A revision of paleotropical Plukenetia (Euphorbiaceae

A Revision of Paleotropical Plukenetia (Euphorbiaceae) Including Two NewSpecies from Madagascar

LYNN J GILLESPIE

Research Division Canadian Museum of Nature PO Box 3443 Station D Ottawa Ontario K1P 6P4Canada (lgillespiemus-natureca)

Communicating Editor Gregory M Plunkett

ABSTRACT A revision of the seven paleotropical species of Plukenetia (Euphorbiaceae) is given Three sections orspecies groups are recognized Two genera Tetracarpidium (synonym Angostylidium) and Pterococcus are treatedhere as sections of Plukenetia The monotypic P sect Angostylidium includes the African species P conophora whileP sect Hedraiostylus (synonym P sect Pterococcus) comprises two African species (P africana and P procumbens) andone Asian species (P corniculata) A third species group restricted to Madagascar is distinguished by an androeciumof sessile anthers on an elongate receptacle and comprises three species two described here Plukenetia deciduafrom southeastern Madagascar is close to P madagascariensis and shares styles partly fused into a cylindricalcolumn but differs in its narrower ovate or triangular-ovate leaf blades smaller eglandular bracts racemes withflowers single per node and fewer anthers on a shorter ellipsoid receptacle Plukenetia ankaranensis from northernMadagasar is distinguished from the previous two species by its styles entirely fused into an enlarged obovatestylar column A key to the seven paleotropical species is provided and their relationship to neotropical species isdiscussed

KEYWORDS Pterococcus taxonomic revision Tetracarpidium

Plukenetia L (Euphorbiaceae ss) is a pantropicalgenus of 19 species belonging to tribe Plukenetieaeof subfamily Acalyphoideae The genus is unusualin the Euphorbiaceae for its 4-carpellate ovary andvine or liana habit Twelve species are known fromthe Neotropics and a synopsis of eleven of thesewas previously presented (Gillespie 1993) In thesame year one new species P carabiasiae Jimenezwas described from Mexico (Jimenez 1993) Arevision of the seven paleotropical species in-cluding two new species from Madagascar ispresented here One of these new species Pankaranensis was first collected by the authorduring a field trip to Massif Ankarana in northernMadagascar

Plukenetia species are twining vines lianas orrarely sprawling perennial herbs found in tropicalwet forest to seasonally dry forest or scrubExcellent field identification characters for thegenus are the presence of distinct paired circularor elliptic basilaminar glands on the adaxialsurface of the leaf blade and 4-parted fruit

Taxonomic History and Generic CircumscriptionPlukenetia is considered here in the broad sense tocomprise all members of tribe Plukenetieae havinga 4-carpellate ovary (Gillespie 1993) This includesthe segregate genera Apodandra Pax amp K HoffmEleutherostigma Pax amp K Hoffm Fragariopsis A StHil Pterococcus Hassk and Tetracarpidium Pax[syn Angostylidium (MullArg) Pax amp K Hoffm]but excludes the 3-carpellate genus Romanoa TrevSt Leon (syn Anabaena Adr Juss) This circum-scription with minor exceptions follows closelythat of the major 19th century works on Euphor-

biaceae (Baillon 1858 Muller 1866 Bentham 1880Pax 1890) In contrast Pax and Hoffmann (19191931) in the most recent revision of Plukenetia andtribe Plukenetieae used a narrow delimitation ofthe genus They elevated two paleotropical sectionsof Plukenetia to generic rank (Angostylidium andPterococcus) described two new neotropical genera(Apodandra and Eleutherostigma) and recognizedthe neotropical genus Fragariopsis as distinct(sometimes also treated as a section of Plukenetia)A more detailed history of the taxonomy ofPlukenetia and its segregate genera particularlythe neotropical ones is described in Gillespie(1993 1994)

Pax and Hoffmann (1919 1931) treated the fourpaleotropical species of Plukenetia sl known at thattime in Angostylidium (as Tetracarpidium in 1931)and Pterococcus All of these species had originallybeen described in Plukenetia Leandri (1938) de-scribed a fifth paleotropical species from Mada-gascar placing it provisionally in Plukenetia as Pmadagascariensis Leandri but mentioned that it isvery near to Tetracarpidium Including this speciesin Plukenetia ss would have been a large rangeextension for the then narrowly defined neotrop-ical genus This treatment of three genera in thePalaeotropics Tetracarpidium (syn Angostylidium)Plukenetia ss and Pterococcus has been largelyfollowed to the present day (eg Airy-Shaw 19711975 1981 1982 1983 Dyer 1975 Grierson amp Long1987 Radcliffe-Smith 1996 Govaerts et al 2000)

Pax and Hoffmannrsquos (1919 1931) treatment ofthe Plukenetia complex was first challenged byCroizat (1941) and more recently by Webster

Systematic Botany (2007) 32(4) pp 780ndash802 Copyright 2007 by the American Society of Plant Taxonomists

780

(1975 1994) and Gillespie (1993 1994) Croizatconsidered Pterococcus as part of Plukenetia becausehe could find no differences to separate the twogenera Croizat also remarked that the wingedovaries considered diagnostic for Pterococcus couldbe found in Plukenetia ss Likewise he did notconsider Angostylidium (5 Tetracarpidium) as a dis-tinct genus and pointed out that Pax andHoffmann (1919) although recognizing the genusalso considered it little distinct from PlukenetiaWebster (1975 1994) included both Tetracarpidiumand Pterococcus as synonyms under Plukenetiaa view followed by Gillespie (1993 1994) andRadcliffe-Smith (2001)

Pterococcus was described by Hasskarl (1842) forthe single species Plukenetia corniculata Sm andnamed after its conspicuously 4-winged capsule(Figs 1I J) Subsequently the genus was treatedmostly as a section of Plukenetia but its circum-scription was controversial Baillon (1858) includedonly P corniculata in the genus (which he calledHedraiostylus Hassk) and considered P africanaSond as a member of Plukenetia ss while Muller(1866) included P corniculata P africana and thenewly described species P hastata MullArg (5 Pafricana) in P sect Hedraiostylus (Hassk) MullArgBentham (1880) expanded the circumscription of Psect Pterococcus (Hassk) Benth amp Hook tocomprise in addition to these three species twoneotropical species P penninervia MullArg and Pverrucosa Sm a treatment followed by Pax (1890)In 1919 Pax and Hoffmann resurrected the genusPterococcus and considered it to comprise Pcorniculata P africana (including P hastata) andthe recently described species P procumbens Prain

Pax and Hoffmann (1919) characterized Ptero-coccus based on both style morphology andpresence of wings on the ovaries Likewise Airy-Shaw (1975) considered the genus to be lsquolsquocloselyrelated to Plukenetia differing principally in thevery short thick stylar column and in the conspic-uously 4-winged or 4-horned capsulersquorsquo While Pcorniculata does have a conspicuously four-wingedcapsule (Figs 1I J) P africana has a capsule withfour shorter wings and P procumbens a capsulewith four small tubercles the latter similar to manyneotropical species including P penninervia Psupraglandulosa LJ Gillespie and P verrucosa(Gillespie 1993 Figs 1D 10I) Although not ashighly developed as in P corniculata distinctly 4-winged ovaries and fruit are also found inMadagascan and neotropical species of Plukenetia(eg P ankaranensis Figs 2E F and P loretensisUle) Styles of Pterococcus were described by Paxand Hoffmann (1919) as connate into a short thickcolumn with sessile stigmas as distinct from styles

connate into an urceolate or cylindrical columnwith free arms short or absent (AngostylidiumApodandra Fragariopsis and Plukenetia ss) or stylesfree above (Eleutherostigma) In fact styles are notuniform within Pterococcus but vary from com-pletely connate into a very short stout stylarcolumn (P corniculata Figs 1G H) to partlyconnate into a short narrower cylindrical stylarcolumn with free recurved arms (P procumbens)The latter species does not key out under Pter-ococcus in Pax and Hoffmannrsquos (1919) key butinstead under Plukenetia Therefore neither thestyle nor winged ovary characters used by Pax andHoffmann (1919) to distinguish the genus Ptero-coccus appear to be justified

The only character that could be used to separatePterococcus from other members of Plukenetia sl isstyle length with Pterococcus having styles shorterthan or equal in length to the ovary (Figs 1G H)All other Plukenetia sl (except some collections ofP brachybotrya MullArg) have styles longer thanthe ovary (Figs 2E 3B D G 4F Gillespie 1993Figs 1 9C 10H) Given the extreme variablity ofstyle morphology (size shape and degree ofconnation) in Plukenetia sl (Gillespie 1993) distin-guishing a genus based only on style size does notseem justified Nevertheless the three species ofPterococcus appear to form a natural group whichis recognized here as P sect Hedraiostylus (seediscussion under this section)

Plukenetia sect Angostylidium MullArg wasdescribed by Muller (1864) for his new species Pconophora and was subsequently consistently rec-ognized as a monotypic section (eg Bentham1880 Pax 1890) In 1919 Pax and Hoffmann raisedthe section to generic rank as Angostylidium butsoon after Hutchinson and Dalziel (1928) correctlyrecognized that Tetracarpidium has priority overAngostylidium a treatment later followed by Paxand Hoffmann (1931) Pax and Hoffmann (19191931) distinguished their monotypic genus fromPlukenetia ss based on its high stamen number(40 versus 12ndash30) and presence of intrastaminaldisc segments but also considered it to be littledistinct from the latter Neither character can beused to separate these two genera Stamen numberwas found to range from 25 to 40 in P conophoraMullArg and some Plukenetia ss species havemore than 30 stamens (eg P stipellata LJGillespie has 25ndash40 stamens Gillespie 1993 Fig9D P madagascariensis has 35ndash60+ stamens) Whilespecies of Plukenetia ss either lack a staminate discor have an annular disc (P penninervia speciescomplex Gillespie 1993 Fig 10G) intrastaminaldisc segments similar to those of Tetracarpidium arepresent in the neotropical species P lehmanniana

2007] GILLESPIE PALEOTROPICAL PLUKENETIA 781

782 SYSTEMATIC BOTANY [Volume 32

(Pax amp K Hoffm) Huft amp LJ Gillespie (previouslyrecognized as the monotypic genus Eleutheros-tigma) When considered in the context of Plukene-tia worldwide neither Tetracarpidium nor Pterococ-cus can be maintained as distinct genera

Species Relationships Among paleotropicalPlukenetia there are three well delineated speciesgroups Plukenetia sect Hedraiostylus comprises twosouthern African species P africana and Pprocumbens and the only known Asian species Pcorniculata The second group P sect Angostyli-dium comprises the single species P conophora oftropical central and west Africa The third speciesgroup comprises three species endemic to season-ally dry areas of Madagascar P ankaranensis Pdecidua and P madagascariensis

Paleotropical species do not form a groupdistinct from neotropical species and do notappear to share any unique characters For themost part the three paleotropical species groupsappear to have closer affinites to neotropicalgroups than to each other and are linked morpho-logically via neotropical species

The precise relationship of these three paleotro-pical species groups to the two neotropical speciesgroups described previously (Gillespie 1993) is notentirely clear All paleotropical species share three-nerved to weakly palmate leaves foveolate pollenand the lack of scattered laminar glands on the leafabaxial surface (rarely present in two species) withP sect Plukenetia (5 neotropical species group 1Gillespie 1993) In addition P sect Angostylidumalso shares stamens with filaments styles partlyconnate into a cylindrical column and large fruitand in fact seems to be little differentiated from Psect Plukenetia In contrast the other two paleotro-pical species groups share characters with bothneotropical species groups Plukenetia sect Hedraios-tylus shares a similar androecium of all stamenswith filaments with P sect Plukenetia and smallcapsules and small lenticular seeds with neotropicalspecies group 2 On the other hand the Madagascanspecies group shares an androecium of sessileanthers with species group 2 and medium-sizedfruit and subglobose seeds with P sect Plukenetia

Similar character trends are seen in both the Oldand New Worlds such as the trends towardentirely fused styles winged ovaries sessileanthers and highly expanded staminate recepta-

cles (characters polarized using Romanoa as sistergroup Gillespie 1993) Some of these sharedcharacters appear to be shared due to commonancestry while others may represent parallelevolution For example the character trend frompartly to entirely fused styles occurs within boththe Madagascan species group and P sect He-draiostylus (although less pronounced in the latter)while in the Neotropics style fusion distinguishesthe two neotropical groups with P sect Plukenetiahaving styles partly fused and species group 2having styles entirely fused (Gillespie 1993) Thepresence of both states in two morphologicallydistinct species groups and the derived statecharacterizing a third group suggests that entirelyfused styles arose independently at least twice andpossibly three times In contrast the long partlyfused style of the African P sect Angostylidium issuggested to be a plesiomorphic character sharedwith P sect Plukenetia via common ancestry basedon a close similarity in overall morphology

Detailed discussions of species relationships andmorphological character evolution and a revisedsectional classification will await the completion ofmolecular and phylogenetic analyses (Gillespiework in progress) In the meantime the existingsubgeneric classification will be maintained for thepaleotropical species

Taxonomic Characters Paleotropical species ofPlukenetia may be distinguished by floral fruit andleaf characters similar to those used in character-izing neotropical species (Gillespie 1993) with theexception of leaf architecture and pollen morphol-ogy As in the neotropics floral and fruit mor-phology is surprisingly diverse Useful staminateflower characters include presence of filaments anddisc and receptacle shape Style morphology andsize vary considerably with styles ranging frompartly to entirely fused and the stylar columnssometimes enlargened into a variety of shapesFruits range from small dehiscent capsules to largeindehiscent fruit and usually have a variouslyshaped protuberance (wing horn tubercle) oneach carpel Useful foliar characters include bladeshape and size presence of stipels and numbersize and position of basilaminar glands

Leaf architecture and pollen morphology bothuseful in characterizing the two main neotropicalspecies groups are insufficiently variable to distin-

r

FIG 1 Plukenetia corniculata A Twining stem with mature leaves and inflorescences B Close-up of leaf blade base showingpaired basilaminar glands and stipels C Branch showing leaf-opposed inflorescence with immature fruit and staminateflowers D Part of inflorescence with staminate flower buds E Staminate flower F Stamen G Young pistillate flower HPistillate flower in post anthesis stage I Capsule side view J Capsule top view K Seed side view A B D H based on Brink5771 C on Koorders 41509 E F on Chin See Chung 2712 G on Lorzing 7821 IndashK on Meijer 7309 Drawing by Anita Walsmit Sachsused with permission from the Nationaal Herbarium Nederland



guish among paleotropical species Paleotropicalspecies share medium-sized pollen grains (equatorialaxis 35ndash51 mm long) with a foveolate exine (Gillespie1994 and results presented here) and leaves withthree-nerved to weakly palmate venation

Paleotropical species appear to be reasonablywell defined in contrast to several neotropicalspecies complexes such as the poorly understoodP penninervia and P brachybotrya complexes in theAndean and Amazonian regions (Gillespie 1993)

TAXONOMIC TREATMENT

PLUKENETIA L Sp Pl 1192 1753mdashTYPE Plukene-tia volubilis L

Vigia Vell Conc Fl Flumin 9 t 128 1832mdashTYPEVigia serrata Vell Conc [5 Plukenetia serrata(Vell Conc) LJ Gillespie]

Fragariopsis A St-Hil Lecons Bot 426 1840mdashTYPE Fragariopsis scandens A St-Hil [5Plukenetia serrata (Vell Conc) LJ Gillespie]

Pterococcus Hassk Flora 25 (2 Bleibl) 41 1842nom cons non Pall 1773mdashTYPE Pterococcusglaberrimus Hassk nom illeg [5 Plukenetiacorniculata Sm]

Hedraiostylus Hassk Tijdschr Natuurl GeschPhysiol 10 141 1843mdashTYPE Hedraiostylusglaberrimus (Hassk) Hassk [5 Plukenetia cor-niculata Sm]

Sajorium Endl Gen Pl Suppl 3 98 1843mdashTYPESajorium corniculatum (Sm) Dietr [5 Plukenetiacorniculata Sm]

Tetracarpidium Pax in Engl Bot Jarhb Syst 26 3291899mdashTYPE Tetracarpidium staudtii Pax [5Plukenetia conophora MullArg]

Pseudotragia Pax in Engl Bull Herb Boissier ser2 8 635 1908mdashTYPE Pseudotragia scandensPax [5 Plukenetia africana Sond] (according toWebster 1994)

Eleutherostigma Pax amp K Hoffm PflanzenrIV147IX (Heft 68) 11 t3 1919mdashTYPEEleutherostigma lehmannianum Pax amp K Hoffm[5 Plukenetia lehmanniana (Pax amp K Hoffm)Huft amp LJ Gillespie]

Angostylidum (MullArg) Pax amp K Hoffm Pflan-zenr IV147IX (Heft 68) 17 1919mdashTYPEAngostylidum conophorum (MullArg) Pax amp KHoffm [5 Plukenetia conophora MullArg]

Apodandra Pax amp K Hoffm Pflanzenr IV147IX(Heft 68) 20 1919mdashTYPE Apodandra loretensis

(Ule) Pax amp K Hoffm [5 Plukenetia loretensisUle] (according to Webster 1994)

Elaeophora Ducke Arch Jard Bot Rio de Janeiro 4112 1925mdashTYPE Elaeophora abutifolia Ducke[5 Plukenetia polyadenia MullArg]

Lianas vines or rarely perennial herbs (Pprocumbens) monoecious or rarely dioecious latexabsent stems twining sometimes initially erect orrarely procumbent and sprawling Leaves simplealternate evergreen or deciduous petiolate sti-pules small deciduous blade chartaceous pin-nately palmately or 3-veined margins subentire toserrulate or rarely serrate often with small orminute glandular setae pair of stipels or smallknob sometimes present at petiole-blade junction1ndash6(ndash10) pairs of flat usually conspicuous basila-minar glands present near base on blade adaxialsurface scattered smaller laminar glands some-times present on abaxial surface usually nearmargin rarely present on adaxial surface (only inP supraglandulosa) Inflorescence a racemosethyrse or rarely a raceme axillary or terminalbisexual with pistillate flower(s) at base andstaminate flowers above or rarely unisexualflowers in condensed cymules rarely in lax cymesor single per node bracts triangular smalleglandular or rarely larger and biglandular (Pmadagascariensis) Staminate flowers small typi-cally green greenish yellow or cream pedicelspresent usually articulated (comprising cyme axisplus true pedicel) sepals 4 or 5 valvate petalsabsent glandular disc absent or present inter-staminal segmented or annular stamens 8 to 60free on convex subglobose or elongate receptaclefilaments short to elongate or anthers sessilepistillode absent pollen tricolpate suboblate tooblate-spheroidal amb obtuse-triangular to sub-circular angulaperturate exine tectate-perforatetectum foveolate to reticulate Pistillate flowers

pedicellate sepals 4 petals and disc absent ovary4-locular with locules uniovulate 4-angled todeeply 4-lobed each angle or lobe carinate andor with a tubercle horn or laterally compressedwing styles partly to completely connate columncylindrical to globose or obovoid free style armsabsent or present entire to obscurely bifid Fruit

a 4-seeded capsule (usually schizocarpous) orberry deeply 4-lobed to subglobose each carpelcarinate andor with central tubercle or wing

r

FIG 2 Plukenetia ankaranensis sp nov A Branch with mature leaves and inflorescence B New shoot with younginflorescence C Closeup of leaf showing glands at base of blade D Staminate flower E Pistillate flower with view of apexshown above F Capsule G Seed ventral view (left) lateral view (right) A B D E based on Gillespie 4074 C on Gillespie 4088F G on Gillespie 4076


Seeds subglobose ovoid or lenticular and thenlaterally compressed ecarunculate surface

smooth rough or verrucate outer seed coat (testa)thin persistent or not

KEY TO THE PALEOTROPICAL SPECIES OF PLUKENETIA

1 Fruit $ 5 cm wide seeds broadly ovoid 25 cm long staminate disc segments numerous slender interstaminalstamens 25ndash40 with filaments on small subglobose receptacle inflorescence axillary styles 4ndash8 mm long longer thanthe ovary mostly connate into a thick cylindrical column free style arms short conspicuously dilated and spreadingtropical Africa (P sect Angostylidium) 1 P conophora

1 Fruit 4 cm wide seeds lenticular broadly ellipsoid or subglobose 2 cm long staminate disc absent stamens 8ndash60if 20 then anthers sessile on elongate receptacle inflorescence position various styles not as above

2 Androecium of sessile anthers on prominent elongate receptacle styles 3 mm long longer than the ovarycapsules 2ndash4 cm wide seeds broadly ellipsoid to subglobose 13ndash18 cm long (capsule and seeds unknown for Pmadagascariensis) inflorescence position various Madagascar (Madagascan species group)

3 Styles entirely connate 6 mm long stylar column obconic or obovoid free style arms absent androecium

1 mm long anthers 15ndash20 inflorescence a thryse terminal and appearing leaf-opposed staminate flowers indistinct cymules glandular knobs absent at petiole apex 5 P ankaranensis

3 Styles partly connate (12 of length) 7 mm long stylar column cylindrical free style arms slender taperedandroecium 15 mm long anthers 18ndash60+ inflorescence a very narrow thyrse or raceme axillary orterminal staminate flowers single per node or in condensed cymules glandular knobs 1ndash2 at petiole apexsometimes minute

4 Inflorescence a terminal raceme flowers single per node bracts triangular 2 mm long epetiolateeglandular androecium 16ndash18 mm long anthers 18ndash30 on oblong-ellipsoid receptacle leaf bladestriangular-ovate or ovate 7 P decidua

4 Inflorescence an axillary thryse flowers in condensed cymules bracts lanceolate 3ndash8 mm long usuallypetiolate and biglandular androecium 3ndash4 mm long anthers 35ndash60+ on narrowly conical receptacle leafblades broadly ovate or orbicular 6 P madagascariensis

2 Androecium of stamens with filaments on small convex to globose receptacle styles 2 mm long shorter than orlength equal to the ovary capsules 2 cm wide seeds broadly lenticular laterally compressed 12 cm longinflorescence terminal appearing leaf-opposed southern Africa and southeast Asia (P sect Hedraiostylus)

5 Petioles (1ndash)3 cm long leaf blade (2ndash)4 cm wide deeply cordate at base with 2 conspicuous basilaminarglands and 2 stipels at base capsule with strap-shaped wing 6ndash12 mm long on each carpel lobe Asia 3 P corniculata

5 Petioles 1 cm long leaf blade 35 cm wide obtuse to truncate hastate or rarely sagittate at base withbasilaminar glands 2ndash12(ndash20) sometimes minute or absent and stipels minute or absent capsule withtubercle or wing 3 mm long on each carpel lobe southern Africa

6 Leaf blade elliptic or ovate (LW 06ndash08) 2ndash45 cm long obtuse or rounded at base basilaminar glands2ndash12(ndash20) 4 P procumbens

6 Leaf blade narrowly triangular lanceolate or linear-lanceolate [LW 005ndash04 (ndash06)] 3ndash8 cm long oftenhastate at base basilaminar glands 2 often minute or absent 2 P africana

PLUKENETIA sect ANGOSTYLIDIUM MullArg Flora47 530 1864 Angostylidium (MullArg) Pax ampK Hoffm Pflanzenr IV147IX(Heft 68) 171919mdashTYPE Plukenetia conophora MullArg

1 PLUKENETIA CONOPHORA MullArg Flora 47530 1864 Angostylidium conophorum (Mul-lArg) Pax amp K Hoffm Pflanzenr IV147IX(-Heft 68) 17 1919 Tetracarpidium conophorum(MullArg) Hutch amp Dalziel Fl W Trop Afr1 307 1928mdashTYPE lsquolsquoCameroon Riverrsquorsquo [Ca-meroun] Mann 2202 (lectotype here desig-nated K [sheet annotated as lsquolsquooriginalisrsquorsquo byMuller Arg] isolectotypes K-2 sheets)

Tetracarpidium staudtii Pax Bot Jahrb Syst 26 3291899mdashTYPE Cameroun Station Johann-Al-brechtshohe 15 Jan 1897 Staudt 802 (holotypeB destroyed illustration at K fragment ofholotype at K isotypes BM MO)

Mallotus preussii Pax Bot Jahrb Syst 23 525 1897Cleidion preussii Baker Kew Bull 143 1910mdashTYPE Cameroun Barombistation 25 Aug

1890 Preuss 420 (holotype B destroyedillustration at K isotypes BM K)

Cleidion mannii Baker Kew Bull 58 1910mdashTYPElsquolsquoUpper Guinea Cameroon Riverrsquorsquo Mann 1202(holotype K)

Monoecious (functionally dioecious ) liana to30 m high trunk to 14+ cm in diam stemssometimes twining glabrescent Stipules triangu-lar 05 mm long Leaves evergreen petioles(15ndash)3ndash6 cm long glabrous or very sparselypubescent with apex pubescent blades elliptic orovate 5ndash12 3 35ndash9 cm chartaceous mostlyglabrous with base pubescent and major veinsvery sparsely pubescent acute to obtuse at apexwith acumen 05ndash1 cm long broadly obtuserounded or rarely shallowly cordate at basemargins serrulate or crenulate-serrulate venation3-nerved at base sometimes weakly palmatesecondary veins mostly brochidodromous orsometimes semi-craspedodromous 3ndash4 on eachside of midrib tertiary veins percurrent or


FIG 3 Plukenetia ankaranensis and P madagascariensis AndashD P ankaranensis A Liana climbing up small tree on erodedlimestone pavement B Flowering branch showing single pistillate flower at base of inflorescence and staminate flowers in budabove C Liana with inflorescences D Close-up of short inflorescence with two basal pistillate flowers and open staminateflowers EndashG P madagascariensis E Liana in dry forest F Vegetative branch G Inflorescences the basal part of threeinflorescences is shown the lower-most one has a single basal pistillate flower the one on the upper left shows the staminateflower buds and the one on the right has an open staminate flower A D Gillespie 4076 B Gillespie 4074 C Gillespie 4088 EndashFGillespie 4125 G Andrianjafy et al 1648 Scale bar in B F 5 2 cm bar in D 5 5 mm bar in G 5 1 cm


FIG 4 Plukenetia decidua sp nov A Branch with inflorescences B Branch with leaves C Staminate flower D Staminatesepal E Androecium F Pistillate flower G Capsule H Seed lateral view (left) ventral view (right) A CndashF based on CapuronSF 18682 B on Capuron SF 27952 GndashH on Decary 3253


sometimes reticulate stipels and glandular knobabsent basilaminar glandular areas 2ndash4(ndash6) irreg-ularly circular to elongate 05ndash4 mm long nearmargin each area usually comprising many smallglands of irregular shape and size laminar glandsabsent Inflorescence unisexual or functionallyunisexual axillary 1(ndash2) per axil main axissparsely puberulous minor axes moderately todensely puberulous peduncle 02ndash24 cm longbracts narrowly triangular very sparsely pubes-cent staminate 1ndash12 mm long pistillate 05ndash1 mmlong staminate inflorescence a narrow thyrse

(rarely with one or two branches near base similarto main axis) 4ndash13 (ndash20) cm long cymulescondensed to somewhat lax with axes to 4 mmlong sometimes with 1 or 2 apparently non-functional pistillate flowers at basal-most nodespistillate inflorescence a raceme 05ndash3 cm longflowers 1 or 2(3) single per node axis mostlyaborted above [rarely with few staminate flowersin cymule(s)] Staminate flower pedicel 2ndash7 mmlong slender glabrous bud globose obtuse atapex sepals 4 or 5 elliptic or narrowly obovoid22ndash36 3 11ndash18 mm mostly glabrous acute at

FIG 5 Distribution of Plukenetia in Africa and Madagascar (inset)


apex androecium subglobose or ellipsoid 12ndash2 mm long comprising 25ndash40 stamens on smallsubglobose receptacle filaments plump-conical02ndash04 mm long disc segments numerous slen-der-cylindrical 01ndash02 mm long interstaminalPistillate flower pedicel 15ndash5 mm long sparselyor very sparsely puberulous sepals narrowlytriangular 06ndash12 3 03ndash07 mm very sparselypuberulous often ciliolate ovary 15ndash3 3 25ndash5 mm (including wings) moderately puberulous4 (rarely 5)-winged wings sparsely puberulousbecoming larger and auriculate in immature fruitstage styles 4ndash75 mm long sparsely puberulousmostly connate into a thick cylindrical column 1ndash25 mm wide conspicuously dilated to 3ndash55 mmwide at apex the 4 free style arms 05ndash2 mm longbroad obscurely bifid divergent their innersurface stigmatic pistillate flowers on staminateinflorescences smaller with ovary lacking wingsstylar column to 45(ndash7) mm long not or variouslydilated at apex free style arms absent or 05 mmlong Fruit 4 (rarely 5)-lobed apparently indehis-cent 25ndash35 3 5ndash7 cm when dry (5 3 75 cmwhen fresh according to Osmaston 2599) glabrouseach carpel subglobose carinate keel to 11(ndash14)mm wide widest at centre and often appearingwing-like pericarp 2ndash3 mm thick when dry Seedsbroadly ovoid 29 3 27 3 25 cm surface(tegmen) medium to dark brown rough withirregular vertical ridges and reticulations testa notpersistent pale brown with fine dark brownmarkings

Pollen Description Gillespie (1994) Fig 19

Illustration Pax and Hoffmann (1919) Fig 5

Distribution and Ecology Sierra Leone (notindigenous) Benin Nigeria Cameroun Equato-rial Guinea Gabon Congo and the DemocraticRepublic of Congo (Zaire) (Fig 5) A liana of terrafirme rainforest growing in primary secondaryand degraded forest gallery forest and in second-ary scrub cacao and coffee plantations near sea-level to 750 m sometimes cultivated In westernAfrica (Sierra Leone to Cameroun) floweringcollections made in November to March andfruiting collections February to June in Nigeriaseeds are recorded to be available from June toSeptember (Egharevba et al 2005) In central Africaflowering apparently throughout the year fruitscollected in February and March

Representative Specimens Examined SIERRA LEONEAllen Town Deighton 3304 (K) Port Loko Deighton 4041 (K)5018 (K)

BENIN Cercle de Porto Novo pres la gare de SaheteChevalier 22870 (K P)

NIGERIA Benin Prov Benin Division Okomu ForestReserve Brenan amp Onochie 8999 (BM K P) Ibadan ProvIgbajo District Aba-panu on mile 8 from Ikirun-Igbajo motorroad Latilo FHI 31764 (K) Calabar Prov Ikpai District

between miles 68 and 69 on Calabar-Mamfe road Latillo FHI53991 (K) near Modakeke Foster 205 (K) Ijebu Prov BaboEko Shasha Forest Reserve Ross 87 (BM MO) Oban DistrictTalbot amp Talbot 1384 (K) Oban District Oban Talbot amp Talbotsn (K MO) Eket District Talbot amp Talbot sn (BM) AgukuDistrict N Thomas 888 (K) Ubuluku N Thomas 2072 (K)

CAMEROUN East Prov Bertoua near catholic missionBreteler 816 (K P) South-West Prov Kumba District by theside of River Kindong NA Forest Reserve MbalangeSouthern Bakundu Binuyo amp Deramola FHI 35499 (K P)Victoria Maitland 375 (K) Konye Bakossi Mbambe ForestMamfe Road 4u569N 9u309E Nemba amp Thomas 434 (MO)Kurume west of Mungo R 4u559N 9u299E D Thomas 5455(MO P) 5456 (MO P) Banyu 5u109N 9u149E Thomas et al5524 (MO) degraded forest at Barombi Kang Kumba4u369N 9u279E Thomas amp Etuge 6984 (MO) forest aroundMasaka-Batanga 5u069N 9u109E Thomas amp Namata 7790(MO) forest along Mungo R by Kurume hammock bridge40 km N of Kumba on Mamfe road Thomas amp Nemba 5419(MO) South Prov Bipinde Zenker 1583 (BM L P) Bipinde1900 Zenker 2234 (A BM COI G L MO O P S) Bipinde1903 Zenker 2551 (BM G L MO P) Bipinde 1907 Zenker3273 (BM G K L MO P S US) Zenker 3311 (BM G L PUS) Zenker 3394 (BM COI G L P US) Bipinde 1912 Zenker4451 (BM G)

CENTRAL AFRICAN REPUBLIC Region de YalingaHaut Oubangui rive droite au Zaes Le Testu 4366 (P)Region de Mbaiki Station Central de Boukoko Tisserant 104(M P) 1261 (BM P) 2045 (BM P)

EQUATORIAL GUINEA Sierra del Crystal Mann 1739(syntypes K P)

GABON Environs de Libreville Klaine 2284 (K P) LeHaute Ogooue Koulamotou Le Testu 8183 (BM P) Oyem LeTestu 9194 (P) Ngounie region Komi near Sindara 1u49S10u489E Thomas amp Wilks 6418 (MO P)

CONGO Lekana village Ebongo Bouquet 2460 (P)Environs de Brazzaville Courtet sn (L)

CONGO DEMOCRATIC REPUBLIC (ZAIRE) EquateurEala Corbisier 1231 (BM K MO P) entre Businga etBanzyville Territoire Businga Lebrun 2013 (MO) entreLibenge et Gemena Territoire Libenge Lebrun 1797 (MO)Orientale Epulu zone de Mambasa (Ituri) 1u259N 28u359EHart 479 (MO) 780 (MO) Bambesa Territoire BambesaGilbert 457 (K) entre Iruma et Mombasa Territoire MombasaLebrun 4165 (K) Yangambi Territoire Isangi Leonard 1283(MO) Malibua route Yangambi-Ngazi Territoire IsangiLouis 1226 (MO) Yangambi plateau de la Lusambila 5 kmau N du fleuve Territoire Isangi Louis 3333 (MO) YangambiReserve Flore Isalowe Territoire Isangi Louis 5945 (MO)Yangambi route Ngazi Territoire Isangi Louis 7980 (MO)Yangambi Plateau de la Luweo Territoire Isangi Louis 9876(MO) Yangambi Territoire Isangi Louis 10259 (MO)Yangambi Plateau de LrsquoEtchwa Territoire Isangi Louis13218 (K)

Vernacular Names and Uses Sierra LeoneAwusa (Creole Deighton 4041 5018) Owusia (Elliot4118) Nigeria African walnut (Lowe 1738) Akanotoli (Thomas 1997) Asala (Yoruba Lowe 1738Osseyemeh 3342) Awusa (Yoruba Kennedy 1731Lowe 1738) Okprsquoa (Thomas 1088) Omumu (Thomas2072) Cameroun Casso (Yaounde Hedin 2101)Casu (Thomas et al 5524) Kaso (Balondo Nemba ampThomas 434) Central African Republic Kaso(Lissongo Tisserant et al 104 1261 2045) Demo-cratic Republic of Congo Tobe (Lunandi Hart 497780 1621 Omaston 2599) Gabon (Raponda-Walker


and Sillans 1961) Bekase (Bakele) Bekasi (Beseki)Bokasu (Bakota) Dugasu (Bapunu) Kasu (fruitnumerous dialects) Mugasa (Eshira BavaramaBavungu) Mukasu (Avili Banzabi Baduma)Muti-a-kasu (Baduma) Nsinga-makasu (Loango)Nwisi-wa-kasu (Ngowe) Ogasu (MpongweGaloa Nkomi Orungu Mitaogo Apindji SimbaIvea Bavove) Known as African walnut Nigerianwalnut and Conophor nut and the extracted oil asConophor oil in the agricultural and biochemicalliterature (eg Sato et al 1991 Animashaun et al1994 Odoemelam 2003 Egharevba et al 2005)

Seeds are reported to be edible and are eatenthroughout the species range in Sierra Leone(Elliot 4118) Nigeria (Thomas 1088 Osseyemeh3342) Cameroun (Thomas amp Namata 7790 Thomasamp Nemba 5419 Hedin 1929) Central AfricanRepublic (Tisserant et al 104 1261) the DemocraticRepublic of Congo (Hart 780) and Gabon (Ra-ponda-Walker and Sillans 1961) Seeds are re-ported to be eaten boiled in Cameroun (Thomas etal 5524) cooked in the Democratic Republic ofCongo (Osmaston 2599) and grilled in Gabon(Raponda-Walker and Sillans 1961) and are soldin markets in Sierra Leone (Deighton 4041 5018) andCameroun (Hedin 2101) Ethnomedicinal use in-cludes macerations of leaves and roots for treatingasthma and hypertension (Okafor and Okorie inEgharevba et al 2005) The uncommon 5-lobedfruits are considered to be good luck charms inCameroun (Thomas et al 5524)

Plants are both cultivated in village farms andencouraged to persist in a semi-wild state onfarmland for their edible seeds and for the oilextracted from the seeds (Hedin 1929 Chevalier1948 Okigbo 1977 Egharevba et al 2005) Thepresence of 5-lobed fruits may provide evidencefor selection by local farmers (the one seen alsolacks the broad keel typically present on eachcarpel) Recent biochemical research has shown theseeds to have high protein and oil content(Akpuaka and Nwankwor 2000 Akintayo andBayer 2002 Odoemelam 2003 Oboh and Ekperigin2004) a similar large-seeded neotropical species Pvolubilis L has attracted interest for its storageprotein containing all essential amino acids (Satheet al 2002) The species has recently been the focusof agricultural research such as characterizing andimproving seed germination and seedling growth(Egharevba et al 2005 Jiofack and Dondjang2006ab)

Discussion Plukenetia conophora is distinctamong paleotropical species and has previouslybeen treated as a separate monotypic genus eitherAngostylidium (Pax and Hoffmann 1919) or Tetra-carpidium (Hutchinson and Dalziel 1928 1958 Pax

and Hoffmann 1931) First described under Pluke-netia (Muller 1864) the species is once again treatedin this genus based on many shared characterswith neotropical species The species most closelyresembles species of neotropical P sect Plukenetia(species group 1 Gillespie 1993 lsquolsquoCylindrophorarsquorsquoPax and Hoffmann 1919) differing only in minorcharacters such as smaller pollen grains and lack ofstipels or glandular knob at the petiole apex andmay indeed belong to this group The monotypicsection P sect Angostylidium is provisionallyretained here for this species pending results ofa phylogenetic study

The species may be distinguished from otherpaleaotropical species by staminate flowers witha disc and either four and five sepals large fruitsand seeds and a conspicuous gynoecium witha thick stylar column and four very shortspreading dilated arms In addition liana habitaxillary inflorescences a greater number of sta-mens and ovoid seeds distinguishes it from P sectHedraiostylus and a little expanded staminatereceptacle and stamens with filaments from theMadagascan species group All of these charactersdistinguishing P conophora in the Palaeotropics areshared with species in the Neotropics (eg habitand aspect with P polyadenia MullArg intrastam-inal disc segments with P lehmanniana gynoeciumsimilar to P stipellata)

The species exhibits two features unusual inPlukenetia irregular and indistinct basilaminarglands and functionally unisexual inflorescencesAt the base of the leaf blade on the adaxial surfacethe species has two or more glandular areas thatare irregular in shape and size and usuallycomposed of many tiny glands In contrast otherspecies generally have two distinct circular orelliptic basilaminar glands

The species appears to have unisexual or at leastfunctionally unisexual inflorescences and may befunctionally dioecious The majority of collectionsexamined have staminate or similar apparentlybisexual inflorescences while others have muchshorter pistillate inflorescences The lsquolsquobisexualrsquorsquoinflorescences have one or two pistillate flowersat their base but these flowers are smaller andmorphologically different (eg ovary not or onlyslightly winged style not or less dilated at apex)from those on pistillate inflorescences and theyappear to be non-functional as no typical maturepistillate flowers or immature fruit have been seenon these lsquolsquobisexualrsquorsquo inflorescences Pistillate inflor-escences may occasionally terminate in a short axiswith small staminate buds but these rarely de-velop into mature flowers Only two pistillatecollections were seen with staminate flowers


approaching mature size one with few separatestaminate cymules above the pistillate flowers(Talbot amp Talbot 1371) the second with a singlestaminate flower in a cymule with a pistillateflower (Talbot sn)

Unisexual inflorescences and dioecism areunusual in the genus the majority of species aremonoecious with bisexual (and sometimes stami-nate) inflorescences Apart from P conophora theonly other species with pistillate and staminateinflorescences is the South American P polyadeniathis latter species appears to be either dioecious ormonoecious with unisexual or occasionally bi-sexual inflorescences (Gillespie 1993) Dioecismand unisexual inflorescences appears at least inpart to be correlated with habitat and habit bothspecies are canopy lianas of rainforest with largefruit and similar leaf morphology

Although the species is for the most partmorphologically rather uniform across its rangethere is some geographical variation in leaf shapeCollections from western Africa (including allcollections from Sierra Leone and Benin) generallyhave ovate leaf blades rounded to shallowlycordate bases and triplinerved to weakly palmatevenation while those from central Africa mostoften have elliptic blades obtuse to rounded basesand triplinerved venation

The small population in Sierra Leone is highlydisjunct from the main range of the species fromBenin to central Africa The species is likely notindigenous in Sierra Leone but appears to havebeen introduced for its edible seeds Two of fourcollections indicate cultivation and Deighton(5018) comments that it is apparently not in-digenous in Sierra Leone The population is alsonot morphologically distinct from other westAfrican collections

Both syntype collections have staminate inflor-escences with small apparently sterile pistillateflowers The collection having the most duplicatesMann 2202 is selected as lectotype Two of thethree duplicates appear to have been annotated asPlukenetia conophora by Muller the one annotatedas lsquolsquooriginalisrsquorsquo is selected as lectotype

PLUKENETIA sect HEDRAIOSTYLUS (Hassk) Mul-lArg DC Prodr 15 (2) 772 1866 Hedraios-tylus Hassk Tijdschr Natuurl Gesch Physiol10 141 1842mdashTYPE Hedraiostylus glaberrimusHassk nom illeg [5 Plukenetia corniculataSm]

Plukenetia sect Pterococcus (Hassk) Benth amp HookGen Pl 3 1 327 1880 Pterococcus HasskFlora 25 (2) Beibl 3 41 1842 nom consmdash

TYPE Pterococcus glaberrimus Hassk nomilleg [5 Plukenetia corniculata Sm]

This species group previously most commonlytreated as the genus Pterococcus (eg Govaerts et al2000) is treated here as Plukenetia sect Hedraiosty-lus which takes precedence over P sect Pterococ-cus This paleotropical section comprises twoAfrican species P africana and P procumbens andone Asian species P corniculata Most recent florasconsidered these species as members of Pterococcus(Hutchinson and Dalziel 1958 Backer and Bakhui-zen van den Brink 1963 Airy-Shaw 1971 19751981 1982 1983 Whitmore 1973 Dyer 1975Grierson and Long 1987 Radcliffe-Smith 1996)few treat the species under Plukenetia (Phillips1951 Leistner 2000 Germishuizen and Meyer2003) The species of this section may be distin-guished from other paleotropical species by veryshort styles that are shorter than or equal in lengthto the ovary small capsules and small lenticularseeds In addition they may be distinguished fromP sect Angostylidium by their terminal leaf-opposed inflorescences fewer stamens absenceof a staminate disc and from the Madagascanspecies group by their stamens with filaments andlittle expanded staminate receptacles

The three species of P sect Hedraiostylus may bemost easily distinguished from each other on thebasis of foliar characters such as leaf blade size andshape petiole length basilaminar gland presencesize and number and stipel presence Stylemorphology (including shape and degree of con-nation) and fruit morphology (particularly the sizeand shape of the appendage on each carpel lobe)provide additional distinguishing characters

2 PLUKENETIA AFRICANA Sond Linnaea 23 1101858 Pterococcus africanus (Sond) Pax amp KHoffm Pflanzenr IV147IX(Heft 68) 221919mdashTYPE South Africa MegalisbergOct Zeyher 1522 (holotype S isotypes BMG K)

Plukenetia hastata MullArg Flora 47 469 1864mdashTYPE Mozambique lsquolsquoZambesica on the Low-er Zambezirsquorsquo Kirk sn (holotype K)

Pseudotragia schinzii Pax Bull Herb Boissier ser 28 635 1908mdashTYPE Namibia lsquolsquoAmbolandOtjihevetarsquorsquo Schinz 895 (holotype G not seenisotype K)

Pseudotragia scandens Pax Bull Herb Boissier ser2 8 636 1908mdashTYPE Namibia lsquolsquoAmbolandOohamarsquorsquo Mar 1886 Schinz 894 (holotype Gnot seen isotype K)

Monoecious vine or sometimes perennial herbwith thick woody rootstock stems twining scan-


dent or sometimes trailing on ground slendersparsely to moderately pubescent new shoots fromrootstock initially erect Stipules linear-lanceolateor linear-triangular ca 1 mm long Leaves withpetioles 02ndash09 cm long sparsely to denselypubescent blades narrowly triangular lanceolateor linear-lanceolate 3ndash8 3 (015ndash)03ndash15(ndash3) cmoften with hastate basal lobes and then to 3(ndash45)cm wide thin to thick-chartaceous glabrescent tosparsely pubescent with major veins usuallysparsely or moderately pubescent narrowly acuteor rarely broadly acute at apex hastate truncateshallowly cordate rarely sagittate or obtuse atbase basal lobes when present usually triangularto 1(ndash15) cm long pointed or rounded andoriented horizontally (rarely directed downwardsand then base sagittate) margins remotely serru-late to serrate especially along basal half venationweakly palmate or 3-nerved at base secondaryveins brochidodromous 2ndash8 on each side of mid-rib tertiary veins weakly percurrent or reticulatestipels usually present 2 01ndash06 mm long basila-minar glands 2(ndash4) circular or elliptic 02ndash03 (ndash04) mm in diam marginal often obsure orsometimes absent laminar glands absent or rarely1ndash2(ndash8) minute Inflorescence a narrow thyrse 1ndash7(ndash10) cm long bisexual terminal but appearingleaf-opposed or rarely terminal on short shootsaxis sparsely to moderately pubescent basally toglabrescent distally peduncle 1ndash8 mm long pistil-late flower 1 at basal-most node staminate flowersnumerous above in small very condensed cym-ules bracts narrowly elliptic or lanceolate 15ndash3 mm long glabrous Staminate flower pedicel 1ndash17 mm long glabrous to very sparsely pubescentespecially near base bud globose broadly obtuseto rounded at apex sepals 4 ovate or elliptic 11ndash15 3 06ndash1 mm glabrescent acute at apex withthickened tip androecium subglobose 1 mm indiam comprising 15ndash20 stamens densely packedon small convex to subglobose receptacle fila-ments conical 02ndash04 mm long disc absentPistillate flower pedicel 3ndash10(ndash14) mm long verysparsely to moderately pubescent sepals ovate orelliptic 2ndash3 3 08ndash15 mm glabrescent to sparselypubescent ovary 1ndash2 3 15ndash3 mm (excludingwings) hirsute 4-winged wings 08ndash15 mm longrounded sparsely hirsute styles 1ndash14 mm longmostly connate into a stout hirsute column 07ndash1 mm long the 4 prominent stigmas forminga cross 12ndash22 mm across at apex each stigmaobtusely obtriangular glabrous Fruit a 4-lobedcapsule 6ndash8 3 11ndash17 mm sparsely pubescentsurface irregularly angular-verrucose each carpellobe with wing 1ndash3 mm long pedicel 06ndash15(ndash25)cm long Seeds broadly lenticular 65ndash8 3 55ndash75

3 25ndash4 mm (including keel) obtuse-triangular inoutline laterally compressed with radial keel 05ndash12 mm wide surface coarsely and irregularlyreticulate pale brown or pale greenish-brownoften with darker orange-brown markings testapersistent


Illustrations Pax and Hoffmann (1919) Fig 7Radcliffe-Smith (1996) Tab 44

Distribution and Ecology Zambia Mozambi-que Zimbabwe Botswana Namibia and SouthAfrica (Fig 5) A vine or perennial herb with twiningor trailing stems growing in open or woodedsavanna woodland or open forest on sand orsandy soil mostly 900ndash1200 m 100ndash500 m in Mo-zambique and SE Zimbabwe Flowering and fruitingcollections made between September and April

Representative Specimens Examined ZAMBIA MachiliDistrict Machili Fanshawe 5957 (K)

MOZAMBIQUE entre Mocubela e Bajone a 155 km deMocubela Grandvaux Barbosa amp Carvalho 4267 (K) LugelaDistrict Moebede road Faulkner 299 (K) MaricaE SofalaDistrict 10 km N of Mwanza where the road crosses therailway line between Dondo and Inhaminga Pope amp Muller 517(K) Gaza Dist Border Post Malvernia Thompson 2 (K MO)

ZIMBABWE ShanganiBubi District Gwampa ForestReserve Goldsmith 6556 (K MO) Chipinga District E sideof Sabi R at Dotts Drift Goodier 661 (K) NyamandhovuDistrict Pasture research station Plowes 1928 (K) WankieDistrict main camp along Dopi Pan road Rushworth 1308 (K)Wankie District Game Reserve Ngwashla Rd Wild 4759(COIMB K S)

BOTSWANA Kweneng District 15 mi from turn offtowards Ngware Hansen 3251 (K) Mochudi Harbour 6333(BM) North District Chobe Game Scout Camp on Savuti RPope et al 834 (K) Francistown Rand 5 (BM) 162 km N ofNata on road to Kasane Vahrmeijer Stee 3164 (K) NorthernDivision Khardoum Valley 15 km E of SW Africa borderWild amp Drummond 7049 (K)

NAMIBIA Dikundu Gless 11378 (K) c 10 mi from Katimaon road (Finaughtyrsquos) to Singalamwe Killick amp Leistner 3188(K) 3 mi S of Omuramba Mpungu on rd to Tsinsabis17u509S 18u409E Winter 3908 (K) Ndonga Camp at junctionof Omuramba Omatako and Okavango River Winter ampMarais 4618 (K)

SOUTH AFRICA lsquolsquoSouth African Gold-fieldsrsquorsquo 1840Baines sn (K) Zoutpansberg E of Pan Schweickerdt ampVerdoom 552 (K)

Vernacular Name Mozambique Nama-han-ham (Quelimane Grandvaux Barbosa 4267)

Discussion This species may be easily distin-guished by its narrow leaves which often havea distinctly hastate base Basilaminar glands aretypically very small and sometimes absent incontrast to the more conspicuous glands of otherspecies A pair of very small stipels or glandularknobs is usually present at the petiole apex Thespecies has a short wing on each carpel lobe andshort styles that are mostly connate and obsurely 4-lobed at the apex both states intermediate betweenthose found in the other two species of the sectionP corniculata and P procumbens


The species shows considerable variation in leafblade shape and degree of development of thehastate base Leaf blades typically vary from linearto lanceolate or narrowly triangular with anon-hastate or obscurely to prominently hastatebase and a remotely serrulate to distinctly serratemargin Although there is some variation withina collection such as short- versus long-hastate orhastate versus non-hastate base most variationappears to be among collections and some appearsto be geographically based In Botswana Zambiaand Zimbabwe leaf blades are mostly very narrowand linear narrowly lanceolate or very narrowlytriangular in shape while in South Africa andNamibia blades tend to be narrowly triangularLeaf variation is most extreme in Mozambiquewhere collections may have relatively broad leafblades (eg Pope and Muller 517) very large hastatelobes scattered laminar glands on the abaxialsurface large and sometimes up to 4 basilaminarglands and in one collection (Pope and Muller 517)numerous basilaminar glands Basal leaf lobes mayoccasionally be oriented downwards rather thanhorizontally and thus appear sagittate (eg Pope458)

Some of the leaf variation both within andbetween collections may be attributable to ageand position on the plant Stems appear to beintially erect or sprawling from a perennial root-stock becoming long and twining There appearsto be some tendancy for leaves near the plant baseto be broader and those on long twining shoots tobe narrower Basal-most leaves often lack a hastatebase (eg Pope and Muller 517 with elliptic obtuse-based basal leaves and Schinz 895)

The species P hastata was described by Muller(1864) and distinguished primarily based ona scandent habit from P africana then thoughtto have an erect habit Prain (1913 1925)following Muller continued to recognize P hastatabased on scandent stems finely dentate leavessometimes deeply sagittate at the base and eightstamens as distinct from P africana considered atthe time to be an erect plant with coarsely toothedleaves that are never sagittate at their base and 12stamens Pax (1919) correctly recognizing thatthese differences do not hold treated P hastata asa synonym of P africana and the species as havingtwining stems and typically hastate leaf bladesIn addition examination of staminate flowers inthe present study revealed a stamen number(15ndash20) greater than that recorded for eitherspecies

3 PLUKENETIA CORNICULATA Sm Nova ActaRegiae Soc Sci Upsal 6 4 1799 Hedraiostylus

corniculatus (Sm) Hassk Cat Hort Bogor Alt234 1844 Sajorum corniculatum (Sm) DietrSynops Pl 5 331 1852 Pterococcus corniculatus(Sm) Pax amp K Hoffm Pflanzenr IV147IX(-Heft 68) 22 1919mdashTYPE Indonesia Moluc-cas Ambon Island lsquolsquoBagulae Regione Amboi-naersquorsquo Rumphius (specimen presumably lostillustration Tab 79 Fig 2 in Rumphius Herbamboin 1 1741 between pages 193 and 194)Fig 1

Monoecious vine or slender liana stems twin-ing sparsely to moderately pubescent becomingwoody Stipules narrowly triangular or lanceolate12ndash2(ndash28) mm long Leaves evergreen petioles [1- square brackets refer to collections from Timor]2ndash8(ndash13) cm long moderately or sparsely pubes-cent blades ovate triangular-ovate oblong-ovateor elliptic [4ndash] 7ndash16 3 [2ndash] 4ndash11 cm wide thin-chartaceous very sparsely pubescent or glabres-cent with major veins usually sparsely or moder-ately pubescent above acute-caudate at apex withtip 06ndash23 cm long cordate at base with U-shapedsinus [01ndash] 05ndash23 cm long margins finely serrateor rarely serrate venation palmate secondaryveins semicraspedodromous 2ndash3(4) [ndash5] on eachside of midrib tertiary veins percurrent orsometimes reticulate stipels 2 05ndash1 mm longusually strongly curled adaxially basilaminarglands 2 circular or elliptic [03ndash] 05ndash1 mm indiam marginal laminar glands absent Inflores-

cence a narrow raceme 15ndash5 cm long bisexualterminal but appearing leaf-opposed sometimesterminal on short shoots or rarely axillary (someThailand collections) axes puberulous peduncle

FIG 6 Distribution of Plukenetia corniculata


1ndash8 mm long flowers 1 (rarely 2) per nodepistillate flower 1 at basal-most node staminateflowers numerous above bracts narrowly triangu-lar 1 mm long Staminate flower pedicel 13ndash25 mm long glabrescent bud globose obtuse atapex sepals 4 elliptic or ovate 07ndash11 3 05ndash08 mm sparsely pubescent near base acute atapex androecium globose 04ndash06 3 06ndash08 mmcomprising 8ndash14 stamens on small globose orconvex receptacle filaments conical 01 mmlong disc absent Pistillate flower pedicel 15ndash9 mm long sparsely puberulous to glabrescentsepals narrowly triangular or lanceolate 12ndash22 3

05ndash1 mm glabrescent ovary 1ndash25 mm in diam(excluding wings) sparsely or very sparsely pub-erulous with puberulent medial line on each carpelextending along wing margin 4-winged wings 05ndash2 mm long elongating to 13 mm and becomingglabrescent in immature fruit stage styles com-pletely connate into a small depressed-globosecolumn 05ndash07 3 08ndash11 mm glabrous withcross-shaped stigmatic surface at apex Fruit a 4-lobed capsule [5ndash] 7ndash11 3 15ndash20 mm glabrescentsurface irregularly verrucate each carpel lobe withcentral strap-shaped wing 6ndash12 3 2ndash3 mm pedicel[06ndash] 2ndash5 cm long Seeds broadly lenticular 8ndash11 3

65ndash8 3 5ndash65 mm obtuse-triangular in outlinelaterally compressed with radial keel 03ndash07 mmwide surface smooth cream pale orange-brown orpale to medium brown with dark orange or darkbrown irregular markings testa persistent

Pollen Description Gillespie (1994)

Illustration Narasimhan et al (1989) Fig 1

Distribution and Ecology Widespread butuncommon throughout SE Asia and Malesia inNE (Assam) and SE (Andhra Pradesh) India(Narasimhan et al 1989) Burma () ThailandPhilippines Malaysia and Indonesia (Fig 6) Alsorecorded but not confirmed from Sikkim in NEIndia (Grierson amp Long 1987) and Bangladesh(Huq 10780 specimen recorded but not found atL) A slender twining liana found in clearings andother disturbed areas of lowland wet (evergreen)and moist (deciduous) forest 50ndash550 m some-times cultivated Flowering collections were madein June in Bangladesh August in India Decemberin the Philippines July and August on Borneoand throughout the year on Java and SumatraFruiting collections were made in August in IndiaJuly and September in Thailand June on theMalay Peninsula December in the PhilippinesJune and August in Sarawak October in Sulawesiand April to August in Java and SumatraReported to be the larval foodplant of theNymphalid butterfly Ariadne isaeus in Malaysia(Kirton T22)

Representative Specimens Examined INDIA AssamCherrapunjee Khasi Hills Koelz 31112A (L) lsquolsquoUpper AssamrsquorsquoJenkins 517 (K)

THAILAND Northern Phitsanulok Tung Salaeng LuangLarsen 552 (L) Southeastern Chanthaburi Klawng Kloi Kerr9241 (BM K L)

PHILIPPINES Luzon Bataan Prov Ayam River LamaoEdano PNH 4157 (A L) Rizal Prov Ramos Bur Sci 24086(BM BO K L MO NY US)

MALAYSIA PahangTrengganu Kuantan District BukitGaling Forest Reserve Kirton T22 (K) Sabah TamkunanDistrict Gn Trusmadi Mantor SAN 125574 (KEP SAN)Sarawak Baram (4th Division) house compound of LongSelstong Ulu cultivated Chin 2712 (A L) Kapit upperRejang River Clemens amp Clemens 21257 (BO K MO NY)without precise locality Smythies 14073 (K-2)

INDONESIA Java Batavia Depok Bakhuizen van denBrink 5490 (BO G K L) 5771 (BO K L) GoenoengkantjanaDistr Lobakkidoel res Bantam Koorders 40372b (BO L)Koorders 41507b (BO L) Koorders 41509b (BO L) Koorders41720B (BO K L UC) without precise locality Zollinger 3167(BM G G-DC) Kalimantan Hayup Winkler 2295 (L)Kwaru Suwaring Winkler 3096 (L) Sulawesi LelewaoDistrict Preho Kjellberg 2544 (S) Sumatra Sumatera SelatanKomering Oeloe Grashoff 587 (BO L) Sumatera UtaraMiddle Habinsaran near Parsoboeran Lorzing 7821 (L)Sumatera Barat Kampong Sabalading on slope of GunongSago near Pajakumbuh cultivated Meijer 7309 (K L) SoengaiLesing Posthumus 994 (BO L) vicinity of Loemban RiaAsahan (east coast) Rahmat Si Boeea 7593 (A) 7827 (A) SiMandi Angih on the Soengei Kanan Subdiv LaboehanBatoe District Kota Pinang Rahmat Si Toroes 3948 (NY)Timor precise locality and collector unknown (L-0023061 L-0023062 L-0023063 L-0023064 L-0023065 L-0023066 L-0023067)

Vernacular Names and Uses India AndhraPradesh Kodigandlam (Narasimhan et al 1989)Malaysia Sarawak Buah andu (Iban Smythies14073) Buah palidung (Kelabit Christensen amp Apu318) Laot (Kenyah Chin See Chung 2712) Indone-sia Sumatra Gandi riman (Posthumus 994) Painapaina (Grashoff 587) Pepina (Bequin 407) Pina-pinamakanan (Rahmat Si Toroes 3948) Java Arojtangtang anging (Koorders 40372b 41509b) FloresLanteng wase (Schmutz 4178) Sunda region Aroytangtang angien (Hasskarl 1842)

The species is recorded as cultivated in Sarawak(Chin See Chung 2712 Christensen amp Apu 318Smythies 14073) and Sumatra (Meijer 7309) InSarawak young shoots leaves and young fruitsare recorded to be eaten as a vegetable while themature seeds are eaten as nuts and are said to tastesimilar to peanuts (Chin See Chung 2712 Christensenamp Apu 318) It is reported to be cultivated asa vegetable by Dayaks and the distinctive form ofthe fruit is sometimes seen tatooed 5 or 6 ina vertical row on the back of an Iban (Smythies14073) In India a paste made from young leaves isused as an oral laxative medication (Narasimhanet al 1989) In Australia the species was collectedunder cultivation in a garden where it was calledBorneo pea (Waterhuse 5980)

Discussion The species may be easily distin-


guished from other members of P sect Hedraios-tylus by its prominently 4-winged capsule (Figs 1IJ) and larger leaf blades with a distinctly cordatebase (Fig 1A) Leaves are also distinct in their longpetioles (2ndash8 cm long) caudate apex and promi-nent basilaminar glands and stipels Styles arecompletely connate and both stylar column andfilaments are shorter than in the other two speciesThis is the only species of Plukenetia known fromAsia

Plukenetia corniculata was the second species ofPlukenetia after the neotropical P volubilis to beillustrated (Rumphius 1741 as lsquolsquoSajor volubilisrsquorsquo)and formally described (Small 1799) However formuch of its history the species has been knownunder a variety of other names Hasskarl (1842)based his new genus Pterococcus on this speciesbut renamed the species lsquolsquoPterococcus glaberri-musrsquorsquo an illegitimate name being based on Pcorniculata Soon after Hasskarl (1843) createda second genus Hedraiostylus and transferred thespecies naming it lsquolsquoHedraiostylus glaberrimusrsquorsquoagain an illegitimate name Later legitimate com-binations include Hedraiostylus corniculatus Sa-jorum corniculatum and Pterococcus corniculatusthe last used by Pax and Hoffmann (1919) and bymost subsequent authors

The species is most densely distributed on theisland of Sumatra and less so on BorneoDistribution outside this area appears to bescattered and sparse The reason may be that itis rarely collected due to its naturally sparsedistribution in these peripheral areas and its vinehabit An alternative explanation may be that itwas introduced and cultivated andor naturalizedin at least some of these areas for its edible seedMany of the collections from these areas are oldand some are recorded as cultivated All collec-tions seen from Timor (pre-1829 based on deter-minations by A Zippelius) Java (numerouscollections all pre-1923 and all but one likelycultivated) Sulawesi (one collection 1929) andthe Moluccas (type collecton only pre 1800) areold as are most from Assam India (all but onepre-1890 or undated) The record from Burmacould not be verified since contradictory localityinformation is given on the only collection seenGriffith 4716 (Mergui [Burma] on one sheetMalakka [Malaysia] on two sheets at K undated)Four of five collections from Sarawak wererecorded as cultivated or from gardens or housecompounds

4 PLUKENETIA PROCUMBENS Prain Bull Misc In-form Kew 1912 240 1912 Pterococcus procum-bens (Prain) Pax amp K Hoffm Pflanzenr

IV147IX(Heft 68) 23 1919mdashTYPE AngolaBenguella Ganguella on the Cubango River atPrinceza Amelia 1520 m 27 Jan 1907 Goss-weiler 2540 (holotype BM fragment at Kisotype COI)

Monoecious perennial herb with woody root-stock stems procumbent sprawling pubescentStipules small Leaves with petioles 02ndash05 cmlong moderately pubescent blades elliptic orovate 2ndash45 3 13ndash33 cm chartaceous sparselypubescent below glabrous or very sparsely pubes-cent above with major veins sparsely pubescentdrying grey-green with purplish tinge obtuse toshort acuminate at apex rounded to broadlyobtuse at base margin finely serrate venation 3-nerved at base secondary veins brochidodromous2ndash5 on each side of midrib tertiary veins weaklypercurrent or reticulate stipels absent basilaminarglands 2ndash12(ndash20) somewhat irregular in size andshape usually circular or elliptic 02ndash08 mm longoften dark purple usually in an irregular line nearmargin laminar glands absent Inflorescence

a narrow raceme 3ndash8 cm long bisexual terminalbut usually appearing leaf-opposed axis moder-ately pubescent peduncle 2ndash5 cm long flowers 1node pistillate flower 1 at basal-most nodestaminate flowers numerous above on axis 1ndash4 cm long bracts lanceolate 2ndash3 mm long gla-brous or very sparsely pubescent Staminate

flower pedicel 25ndash35 mm long pubescent budglobose broadly obtuse to rounded at apex sepals4 ovate 2 3 1 mm acute at apex androecium1 mm in diam comprising 12 stamens denselypacked on small convex to globose receptaclefilaments slender dilated at base 04 mm longdisc absent Pistillate flower pedicel 3ndash10 mmlong pubescent sepals ovate or lanceolate 25ndash43 1ndash2 mm sparsely pubescent medially ovary1ndash2 3 15ndash25 mm (excluding wings) denselypubescent 4-winged wings to 1 mm long sparselyto moderately pubescent styles 12ndash18 mmlong glabrous partly connate into a column1 mm long 05ndash06 mm wide at base the 4 stylearms divergent 2 mm across at apex each armdilated and recurved with the adaxial surfaceentirely stigmatic Fruit a 4-lobed capsule 6 3

12ndash20 mm sparsely pubescent surface denselypapillose each carpel lobe with central tubercle1 mm long pedicel 11ndash13 mm long Seeds notseen

Distribution and Ecology Angola known onlyfrom the type collection (Fig 5) A perennialwoody-based mostly herbaceous plant with de-cumbent stems collected on clayey soils in shortthickets in the interior highlands 1500m The


single collection with flowers and immature fruitwas collected in January

Discussion This species may be distinguishedby its small elliptic leaf blades with a rounded orbroadly obtuse base Leaves are mostly purplishtinged with small often numerous basilaminarglands and lack stipels Compared with the othertwo species of P sect Hedraiostylus the styles arelonger and only partly connate the inflorescenceshave longer peduncles (2ndash5 cm versus 1 cm) thatare longer than the flower-bearing part of the axisand each capsule lobe bears a rounded tuberclemuch shorter than the wing of the other twospecies

The species was described as a perennial herbwith a woody base and has numerous highlybranched radiating prostrate stems If indeed thespecies has a non-scandent habit this would bedifferent from the twining vine or liana habit of allother species of the genus While stems may besprawling there is no evidence that they aretwining on the single collection known The closestspecies in habit and also likely the most closelyrelated is P africana which has a perennial woodyrootstock and stems that appear to be initially erector sprawling becoming long and twining

Prain (1912) distinguished his new species basedon prostrate habit leaf blades with broadlyrounded base and short petioles He later alsoused inflorescence position terminal versus lateralracemes to distinguish the species from P africana(Prain 1913) However neither inflorescence posi-tion nor petiole length can be used to distinguishthe two species Inflorescences of both species areterminal and may become leaf-opposed althoughinflorescences of P africana are more typically andprominently leaf opposed while those of Pprocumbens appear to become so only at a laterstage and not prominently so This difference islikely due to a difference in habit few long twiningsympodial stems each with numerous inflores-cences in P africana versus numerous shorterhighly branched stems each with few inflores-cences in P procumbens Pax and Hoffmann (1919)used habit as the main character to distinguish thetwo species

Information on the locality of the type collectionis somewhat contradictory though appears to referto the same location The BM specimen gives thelocality as lsquolsquopraesidium of the Princeza AmeliaCubangorsquorsquo the K specimen as lsquolsquopraesidium of thePrinceza Amelia ndash Cubango Ganguellasrsquorsquo the COIspecimen as lsquolsquoVila da Ponte Ganguelas Huilarsquorsquowhile the type description gives lsquolsquoBenguella Gang-uella on the Cubango River at Princeza Amelia1520 mrsquorsquo

MADAGASCAN SPECIES GROUP

The three Madagascan species form a distinctgroup that has yet to be assigned to a section thesingle species known up to now has always beenincluded within Plukenetia Unique among paleo-tropical species they share an androecium ofsessile anthers on a prominent elongate receptacleIn addition the group differs from P sectHedraiostylus in their prominent styles (35ndash16 mmlong) longer than the ovary distinctly carinatefruit and broadly ellipsoid to subglobose seedsand from P sect Angostylidium in the absence ofa staminate disc Their medium-sized fruit (2ndash4 cmwide) and seeds (both unknown for P madagascar-iensis) are intermediate in size between the smallcapsules of P sect Hedraiostylus and large fruit ofP conophora All three species have two conspicu-ous basilaminar glands on the leaf blade adaxialsurface as typical of most Plukenetia species

Characters variable within the Madagascanspecies group include style connation shape andlength androecium shape and size anther numberinflorescence type length and position leaf bladeshape and presence of glandular knobs at thepetioleblade junction

5 Plukenetia ankaranensis LJ Gillespie spnovmdashTYPE Madagascar Prov AntsirananaSpecial Reserve 3 Ankarana 7 km SE ofMatsaborimanga trail between Camp Anglaisand river 3 km SW of Camp Anglais12u559S 49u069E 150 m 28 Nov 1990 Gille-spie 4076 (holotype MO isotypes CANDAV G K L NY P TAN US FAApreserved material at MO) Figs 2 3AndashD

Plukenetia madagascariensis Leandri et P deciduaLJ Gillespie affinis a quibus differt stylis brevior-ibus connatis omnino columna obovata

Monoecious liana base often expanded andbulbous stems glabrous or sparsely pubescentdistally sometimes glaucous older branches be-coming thick woody and often twining Stipulestriangular 05 mm long Leaves with petioles(08ndash)15ndash6 cm long moderately to densely puber-ulous blades ovate oblong-ovate or suborbicular(3ndash)5ndash10 3 (2ndash)4ndash6 cm chartaceous glabrescent tosparsely puberulous with major veins puberulousacuminate at apex with tip 3ndash12 mm long truncateto shallowly cordate at base with sinus to 9 mmdeep margins crenate-serrate venation 3-nerved atbase or weakly palmate secondary veins semicras-pedodromous or sometimes craspedodromous 2ndash4on each side of midrib tertiary veins percurrent orsometimes reticulate stipels and glandular knobabsent basilaminar glands 2 circular (02ndash)04ndash


08 mm in diam marginal laminar glands absentInflorescence a racemose thryse (2ndash)5ndash16 cm longbisexual or sometimes staminate terminal butbecoming leaf-opposed axes puberulous to gla-brescent peduncle usually absent (and thenpistillate flower basal) or sometimes up to 02 cmlong pistillate flowers (0)1ndash2(ndash3) 1 per node atbasal-most node(s) (rarely in basal bisexual cymuleor emerging from stem below inflorescence)staminate flowers numerous above in cymulescymules lax to moderately condensed primarycyme axes (15ndash)5ndash12(ndash22) mm long the longestones rarely subtended by a reduced leaf secondaryaxes to 3 mm long bracts linear-lanceolate linear-triangular or lanceolate sparsely puberulentstaminate 15ndash3 mm long pistillate 15ndash7 mm longStaminate flower pedicel 5ndash11 mm long sparselypuberulous bud ovoid bluntly acute at apexsepals 4 narrowly elliptic or lanceolate 12ndash22 3

04ndash1 mm glabrous or sparsely puberulous whit-ish or pale yellowish green acute and thickened atapex reflexed at anthesis androecium oblong-cylindrical 06ndash1 3 05ndash08 mm comprising 15ndash20sessile yellow anthers densely packed on a stalkedcylindrical receptacle Pistillate flower pedicel 2ndash12 mm long sparsely puberulous sepals narrowlylanceolate 15ndash24 3 04ndash05 mm green glabres-cent ovary 1ndash25 3 15ndash53 mm 4-horned horns05ndash2 mm long styles completely connate into anobovoid or obconical column 35ndash55 mm long06ndash1 mm wide at base dilated to 18ndash25 mm atapex green often somewhat 4-lobed distallystigmas forming a raised circular often somewhatlobed disc at apex Fruit a 4-lobed capsule 16 3

3ndash4 cm green surface verrucose and minutelypapillose with papillae bearing minute hairs eachcarpel lobe carinate keel widened centrally intohorn-like wing 08ndash1 cm long pedicel 09ndash13 cmlong Seeds subglobose 15ndash18 3 16ndash17 3 14ndash17 cm ventral surface distinctly angular dorsalsurface rounded surface (tegmen) smooth tosomewhat verrucate pale brown testa whitishnot or partly persistent

Pollen Tricolpate suboblate (PE 5 071ndash080)polar axis 28ndash33 mm equatorial axis 35ndash45 mmamb subcircular angulaperturate colpus broadwith margins uneven and jagged exine 15ndash2 mmthick evenly thickened or slighter thinner towardsmargin tectum foveolate (vouchers Gillespie 40744076)

Distribution and Ecology Known only fromthe Ankarana Massif in northern Madagascar(Fig 5) where it grows in openings within dryforest on eroded limestone pavement 100ndash200 mA liana with branches that twine on shrubs orsmall trees or sprawl over bare rock pavement

(Figs 3A C) Flowering and fruiting specimenscollected in late November and early December

Specimens Examined MADAGASCAR AntsirananaSpecial Reserve 3 Ankarana 7 km SE of Matsabori-manga trail between Camp Anglais and river 3 km SW ofCamp Anglais 12u559S 49u069E Gillespie 4074 (CAN K MOP TAN US) Gillespie 4075 (CAN MO) Gillespie 4088 (CANK MO TAN) Reserve Speciale Ankarana Lac Vert 7 kmSE of Matsaborimanga 12u559S 49u069E Lees sn (CAN US)southwest of Antsiranana Reserve Speciale de Ankarana12u519S 49u049E Malcomber 1877 (CAN DAV G K MO PTAN US)

Discussion This species was discovered by theauther during a fieldtrip to the Ankarana Massifnear the northern tip of Madagascar The specieswas immediately distinguishable from the onlyspecies then known in Madagascar P madagascar-iensis by its distinct gynoecium with styles entirelyfused into an enlarged obovoid-obconic stylarcolumn (Figs 2E 3B D) Both P madagascariensisand P decidua have partly fused styles anda cylindrical stylar column (Figs 3G 4F) Inaddition P ankaranensis may be distinguished byits terminal leaf-opposed thyrse with moderatelyopen cymes distinctly winged ovaries shorterstyles ( 6 mm long) smaller staminate flowerswith a shorter androecium ( 1 mm long) andabsence of stipels or glandular knobs at the petioleapex Capsules and seeds are larger than those ofP decidua (unknown for P madagasarensis) Pollengrains are smaller than all other examined speciesof Plukenetia The species shares with P madagas-cariensis a similar leaf blade shape inflorescencetype (thryse) and length (5ndash19 cm) and with Pdecidua terminal inflorescences and stamens 30 orfewer

6 Plukenetia decidua LJ Gillespie sp novmdashTYPE Madagascar Toliara entre Ampanihyet Itrobiky route Ampanihy-Androka 4 Jul1958 Capuron SF 18682 (holotype P isotypesP-2 sheets) Fig 4

Plukenetia madagascariensis Leandri affinis a qui-bus differt foliis ovatis vel triangularibus ovatisracemis brevioribus terminalibus floribus nodisinguli bracteis parvioribus eglandulatis antherispaucioribus

Monoecious liana stems glabrous distinctlyglaucous sometimes twining older branches slen-der woody greyish or mottled pale grey and tansomewhat crooked and knobby with raised leafscars Stipules triangular 04ndash1 mm long Leaves

deciduous petioles 15ndash5 cm long glabrous dis-tinctly glaucous blades ovate or triangular-ovatesometimes narrowly so 3ndash6 3 1ndash4 cm chartac-eous glabrous acuminate at apex obtuse truncateor shallowly cordate at base with sinus to 05 cm


deep margins distinctly serrulate venation 3-nerved at base secondary veins mostly semi-craspedodromous 4ndash8 on each side of midribtertiary veins percurrent small glandular knob to06 mm long usually present between and justbelow basilaminar glands sometimes absent basi-laminar glands 2 suborbicular 08ndash13 mm indiam prominent above petiole apex laminarglands absent or occasionally 1ndash3 per side nearmargin towards apex Inflorescence a narrowraceme 3ndash7 cm long bisexual terminal at branchapex or sometimes on very short shoots axesglabrous peduncle usually absent (and pistillateflower basal) flowers 1 per node pistillate flower 1at basal-most node staminate flowers numerousabove bracts triangular-ovate 1ndash2 mm long oftenwith a small lobe on each side margin whitishalong lower half Staminate flower pedicel 4ndash7 mm long glabrous bud ellipsoid obtuse at apexsepals 4 narrowly elliptic 3ndash38 3 15 mm wideacute at apex glabrous open wide at anthesisandroecium ellipsoid or oblong-ellpsoid 16ndash18 3

13ndash15 mm comprising 18ndash30 sessile anthers ona narrowly stalked ellipsoid receptacle anthersdensely packed proximally sparser distally withapex often bare Pistillate flower pedicel 4ndash5 mmlong glabrous sepals narrowly triangular 2ndash25 3

06ndash07 mm glabrous ovary 15ndash25 3 2ndash4 mm(including wings) glabrous 4-winged wings05 mm long styles 8ndash11 mm long glabrouspartly connate into a cylindrical column 4ndash6 3

1 mm the 4 free style arms 37ndash5 mm longtapered apically spreading with their innersurface entirely stigmatic Fruit a (1ndash)4-lobedcapsule 11ndash13 3 23ndash35 cm (immature) gla-brous surface smooth and faintly reticulate eachcarpel lobe carinate keel widened centrally intohorn-like wing to 10 mm long in immature fruit2ndash25 mm long in mature fruit styles to 12 mmlong persistent at least in immature fruit pedicel04ndash08 cm long Seeds broadly ellipsoid 13 3

11ndash12 3 11ndash12 mm ventral surface distinctlyangular dorsal surface rounded surface (tegmen)densely minutely verrucate dull pale orange-brown testa not or partly persistent white topurplish

Pollen Tricolpate suboblate (PE 5 071ndash075)polar axis 32ndash35 mm equatorial axis 44ndash51 mmamb subcircular colpus broad with margins un-even and jagged exine 25 mm thick evenlythickened or somewhat thinner towards margintectum foveolate smooth (voucher Capuron SF18682)

Distribution and Ecology Restricted to thesouthern tip of Madagascar (Fig 5) A slenderdeciduous liana of dry scrub 20ndash250 m Flowering

when leafless or with young leaves Floweringcollections made in July August September andNovember fruiting collections made in August andDecember

Specimens Examined MADAGASCAR Toliara Sourcede Ranomay Tangobory Capuron SF 27952 (P) AntanimoraDistrict drsquoAmbovombe Decary 2929 (P) Ambovombe Decary3253 (P-2 sheets) Bassin inferieur du Mandrare environs deBehara Humbert amp Swingle 5664 (P-2 sheets) ReserveNaturelle Integrale drsquoAndohahela ENE Ihazofotsy24u499000S 46u369360E Rakotomalaza 597 (CAN MO PTAN US)

Discussion This is a distinctly deciduousslender liana (also described as arbuste sarmen-teux) that appears to flower prior to or simulta-neously with the new flush of leaves The distinctlyglaucous branches and narrower ovate or triangu-lar-ovate leaf blades are distinct from the other twospecies in Madagascar The more prominentbasilaminar glands are positioned together at thepetiole apex and usually have a small glandularknob between Shorter narrow racemes with onlyone flower per node and glabrous axes and flowersalso contrast with the longer thyrses staminateflowers in cymules and usually puberulous axesand flowers of the other two species This speciesappears to be most closely related to P madagascar-iensis sharing a very similar style morphology

Inflorescences are generally terminal at the endsof branches but in one collection (Decary 2929) ofa long twining branch they are terminal on veryshort shoots and appear almost axillary (with a leafscar the only evidence of the subtending deciduousleaf)

The collection Rakotomalaza 597 with immatureand almost mature capsules is assigned to thisspecies but differs in two characters from othercollections seen The collection lacks the whitishwaxy bloom characteristic of other collectionsexamined (but preservation in alcohol may havedissolved the bloom) The mature leaves havetruncate to cordate blade bases compared to othercollections which have obtuse-based leaves thatappear to be immature or expanding Otherwisethe collection fits well within this species bothmorphologically and geographically

7 PLUKENETIA MADAGASCARIENSIS Leandri BullSoc Bot France 85 527 Figs 113ndash14 1938mdashTYPE Madagascar Prov Mahajanga Bois deMorataitra rive droite de la Betsiboka E deMaevatanana Mar 1899 Perrier 848 (lectotypedesignated here P isolectotypes L P-2sheets) Fig 3EndashG

Monoecious liana stems glabrous often twiningolder branches thick woody Stipules triangular05ndash08 mm long Leaves apparently deciduous


petioles 25ndash8 cm long glabrous or very sparselypubescent blades ovate or sometimes suborbicu-lar 45ndash105 3 4ndash75 cm chartaceous glabrous orglabrescent slender acuminate to caudate at apexwith tip 07ndash2 cm long cordate truncate orrounded at base with sinus to 1 cm deep marginscrenulate-serrulate venation 3-nerved at base orweakly palmate secondary veins mostly craspe-dodromous 3ndash5(ndash6) on each side of midribtertiary veins percurrent knobs 1 or 2 oftenglandular located between basilaminar glandsbasilaminar glands 2 circular 04ndash1 mm in diammarginal or just above petiole apex laminarglands absent Inflorescence a spicate thryse 6ndash19 cm long bisexual or staminate axillary axesglabrous to sparsely pubescent peduncle 05ndash4 mm long pistillate flowers (0)1ndash2(3) 1 per nodeat basal-most node staminate flowers numerousabove in few-flowered cymules cymules con-densed primary cyme axes 02ndash15 mm longbracts lanceolate or linear-lanceolate 3ndash8 mmlong glabrescent to moderately pubescent atten-uate at apex petiolate biglandular or rarely 4-glandular bracteoles lanceolate or linear-lanceo-late to 3 mm long eglandular or sometimesminutely biglandular Staminate flower pedicel5ndash10 mm long glabrescent to moderately pubes-cent buds narrowly ovoid or elliptic bluntlyacuminate bluntly acute or obtuse at apex sepals4 ovate or lanceolate 3ndash6 3 15ndash25 mm paleyellowish green to cream very sparsely tomoderately pubescent attenuate and distinctlythickened at apex androecium narrowly conicalor narrowly ovoid-conical 3ndash4 3 09ndash12 mmcomprising 35ndash60+ sessile yellow anthers looselypacked on shortly stalked narrowly conical palegreen receptacle Pistillate flower pedicel 2ndash25 mm long sparsely to moderately pubescentsepals lanceolate or ovate 25ndash4 3 14 mmgreen sparsely to moderately pubescent ovary2 mm in diam verrucose pubescent not dis-tinctly winged or horned styles 10ndash16(ndash22 accord-ing to Perrier) mm long green partly connate intoa cylindrical column 6ndash8(ndash12) 3 1 mm the fourfree style arms 4ndash8(ndash10) mm long tapered apicallyFruit and seeds not seen


Illustration Leandri (1938) Figs 113ndash14

Distribution and Ecology Widespread butwith a scattered distribution in western Madagas-car from east of Analalava to just north of Toliara(Fig 5) A slender liana found in open areas in dryforest on limestone or sandy soils (Fig 3E) 20ndash600 m Plants are apparently deciduous during thedry season with new leafy shoots produced inDecember (November to January) at the beginning

of the wet season Flowering specimens werecollected between January and April when eithernew or mature leaves were present

Specimens Examined MADAGASCAR MahajangaTsingy de Bemaraha E Antsalova Cremers 3771 (P-5 sheets)Reserve Naturelle Bemaraha Ambodiriana (campement atriver crossing along Antsalova-Ambondro track) 9 km E ofAntsalova 18u399S 44u439E Gillespie 4144 (CAN MO) 1ndash3 km SW of Antseranandraka along road to Ambarenybetween Lake Bemaraha and Lake Masama 30 km SW ofAntsalova 18u509S 44u249E Gillespie 4175 (CAN K MOTAN) Antsingy drsquoAntsalova Reserve Natural Bemaraha(RN 9 Morat 4893 (P TAN) Haute Sofia pres drsquoAntoakab-ary 600 m Nov 1922 Perrier 15078 (syntype P) ToliaraLambobe River 6 km N of Beroboka 55 km NE ofMorondava 19u549S 44u369E Gillespie 4125 (CAN K MONY P TAN US) Route N9 km 37ndash39 near Andrevo23u039S 43u359E Mabberley 965 (K) Belalanda CommuneRanobe 23u009460S 43u399090E Andrianjafy et al 1648 (Pphotos seen)

Discussion This species may be distinguishedby its elongate androecium (25ndash4 mm long) withmany anthers (35ndash60) and relatively large (3ndash8 mmlong) inflorescence bracts which are often petiolateand biglandular The presence of petiolate bigland-ular bracts is unique in Plukenetia all other specieshaving smaller eglandular usually narrowly tri-angular bracts The species may also be distin-guished from other members of the Madagascanspecies complex by its axillary inflorescences andspecifically from P ankaranensis by its longerpartly connate styles (Fig 3G) and one or twosmall glandular knobs at the petiolebladejunction and from P decidua by its broader leafblades (Fig 3F) longer inflorescences and longerstyles

The species shares with P decidua a very similarstyle morphology of styles connate for one half totwo thirds of their length into a cylindrical stylarcolumn with tapered free style arms (Figs 3G4F) Similar style morphology is present only inthe neotropical species P lehmanniana (styles armsare less tapered and blunt-tipped) (Gillespie 1993Fig 1A) Other paleotropical species withpartly connate styles have either much shorterstyles or mostly connate styles with short dilatedarms

Several characters appear to be somewhat vari-able in the species Inflorescence axes may besparsely pubescent (collections from Bemerahaarea) or glabrous The two basilaminar glands onthe adaxial surface of the leaf blade may be locatedeither along the margin near the base or just abovethe petiole apex with either one or two knobsbetween Further collections are needed todetermine if this variation has a geographicalbasis and to ascertain if this is a single variablespecies or if more than one taxon should berecognized


EXCLUDED SPECIES

PLUKENETIA ZENKERI Pax Bot Jahrb Syst 43 831909 5 Hamilcoa zenkeri (Pax) Prain BullMisc Inform Kew 1912 107 Fig p984 1912

ACKNOWLEDGEMENTS The author thanks the curators atA BM BO CO DAV G G-DC K KEP L MO NY P SSAN TAN UC and US for loans of specimens andor forfacilitating herbarium visits MO and TAN for facilitatingfieldwork in Madagascar Peter Phillipson for providing thephotograph in Fig 3G and for bringing to my attention recentcollections from Madagascar Peter van Welzen forfacilitating my visit to Leiden and helpful comments on aprevious version of the manuscript Geoff Levin and GordonMcPherson for valuable reviews Susan Laurie-Bourque andAnita Walsmit Sachs for providing the line illustrations andthe National Herbarium Netherlands for permission to useAnita Walsmit Sachsrsquo illustration The Netherlands Organi-zation for Scientific Research (NWO visitorrsquos grant B85-369)and the Canadian Museum of Nature are gratefullyacknowledged for financial assistance

LITERATURE CITED

AIRY-SHAW H K 1971 The Euphorbiaceae of Siam KewBulletin 26 191ndash363

mdashmdashmdash 1975 The Euphorbiaceae of Borneo Kew BulletinAdditional Series IV London Her Majestyrsquos StationeryOffice

mdashmdashmdash 1981 The Euphorbiaceae of Sumatra Kew Bulletin 36239ndash374

mdashmdashmdash 1982 The Euphorbiaceae of Central Malesia (CelebesMoluccas Lesser Sunda Is) Kew Bulletin 26 191ndash363

mdashmdashmdash 1983 An Alphabetical Enumeration of the Euphorbiaceaeof the Philippine Islands Kew Royal Botanic Gardens

AKINTAYO E T and E BAYER 2002 Characterization andsome possible uses of Plukenetia conophora and Adenopusbreviflorus seeds and seed oils Bioresource Technology 8595ndash97

AKPUAKA M U and E NWANKWOR 2000 Extractionanalysis and utilization of a drying-oil from Tetracarpi-dium conophorum Bioresource Technology 73 195ndash196

ANIMASHUAN T R A TOGUN and R C HUGHES 1994Characterization of isolectins in Tetracarpidium cono-phorum seeds (Nigerian walnut) Glycoconjugate Journal11 299ndash303

BACKER C A and R C BAKHUIZEN VAN DEN BRINK 1963 Floraof Java Groningen Netherlands N V P Noordhoff

BAILLON H E 1858 Etude general du group des EuphorbiaceesParis Victor Masson

BENTHAM G 1880 Euphorbiaceae Pp 239ndash340 in GeneraPlantarum vol 3 eds G Bentham and J D HookerLondon Lovell Reeve and Co

CHEVALIER A 1948 Une plante oleagineuse africaine (Tetra-carpidium conophorum Hutch et Dalziel) Revue interna-tional de botanique appliquee et drsquoagriculture tropicale 28465ndash466

CROIZAT L 1941 The tribe Plukenetiinae of the Euphorbia-ceae in eastern tropical Asia Journal of the ArnoldArboretum 22 417ndash431

DYER R A 1975 The genera of Southern African floweringplants Pretoria Botanical Research Institute Depart-ment of Agricultural Technical Services

EGHAREVBA R K M I IKHATURA and C KALU 2005 Theinfluence of seed treatments and growing media onseedling growth and development of African walnut

Plukenetia conophorum African Journal of Biotechnology 4808ndash811

GERMISHUIZEN G and N L MEYER 2003 Plants of southernAfrica an annotated checklist Pretoria National BotanicalInstitute

GILLESPIE L J 1993 A synopsis of neotropical Plukenetia(Euphorbiaceae) including two new species SystematicBotany 18 575ndash592

mdashmdashmdash 1994 Pollen morphology and phylogeny of the tribePlukenetieae (Euphorbiaceae) Annals of the MissouriBotanical Garden 81 317ndash348

GRIERSON A J C and D G LONG 1987 Flora of Bhutanvol 1 Edinburgh Royal Botanic Garden

GOVAERTS R D G FRODIN and A RADCLIFFE-SMITH 2000World Checklist and Bibliography of Euphorbiaceae (andPandaceae) Kew The Royal Botanic Gardens

HASSKARL J K 1842 Plantarum genera et species novae autreformatae javenses Flora 25(2) Beiblatt 1ndash56

mdashmdashmdash 1843 Adnotationes de plantis quibusdam Javanicisnonnullisque Japonicus haud rite cognitis e CatalogoHorti Bogoriensis excerptae Tijdschrift voor natuurlijkegeschiedenis en physiologie 10 115ndash150

HEDIN L 1929 Une plant oleagineuse peu connue d lrsquoOuestAfricaine le Tetracarpidium conophorum Revue interna-tional de botanique appliquee et drsquoagriculture tropicale 9752ndash753

HUTCHINSON J and J M DALZIEL 1928 Flora of West TropicalAfrica vol 1 (2) 1st ed London The Crown Agents forthe Colonies

mdashmdashmdash and mdashmdashmdash 1958 Flora of West Tropical Africa vol 1(2) 2nd ed revised by R W J Keay London The CrownAgents for the Colonies

JIMENEZ RAMIREZ J 1993 Especie nueva de Plukenetia(Euphorbiaceae) del estado de Oaxaca Mexico Analesdel instituto de biologia de la universidad nacional autonomade Mexico serie botanica 64 55ndash58

JIOFACK T and J P DONDJANG 2006a Characterisation desdiapores de Tetracarpidium conophorum et effet du modeet de la duree de conservation des diaspores sur lagermination Tela Botanica (on-line articles) httpwwwtela-botanicaorgpagemenu_381

mdashmdashmdash and mdashmdashmdash 2006b Technique de micropropagationde Tetracarpidium conophorum le bouturage Tela Botanica(on-line arcticles) httpwwwtela-botanicaorgpagemenu_382

LEANDRI J 1938 Euphorbiacees malgaches nouvelles recol-tees par M H Perrier de la Bathie Bulletin de la societebotanique de France 85 523ndash533

LEISTNER O A 2000 Seed plants of southern Africa PretoriaNational Botanical Institute

MULLER J 1864 Neue Euphorbiaceen des Herbarium Hookerin Kew auszugsweise vorlaufig mitrgetheilt aus demManusript fur De Candollersquos Prodromus Flora 47465ndash471 529ndash540

mdashmdashmdash 1866 Euphorbiaceae Pp 189ndash1260 in ProdromusSystematis Naturalis Regni Vegetabilis vol 15 (2) ed A Pde Candolle Paris Victor Masson

NARASIMHAN D N RAMA RAO and T RAVISANKAR 1989Two rare interesting taxa of Euphorbiaceae fromAndhra Pradesh India Journal of Economic and Taxo-nomic Botany 13 56ndash59

OBOH G and M M EKPERIGIN 2004 Nutritional evaluationof some Nigerian wild seeds Food 48 85ndash87

ODOEMELAM S 2003 Chemical compositon and functionalproperties of conophor nut (Tetracarpidium conophorum)flour International Journal of Food Science and Technology38 729

OKIGBO B N 1977 Neglected plants of horticultural


importance in traditional systems of tropical Africa ActaHorticulturae 53 130ndash150

PAX F A 1890 Euphorbiaceae Pp 1ndash119 in Die NaturlichenPflanzenfamilien III 3(5) 1st ed eds A Engler and KPrantl Leipzig W Engelmann

mdashmdashmdash and K HOFFMANN 1919 Euphorbiaceae-Plukenetii-nae Pp 1ndash108 in Das Pflanzenreich IV147XI (Heft 68)ed A Engler Liepzig W Engelmann

mdashmdashmdash and mdashmdashmdash 1931 Euphorbiaceae Pp 11ndash233 in DieNaturlichen Pflanzenfamilien vol 19c 2nd ed eds AEngler and K Prantl Liepzig W Engelmann

PHILLIPS E P 1951 The Genera of South African FloweringPlants 2nd ed Department of Agriculture BotanicalSurvey Memoir 25 Pretoria Government Printer

PRAIN D 1912 Diagnoses africanae XLVIII Bulletin ofMiscellaneous Information Kew 1912 224ndash240

mdashmdashmdash 1913 Plukenetia Pp 949ndash952 in Flora of Tropical Africavol VI (1) ed W T Thiselton-Dyer London Reeve amp Co

mdashmdashmdash 1925 Plukenetia Pp 496ndash497 in Flora Capensisvol V (2) ed W T Thiselton-Dyer London Reeve ampCo

RADCLIFFE-SMITH A 1996 Euphorbiaceae Part 1 Pp 1ndash333 inFlora Zambesiaca vol 9 (4) ed G V Pope Kew RoyalBotanic Gardens

mdashmdashmdash 2001 Genera Euphorbiacearum Kew Royal BotanicGardens

RAPONDA-WALKER A and R SILLANS 1961 Les Plantes Utilesdu Gabon Paris Editions Paul Lechevalier

RUMPHIUS G E 1741 Herbarium Amboinense vol 1 Amster-dam apud Franciscum Changuion

SATHE S K B R HAMAKER K W SZE-TAO and MVENKATACHALAM 2002 Isolation purification andbiochemical characterization of a novel water sol-uble protein from Inca Peanut (Plukenetia volubilis L)Journal of Agricultural and Food Chemistry 50 4906ndash4908

SATO S T ANIMASHYUAN and R C HUGHES 1991Carbohydrate-binding specificity of Tetracarpidium con-ophorum lectin The Journal of Biological Chemistry 26611485ndash11494

SMALL J E 1799 Dissertatio botanica de Plukenetia Nova ActaRegiae Societatis Scientiarum Upsaliensis 6 1ndash4

WEBSTER G L 1975 Conspectus of a new classification of theEuphorbiaceae Taxon 24 593ndash601

mdashmdashmdash 1994 Classification of the Euphorbiaceae Annals ofthe Missouri Botanical Garden 81 3ndash32

WHITMORE T C 1973 Tree Flora of Malaya London Long-man


(1975 1994) and Gillespie (1993 1994) Croizatconsidered Pterococcus as part of Plukenetia becausehe could find no differences to separate the twogenera Croizat also remarked that the wingedovaries considered diagnostic for Pterococcus couldbe found in Plukenetia ss Likewise he did notconsider Angostylidium (5 Tetracarpidium) as a dis-tinct genus and pointed out that Pax andHoffmann (1919) although recognizing the genusalso considered it little distinct from PlukenetiaWebster (1975 1994) included both Tetracarpidiumand Pterococcus as synonyms under Plukenetiaa view followed by Gillespie (1993 1994) andRadcliffe-Smith (2001)

Pterococcus was described by Hasskarl (1842) forthe single species Plukenetia corniculata Sm andnamed after its conspicuously 4-winged capsule(Figs 1I J) Subsequently the genus was treatedmostly as a section of Plukenetia but its circum-scription was controversial Baillon (1858) includedonly P corniculata in the genus (which he calledHedraiostylus Hassk) and considered P africanaSond as a member of Plukenetia ss while Muller(1866) included P corniculata P africana and thenewly described species P hastata MullArg (5 Pafricana) in P sect Hedraiostylus (Hassk) MullArgBentham (1880) expanded the circumscription of Psect Pterococcus (Hassk) Benth amp Hook tocomprise in addition to these three species twoneotropical species P penninervia MullArg and Pverrucosa Sm a treatment followed by Pax (1890)In 1919 Pax and Hoffmann resurrected the genusPterococcus and considered it to comprise Pcorniculata P africana (including P hastata) andthe recently described species P procumbens Prain

Pax and Hoffmann (1919) characterized Ptero-coccus based on both style morphology andpresence of wings on the ovaries Likewise Airy-Shaw (1975) considered the genus to be lsquolsquocloselyrelated to Plukenetia differing principally in thevery short thick stylar column and in the conspic-uously 4-winged or 4-horned capsulersquorsquo While Pcorniculata does have a conspicuously four-wingedcapsule (Figs 1I J) P africana has a capsule withfour shorter wings and P procumbens a capsulewith four small tubercles the latter similar to manyneotropical species including P penninervia Psupraglandulosa LJ Gillespie and P verrucosa(Gillespie 1993 Figs 1D 10I) Although not ashighly developed as in P corniculata distinctly 4-winged ovaries and fruit are also found inMadagascan and neotropical species of Plukenetia(eg P ankaranensis Figs 2E F and P loretensisUle) Styles of Pterococcus were described by Paxand Hoffmann (1919) as connate into a short thickcolumn with sessile stigmas as distinct from styles

connate into an urceolate or cylindrical columnwith free arms short or absent (AngostylidiumApodandra Fragariopsis and Plukenetia ss) or stylesfree above (Eleutherostigma) In fact styles are notuniform within Pterococcus but vary from com-pletely connate into a very short stout stylarcolumn (P corniculata Figs 1G H) to partlyconnate into a short narrower cylindrical stylarcolumn with free recurved arms (P procumbens)The latter species does not key out under Pter-ococcus in Pax and Hoffmannrsquos (1919) key butinstead under Plukenetia Therefore neither thestyle nor winged ovary characters used by Pax andHoffmann (1919) to distinguish the genus Ptero-coccus appear to be justified

The only character that could be used to separatePterococcus from other members of Plukenetia sl isstyle length with Pterococcus having styles shorterthan or equal in length to the ovary (Figs 1G H)All other Plukenetia sl (except some collections ofP brachybotrya MullArg) have styles longer thanthe ovary (Figs 2E 3B D G 4F Gillespie 1993Figs 1 9C 10H) Given the extreme variablity ofstyle morphology (size shape and degree ofconnation) in Plukenetia sl (Gillespie 1993) distin-guishing a genus based only on style size does notseem justified Nevertheless the three species ofPterococcus appear to form a natural group whichis recognized here as P sect Hedraiostylus (seediscussion under this section)

Plukenetia sect Angostylidium MullArg wasdescribed by Muller (1864) for his new species Pconophora and was subsequently consistently rec-ognized as a monotypic section (eg Bentham1880 Pax 1890) In 1919 Pax and Hoffmann raisedthe section to generic rank as Angostylidium butsoon after Hutchinson and Dalziel (1928) correctlyrecognized that Tetracarpidium has priority overAngostylidium a treatment later followed by Paxand Hoffmann (1931) Pax and Hoffmann (19191931) distinguished their monotypic genus fromPlukenetia ss based on its high stamen number(40 versus 12ndash30) and presence of intrastaminaldisc segments but also considered it to be littledistinct from the latter Neither character can beused to separate these two genera Stamen numberwas found to range from 25 to 40 in P conophoraMullArg and some Plukenetia ss species havemore than 30 stamens (eg P stipellata LJGillespie has 25ndash40 stamens Gillespie 1993 Fig9D P madagascariensis has 35ndash60+ stamens) Whilespecies of Plukenetia ss either lack a staminate discor have an annular disc (P penninervia speciescomplex Gillespie 1993 Fig 10G) intrastaminaldisc segments similar to those of Tetracarpidium arepresent in the neotropical species P lehmanniana












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Documents

A revision of paleotropical Plukenetia (Euphorbiaceae