BLUMEA 38 (1993) 1-24 A taxonomic revision of WillughbeiaRoxb. (Apocynaceae) D.J. Middleton DepartmentofBotany,TrinityCollege,Dublin 2,Ireland Summary Thegenus WillughbeiaRoxb. isrevised.A totalof 15speciesare recognised.One newcombination is made. Urnularia Stapfisreduced tosynonymy within Willughbeia.Speciesexclusae have been givenaswell asanindex ofexsiccatae. Introduction Willughbeia was first describedby Roxburgh (1819) froma specimen collectedin Chittagong, Bangladesh. However, thename turnedoutto be a laterhomonym of Willughbeia Scop. (1789) andtheorthographic variantWillughbaeya Necker(1790). Thelatter is asynonymof, andhasbeenrejected infavourof, Mikaniain theCom- positaeandtheformeris asynonymofAmbelaniaAubletin theApocynaceae. Wil- lughbeia Scop, has subsequently beenrejected infavour ofWillughbeia Roxb. fol- lowing therecommendationby Green (1935). The 'genus' Willughbeia Klotzsch (1861) hasnobasis as theauthorpublished threenew species whichheerroneously referredto Willughbeia and did not give arevised description ofthe genus.These havesubsequently beenreducedto species ofSabaPichonand Ancylobothrys Pierre. Wallich(1832) proposed thenameAncylocladus toencompass a possible new genuscontaining both Willughbeia andPacouria.As thesegeneraremaindistinctand therulesofprioritydonotallowforthereplacement ofoldernamessimply because generahavebeen united, thenameAncylocladus is superfluous. Urnulariais not sufficientlydistinctfromWillughbeiatoremainseparate. Markgraf (1972) distinguished thetwogeneraonthe grounds thatWillughbeia hasacylindric corollatubeand congested inflorescenceswhichare shorterthan thepetioles whilst Urnulariahasaninflatedtubeandelongated inflorescences, normally longer than the petiole. However,the inflatedcorollatubeis partly aresultofthe shorterlength in Urnulariaas thespeciesofbothgenerahavetubes inflatedaroundthestamens. Wil- lughbeia anomalaMarkgrafhasinflorescencesconsiderably longer thanthepetioles although Markgrafsuggests thattheinflorescenceis morerobustinthisspecies com- pared to thosespecies normally includedin Urnularia.However,thisisnotparticu- larlymarked. There arealsoanumberofotherspecies wheresomeindividualshave elongated inflorescences, particularly in W. tenuiflora Dyer ex Hook, f., W.cirrhi- fera Abeywickr. andW. coriacea Wallich. Willughbeiaangustifolia (Miq.) Markgraf 2 BLUMEA Vol. 38, No. 1, 1993 has thedelicateinflorescencesand shortinflatedcorollatubesnormally characteristic ofUrnulariaspecies. However, it also has a very shortinflorescence. Willughbeia cirrhiferaalso hasaninflatedcorollatubebut usuallyhas therelatively shortrobust inflorescenceofWillughbeia sensu stricto. Otherdifferences, mentionedbyvarious authors, instigma shape, stigmoid apexlength andcalyx dissection, were all found not tobe differentbetweenthetwo genera.Thestigmaand stigmoid apex characters are sharedby W.angustifolia andthecalyx charactersimply doesnot holdup. Urnu- lariaStapfis a conservednameagainstthefungus genus UrnulariaP. Karsten. The generic nameWillughbeiopsis wasproposed as asubstitute byRauschert (1982) be- foreUrnulariaStapfwas conserved. Willughbeia hasconsistently beenplaced inthetribeCarisseae(Bentham& Hooker, 1873;Hooker, 1882; Pichon, 1948, 1950, 1953; Leeuwenberg, 1983). In addition Pichon(1948)placed itinthesubtribeWillughbeiinae, initiallyonitsown (although herecognised bothWillughbeia andUrnularia),butlater(Pichon, 1953)includingthe monotypic African genusCylindropsis whichheremoved fromtheLandolphiinae. Leeuwenberg (1983) maintainedthis arrangement. Thegreatestproblem inthegroupis amongstthosespecies formerly in Urnularia. Willughbeia javanica, W.lanceolata, W.ovatifolia, W. beccarianaandW. flavescens areclosely relatedandidentificationofsterilematerialis notalways easy. Thereare only limitedcollectionsoffertilematerialforthesespecies andW.ovatifolia isknown fromonly onecollection. Furthercollections, particularly inBorneo, Sulawesi, Su- matraand Java,are needed. No chromosomenumbers are reported for species ofWillughbeia but, where known,allothermembersoftheCarisseaearex= 11(VanderLaan&Arends, 1985). MATERIALS AND METHODS Herbariummaterialwas studiedfromthefollowing herbaria:A,AAU,ABD,B, BKF, BM, BO,BR, BRI,C, CAL,E,FI, G, GH,K, K-W, KEP,L, M, MEL,MO,P, PNH, S, SING,TCD, U,US,Z(Holmgren etal„ 1990). Thedimensionsgiven inthe descriptions arefordriedmaterialexceptforandroe- ciumand gynoecium characterswhichare measuredfromflowersboiledin water to reconstitutethem. SYSTEMATIC TREATMENT WILLUGHBEIA WillughbeiaRoxb.,PI.Corom. 3 (1819) 77,t. 280,nom. cons., non Scop. (=Ambelania Aublet); G.Don,Gen. Syst. 4 (1837) 101; A.DC.,Prod.8 (1844) 321;Benth. & Hook.f., Gen. PI.2 (1873) 691;K. Schum. in Engler& Prantl, Nat. Pflanzenfam. 4, 2(1895) 130;Pichon,Mem. Mus.Nat. Hist. Nat. n.s. 24 (1948) 153. WilloughbeiaHook. f„Fl.Brit. India 3 (1882)623, orth. var. Willoughbyaauct.plur., orth. var. Willughbya Merr.,J.Str. Br.Roy. As.Soc. spec, no.(1921)495,orth. var.Willughbeiasubg.EuwillughbeiaKing& Gamble,J. As.Soc.Beng. 74 (1907)391. Pacuria sect. Willoughbya(auct.) Kuntze inPost& Kuntze,Lex. (1904)412. Ancylocladus Wallich,PI.As. Rar. 3(1832) 45; Kuntze, Rev. Gen. PI. 1 (1891)412; Pierre, Bull. Soc. Linn. Paris II, 1(1898)94.AncistrocladusPierre, Bull. Soc. Linn. ParisII, 1(1898) D.J. Middleton: Revision of Willughbeia 3 100,lapsunonWallich. Ancylocladussect.EuancylocladusPierre,Bull. Soc. Linn. ParisII, 1 (1898) 94. Willughbeiasect.Euancylocladus (Pierre)K. Schum. in Engler& PrantI,Nat. Pflanzenfam.Nachtr. 2(1900) 54. — Ancylocladus sect.HypoancylocladusPierre,Bull. Soc. Linn. ParisII, 1(1898)97. Willughbeiasect. Hypoancylocladus(Pierre)K. Schum. in Engler &PrantI,Nat. Pflanzenfam.Nachtr. 2(1900) 55. Pacuria sect. HypoancylocladusKuntze in Post& Kuntze,Lex. (1904) 412. —Ancylocladus sect.CyclopholisPierre,Bull.Soc.Linn. Paris II, 1 (1898)98. Willughbeiasect.Cyclopholis (Pierre)K.Schum.in Engler&PrantI,Nat. Pflanzenfam.Nachtr. 2(1900) 55. Urnularia sect. Cyclopholis (Pierre)Pichon, M6m.Mus. Nat. Hist. Nat. n.s. 24 (1948) 155. —Urnularia Stapf,Hook. Ic. PI. (1901) t.2711,nom. cons.;Pichon, Mem. Mus.Nat Hist. Nat. n.s. 24 (1948) 154. Willughbeiasubg.Urnularia (Stapf)King& Gamble,J.As. Soc.Beng.74(1907)391. Urnularia sect. UrnulaPichon,Mem. Mus.Nat. Hist. Nat. n.s. 24 (1948) 155. WillughbeiopsisRauschert, Taxon 31 (1982) 556, nom. rejec.—Typespecies: Willughbeiaedulis Roxb. Woody climbers, producing latex. Branches lenticillate, glabrous or puberulent, bearing tendrils formedfrom modifiedinflorescences.Leaves opposite, thoseofa pair equal,entire, petiolate; colletersabsentfromleafaxils; bladepaperytocoriace- ous. Inflorescence ofaxillary and,rarely, terminalcymes,often very short, appear- ing fasciculate; axes puberulent or glabrous; bracts small, ovateoroblong. Flowers 5-merous, actinomorphic. Sepals connate atthebase; lobesfreeand erectorslightly reflexed; colleters absent. Corolla:lobesinbud overlapping to theleftforminga cone or cylinder oferectlobes;tube cylindric, somewhatinflatedaroundthestamens,or shortand inflated; lobes spreading and ovate, elliptic or oblong. Stamen insertion variable,completely includedwithinthetube;freefromthepistilhead;filamentvary- ing in length; anthersovateto lanceolate, apex acute or obtuse,baserounded.Disk absent.Ovary single, unilocular,with 2parietal placentas; glabrous; superior to semi- inferior; style columnar; stigma ellipsoid or cylindric; stigmoid apex as long as or longerthanthestigma. Fruitafleshy berry; spherical, ellipsoid orpear-shaped; few tomanyseeded; indehiscent.Seedcompressed ovoid; withoutacoma; smooth, with avery thinendosperm andthick homy cotyledons. KEY TO THE SPECIES la. Corollatubeinflated, usually <3mmlong 2 b. Corollatubecylindric, inflatedonly around stamens,> 3mmlong 8 2a. Peduncle brownpuberulent; Java 10. W. javanica b. Peduncle glabrous orsparsely whitepuberulent; not inJava 3 3a. Corollatubepuberulent within,only weakly inflated,lobescoming toapoint in bud; stigma well developed. Sri Lanka 4. W. cirrhifera b. Corollatubeglabrous within, strongly inflated,lobesforming acylinder inbud; stigma inconspicuous. Notin Sri Lanka 4 4a. Inflorescenceaxis shorterthanoras long as subtending petiole 1. W. angustifolia b. Inflorescenceaxis longerthansubtending petiole 5 5a. Tertiary leafvenationlargely obscure; leaves glaucous beneath; corollalobes 3-3.5 mm long 13. W. ovatifolia b. Tertiaryleafvenationclearly visible; leavesnot glaucous beneath; corollalobes 1.4-2.8 mm long 6 4 BLUMEA Vol. 38, No. 1, 1993 6a. Lateralnervesinleafalmostperpendicular tomidribwith1-3intercalatedveins between; leaves dullabove andbeneath 7. W. flavescens b. Lateral nerves in leafbent strongly forwards or, ifnot, then without inter- calatednerves andshiny above 7 7a. 18-30flowersperinflorescence; leaves dullabove, 1.4-2.8x as long as wide 3.W. beccariana b. 8-15flowersperinflorescence; leavesshiny above, 2.3-3.8x as long as wide 11. W. lanceolata 8a. Inflorescencebrownpubescent; calyx lobesbrownpubescent 9 b. Inflorescenceglabrousorwhitepuberulent; calyx lobesnormally glabrous 12 9a. Inflorescenceaxis longerthanthesubtending petiole; leavesdullochrebeneath when dry 2. W. anomala b. Inflorescenceaxis shorteror as long as the subtending petiole; leavesnot dull ochrebeneathwhen dry 10 10a. Leaves with 1 or2 intercalcatedveins parallel to the lateralnerves; stamens inserted in middle oftube 12. W. oblonga b. Leaves withobscure tertiary venationorwithveins perpendicular tothelateral nerves; stamens insertedbelowmiddleoftube 11 11a. Corolla tube> 7.5 mm long, lobes > 8 mm long; stamens inserted in lower quarteroftube; fruitpear-shaped; leavesshiny above,venationmostly obscure 15. W. tenuiflora b. Corollatube <6.5mmlong, lobes<6.5 mmlong; stamens insertedatabouta thirdoftheway upthetube;fruit spherical; leaves dullabove, venationclearly visible 14. W. sarawacensis 12a. Leaves withdistinctintercalcatedveins betweenthe lateralnerves 13 b. Leaves withindistinctvenationorwithveinsperpendicular tolateralnerves and nointercalcatedveins 14 13a. Inflorescenceaxis as long as or longer than the subtending petiole; pedicels glabrous; corollatube < 3.5mm long, lobes< 4.5 mmlong; leafapex long acuminate or subcaudate.Sri Lanka 4. W. cirrhifera b. Inflorescenceaxis mostly shorterthan subtending petiole; pedicels puberulent; corolla tube > 3.8 mm long, lobes > 4.5 mm long; leafapex acuminate to rounded. India to SE Asia 6. W. edulis 14a. Leaves glaucous beneath;corollatube9.7-15mm long, lobes 14-24mmlong 9. W. grandiflora b. Leaves notglaucous beneath;corollatube3-8 mmlong, lobes4-10 mmlong 15 15a. Leaves thickly coriaceous;tertiary venationclearly visibleperpendicular to and connectedbetween nerves; calyx lobes< 1 xas long as wide 8. W. gigantea b. Leaves paperytocoriaceous; tertiary venationfaintorobscure; calyx lobes> 1 x as long as wide 5. W. coriacea 1. Willughbeia angustifolia (Miq.) Markgraf Willughbeiaangustifolia(Miq.)Markgraf, Blumea 20 (1972)414.—VaheaangustifoliaMiq.,Fl. Ind. Bat.2(1857)394.—Type:Diepenhorst2088(Ulecto;L iso). D.J. Middleton: Revision of Willughbeia 5 Willughbeia apiculataMiq.,Sum. (1861)227,551;Boerl., Handl. 2(1899) 392; Bull. Inst. Bot. Buitenzorg 5 (1900) 6; Hallier,Jahrb. Hamb. Wiss. Anst. 17, Beih. 3 (1900) 144; Posth., Leidsche Geol. Meded. 5(1931) 503;Heyne,Nutt.PI.Ned. Indie ed.2(1927) 1272.—Type: Diepenhorst2140 (Ulecto;Liso). Willughbeia rufescens DyerexHook, f.,Fl. Brit.India 3 (1882)326, syn. nov.;King& Gamble, J.As.Soc. Beng.74 (1907) 398;Ridley, Fl.Mai. Pen. 2(1923)325. —Ancylocladus rufes- cens(Hook, f.)Kuntze,Rev.Gen. PI. 1 (1891)412.—Urnularia rufescens (Hook,f.)Stapfex S.Moore,J. Bot.63,Suppl. (1925)67; Markgraf, Blumea 20 (1972)409. —Willughbeiafla- vescensvar.rufescensRidley, Fl. Mai. Pen. 2(1923)325.—Willughbeiopsisrufescens(Hook,f) Rauschert,Taxon 31 (1982)556. —Type: Maingay1092 (Klecto;Kiso). Willughbeia elmeriMerr.,Univ. Calif.Publ. Bot. 15 (1929)253,syn. nov.;Tsiang,Sunyatsenia2 (1934)94; Markgraf,Blumea 20 (1972)414.—Type:Elmer21038(BM lecto;A,BR,G,GH, K,L,M, MO,P,S, U, Ziso). Willughbeiaangustifoliavar. graciliorMarkgraf,Blumea 20 (1972)414,syn.nov. —Type:Endert 3562(L lecto;A,Kiso). Woody climberto60m. Branchletsglabrous, veryrarely minutely puberulent; lenti- cillate.Leaves', petiole 0.4-1.7cm long; bladeelliptic, ovate oroblong, apexobtuse to acuminate, baseroundedto cuneate; 1.5-4.2xas long as wide,2.6-14.3 x0.9- 4.4cm; subcoriaceaoustothickly coriaceous; glabrous; 9-24pairsoflateralnerves at60-85°,reaching margin oranastomosing shortly beforeit, tertiary venationof1, rarely to3, intercalcatedveinsandthen withfurtherreticulatevenationoralmostob- scure. Inflorescence axillary, rarely upto 3inoneleafaxil; axisshorteror as long as subtending petiole, to 1.7 cm long; axes glabrous; 5-19 flowersperinflorescence; pedicel 0.8-3.7 mm long. Sepals ovate, apex obtuse to acuminate; 0.9-1.8 mm long, lobes0.5-1x0.4-0.7mm, 1-2.5 xas long as wide; glabrous, ciliate.Corolla whiteorgreenish; tubeinflated, 1.2-3mmlong,outside andinsideglabrous; lobes oblong or elliptic, 1.7-4.5 mm long, glabrous. Stamens inserted at 0.6-1.7 mm frombase, 0.40-0.63of tube length; filaments0.3-0.6 mm long; anthers 1.3-1.8 x as long as wide,0.4-0.9 x0.3-0.5 mm, ovate to elliptic. Ovary 0.5-0.8 mm long; style 0.3-0.5 mm long, not impressed on ovary; stigma 0.1-0.2 mm long; stigmoid apex0.1-0.2mmlong.Fruitspherical; 1.9-9.9cm diameter; pale green, yellow, orangeorreddish. Seed 1.2-2.4x0.6-1.4 x0.6-1.1 cm. Distribution- SouthernThailand,MalayPeninsula,Borneo,Sumatra,Bum,Nico- barIslands. Notes- Thisspecies is very variableinleafshape, sizeand venation.Itcanreadi- ly be distinguished fromtheother species of Willughbeia by its combinationofa small inflatedcorollatubeandshort delicateinflorescence. Fourspecimens, two in Land two in U, havethe same collection number(HB 990). They represent, however, two distinct collections from Sumatra, one ofW. angustifolia andoneofW. coriacea.FurthermoreBlumehimselfis not reported to havecollectedinSumatra(Van Steenis-Kruseman, 1950). Geographicalselection ofthec.80 specimensstudied: INDIA.GreatNicobar Island:LafulForest,Hore 8720(L). THAILAND. Pattani,Betong,Kerr7683 (BM,E,K,L). MALAYSIA. Peninsula: s.l..Maingay 1092 (K, typeof W. rufescens). Johor: Tg. Penawar, Cockburn FRI7626 (KEP,L). Kelantan: Ulu SungeiAring, SungeiTapah, Cockburn FRI7175 (L, SING). Negri Sembilan: Tampin,Burkill 1304,3206(K,SING).Pahang: Bukit Teresch,Ta- moraNegara,Kengetal. 42 (SING).Penang:Balik Pulau,Ridley s.n.(SING). Perak: King'sCol- 6 BLUMEA Vol. 38, No. 1, 1993 lector 7848 (K). Selangor: BukitLagong,Jaamat 47060 (KEP). —Sarawak: Bintulu, Similajau ForestReserve,BrunigS 8637(L); Baram,SungeiTutoh,Chew WeeLek1089 (A,SING);Laiiibir NationalPark,Miri,Lee S46452 (L); UluMayeng, Kakus,LuangS21822 (SING); Kalabit High- lands,Hills nearKampongPaLungan,Nooteboom & Chai 2087(L, US);Kapit, SungeiMengiang Balleh,Ismawi & JugahS 29608 (L). Sabah: TawauDist.,Kalabakan ForestReserve,Bakar SAN 24957(L);Tawau. Dist.,Elphinstone Prov., Elmer 21038 (A,BM, BR,G, GH,K.L,M. MO, P,S, U, Z; type of W. elmeri),Elmer 21486 (A,B, BM,BR, GH, L, MO,P,S, U, Z); Lamag Dist.,Abai Kinabatangan,BanangSAN 51951 (SING); LamagDist.,Lake below GunongLotung Inorat,Gibot SAN83410(L); Kuala Penyu, KepaganForest Area,Amin& HeyaSAN86325(L); Sandakan Dist.,SepilokForestReserve, HujongTaping,Ampon& Aban SAN 73681 (K,L); San- dakan Dist., Bukit Senilakan,Ulu Mananam,Meijer 51234 (K,L); Sandakan Dist.,Kabili Forest Reserve, Puasa 4856(A, SING,US); Sandakan Dist., Jalan HujongTanjong,Sikar SAN 39803 (L);TongodDist., BukitTinker, Kuala Keramut,Sundaling SAN96666 (A,L);PaparDist.,Man- dahan,Talib Bidin 80658(C); KualaPengu Dist.,Kepayan,TalipSAN50985 (L);SipitangDist., SibuluRiver,3.5miles SSWofSipitang, WoodSAN 15258(A,L,SING). SINGAPORE.Changi,Ridley 3996(K,MEL);SungeiJurong,Ridley6047(BM); Chanchu Kang, Ridley6143 (BM); MandaiRoad,Kiah37135(A,KEP). BRUNEI.AshtonBRUN845 (L,SING); Ulu Belalong,Temburong,AshtonBRUN440,447(L, SING); Belait Dist.,Sena, vanNiel4274 (L); AndalauForestReserve, Sinclair & Kadim 10449 (A,E,L,SING); BadasForest Reserve, Vermeulen1240(L). INDONESIA.Kalimantan: Tengah:BukitRaya, Veldkamp8594(L,US).Timur: Longtesak,Wiri- adinata 1215 (AAU,L). WestKutai,nearMtKemul,Endert3562(A,K,L,type ofW.angustifolia var. gracilior),3854 (B,L); EastKutai,G.TepianLobang,NE ofSangkulirang, Kostermans6037 (L). —Sumatra. Atjeh: GunungLeuser Nature Reserve, de Wilde & de Wilde-Duyfjes13635 (K). Bangka:Lobok-besar,Kostermans & Anta 129 (A, K,L,SING), 234 (A,L), 404 (A,K,L,SING). Barat:Padang,SungeibulaBeccari 938(BM,FI,L, MEL);Priaman,Diepenhorst2140(L,U, type ofW.apiculata),2088(L, U,typeofVaheaangustifolia). Selatan:Palembang,RawasRiver,Forbes 3232(A,L).—Buru.WaeLang,Nooteboom 5180 (L). —Ambon. Hitoe,Warburg17462(E). CULTIVATED. Java:BogorBotanicGarden X.C.42.Aa(L). 2. Willughbeia anomala Markgraf WillughbeiaanomalaMarkgraf, Blumea20(1972)415. —Type: Wood,Smythies& Ashton SAN 17525 (Lholo;BRI,Kiso). Woody climberto20m. Branchlets glabrous orsparselyrufouspubescent; branches lenticillate. Leaves: petiole 0.7-1.6 cm long;bladeelliptic oblong, apexacuminate, base cuneate;2.6-4.2xas long as wide,6-18.5x2.1-4.9 cm; papery to subcoria- ceous; glabrous or witha few brownhairson midribbeneath, dullochre beneath whendry; 12-23pairs oflateralnerves at 55-80°, reaching margin, tertiary vena- tionfaintandperpendicular tonerves, occasionally withoneintercalcatedvein.In- florescence axillary and terminal, axis longer than subtending petiole, to4cm long; 4-19flowers perinflorescence; axes rufouspubescent; pedicels 1.7-3.8mmlong. Sepals ovate, apex rounded, thick; 1-1.4mm long, lobes 0.5-0.7 x0.3-1 mm, 0.6-2x as long as wide; rufouspubescent, ciliate.Corollatube cylindric, 10-14 mmlong, rufouspuberulent in 5rows down tubeor glabrous, pubescent inthroat; lobes oblong orelliptic 6-8.2 mmlong, ciliateor not.Stamensinsertedat 1.1-1.7 mm frombase, 0.11-0.14oftube length; filaments0.4-0.8 mmlong;anthers 2.6- 2.8 x as long as wide, 1.3—1.4 x 0.5 mm, lanceolate. Ovary 0.6-0.9 mm; style 0.1-0.2 mmlong, impressed on ovary; stigma 0.3-0.4 mm long; stigmoid apex 0.5-0.6mmlong.Fruit spherical; 6.6 cm diameter. Seed2.6 x 1.5 x 1.3cm. Distribution- Borneo,Philippines(Mindanao). D.J. Middleton: Revision of Willughbeia 7 Specimensstudied: MALAYSIA.Sarawak: Santubang,Sinclair 5578 (E), 38326(K); Bau,KampongKerokong,Oth- manet al.S 37544(L, MO);Kuching,G. Matang,Smythies 12502 (L,SING). BRUNEI.KualaBelait,AndalauFor. Res. (mislabelledSabah),Woodetal.SAN 17525(BRI,K, L, type),NangkatNN175 (K),AshtonBRUN 600 (K,L). PHILIPPINES. Mindanao: ZamboangaLay s.n. [BM;the label saysonly 'Philippines' but Van Steenis-Kruseman (1950) suggeststhat theabove site was hisonly collecting locality in thePhilip- pines]. 3. Willughbeia beccariana (Kuntze ex Pierre) K. Schum. Willughbeiabeccariana (KuntzeexPierre)K.Schum. inEngler&Prantl,Nat. Pflanzenfam.Nachtr. 2(1900)55.—Ancylocladusbeccarianus Kuntze,Rev.Gen.PI. 1(1891)412,nom. nud.;Pierre, Bull. Soc. Linn.Paris II, 1(1898)98.—Urnularia beccariana (KuntzeexPierre) Stapf,Hook. Ic.(1901)t. 2711: 1; Markgraf,Blumea 20 (1972)409. —Willughbeiopsisbeccariana (Kuntze exPierre) Rauschert,Taxon 31(1982)556. —Type:Beccari 3764 (Klecto;FI,Piso). Woody climberto 35 m.Branchletsglabrous; densely lenticillate.Leaves: petiole 0.8-2.4 cm long; bladebroad elliptic, short acuminate, base cuneate to rounded; 1.4-2.8 xas long as wide,7.7-17.5 x 2.5-7.2 cm; subcoriaceous to coriaceous; dullabove andbeneath; glabrous; 8-15pairs ofstrong lateralnerves at 40-70° reaching margin, tertiary venationalmostperpendicular tothemidrib, oblique to the nerves, several in each space. Inflorescence axillary, sometimes2 inan axil; axis longer thanthe subtending petiole; to5 cm long; 18-30flowers perinflorescence; axes glabrous or sparsely and minutely puberulent; pedicel 1.4-5.5 mm long. Sepals ovate, apex rounded; 1-2 mmlong, lobes0.6-1 x0.9-1 mm, 0.67-1 x as long as wide; glabrous, ciliate.Corollatube inflated, 1.8-2.8mmlong, glabrous outsideandinside; lobesovate,obtuse,2-2.8mmlong, glabrous. Stamensinserted at 1.3-2.2mm, 0.57-0.71of tube length; filaments0.3-0.9 mm long; anthers 1.4-3x as long as wide, 0.7-0.9 x0.3-0.5 mm,lanceolate to ovate. Ovary 0.9- 1.1 mm long; style0.5-0.9mmlong, not impressed onovary; stigma0.1 mmlong, inconspicuous; stigmoid apex 0.1-0.2mmlong.Fruitspherical; 3-7 cm long,2.2- 7 cm wide; green,yellow ororange.Seed 1.1-1.7x0.7-1.1x0.6-0.7 cm. Distribution- Borneo,Sulawesi. Specimens studied: MALAYSIA.Sabah: MtKinabalu: TenomPok,Clemens 26209A (BM,G,L); MtKinabalu,Fos- ter-Puasa3607 (K); Penatoram basin,Clemens 40197(L); Lubok Dorat,Weron,SandalingSAN 78124 (K); Kota Kinabalu Dist.,Meijer33528(K,L); Tawau Dist., Mile 13,A.PasRoad,Singh & Nardin 48484(a)(L); Sandakan,Kretam For. Res.,Amin et al.SAN 96750 (K).—Sarawak: s.l.Beccari 3764(FI, K, P,type);AlongSungeiNiah,NiahNat. Park,4th Div., Yii Puan Ching S 40112 (K, MO); Ulu Kelawit,Tatau,Ashton S 16468(K,SING); Baram,Hose 24 (K,P);Ulu Sungei Belaga,7th Division,Othmanetal. S43489 (K). BRUNEI. Simpang 370,Jalan Muara, Wong& Siow 557 (K,L); TemburongDist.,Forman & Blewett 956 (K); BukitPuan,Sungei Belait,Sinclair 10518(E,K,L, SING); Belait Dist.,Labi, KampungTeriam,NangkatNN164 (K). INDONESIA.Kalimantan:Timur: Berau,MtNjapaonKelaiRiver,Kostermans 21354(A.D.- Sulawesi: Utara: BolaangMongondow,DumogaBone N.P.,Toraut Dam,de Vogel& Vermeulen 6511 (K,L); DumogaBoneN.P.,Doloduo,TumokangLama Whitmore & Sidiyasa3462(K); Du- moga BoneNationalPark,GunongMogogoipa,deVogel& Vermeulen 7008(K). CULTIVATED. Java:BogorBotanic Garden,vanRomburgh 10(P). 8 BLUMEA Vol. 38,No. 1, 1993 4. Willughbeia cirrhifera Abeywickr. Willughbeia cirrhifera Abeywickr., Ceylon J.Sci.,Biol. Sci. 2 (1959) 84;Huber,Fl. Ceyl. 1 (1) (1973)8; ibid. 4(1983)33.—Chilocarpus ceylanica Wight,Ic. PI.Ind. Or. 4(2) (1848) 1, t 1288.—Willughbeia ceylanica Thw. non Sprengel,Enum. PI. Zeyl.(1860)191;Hook, f., Fl. Br.India3 (1882)624;Trimen,Handb. Fl.Ceyl. 3(1895) 123 (as W. zeylanica).—Ancy- locladus ceylanicusKuntze,Rev.Gen.PI. 1(1891)412.—WinchiacirrhiferaGardnerin Thw., Enum. PI.Zeyl.(1864) 191,in synonymy ofWillughbeiaceylanica. —Type:Gardner550(K lecto; BM,BR,K, Piso). Woody climber.Branchesglabrous,lenticillate.Leaves:petiole 0.7-1.5cm long; blade broadelliptic toobovate, apexlong acuminatetosubcaudate,basecuneate toobtuse; 1.6-2.8xas long as wide,5.4-15 x2-6.9cm; coriaceous; glabrous; 22-34pairs oflateralnerves, almostreaching margin,notparticularly distinctfromslightly weaker parallel secondary veins andfurtherreticulate venation.Inflorescence axillary, axis as long as or longer than subtending petiole, to 4.5 cm long; 3-10flowers perin- florescence; axes glabrous; pedicels 2-4.4 mmlong. Sepals ovate,apexroundedor obtuse; 1.8-2.5mm long, lobes 1.1-2x 1.1-1.3 mm, 0.7-1 x as long as wide; glabrous, ciliate.Corolla tubeweakly inflated, 2.2-3.5 mmlong,outsideglabrous, insidepuberulent; lobesin budacuminate, 3.9-4.5mmlong, endsacute, glabrous. Stamensinserted at 1.3-2.1mm frombase, 0.43 oftube length; filaments0.7-1.2 mm long; anthers 2.3-2.8x as long as wide, 1-1.1 x 0.3-0.4 mm, lanceolate. Ovary 0.7-1 mmlong; style 0.5-1.1 mm long, not impressed on ovary; stigma 0.3-0.5 mm long; stigmoid apex 0.5-0.9 mmlong. Fruit spherical 4.3-15 cm diameter; yellow, tinged red. Seednot studied. Distribution- Sri Lanka. Specimensstudied: SRILANKA. KandyDist., Hantane,Gardner 550 (BM,BR,K,P); Deltota,Thwaites 1829(B, BR,G, K, MEL,P); ErutnagodanearKuruwita,Huber 17(BM, K, US);Galle Dist., Kanneliya Forest,Huber 62 (BM, K, US); Southernslope ofHaycock,Huber 64 (US); SinharajaForest, Kostermans 26688 (G,L); across Kanneliya River near Kitulgalle,Kostermans 28366 (K,L), 28419(L). CULTIVATED.Java:BogorBot.Gard.,Merrills.n.(US),Spire 86(P). 5. Willughbeia coriacea Wallich Willughbeiacoriacea Wallich,PI.As.Rar. 3 (1832)45; G.Don, Gen. Syst. 4(1837) 102;A.DC., Prod.8(1844)321;Miq.,Fl.Ind. Bat.2(1857)391;Hook.f„Fl.Br.India 3(1882)623;Ridley, Fl.Sing. (1900) 82; King& Gamble,J.As. Soc. Beng. 74 (1907) 393; Ridley,Fl. Mai. Pen. 2(1923)323;Tsiang,Sunyatsenia2(1934)93; Markgraf, Blumea 20(1972)412—Ancylocla- dus coriaceus(Wallich) Kuntze,Rev.Gen.PI. 1(1891)412.—Type; Wallich1620(K-Wholo). WillughbeiafirmaBlume,Mus.Bot. Lugd.Bat. 1(1850) 154;Miq.,Fl. Ind.Bat. 2(1857) 390; Hook,f.,Fl.Br.India3(1882)624;Stapf,Trans. Linn. Soc.Lond.,Bot. 4(1894)207;Ridley, Fl. Sing.(1900)82; Boerl.,Bull. Inst. Bot.Buitenzorg5 (1900)4;King& Gamble,J.As. Soc. Beng.74 (1907)394;Ridley, J.Str.Br. Roy. As.Soc. 59 (1911) 129;Fl. Mai. Pen. 2(1923) 323; Tsiang,Sunyatsenia2 (1934)93; Kerr,Fl.Siam. En. 2(1939)424; Backer & Bakh. f., Fl.Java 2(1965)224. —Ancylocladusfirmus (Blume)Kuntze,Rev. Gen. PI. 1(1891)412.— Type:Korthals 1042 (Lholo). Willughbeiafirmavar.oblongifoliaBlume,Mus.Bot.Lugd.Bat. 1 (1850)154.—Type:Korthals s.n.(Lholo). D.J. Middleton; Revision of Willughbeia 9 WillughbeiaburbidgeiDyer,KewGardensReport (1880)44.—Type:Treachers.n.(Kholo). Ancylocladusvriesianus Pierre,Bull. Soc. Linn. Paris II, 1(1898)95. — Willughbeiavriesiana (Pierre)K.Schum. in Engler& Prantl,Nat. Pflanzenfam.Nachtr. 2(1900)55.—Type: de Vriese s.n. (Llecto;Piso). Ancylocladus minutiflorusPierre,Bull. Soc. Linn. Paris II, 1(1898) 95.—Willughbeia minuti- flora (Pierre)K. Schum. in Engler& Prantl,Nat. Pflanzenfam. Nachtr. 2(1900)55.—Type: Beccari 4030 (Klecto;FI,Piso). AncylocladusnodosusPierre,Bull. Soc.Linn. ParisII, 1 (1898)96.—Willughbeianodosa (Pierre) K. Schum.in Engler& Prantl,Nat. Pflanzenfam.Nachtr. 2(1900)55.—Type:Beccari 1530 (Klecto; FI,M,Piso). Willughbeiafirmavar. macrophyllaBoerl.,Bull. Inst.Bot.Buitenzorg5 (1900)4.—Type:vanRom- burgh54(BO holo;TCDphoto). Willughbeiafirmavar.platyphyllaBoerl.,Bull. Inst. Bot.Buitenzorg5(1900)4.—Type:vanRom- burgh9(BO lecto;BOiso;TCDphoto). Willughbeiafirmavar.obtusifolia Boerl.,Bull. Inst. Bot.Buitenzorg 5(1900)4.—Type: vanRom- burgh48(typenotlocated). Large climberto30m.Branches mostlyglabrous, veryrarelypuberulent; lenticillate. Leaves: petiole0.6-4cm long;bladeelliptic, ovate,oblong orobovate,apex shortly acuminatetoacute,base cuneate to rounded; 1.6-5.2xas long as wide,3.3-29 x 1.4-10.4cm; paperytocoriaceous; shiny aboveandbeneath; mostly glabrous, very rarely puberulent onmidribbeneath; 6-20pairsoflateralnerves at50-80° curving towardsbutnot reaching margin, tertiary venationmostly obscure, sometimesfaint andthenperpendicular tothenerves.Inflorescence axillary,veryrarely terminal;axis usually shorterthan thesubtending petiole, veryrarely longer, to 3 cm long; 3-25 flowers per inflorescence; axes mostly glabrous, rarely minutely and sparsely pu- berulent; pedicel 0-5.3cm long. Sepals oblong orovate, apexrounded or obtuse; 1.5-4 mm long, lobes 0.8—3.3x 0.6—1.9 mm, 1.1-2.5 x as long as wide; gla- brous, ciliate.Corollawhite or yellow, rarelytinged withred;tubecylindric, 3.5-8 mmlong,outside glabrous, inside puberulent; lobes oblong, 4-10mm long, ciliate or not. Stamensinsertedat 1.5-2.7mm frombase, 0.31-0.51oftubelength; fila- ments 0.4-0.7 mm long; anthers 2-4x as long as wide,0.8-1.2 x0.2-0.5 mm; lanceolate. Ovary 0.3-1.1 mm long; style 0.4-1.1 mm long, not impressed on ovary; stigma0.2-0.5 mmlong; stigmoid apex 0.3-0.9 mm long. Fruitspherical, pearshaped, oblong orellipsoid; 2.6-12cm long, 1.7-7 cm wide;green,yellow or orange.Seed 1.3-1.8x0.7-1.3x0.7-1.1cm. Distribution- SouthernThailand,MalayPeninsula,Sumatra, Java, Borneo. Ecology - Climber inprimary orsecondary forest to 1600m. Geographicalselection ofthec.220specimensstudied: THAILAND. SuratThani:Khao Nong,Kerr 13234(AAU, BM,E, K, P). Krabi: NongKhon, Sangkhachand1031 (K,L). Nakhon SiThammarat:Khao Luang,vanBeusekom&Phengkhlai793 (E,K,L); Lansagah,Gahrome Falls,Ramsri 102 (A,L). Pattani: BanangSta.,Kerr 7415 (BM.E, K,L), 7386(A, ABD,BM,E, K,L,P,TCD). LAOS.Pak Munung,WiengChan,Kerr21206 (BM,K,L). MALAYSIA.Peninsula:Penang: s.l„Ridley s.n. (Z),Curtiss.n. (SING), 1500(CAL,SING); Wellesley, Askey 1613(SING); GovernmentHill Road, Curtiss.n. (SING); GovernmentHill, Ridley847 (SING); Balik Pulau,Ridley s.n.(SING); PenangHill,Ridley9359 (SING).Kelantan: Tawah Mervah,Muton 94531 ((KEP); Pasir Lalut,K.Lebir, Gimlette s.n. (SING). Perak: s.l., Wray 3028(SING), 4176(CAL,K,SING),King'sCollector10050(SING), 10854 (Z);Lenggong, 10 BLUMEA Vol. 38,No. 1, 1993 Chan FRI 13348 (SING); Kensing, Wray 59 (SING); Larut,Waterfall Hill,Wray s.n. (SING), 5 (K, SING); Larut, King'sCollector 5331 (K,MEL,P);Taiping Hill,Haniff& Nur 2396 (SING); Hutan Simpan TanjongTuallang, T.& P. 1066 (L,SING). Pahang:S.Sat,Ulu Tembeling,Hen- derson21955 (SING).Selangor:SelangorForest,Burkill529(SING);KualaLumpur,BukitLagong, Achmad 99030 (K,KEP,L); Tanah Merah,Kemahan For. Res., SowKEP 94531 (K,L, SING); BulohFor. Res.,Kochummen FRI 16041 (A, K,L,SING); Ulu Langat,BukitTangkol,K. Pan- som,Umbai KL 1585(A,SING). Trengganu:Kemaman,Symington 26968(KEP); Kuala Treng- ganu, David208(P). NegriSembilan: Haynesetal. 1699 (KEP);PortDickson,PursegloveP.4343 (K, L, SING), Rantau,Alvins 2300(SING). Malacca: s.l.,Ridley 399 (K, L). Johor: Summit of GunongPanti,Maxwell 81-127(AAU,L, SING); AyerPanas,Curtis 3485 (K, SING), 3476(K, SING); Ma'okilFor. Res.,Shah MS 3676(KEP,SING).—Sarawak: s.l.Beccari 1530(FI,K, M,P, type ofAncylocladus nodosa),4030 (K, G, P,type ofAncylocladus minutiflora); Lawas River, Menoongan,Treacher s.n. (K, type ofW.burbidgei); DulitRange,S TinjarAwa& Yii S 46789 (K); Kuching,Haviland 2168 (BM,SING), Haviland & Hose3492 (BM,GH, K,L); Kuching, SemengohArboretum,Kudi S 31888(K,L,SING); Similajan For. Res.,Burley& Lee 270 CL, SING);SungeiTanPurseglove P.5422(A,L,SING); BaseofBukitLambir,Miri,OthmanHaron S21383(A, K,L,SING); Path fromKpg SeropaktoBugohRange,Bau,Paie& Mamit S 29599 (A, E,K,L,SING); SantubongFoothills,Chew WeeLek1433 (A,AAU,K,L,SING);KapitDist., BktRaya,Soepadmoet al.S27634(A,K,L,SING);KapitDist.,Foothills ofBukit Batu Tibang, Anderson & Paie S.28509 (A, E,K,L); KapitDist,SungeiBalangAnderson & Paie S 28884 (A, K,L); MlGading, Lundu,Haviland 989 (BM,K,SING); Kalabit Highlands,Batu Lawi,Noote- boom & Chai 2333 (B,L,US); ApaBatu Buli,Nooteboom & Chai 2185 (L,US); Bukit Salong, Ulu Sampuran, Melinau,Kapils, Manis & Salang S 41631 (K, L); SungeiMedamit,Limbang, Wright& Ismawi S 32207 (K, L,SING). —Sabah: Temburong,Kuala Belalong,Smythies et al. SAN 17074 (A,K,L); Tawau, HapSeng,Sri Usukan,Fedilis & SumbingSAN 88475 (L), SAN 89116 (L); Tawau, Kalabakan Wood SAN A4041 (A,L, SING); KgKayo Madang,Evangelista 2534(272) (SING); Lahad Datu,PulauKerning, Kretam Beasr River,Kretam, Wood SANA3500 (L,SING); MtKinabalu, Dallas,Clemens 26063 (A,G,L,SING); Kinabalu,Penibukan,Clemens 31026 (A, B,BM,G,K,L); Kinabalu,below path toRanau,Carr SFN 27086 (SING); Kinabalu, Mesilau River, Chew & CornerRSMB 4888 (K, SING); Sandakan, Ramos 1755 (GH,L, US), 1756 (A,K), 1757 (P),1758(A,BM,SING). SINGAPORE,s.l., Ridley 2712 (BM,L,MEL),Gandogers.n. (MO). MacRitchie Reservoir, Maxwell 76-789 (AAU, L), 77-101 (AAU,L), Sinclair 7997(E),4906 (E), 40219 (E,L); Bukit TimahReserve,Leeuwenberg13343(L, WAG);Changi,Ridley4431 (BM,K, MEL). BRUNEI. Ulu Senuko, Labu,Ashton BRUN 3349 (L,SING). INDONESIA.Kalimantan: Timur:BelayanR. nearLongBleh,Kostermans 10377 (L),10403 (L), BelayanRiver,G.Kelopok,nearTabang,Kostermans 10595(K,L), 10653(A,K,L,SING);7km NofSamarinda,Leeuwenberg&Rudjiman 13032(L).Selatan: BankofR.Lawa,Winkler3152 (K, G,L,P,SING,Z);Tanah LautDist.,Hutan Kintop,Leeuwenberg&Rudjiman 13384(L).Tengah: BukitRaya,Nooteboom 4355(A,L). Barat:Endert 4991 (L);Headwaters ofSKahayan,5km NE ofHaruwu Village,Burleyet al.649 (A, E,L,SING). —Simeulue: Tapah,Achmad1652 (L).— Sumatra: s.l.,Korthalss.n. (L, typeof W.firma var.oblongifolia);Korthals 1042 (L, typeofW. firma); de Vrieses.n.(L, P,typeofAncylocladus vriesianus), 4b (L); IIB990 (seenote under W. angustifolia). Utara: Asahan,HutaPadang,Krukoff4329(A,BR,G,L, MO,SING,US). Riau: P. Lingga,Pasirpangarayan,Biinnemeijer6972 (K,L,P,U); vicinityofTaloen naoeli, Toba,Rahmat si Boeea 11091 (L). Barat:Pajakumbuh, MtSago,Ichlas 43 (K,L),Meijer 4664 (L); Sijunjung, HB 1168(U). Selatan:Palembang,Grashoff886(L);Palembang,Macara Bliti,Karehakar exHeyne s.n.(L);Palembang,LematangVeloe,Lambach1355 (L). Bangka: Oedioel10(L); Kekoelek5(L); G. Maras,Kostermans & Anta 1343(K,L,P,SING).Lampung: WayKambas,Mochtar s.n. (L). — Java:s.L,MEL 1605384.Barat: Banten, Hinterland ofSanghijang,Adelbert 295(G,K,L,P, SING, US). CULTIVATED. Java: cultivated,Schiffner 19(L, Z);Kawakami 20388(A). Sumatra:Lampang, Leembraggens.n. (K).