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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 Stable URL: http://www.jstor.org/stable/2419535 . Accessed: 10/02/2015 22:33 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany. http://www.jstor.org 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 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 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 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 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 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 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 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. 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 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 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 580 SYSTEMATIC BOTANY [Volume 18 lcm u1 ln'm lcm~~~~~~~~~~~~~~c lcm H AS lrnm lmm~~~~~~~~~~~ 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 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). 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 582 [Volume 18 SYSTEMATIC BOTANY 1m j2mm~~~~~~~~m (I ZZNK \ fl,'// ~ ~ ; ~~ F ~ ~ ~ t~ ]2mm ~~~~~~~~~~~~~~~~~~~~~~~~~~ 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 ~ 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). 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 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 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 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 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 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 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 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- 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 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- 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 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 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 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 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 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. 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