia since both characters are found within
the genus. Plukenetia polyadenia and P. lehmanniana also have fleshy fruit. The androecium is
unique in that the sessile anthers are widely
scattered on a greatly enlarged globose receptacle, but this state may be considered derived
from the more common similar state of sessile
anthers densely crowded on a smaller globose
receptacle (as in P. brachybotryaand P. loretensis).
All anthers sessile, styles entirely connate into
an obovoid column, pollen with a reticulate tectum (Gillespie, in press), and pinnately veined
leaves places it in species group 2. In addition
to the unusual androecium and fleshy fruit, P.
serrata may be distinguished by the stipellate
petiole apex and inflorescences with up to 10
pistillate flowers.
As Webster (in press) pointed out, the name
Vigia serrata has precedence over Fragariopsis
scandens, hence a new combination in Plukenetia
must be made.
8. Plukenetia stipellata L. J. Gillespie, sp.
nov.-TYPE: Costa Rica, Heredia, Finca la
Selva, OTS Field Station on the Rio Puerto
Viejo, just E of junction with the Rio Sarapiqui, 13 Sep 1983, Gillespie 413 (holotype:
US!; isotypes: CR!, MO!; FAA preserved material at US). Figs. 3, 4-6, 9.
A P. volubili L. petiolo apice stipellato, stylis
brevioribus (8-10 mm), filamentis gracilibus
longioribus (1-1.2 mm) at sepalis florum staminatorum plerumque 5 differt.
Monoecious twining vine; branches slender,
branchlets glabrescent or sparsely puberulous.
Leaves alternate, simple; stipules small, deciduous; petiole 2.5-7 cm long, sparsely puberulous; blade chartaceous, triangular-ovate or
broadly triangular-ovate, 8-15 cm long, (4) 613 cm wide, slender-acuminate at apex with
[Volume 18
acumen 0.8-2.0 cm long, cordate at base with
sinus 0.6-1.5 (2.5) cm deep, serrulate,glabrescent or sometimes sparsely puberulous below,
major veins puberulous below; venation palmate, primaryveins 3, secondary veins 2-3 on
each side of central primaryvein and 2-4 on
lower side of lateral primaryveins, brochidodromous, tertiaryveins percurrent;quaternary
veins reticulate;basilaminarglands 2, narrowly
transverse-ellipticto obovate, 0.4-1.8 mm long,
1-3.2 mm wide, marginal, adaxial; laminar
glands absent; pair of stipels at petiole apex
between basilaminarglands,stipelsconical,0.31.0 mm long, pointed or rounded at apex, rarely
with minute glandular knob between. Inflorescence slender racemose, (2) 6-10 (23 in fruit)
cm long, bisexual, terminal on short shoots
bearing 1-2 leaves or rarely axillary,axes puberulous, pistillate flower single at basal-most
node, staminate flowers numerous in condensed cymes at distal nodes, cyme axes to 1
mmlong; bractstriangular,0.6-1 (1.7) mm long.
Staminatepedicel 0.3-0.8 mmlong, puberulous;
bud broadly elliptic, 1.8-2.5 mm long, obtuse
at apex; sepals (4) 5, narrowly ovate, 2-3 mm
long, ca. 1 (1.5) mm wide, sparsely puberulous
especially at base, valvate; corolla and disc absent; stamens25-40 on convex receptacleca. 0.5
mm in diameter, filamentsslender, 1-1.5 mm
long; pollen oblate-spheroidal,53-60 ,umin polar diameter,58-69 ,umin equatorial diameter,
tricolpate,amb obtuse-triangular,colpus broad
with uneven margins,tectumfoveolate,surface
smooth to minutelyscabrate. Pistillate pedicel
2.5-5 mm long, puberulous; sepals 4, triangular
or narrowlytriangular,1-1.5 mm long, acute or
attenuateat apex, glabrescent;corolla and disc
absent; ovary4-locular,1-3 mm long, 2.2-7 mm
wide, puberulous, 4-winged, wings rounded,
each locule uniovular; stylesconnate into a cylindrical column, 6-11 x 0.8-2.0 mm, sparsely
puberulous, the 4 freestylearms 1-2 mm long,
divergent and appearing cross-shaped when
mature. Fruitingpedicel 1.5-2.5 cm long; capsule 4-lobed, 1.2-2.0 x 2.3-3.2 cm, glabrous,
each carpel lobe carinatewith centralrounded
horn.Seeds lenticular,laterallycompressed,1.11.4 cm long, 0.3-0.5 cm wide, 0.9-1.4 cm thick,
angular-circularin outline, brown with irregular dark brown patches, ecarunculate.
Distributionand habitat. Found from Veracruz and Chiapas, Mexico to Panama and pos-
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GILLESPIE: PLUKENETIA
1993]
589
flowershaving stamenswith longer slender filaments and usually five sepals (Fig. 9D). The
two stipel-likeappendages at the petiole apex
are conical in shape, either pointed or someRepresentativespecimensexamined. COSTA RICA.
what rounded at the apex and at least sometimes
Alajuela: N of San Ram6n, kms 15-35 of road to La
appear to be glandular.
Tigre and Fortuna,Tayloretal. 4193 (MO). Alajuela/
Several collections from north central VenGuanacaste: slopes of Miravalles, above Bijagua, Gomez et al. 19184 (MO). Cartago: Atirro,Turrialba, ezuela appear to be morphologically intermeZamora 1306 (F, MO). Heredia: 1 km W of Puerto diate between this species and P. volubilisand
3611 (F, MO, NY). Limon: will be discussed in greater detail under the
Viejo de Sarapiqui,Jiminez
29 air kmW ofTortuguero,10?30'N,83?47'W,Davidson latterspecies. Two fruitingcollections fromCoet al. 6734 (F, MO, NY, US); S end of Lomas de Sierpe, lombia, Pennell4584 from the state of Bolivar
NE of terminus of road fromVilla Franca, 10?19'N, and Duke 15375 fromChoc6, have leaves with
83?34'W, Grayumet al. 3511 (DAV, F, MO). Puntapaired stipels at the petiole apex and may berenas: Reserva Forestal Golfo Dolce, Osa Peninsula,
long to this species. However, because of the
Rancho Quemado, ca. 15 km W of Rinc6n, Hammelet
al. 16933 (MO); Rinc6n de Osa, slopes above airport, lack of flowersit is not possible to determine
whetherthese collections representthis species
Liesner2068 (MO, US).
or are more similar to the intermediatepopuAlta Verapaz: Secanquim, Cook &
GUATEMALA.
Griggs265 (US); between Secoyocte and Sepacuite, lation fromnorth central Venezuela.
sibly northern Colombia (see discussion below).
A twining vine of disturbed sites in moist to
wet forest, sea level to 1200 m.
Pittier324 (US), 342 (BM, NY, US).
MEXICO.Chiapas:ca. 1 miNE ofRuinasde Palen- 9. Plukenetia supraglandulosa L. J. Gillespie,
78420(DAV). Veracruz:Estacionde
que,Armbruster
sp. nov.-TYPE: FrenchGuiana, SommetTaSan AndresTuxtla,CedBiologiaTropicalLosTuxtlas,
bulaire, zone centrale,versant occidental,
illo & Calzada 56, MEXU 41650 (BM, F, K, MO); La
ca. 40 km SE de Saul, 27 Aug 1980,Granville
Palma, Sontecomapa, Los Tuxtlas,Sousa 2661 (US); Hi3626 (holotype: US!; isotypes: CAY!, U!).
dalgotitlan,17?18'N,94?38'W,Vazquezetal. 1253,MEXU
Figs. 2, 7-8, 10.
44836 (BM, MO).
Matagalpa: Cerro Musuin, 8 km de la
NICARAGUA.
Poblacion Wanawas, Araquistain& Moreno2360 (DAV,
MO). Rio San Juan: 1 km E of Rio Sabalos, 11?2'N
84?27'W,Moreno23204 (MO). Zelaya: Monkey Point,
11036'N, 83039'W,Moreno & Sandino 12112 (DAV, F,
MO); Rio Punta Gorda, 11032'N, 84005'W,Moreno &
Sandino 13085 (K, MO); ca. 3-4 km SW of Colonia
Naciones Unidas on road to Colonia Nuevo Leon, ca.
11042'N,84019-20'W,Stevenset al. 5088 (F, MO).
PANAMA.
Cocle: ca. 12 mi from Llano Grande,
8047'N,80028'W,Churchillet al. 4149 (MO). Col6n: S
of Porto Bello, along streamrunning into Rio Buena
Ventura,Foster2060 (F, GH, MO); along Rio Guanche,
8686 (MO). Darien:
6 km S ofPortobelo,Nee & Gentry
Cerro Pirre, valley between Pirre and next most
southerlypeak, Folsomet al. 4426 (F, MO). Los Santos: between Los Santos and Guanare, Woodsonet al.
1201 (A, F, MO, NY). Panama: 4-5 hrs walk upriver
fromTorti Arriba,Folsomet al. 6845 (F, MO).
P. penninerviae
Muell. Arg. et P. multiglandulosae Jabl.affinis,a quibus differtinflorescentia
longissima et foliis supraglandulosis glandibus
numerosis dispersis.
Monoecious liana; branches slender, branchlets puberulous. Leaves alternate,simple; stipules small, deciduous; petiole 0.5-2.0 cm long,
puberulous; blade chartaceous,ellipticor ovateelliptic, 7-13 x 3-8 cm, slender-acuminate at
apex with acumen 0.5-1 cm long, acute, cuneate
or sometimesobtuse at base, serrulate,glabrescent except basal part of midrib puberulous,
majorveins and midribsometimessparselypuberulous below; venation pinnate, secondary
veins 6-10 on each side of primaryvein, distinctlybrochidodromous,tertiaryveins percurrent, quaternary veins reticulate; basilaminar
glands 2-4 (6), circular or elliptic, 0.6-1.9 mm
This species, widespread and moderately long,0.5-0.7 mmwide, adaxial,arrangedin pairs
common in Mesoamerica, has previously been on either side of midrib; laminar glands cira species restrictedto cular,0.4-0.7 mm in diameter,on both surfaces,
confused with P. volubilis,
South America and the Lesser Antilles. Al- 10-20 mostly 1-5 mm from margin below, 8though similarto P. volubilisin habit and overall 18 scatteredbetween midriband marginabove.
appearance, P. stipellatadiffersin having a pair Inflorescence slender racemose, 2-8 cm long,
of minute stipels at the petiole apex (Fig. 9B), bisexual, axillary, axes puberulous; pistillate
a shorterstylarcolumn (Fig. 9C), and staminate flowersingle at base, staminateflowersnumer-
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590
SYSTEMATIC BOTANY
ous in condensed cymes above; bracts triangular, ca. 1 mm long. Staminatepedicel 3.5-4.5
cm long, densely puberulous; bud subglobose,
ca. 1 mmlong or less, rounded or broadlyobtuse
at apex; sepals 4, elliptic, 1-1.3 mm long, acute
and thickenedat apex, reflexed;corolla absent;
disc 4-lobed, located between outer stamen
whorl and inner stamen cluster;androecium of
6-9 sessile antherson central,shortlystipitate,
globose receptacle and 4 outer stamens with
filaments0.3-0.4 mm long arising fromunder
disc and archingupwards; pollen suboblate,3133 ,umin polar diam., 39-45 ,umin equatorial
diam., tricolpate,amb obtuse-triangular,colpus
broad with uneven margins,tectumreticulate,
muri crenate. Pistillate pedicel 6-12 mm long,
puberulous; sepals 4, triangular,ca. 1 mm long,
puberulous; corolla and disc absent; ovary
4-locular,each locule uniovular, ca. 1 mm long,
ca. 2 mm wide, densely puberulous, 4-winged,
wings rounded; styles entirelyconnate into a
stout-cylindricalcolumn, ca. 1.6 x 0.6 mm,
sparselypuberulous. Fruitingpedicel ca. 25 mm
long; capsule 4-lobed, ca. 7 x 15 mm, sparsely
puberulous, pendent, each carpel lobe with
conical tubercleca. 1 mm high. Seeds lenticular,
laterallycompressed,ca. 7 mm long, 4.6-4.8 mm
wide, circularto broadly elliptic in outline,reddish brown with raised reticulatepattern,ecarunculate.
[Volume 18
P. multiglandulosa
by the scatteredlaminarglands
on the upper leaf surface, puberulous indumentum,and longerinflorescences.While many
species of Plukenetiahave scattered laminar
glands on the lower leaf surface mostly near
the margin and all have one to several pairs of
basilaminar glands on the upper surface,P. suis the only species to have laminar
praglandulosa
glands scatteredon the upper surface.
10. PLUKENETIA
VERRUCOSA
Smith, Nova Acta
Regiae Soc. Sci. Upsal. 6: 4. 1799.-TYPE:
Surinam, Herb. Linnaeus(LINN, not seen,
microficheIDC 5073.1489.2!). Fig. 1D.
Plukenetiaintegrifolia
Vahl, Eclog. Amer. 3: 43.
1807.-TYPE: Guyana, "Demerari", vonRohr
s.n. (holotype: C, not seen, photo A!).
Distribution
and habitat. Restrictedto and apparently rare in Trinidad, the Guianas, and
northernAmazonian Brazil (Amapa, northeast
Amazonas, Para, and Roraima). A slender twining vine found mostly in disturbed areas or
forestedge of lowland wet forest.
Representative
specimens
examined. BRAZIL. Amapa: on lower slopes of Mt. Tipac, 3?36'N, 51?19'W,
Irwin48796 (NY, US). Amazonas: Janauari,Rio Negro opposite Manaus, Pranceet al. 11255(DAV). Para:
right branch of Rio Moju, Burchell9369 (K, NY,
P). Roraima: Rio Branco, Kuhlmann252, HJBR2996
(DAV).
FRENCHGUIANA. St. Jean du Maroni, Benoist893
Distribution
and habitat. Endemic to French
Guiana and the Brazilian stateofAmap'a.A slen- (P); Karouany (also Maroni), Sagot 22 (BM, G-BOIS,
der liana known from submontane forest on G-DC, K, MPU, P, U; includes several collections).
GUYANA. Camp Macaw Falls,Waini R.,Beckett
8455
lateriticplateau summitsat 500-800 m in French
(BRG, NY).
Guiana and along roadside throughheavily forSURINAM. Nickerie R., near Awawara, BW 893 (K,
ested hills at 70-300 m in Amap'a. Flowers re- U); Voltzberg,Pulle 221 (U).
portedin January,Augustand November; fruits
TRINIDAD. Catham, Cedras, Broadway,TRIN 7236
reportedin Januaryand August.
(NY).
Additional
specimens
examined.BRAZIL. Amapa:
Serrado Navio,alongmainroadbetweenIgapo Baixinhoand roadto PortoTerezinha,70-300m,Cowan
38204(K, NY, US).
FRENCH GUIANA: Massif du Decou Deou, E summit,
Granville
5315 (U); MontAtachiBacca,S of summit
plateau,12 kmSE ofGobayaSoula,3?33'N,53055'W,
Granville
etal. 10783(CAY,US).
Despite its cordiformor ovate, triplinerved
leaves, this species appears to be most closely
related to the neotropical species having elliptic,pinnatelyveined leaves. The species fitswell
into species group 2, being characterizedby an
androecium of dimorphic stamens, virtually
identical to that of the P. penninerviaspecies
complex (Fig. lOG), stylesthatare entirelyfused
The species belongs to the P. penninervia
spe- into an obovate or subglobose column (Fig. 1D)
cies complex and may be distinguishedfromP. similar to that of P. brachybotrya,
a small fourpenninervia
by the leaves having numerous lam- lobed capsular fruit,and pollen with a reticulate
inar glands scatteredon both upper and lower tectum(Gillespie, in press). Plukenetiaverrucosa
leaf surfaces(Fig. 2), the minutelyserrulateleaf may be distinguished vegetatively from the
margin,and the longer inflorescences,and from other species having triplinervedor palmately
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1993]
GILLESPIE: PLUKENETIA
veined leaves (i.e., species group 1) by its less
robusthabit and thinner,narrowerleaves with
a stipellate petiole apex.
11. PLUKENETIA
VOLUBILIS
L., Sp. pl. 1192.1753.TYPE:West Indies, Illustrationt. 13 (lower
half) in Plumier, Nov. Pl. Amer. 47. 1703.
Figs. 1C, 9E.
Plukenetia
peruvianaMuell. Arg.,Linnaea 34: 157.
1865.-SYNTYPES:Peru, Herb.Pavon (G-DC!,
photo F 7111!, G!); Peru, "prov. Maynas",
Poeppig2110 (not seen).
PlukenetiamacrostylaUle, Verh. Bot. Vereins
Prov. Brandenburg 50: 80. 1908.-TYPE:
Brazil, Amazonas, Rio Jurua,near Jaburiu,
Ule 5864 (isotype: G!, photo F 24573!, fragment at F!).
?Fragariopsis
paxiiPittier,J.Wash. Acad. Sci. 19:
351. 1929.-TYPE: Venezuela, Federal District,Loma de En Medio, Valley of Puerto
la Cruz, 1000 m, Pittier8109 (isotypes: GH!,
US!) [appears to be intermediatebetween
P. volubilisand P. stipellata;see discussion
below].
Manu, Vargas11616 (US).
591
San Martin: JuinJui,alto
Rio Huallaga,Klug3901(F, K, MO, US).
ST. VINCENT. In forest,
Smith& Smith422 (BM,GH,
NY, US).
SURINAM. ValleyofWestRiver,2-5 kmSW ofJuliana Top,Irwinetal. 54897(F, GH, MO, NY, U, US).
VENEZUELA. Amazonas:0-1 kmS ofRioMawarinuma,3 kmbyairE ofCerrode la NeblinaBaseCamp,
0?50'N,66?09'W,Liesner
16197(DAV, MO, NY).
This species may be distinguished fromthe
closely related species, P. stipellata,by a single
glandular knob at the petiole apex, longer stylar
column (Fig. 1C), and staminate flowers with
shortconical filamentsand foursepals (Fig. 9E).
The stylarcolumn is the longest of any species
of Plukenetiaand may attain a length of 3 cm.
An isolated population (represented by the
collections,Agostini& Farinas54 and Steyermark
89966) in the coastal mountainsof northcentral
Venezuela (Estado Miranda and Distrito Federal) appears to be morphologically intermediate between P. volubilisand P. stipellata.AlthoughidenticaltoP. volubilis
in staminateflower
morphology including sepal number, stamen
number,and filamentsize and shape, these colDistribution.Found in the Lesser Antilles, lections have leaves with paired stipels at the
Surinam, and along the northernand western petiole apex typical of P. stipellata.Stylar coledge of the Amazon basin in Venezuela (Ama- umn length is intermediatebetween the two
zonas), Colombia (Meta), Ecuador, Peru, Boliv- species (10-15 mm long). The type collection of
ia, and Brazil (western Amazonas). A twining Fragariopsis
paxii,Pittier8109, appears to belong
vine found in disturbedareas or forestedge of to this population. Pittierconsidered his new
moist or wet lowland forestup to 900 m.
species to belong "without any possible doubt"
to Fragariopsis,
presumably on the basis of the
Representative
specimens
examined. BOLIVIA. La Paz:
paired stipels (unknown in Plukenetiaat that
Basin of Rio Bopi, San Bartolome,Krukoff
10082 (A, F, time) and the stamenswhich he misinterpreted
G, NY, US). Santa Cruz: 4 km (by air) WSW of Buena as lacking filaments.The
style was incorrectly
Vista, 17?28'S,63?42'W,Nee 35694 (MO, NY).
described as 5-6 mm in length, much shorter
BRAZIL. Amazonas: Mun. Eirunepe, basin of the
than the stylarcolumns of eitherP. stipellataor
upper Jarua,Froes44 (US), 48 (US). Para: vicinityof
P.
volubilis.Pittiermost likely examined speciPara, Baker153 (BM, G).
mens having immatureor incomplete flowers
COLOMBIA. Meta: Cano Yerli, entre el rio Guelar y
such as the specimen at US which has only a
el canioGuapayita, Cordillera La Macarena, Idrobo&
Schultes758A (F, GH, MO, NY, US). Putumayo: single pistillateflowerwith a stylarcolumn broPuerto Limon, cultivated,Plowman2191 (ECON).
ken near itsbase. However, the specimen at GH
DOMINICA. Rosehill, Eggers580 (G, GH).
includes pistillate flowerswith stylarcolumns
ECUADOR. Napo: 4 km S de Puerto Napo en el Rio 14-15 mm long, congruent with other collecNapo, Dodson et al. 14926 (K, MO, NY); Limoncocha, tions of this
morphologicallyintermediatepopZarucchi2369 (GH, MO, US), 2378 (DAV, GH, US).
ulation.
GRENADA. Concord R., Eggers6470 (P, US).
The greatestdegree of variation in P. volubilis
MARTINIQUE. Near Lamentin, Hahn 1278 (P).
is found in collections fromthe eastern slopes
PERU. Amazonas: Lugar Huampami, Ancuash 248
(GH, MO). Cuzco: Atalaya,Kosaipata, prov. Pancas- of the Andes bordering the Amazon basin in
tambo,Vargas13994 (US). Junin:Satipo, Woytkowski Peru. While the majorityof collections may be
5891 (MO, US). Loreto: Mishuyacu,near Iquitos,Klug considered as typical of the species, there are a
1438(F, US). Madre de Dios: Piaipiai, Defensa, Prov. numberof morphologicallyunusual ones. Two
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592
SYSTEMATIC BOTANY
[Volume 18
collections (Klug 3846, 3901) fromSan Martin, BENTHAM, G. 1880. Euphorbiaceae. Pp. 239-340 in
GeneraPlantarumad ExemplariaImprimisin HerPeru have flowerswith somewhat longer (0.8
bariisKewensibusServata Definita,vol. 3, eds. G.
mm) slender-conical filaments;Klug 3846 also
Benthamand J.D. Hooker. London: Lovell Reeve
has flowerswith stylearms 3-4 mm long rather
and Co.
than the typical length of 1-2 mm. Collections
CROIZAT, L. 1944. Euphorbiaceae Colombianae, III:
from Cuzco, Junin,and Pasco, Peru at 1600PlukenetiaLinnaeus. Caldesia 11: 431-432.
2100 m, elevations much higher than either P. DODSON, C. H. and A. H. GENTRY. 1978. Flora of the
the only othermembers
volubilis
or P. polyadenia,
Rio Palenque Science Center. Selbyana 4: 1-628.
of species group 1 known fromPeru, may rep, and F. M. VALVERDE. 1985. La Flora
f
resent a distinctspecies (Gentry& Smith35906,
de Jauneche.Quito: Banco Central de Ecuador.
Solomon3166, Woytkowski
6670). While labeled DUCKE, A. 1925. Plantes nouvelles ou peu connues
de la region amazonienne III, Euphorbiaceae. ArP. volubilis,
the collectionsmostclosely resemble
chivos do JardimBotanico do Rio de Janeiro4:
P. stipellatain floralmorphologyand P. lehman107-115.
nianain aspect including the more robustdens1930. Plantes nouvelles ou peu connues de
er inflorescenceswith larger flowers.
la region amazonienne IV, Euphorbiaceae. ArSeveral collections have unusually large capchivos do JardimBotanico do Rio de Janeiro5:
sules withfiveor six carpels(Killip& Smith29927,
145-157.
McDaniel & Rimachi30269, Vargas 13994, and GILLESPIE, L. J. In press. Pollen morphologyand phyVdsquez& jaramillo6167 fromPeru, Mexia 7265
logeny ofthe tribePlukenetieae (Euphorbiaceae).
fromEcuador, and Plowman2191 fromColomAnnals of the Missouri Botanical Garden.
bia). Three of these collections are recorded as MAcBRIDE, J.F. 1951. Flora of Peru: Euphorbiaceae.
Field Museum of Natural History,Botanical Secultivated,while labels of fourmentionthatthe
ries 13(3a): 3-200.
seeds are edible and are eaten roasted like peanuts (several of the common names recorded, MUELLER, J. 1866. Euphorbiaceae. Pp. 189-1260 in
Prodromus
Systematis
NaturalisRegniVegetabilis,
vol.
e.g., sacha-inchik and sacha mani, translateas
15 (2), ed. A. P. de Candolle. Paris: VictorMasson.
wild peanut). Thus the higher than normal car- PAX,F. A.
1890. Euphorbiaceae. Pp. 1-119 in Die
pel number appears to be a characterselected
Natuirlichen
1st ed., III. 3(5), eds.
Pflanzenfamilien,
forundercultivation.Frequentlycorrelatedwith
A. Engler and K. Prantl.Liepzig: W. Engelmann.
a carpel number of five or six is a more promand K. HOFFMANN. 1919. Euphorbiaceae-Pluinently serrate leaf margin, as opposed to the
kenetiinae. Pp. 1-108 in Das Pflanzenreich,
IV.147.XI.(Heft 68), ed. A. Engler. Liepzig: W.
typical serrulatecondition.
Engelmann.
and
. 1931. Euphorbiaceae. Pp. 11-233
ing withme his knowledgeofthePlukenetieaeand
in Die NatiirlichenPflanzenfamilien,
2nd ed., 19c,
forcommentson the manuscript;
Dan Nicolsonfor
eds. A. Engler and K. Prantl. Liepzig: W. EngeltheLatindiagnoses;and GoeffLevin,Gracorrecting
mann.
dy Webster,and Anna Weitzmanforhelpfulcom- RADCLIFFE-SMITH, A. 1980. A note on Romanoa(Eumentson themanuscript.
Thanksalso to thecurators
phorbiaceae). Kew Bulletin 34: 591-592.
of thefollowingherbariaforfacilitating
visitsand/ WEBSTER, G. L. 1975. Conspectus of a new classifior sendingloans: A, BM, BRG,CAY, DAV, F, FDG,
cation of the Euphorbiaceae. Taxon 24: 593-601.
G, GH, K, MO, NY, P, U, UC. The drawingswere
. In press. Synopsis of the suprageneric taxa
made by CathyPasquale and Alice Tangerini.This
of Euphorbiaceae. Annals of the Missouri Botanresearchwas supportedin partby theBiologicalDiical Garden.
ACKNOWLEDGMENTS. I thank Mike Huft for shar-
versityof The GuianasProgramof the Smithsonian
Institution.
This studyis publicationno. 12 in the
series"BiologicalDiversityoftheGuianas."
LITERATURE CITED
BAILLON,H. E. 1858. Etudegeneraldu groupdes Euphorbiacees.
Paris: Victor Masson.
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