A revision of Jatropha (Euphorbiaceae) in Malesia - Naturalis ...

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Blumea 62, 2017: 58–74 ISSN(Online)2212-1676www.ingentaconnect.com/content/nhn/blumea https://doi.org/10.3767/000651917X695421RESEARCH ARTICLE

INTRODUCTION

The genus JatrophaL.hasrecentlygainedincreasedinterestof the general public due to one of its species, J. curcas L.,ofwhichtheoilintheseedsisanewsourceofbiofuel(e.g.,Berchmans&Hirata2008).Jatropha is a widely distributed genus,rangingfromtropicalAmericatoAfricaandIndia(Deh-gan&Webster1979).InMalesiathemembersofthisgenusare introduced and were formerly used for medicine and as hedgerows, nowadays they are mainly used as ornamentals (J. gossypiifoliaL.,J. integerrimaJacq.,J. multifidaL.,J. podagricaHook.)(labelinformation,Burkill1935,Heyne1950),orforbiofuel(J. curcas).Especially,J. curcas and J. gossypiifolia escaped from cultivation and established themselves over thewholeofMalesiaandhavethepotentialtobedamaging,invasivespecies(Sweetetal.,unpubl.ms.).Jatropha gossypiifolia is already considered a medium impact invasive weed in Australia(Thorpe&Lynch2000)andspeciesforitsbiologicalcontrolareevaluated(Heardetal.2002).Webster(1994,2014)andRadcliffe-Smith(2001)classifyJa tropha in the subfamily Crotonoideae tribe Jatropheae.Intheskeleton phylogeny of the EuphorbiaceaebyWurdacketal.(2005)Jatropha ispartofcladeC1(theirf.4),togetherwithmembers of tribe Crotoneae (sensuWebster 1994, 2014,Radcliffe-Smith2001).MüllerArgoviensis(1866)providedafirstinfragenericclassi-fication,basedonworkofBaillon(1864),recognizingthreesec-tionsofwhichtwoweresubdividedintotwosubsections.Hestillincluded Cnidoscolus as the third section, now regarded as a distinctgenus(e.g.,McVaugh1944,Radcliffe-Smith2001,Web-ster2014).Pax(1910)madeamuchmoreelaboratedivisionwith subgenera, sections and subsections, but also included Cnidoscolus.ThemostrecentinfragenericclassificationisbyDehgan&Webster(1979),whichwasmoreorlessconfirmedbyaphenetic (and lessbyan,onmorphologicalcharactersbased,phylogenetic)analysisbyDehgan&Schutzman(1994).Dehgan&Webster(1979)basedthemselvesonPax(1910)and

alsousedsubgenera,sectionsandsubsections(butexcludedCnidoscolus).OftheMalesianspeciesonlyJ. curcas is part of subg.Curcas(Adans.)Paxsect.Curcas(Adans.)Griseb.Theotherfourspeciesareclassifiedinsubg.Jatropha.Withinthelatter subgenus J. gossypiifoliaispartofsect.Jatrophasubsect.AdenophoraePaxexDehgan&G.L.Webster(nom.inval.,mustbesubsect.Jatropha);J. integrifoliaispartofsect.Polymorpha Paxsubsect.Polymorphae(Pax)Dehgan&G.L.Webster,and,finally,J. multifida and J. podagricaareplacedinsect.Peltatae (Pax)Dehgan&G.L.Webster.AfirstattempttocircumscribeMalesiaasaregionwasmadebyZollinger(1857;seealsoLam1937,Raes&VanWelzen2009).VanSteenis (1948,1950)was thefirst toprovideproof thatMalesiaisaphytogeographicalregion,whichwasconfirmedbyVanWelzenetal. (2005). In1951VanSteenis launchedhis ambitious international project to revise all plant genera inMalesia,the‘FloraMalesiana’project,that,todate,isstillrunning.Thepresentrevisionisacontributiontothisproject.

CharactersThe character states mentioned in this chapter only pertain to theMalesiantaxa.Anelaborateandexcellentdiscussionofthemorphology of JatrophacanbefoundinDehgan&Webster(1979).

HabitDehgan&Webster(1979)describeJ. gossypiifoliaasa‘facul-tativelyannual,herbaceoussubshrubwithwoodystem’astheplantmoreorlessdiesbackeveryyearingreenhouses.Outsidegreenhouses, under tropical conditions, the species is truly perennial and just like J. integerrimaashrub.Jatropha podagricaischaracterizedbyDehgan&Webster(1979)asan‘erectfleshysubshrubwithwoody-succulent,above-groundcaudexandbranches’, thisspecies typicallyhasabroad,succulentflask-like stem-base apically tapering into a far more slender upperstempartwithaterminaltuftofleavesandinflorescences.Jatropha curcas and J. multifida can become higher shrubs to smalltreesupto10mhigh.

IndumentumMostofthespeciesareglabrousorhaveinvariouspartsanindumentumofmainlysimplehairs.OnlyJ. gossypiifolia has long and very distinct glandular trichomes all over the plant,

A revision of Jatropha (Euphorbiaceae) in MalesiaP.C.vanWelzen1,2,F.S.T.Sweet1,F.J.Fernández-Casas3

1 NaturalisBiodiversityCenter,researchgroupBiodiversityDiscovery,P.O.Box9517,2300RALeiden,TheNetherlands;

correspondingauthore-mail:[email protected] InstituteofBiologyLeiden,LeidenUniversity,P.O.Box9505,2300RALeiden,TheNetherlands.

3 Campanario,Badajoz; formerly:Real JardínBotánico,CSIC,PlazadeMurillo2,E-28014Madrid,Spain;e-mail:[email protected].

Key words

Euphorbiaceaeintroduced speciesinvasive speciesJatrophaMalesiarevision

Abstract Jatropha, a widespread, species rich genus, ranges from the Americas and Caribbean to Africa and India.InMalesiafivespeciesoccur,allofwhichwereintroducedandoriginatedinCentralandSouthAmerica.Thefivespeciesarerevisedandanidentificationkey,nomenclature,descriptions,distributions,ecology,vernacularnames,usesandnotesareprovided.SpecialattentionisgiventotheusesofJ. curcas, because it is steadily gain-ingpopularityasapotentialbiofuelplantand,becauseofthat,isbeingcultivatedmoreoften.

Published on 20 April 2017

59P.C.vanWelzenetal.:A.revisionofJatropha inMalesia

which come in two types, branching trichomes with an elongated glandular head or mushroom-like, non-branching ones with a broad,horizontalglandular,capitatehead.

Leaf morphologyThe stipules are ovate or triangular in J. curcas and J. integerrima,andcanbeearlycaducous.InJ. multifida and J. poda grica the stipules are branched into narrow lobes, in J. gossypiifolia the stipules are dissected and resemble the branching glandulartrichomes.All the species have simple, alternate leaves that are lobed (3–13lobes),butsometimesthelobesarenotobvious(J. inte gerrima).Jatropha integerrima and J. multifida have small side-lobes, J. integerrima mainly at the base of the blade, and J. mul tifidaalternatelyalong the lobes.Themarginsaregenerallyentire, with J. gossypiifolia and J. multifidabeingexceptionstothat.Jatropha gossypiifoliahasafinelyserratemarginwithsimple and glandular, unbranched hairs, and J. multifida has oneormore side-lobes.The speciesare basally palmatelyveined, thesecondaryveinsalongthemidrib(of thecentrallobe)loopandanastomosenearthemargin,thehigherorderofveinsarereticulate.

InflorescencesThe inflorescences are usually subterminal and cymose, often corymbiform, consisting of usually compound cymes or varia-tionswithreductions.Typicalforthegenusisthateveryprimarybranch ends with a pistillate flower, the lateral flowers in the cymesaregenerallystaminate.The basal bract is by far the largest and often they have similar hairs or glandular hairs as the normal leaves, sometimes they alsohave stipule-like lateral structures.Upwards thebractsdecreaseinsize.

Staminate and pistillate flowersTheflowersare5-merousandactinomorphic.ThesepalsofthestaminateflowersareconnateintoacalyxinallspeciesexceptJ. gossypiifolia;thoseofthepistillateflowersarealsoconnateinallspeciesexceptJ. podagrica.Thesepalscanhaveentireorserratemarginswithglandularhairs.Thepetalmarginsareentire.Thediscis5-lobed.Jatropha curcas and J. integerrima have10stamens(twowhorlsof5),theotherspecieshave8(outer,basalwhorl5,inner,upper3),especiallytheinneronescan be united into an androphore, but the stamens are free in J. multifida and J. podagrica.Theanthersare(dorsi)basifixedand open through a lengthwise slit per theca, either latrorse orextrorse.Therearetwothecae,whichareoftenbasallydi-varicate.Thestaminateflowerslackapistillode.Thepistillateflowersaregenerally(2–)3(–4)-locular,withasingleovuleperlocule, and the style is short and the three stigmas usually have a lower, narrow, unreceptive part and a broadened and largely split,upperreceptivepart.

Fruits and seedsThefruitsareallellipsoid,smoothcapsules(regmas),usuallyslightly3-lobedandoftenwiththreegroovesalongtheseptaandthreeslightgrooves(orthickenings)overthelocules.Thewallsarethin,atmost1mmthick.FruitdehiscenceisloculicidalinJ. curcas, septicidal and partly loculicidal in J. gossypiifolia and J. podagrica, and only septicidal in J. integerrima and J. mul tifida.Theexpulsionoftheseedsfromtheloculeisperformedexplosivelyinsomespeciesbytwistingtheloculewall,aidedbyathickeningoftheradialwall,alsocalledacallus.Thegloboseto ellipsoid seeds are glabrous and possess an apical, often multilobatecaruncle.

TAxONOMIC TREATMENT

JatrophaL.

JatrophaL.(1753)1006;(1754)437;A.Juss(1824)37;Baill.(1858)294;Miq.(1859)391;Müll.Arg.(1866)1076;Kurz(1877)402;Benth.(1880)290;Hook.f. (1887) 382;Pax (1910) 21;Gagnep. (1926) 323;Pax&K.Hoffm.(1931)160;McVaugh(1945)271;Backer&Bakh.f.(1963)494;AiryShaw(1972)283;(1975)137;Dehgan&G.L.Webster(1979)1;AiryShaw (1982) 25;Grierson&D.G.Long (1987) 790;Radcl.-Sm. (1987)343;G.L.Webster(1994)103;Philcox(1997)83;Govaertsetal.(2000)1017;Radcl.-Sm.(2001)288;Chantharaprasong&Welzen(2007)343;LiBingtao&M.G.Gilbert(2008)268;G.L.Webster(2014)125;Fern.Casas(2016) 2.―Jatropha L. subg.Jatropha subsect.Jatropha:Dehgan&G.L.Webster(1979)39.―Conservedtype(Wiersemaetal.2015:238):Jatropha gossypiifoliaL.

CurcasAdans.(1763)356;Baill.(1858)313;Britton&Millsp.(1920)224.―CurcasAdans.sect.EucurcasBaill.(1858)314,nom.inval.―Type:Jatropha curcasL.(Adansondidnotprovideaspeciesname,thecombina-tion Curcas adansoniiEndl.exHeynh.wasmadelater).

CastiglioniaRuiz&Pav.(1794)139.―Type:Castiglionia lobata Ruiz&Pav.[= Jatropha curcasL.]

MozinnaOrtega(1798)104;A.Juss.(1824)35;Hook.(1841)t.357.―CurcasAdans.sect.Mozinna (Ortega)Baill. (1858)315.―Type:Mozinna spathulata Ortega[=Jatropha dioicaSessé].

LoureiraCav.(1799)17.―Lectotype(designatedbyDehgan&Webster1979:47):Loureira glanduliferaCav.[=Mozinna cordataOrtega=Jatropha cordata(Ortega)Müll.Arg.].

Adenoropium Pohl (1827) 12.―Jatropha L. sect.Adenorhopium (Pohl)Griseb(1859)36;Müll.Arg.(1866)1082;Benth.(1880)291.―Lectotype(designatedbyDehgan&Webster1979:39):Adenoropium gossypiifolium (L.)Pohl[=Jatropha gossypiifoliaL.].

ZimapaniaEngl.&Pax(1891)119.―Type: Zimapania schiedeanaEngl.&Pax[=Jatropha dioicaSessé].

CollenuciaChiov.(1929)177.―Type:Collenucia paradoxaChiov.[=Jatropha paradoxa (Chiov.)Chiov.].

(DescriptionbasedonMalesianspeciesonly)Largeherbstoshrubstotreelets,monoecious,protogynous;taprootthick,long. Indumentum absent, of simple hairs or glandular.Stipules distinctornot,simpleorsplitmultiple times.Leaves simple, alternate,eglandularexceptsometimesforhairs;petiolesnotpulvinate;bladesoftenpalmatelylobedto-partitewithlobedsegments,marginentiretoundulatetofinelyserrate,withsim-pleandglandularhairsontipswhenserrate;venationatleastbasally palmate, bronchidodromous, anastomosing, veinlets re-ticulate.Inflorescences usually terminal, cymose, often corymbiform, with a pistillate flower ending every primary branch, more lateral flowers staminate; bracts elliptic or triangular,sometimes narrow, margin entire to serrate, becoming smaller upwards.Flowersunisexual,5-merous,actinomorphic;sepals5, oftenbasallyunited,imbricate;petals5,freeoradnate,contort,glabrous,butinsomespecies(partly)hairyinside;discglands5,alternatingwiththepetals.Staminate flowers: sepal margin serratetoentire;petalmarginentire;stamens8or10intwowhorls, outer5, inner5or3, filaments freeorpartlyunited(especiallyinnerwhorl)inanandrophore,anthers(narrowly)elliptical to triangular, (dorsi)basifixed, opening latrorse toextrorsevia lengthwiseslits,2-thecate, thesebasallyhardlyto distinctly divaricate; pistillode absent.Pistillate flowers: sepals, petals and disc glands like in staminate flower, ovary (2–)3(–4)-locular,withasingleovuleper locule;styleshort,stigmas3,oftenresemblinganthersandusuallydividedintoanarrowunreceptivepart(filament-like)andareceptive,almostcompletelysplit,broadenedandthickenedpart(anther-like).Fruits globulartoellipsoid,capsular,slightly3-lobed,dehiscingeither only septicidally, only loculicidally or completely septici-dallyandpartlyloculicidally;wallthin,atmostc.1mmthick.Seeds glabrous;caruncle2-tomultifid. Distribution―AccordingtoGovaertsetal.(2000)agenusofc.190speciesinthetropicalandsubtropicalregionsoftheAmericas (incl.Caribbean),Africa (incl.Madagascar) up to

60 Blumea–Volume62/1,2017

Fig. 1LeafshapesandflowerdetailsinMalesianJatrophaL.–J. curcasL.:a.Leaf.–J. integerrimaJacq.:b.Leaf.–J. multifidaL.:c.Leaf.–J. podagrica Hook.:d.Leaf.–J. gossypiifoliaL.:e.Habit;f.staminateflower;g.androeciumwithdiscglands;h.pistillateflower;i.baseofovarywithdiscglands;j.fruit,k.columella;l.seedwithapicalcaruncle(a:Bullock 879;b:Noerkas 62;c:Anonymous s.n.,barcodeL0034581;d.M. Reekmans 7729;e–g:F.G. Dickason 8644;h,i:A.M. Huq & M.K. Mia 10498;j– l:A.H.M. Jayasuriya 1248;allL).—Drawing:AnitaWalsmitSachs,2006.

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India.Severalspeciesare introducedthroughout thetropicsworldwide,fivespeciescultivatedinMalesia,oftenescapingandpossiblyinvasive.

KEy TO THE SPECIES

1.Branches,petiolesandleafbladeswithbranchingandun-branched glandular hairs . . . . . . . . . . . . 2.J. gossypiifolia

1.Branches,petiolesandleafbladeswithoutglandularhairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Leafbladepeltate . . . . . . . . . . . . . . . . . . . 5.J. podagrica2. Leafnotpeltate,petiolebasallyattachedtoblade. . . . . . 33.Petiolehairy(usemagnification);bladesnotlobedorwith

small, short basal lobes . . . . . . . . . . . . . . 3.J. integerrima3.Petioleglabrous;blades3–13-lobed . . . . . . . . . . . . . . . . 44.Stipulesveryearlycaducous,broadlyovate,entire,densely

hairy.Leaves3-or5–(7-)palmatilobedlobesjoineduptohalfway. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.J. curcas

4.Stipulespersistent,dissectedintomanyflagelliform,glabrousfilaments. Leaves (6–)9–11(–13)-palmatifid, lobes onlybasally joined . . . . . . . . . . . . . . . . . . . . . . . . 4.J. multifida

1. Jatropha curcasL.―Fig.1a,2;Map1

Jatropha curcasL.(1753)1006;Blanco(1837)759;(1845)522;Miq.(1859)392;Müll.Arg.(1866)1080;Kurz(1877)403;Blanco(1879)160,t.384;Hook.f.(1887)383;Thell.(1908)784;Pax(1910)77,f.30;Merr.(1917)324;(1918)229;(1923)449;Gagnep.(1926)324;McVaugh(1945)283, f.7,8,23;Corner(1951)259,pl.59;Backer&Bakh.f.(1963)494;AiryShaw(1972)283;(1975)137;(1982)25;Mabb.(1983)87;(1984)442;Radcl.-Sm.(1987)356;Grierson&D.G.Long(1987)790;Philcox(1997)85;Chanthara-prasong&Welzen(2007)344,f.11A;LiBingtao&M.G.Gilbert(2008)268;Fern.Casas(2016)5,f.1-4,map1.―Manihot curcas(L.)Crantz(1766)167.―Jatropha acerifoliaSalisb.(1796)389,nom.superfl.―Castiglionia lobataRuiz&Pav.(1798)277,nom.superfl.―Curcas adansoniiEndl.exHeynh.(1846)176(seealsoMabb.1984:435).―Curcas indicaA.Rich.(1850)208,pronom.nov.,nom.superfl.―Curcas curcas(L.)Britton&Millsp.(1920)225,nom.inval.―Curcas lobata(Ruiz&Pav.)Splitg.exLanj.(1931)154,nom.superfl.,comb.inval.,insynon.―Lectotype(des-ignatedbyRadcliffe-Smith1987):Linnaeus,Hort.cliff.(1737,published1738)445:Jatropha no.3,Surinama,Jamaica,Brasilia.(Representativespecimen in BM: http://data.nhm.ac.uk/dataset/collection-specimens/resource/05ff2255-c38a-40c9-b657-4ccb55ab2feb/record/1565052).

Ricinus americanus [Rumph. (1743) 95, nom. inval.;]Mill. (1768) underRicinus.―Type:notindicated(SeeThellung1908:784forsynonymy).

Ricinoides americanaGarsault[(1764)t.67,nom.nud.;](1767)51.―Type:Notindicated.

Curcas purgansMedik.(1787)119.―Type:Notindicated.Jatropha edulisSessé(1794)3.―Type:M. Sessé Lacasta, J.M. Mociño,

J.D. del Castillo & J.M. Maldonado 4233(holoMA;isoF,US),Mexico.Ricinus jarakThunb.(1825)23.―Type:Notindicated.[Curcas drasticaMart.inSchrank&Mart.(1829)50,nom.nud.]SeeMab-berley(1984)442forsynonymy.

[Jatropha moluccanaWall.(nonL.)(1847)nr.7799E,nom.nud.].[Jatropha condorWall.(1847)nr.7799F,nom.nud.].Jatropha tuberosaElliot(1859)85.―Type:Notindicated,growninagarden.SeeMabberley(1983)87forsynonymy.

Jatropha yucatanensis Briq.(1900)230;Pax(1910)77.―Type:Linden s.n., 1840(holoG;isoF),Mexico,Yucatan.

Jatropha afrocurcasPax(1909)83;(1910)79.―Type:P. Jaeger 342(B?†), Deutsch-Ostafrika(Tanzania),Sseke.

Shrubstotreelets,upto7mhigh,stemupto15(–28)cmdiam,many-branched; flowering twigs 4–15 mm diam, ridged when dried,snappingeasily,epidermiseasilypeeling,tan-coloured.Outer barkgreyishgreentogrey,veryrough;underbarkgreen;innerbarklightgreenishcreamtostraw;exudate(whitish)paletranslucent,watery to somewhat sticky sap; sapwoodpink,whitelater;heartwoodpulpy.Indumentum of simple, long villous orarachnoidhairsonvariousparts.Stipules veryindistinct,ex-tremelyearlycaducous,broadlyovate,c.1by1.5mm,denselyvilloushairy.Leaves: petiole6.5–23cmlong,diam0.5–5mm,basally slightly thickened and triangular, above flat or glabrous tosomewhathairy;bladeovateto3-or5–(7-)palmatilobed,widest±inmiddle,7–17by6–16.5cm,0.7–1.5timeslongerthan wide, glossy, smooth, base emarginate to cordate, margin entire,oftensomewhatundulate,somewhatrevolute,apexofcentrallobeacutetoacuminate;lobesshort,atmosttillhalfofleafblade,±triangular;uppersurfacedulltoshinydarkgreen,usually glabrous, sometimes hairs along veins when young, leaving white dots as scars, lower surface dull light green, glabroustomorehairythanuppersurface;venationpalmate,7 veins originating from base, basal 2 weakest developed, up to c. 7 veins alongmidrib, higher order veins reticulate.Inflorescences axillary compound cymes, often several pernode, erecto-patent to patent, green, flowers at end of main axespistillate,othersstaminate;peduncle3.5–4.5cmlong,c.1mmwide;rachis5–20mmlong,(sub)glabroustovilloustoarachnoid;bractselliptic,basaloneup to13by1.3mm,becoming smaller towards top of inflorescence, margin entire, apexacute,(sub)glabroustovilloustoarachnoidonbothsides.

Map 1DistributioninMalesiaofJatropha curcasL.


Fig. 2 Jatropha curcasL.a.Terminalbranchlet;b.abaxialsurfaceleaftip;c.detailofinflorescence;d.staminateflower;e.pistillateflower;f.fruit;g.seed,ventralside;h.seed,dorsalside;i.seed,lateralside(a,b:A. Ibáñez 2024;c-e: E.L. Ekman H8156;f-i: M. Nee 41135;allMA).—Drawing:JoséMaríaPizarroDomínguez,2006.

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Flowerscup-shaped,palegreentoyellowtowhite,fragrant;pedicelsupto9mmlong,(sub)glabroustovilloustoarachnoid,withanabscissionzone,oftensubapical;calyxlobesbasallyconnate,outside(sub)glabroustohairy,insideglabrous;petalswithmarginentire,apexemarginate,outsideglabrous,insidevillous.Staminate flowersc.9mmdiam;sepalsovatetoelliptic-oblong,4–5by2–3mm,marginentire,apexobtusetoacute;petalsobovate-oblong,7–8by2.5–3mm,apexrounded;disclobesverticallytongue-like,1–1.2by0.4–0.5mm,glabrous;stamens 10, outer 5 almost free or adnate to inner united 5, androphoreupto5mmlong,lightgreen,freefilamentsupto4mmlong, lightgreen,anthersnarrowly triangular,1.6–1.7byc.0.5mm,basallycleft,basifixed,extrorseopeningwithlengthslits,lightyellow.Pistillate flowersc.6mmdiam;calyxlobesovate,3.2–5by1.5–2.5mm,marginentire,apexacute; petalslongovate,6.2–7by2–3mm,apexrounded;discglandsobtrapezoid,0.9–1by0.8–1mm;ovaryovoid, 2–3by1.7–2mm,lightgreen;stylec.0.5mmlong,green;stigmasgreen,filament-like,non-receptivepartc.0.5mm long,anther-like,receptivepartbasi-dorsifixed,c.1mmlong,apicallysplit tillhalfway.Fruits long-ovoid,2.3–3by1.8–2.5cm,6-grooved,surfacerugged,glabrous,yellowwhenripe,blackwhendried;wall up to 1mm thick, opening loculicidally only; columellaT-shaped,up to2.2cmlong.Seeds ellipsoid but somewhat flattened dorsiventrally, with a slight sharp ridge on the inside, 16–19by10–12by8.5–9.5mm,whendrydarkwallburstingwithsmallwhitedot-tostripe-like,mainlytransverseopenings;carunclevestigialorpoorlydevelopedinafoldoverthehilum. Distribution―CentralandSouthAmerica,Caribbean,wide-lyintroducedinMalesia. Habitat&Ecology―Aculturefollower,foundnearvillages,on and along roads, in cultivated areas and grounds, in gardens, atlandfills,nearriversintheshade,secondary/opengrassland,open plains, in and around primary, disturbed and secondary forest,nearthecoastandonlimestonehills;soiloftensandy,clay-loam,loam,riverbedswithgravelandsand;oncereportedfromrhyolitebedrock.Altitude:sea-levelupto800(–1200)m.Floweringandfruiting:throughouttheyear. Vernacularnames―MalayPeninsula(Corner1951):Jarak,Jarakbelanda,Jarakpagar(Malay).Sumatra:Djarak,Djira.Java:Jarakpagar(Malay).Borneo:Jarak(Malay);Tangan-tan-gan(Brunei);Brunei:Jarigan-tangan.Philippines(mainlyafterMerrill1923):Galúmbang(Pampángan);Kásla(Bisáya);Kirisól,Taba-tabá, Tañgan-tañgan-túba, Túbang-bakód (Tagálog);Tubanmiyalad (Ifugao);Tuban bakod (Tagbanua);Tubangbakod(Tagkawayan);Tagumbáu,Tagumbau-na-puráu,Tauuá(Ilóko);Takumbau(Sambáli);Taua-tauá(Igorot, Ilóko),Túba(Bíkol,Igorot,Tagálog).Sulawesi:Jarah.Moluccas:Kadatao(Halmahera Island);Kaidoean(SeramIsland).Purgeernoot(Dutch);Physicnut(English). Traditionaluses(partlyafterBurkill1935,Heyne1950)― Tra-ditionally, J. curcashasmanyuses.Thecomplete plant and almosteverypartoftheplanthasbeenfounduseful.Theshrubis often an ornamental, usually used as hedges, but also as firewood.Itisalsoplantedtoreforesthillsandseasides,andinonepartofAfricaitisplantedongraves.Thebark is applied tosnake-bitesandbitesofotheranimalsasanantidote.Thestem juice has piscicidal properties and is used to kill or stu-pefyfish.Thejuiceisalsousedfortheproductionofblueandreddyeandmedicinallyitisappliedagainstsprue(Malaysia,Perak),droppedinwoundsandevenusedbychildrentoblowbubbles.Theleaves are applied to wounds, are used to get ridofitches,andcanbeappliedtobruisesafterbeingcooked.Also, rubbing them on the abdomen of children stimulates the intestines and a decoction of the leaves forms a cure for diar-rhea,aremedyforcoughsandalotionforeczemaandulcers.Whilethematureleavesaretoxicinlargerquantities,theyoungleavesaresometimeseatenorusedasflavouringincooking.

The seeds are used as a medicinal purgative and mashed they areappliedtowoundsasastyptic. InGuam(Marianas)theseedsareeateninsmallquantities.IntheUSAtheyarealsousedforcriminalpoisoning.Theseeds,whenawickisapplied,are burned as lamps, just as the pressed oil, the latter is also usedinsoapproductionandcanbeusedinwoolspinning.Itis said that rubbing the seed oil on your scalp stimulates hair growth, and energetic rubbing of the oil on a female abdomen willproduceabortion.Philippineslabelsindicatethattheplant(notwhichpart)isusedasapainrelieverandforfractures. Modernusesandpossiblefutureuses(afterPandeyetal.2012)― Jatropha curcas has recently caught wide attention asabiofuelplant.Thisisbecausetheseedshaveanoilcon-tentofaround30–40 %, the plant can grow on wasteland and marginal land unsuitable for food crops and is not a food crop itself.AlsoimportantisthattheJatropha bio-diesel is as good as petro-diesel.Thefruit-hullsthatareleftcanbeusedasbioactivecompost.Thecultivationforbio-dieselproducesaseedcakeasaby-product.Thisseedcakecanbeusedasmanure,butcanalsobeusedforbiogasproductionthroughanaerobicdigestion.The cultivation of J. curcasalsohasenvironmentalbenefits.Itis a potential phytoremediator for polluted soil and a sequester ofatmosphericcarbon(tostoreitinthesoil).Itcanalsohelpto control soil erosion because it develops a deep taproot that canstabilize thesoil.Also,newmedicaluses forJ. curcas, which range fromHIV-and tumor-suppressingpropertiesofthewaterextractofthebranches,tousingtheleavesagainstsyphilis,havebeenfound.Thedownsideisthatthespeciesoftenescapescultivationandcanbecomeaninvasivespecies.InMalesiaspecimensarecollectedonallmajorislands,buttherearenoreportsofnegativeaspectsofinvasiveness.

2. Jatropha gossypiifoliaL.―Fig.1e–l,3;Map2

Jatropha gossypiifoliaL.(1753)1006(‘gossypifolia’);Müll.Arg.(1866)1086;Hook.f.(1887)383(‘gossypifolia’);Pax(1910)26;Merr.(1923)449;Gagnep(1926)326;McVaugh(1945)281,f.13,18;Corner(1951)260,text-fig.83;Backer&Bakh.f.(1963)494;AiryShaw(1972)283;(1975)137;(1982)25;Radcl.-Sm.(1987)354;Philcox(1997)83;Chantharaprasong&Welzen(2007)346,f.11E–L;Fern.Casas(2016)18,f.5-8,map2.―Manihot gossypiifolia (L.) Crantz (1766) 167 (‘gossypifolia’).―Adenoropium gossypiifolium (L.)Pohl (1827) 16 (‘gossypifolium’).―Conserved type(Wiersemaetal.2015:395):LinneanHerbariumno.1141.1(LINN).

Jatropha staphysagriifoliaMill.(1768)underJatropha,no.9(‘staphysagrifolia’).―Jatropha gossypiifolia L. var.staphysagriifolia (Mill.)Müll.Arg.(1866) 1087 (‘staphysagriaefolia’), nom. illeg. (autonym rule).―Type:Notindicated.

Adenoropium elegansPohl(1827)15.―Jatropha elegans(Pohl)Klotzsch(1853) 102.―Jatropha gossypiifolia L. var.elegans (Pohl)Müll.Arg.(1866)1087;Pax(1910)26;Backer&Bakh.f.(1963)494;Philcox(1997)83.―Type:Von Martius s.n.(holoM;isoL,barcodeL0034540),Brasilia,VillamdosIlhéos,CapitaniaeBahiae.

Adenoropium jacquinii Pohl (1827) 15.―Jatropha jacquinii (Pohl)Baill.(1864)268.―Type:Notindicated.

Jatropha glanduliferaauct.nonRoxb.:Kurz(1877)403.See www.ipni.org or www.theplantlist.orgformoreinfraspecificnamessyno-nymouswithvar.gossypiifolia.

(Herbsto)shrubstotreelets,atleastupto5mhigh,upto12cmdiam,branching,succulent;floweringbranches 1.5–8mmdiam, round, with branching glandular hairs, especially when young,oftendarkmaroon.Outer barkthin,finelypustular-lenti-cellatetosmooth,grey;sapthin,greyishorwhitish.Indumentum of simple, white, long hirsute hairs and branching glandular trichomes with elongated heads and non-branching glandular mushroom-shaped trichomes, glandular hairs with dark maroon stalkandyellowhead.Stipules dissected and appearing as a bandofbranchingglandulartrichomes(asimilarbandisalsopresentwiththebracts).Leaves: petiole2–15.5cmlong,1–2mm diam, basally thickened, round but above flat to grooved,


Fig. 3 Jatropha gossypiifoliaL.a.Terminalbranchlet;b.leafvenation,abaxialsurface,withmushroom-likeglandularhairsalongmargin;c.detailofinflo-rescence;d.infructescencewithmaturefruit;e,f.staminateflowers;g.discandandroecium;h,i.pistillateflowers;j.seed,ventralview;k.seed,dorsalview; l.seed,lateralview(a-c, e- l: G.G. Hatschbach 64656;d:P.L. Krieger 10422;allMA).—Drawing:JoséMaríaPizarroDomínguez,2006.

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Map 2DistributioninMalesiaofJatropha gossypiifoliaL.

upper surface with hirsute hairs and along ridges branched glandularhairs,maroon;bladeovate,3–5-palmatifid,2–12.5by2.5–16cm,0.6–0.9timesaslongaswide,baseroundedtoslightlyemarginate,marginsubentiretofinely,shallowlyser-rate, teeth ending in unbranched glandular trichomes, ciliate withhirsutehairs,apexofcentral lobeobtusetoacuminate,lobes usually obovate, basally united, surfaces often showing smallglands(hydatodes?),uppersurfaceglossydarkgreento brownish, often, especially when young, with hirsute hairs along major veins, lower surface light green to completely red whenyoung,glabrous;venationpalmate,with7basalveinsofwhichcentral3or5moredeveloped,nervesincentrallobe8–14pairs,anastomosingandloopednearmarginexceptforbasalonewhichendsinsinusbetweenlobes.Inflorescences compound subterminal cymes, erect to erecto-patent, up to 16.5cmlong,laxlyhirsute,greentomaroon;peduncle2.5–20cm long, 1–3mmwide; rachis 0.9–7 cm long; cymeswithcentral flowers pistillate, staminate flowers alongbranches;bractselliptic,basalones10–19by2–4mm,indumentandbasalstructureslikeleavesandstipules,respectively.Flowers cup-shaped;pedicel2–10mmlong,withsimplehairs,subapi-calabscissionzone;sepalsfree,ovatetoelliptictoobovate,green to maroon, margin serrate with simple hairs and teeth ending in a glandular hair, apex cuspidate; petals obovate,apexrounded,maroonwith lightgreentoyellowbasalpart.Staminate flowersc.6mmdiam;sepalsc.4by1.2mm;pet-alsc.4.5by2.5mm;disclobesobtrapezoid,convex,c.0.5 by 0.5mm; stamens 8, 5 in outerwhorl, 3 united in innerwhorl,outeralmostfree,withfreefilamentpartc.1.2mmlong,yellow-red,androphorec.2.5mm long, lightgreen,antherstriangular,0.5–0.6by0.5–0.6mm,orange-red,especiallyin-neronesbasallydivaricating,dorsibasifixed,latrorseopening.Pistillate flowers4.5–6mmdiam;sepals4.5–6by2–2.5mm;petalsc.5by3mm,caducous;disclobes±rectangular,c.1by0.5mm,thick;ovaryovoid1.5–3by1.3–3mm,6-ribbed,green,fewhairs,styleabsenttoveryshort,c.0.1mmlong;stigmaslightgreen,withunreceptivelowerpartc.1mmlong,apically receptive part thickened, split, U-shaped, 0.6–0.8mm long.Fruits oblong, slightly 3-lobed, 8–12 by 7–11.5mm, pendant, sparingly hirsute to subglabrous, dehiscing completely septicidally and partly loculicidally, shiny green whenimmature;wallc.0.5mmthick;columella6.3–8.5mm

long,narrowlyT-shaped,veryslender.Seeds somewhat dor-siventrallycompressed-ellipsoid,8–8.5by4.5–5by3–4mm;carunclemultifid,exceedingtheseedapex. Distribution―MexicotoNSouthAmericaandCaribbeanIslands,introducedandestablishedthroughoutMalesia. Habitat&Ecology―Wetareaslikeswamps,coast,littoraland sublittoral, gradually sloping reef flats, and damaged man-grove ecotone, but also secondary forest, lowland savannah, grassy plains, wasteland, road sides, usually open areas in general.Soil:sandyloam,(white-)sand,loam,clay,rockyclayloam,oftensiltish;bedrock:granite,limestone.Altitude:sea-levelupto750m.Floweringandfruiting:throughouttheyear. Vernacularnames―MalayPeninsula(mainlyafterCorner1951):Jarak,Jarakberemah,Jarakhitam,Jarakkling,Jarakmerah(Malay).Java:Djarak,Djarakkosta,Djaraktjina.Phil-ippines (mainly afterMerrill 1923):Balautandoiong,Tagum-bau-a-nalabága, Taua-tauá (Ilóko); Bongalon (Tagbanua);Lansi-lanináan (Tagálog);Túba-sa-buáia (Bíkol);Tuba-túba(PanayBisáya,CebuBisáya).LesserSundaIslands:Flores:Waru-wégé(Takatunga,Ngadha);Timor:Damarmerah;Paukopna(Dawan);Alor:Arangfai,Iwangbawiw,Trainkenanagar.Cotton-leavedphysic-nut(English). Uses(afterBurkill1935,Heyne1950)―Ornamentalplant,plantedinhedges.Medicinallyusedagainstdiarrhea.Aswal-lowing of a decoction of 7–21 leaves works as a remedy for dry belly-ache.Seedsareusedcriminallyasapoison,butalsoasapurgative;seedoilusedaslampoil,usefulintreatingleprosy.

Note―Ifasubdivisionofthissomewhatvariablespeciesisdesirable,thenthespecimensinMalesiaaregenerallyregardedasbelongingtovar.elegans.

3. Jatropha integerrimaJacq.―Fig.1b,4;Map3

Jatropha integerrimaJacq.(1760)32;(1763)256,t.183,f.47;Pax(1910)50;McVaugh(1945)274,f.5,6,12,16,21,22;AiryShaw(1972)284;(1982)25;Radcl.-Sm.(1987)353;Chantharaprasong&Welzen(2007)347,f.11B,pl.XIX-2;Fern.Casas(2016)28,f.9-13,map3.―Adenoropium integerrimum(Jacq.)Pohl(1827)14. ―Jatropha diversifoliaA.Rich.(1850)207,nom.superfl.;Müll.Arg.inDC.(1866)1094.―Type:Notindicated.

Jatropha hastataJacq.(1760)32;(1763)256,t.173,f.54;Pax(1910)51;Backer&Bakh.f.(1963)494.―Adenoropium hastatum(Jacq.)Britton&P.Wilson (1924)485.―Jatropha integerrimaJacq.var.hastata (Jacq.)Fosberg(1976)102.―Lectotype(designatedhere):Jacquin s.n.(BM).


Fig. 4 Jatropha integerrimaJacq.a.Brancheswith inflorescences;b. leaf;c. leafbasewithsmall lobes;d.bract;e.staminateflowershowingdiscand androecium;f.pistillateflowerinsection;g,h.partlyloculicidallydividedtwo-valvedcocci,outsideandinside,respectively;i.seed,dorsalview;j.seed,ventralview(a-d, f: F.R. Fosberg 59690;e:E.L. Ekman 2369;g- j: E.L. Ekman H4472;allMA).—Drawing:JuanLuisCastilloGorroño,2006.

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Jatropha acuminataDesr.(1797)8;Vent.(1803)t.52.―Type:Herb. de Jussieu s.n.(holoP-JU;IDCmicrofiche6206,box31,fiche1194,no.20),SaintDomingue(SantoDomingo).

Jatropha pandurifoliaAndrews(1802)t.267(‘panduraefolia’);Pax(1910)49;Gagnep.(1926)326.―Adenoropium pandurifolium(Andrews)Pohl(1827)14.―Jatropha diversifoliaA.Rich.var.pandurifolia(Andrews)M.Gómez(1894)51.―Type:Andrews(1802)t.267.

Jatropha coccineaLink(1822)406.―Adenoropium coccinea(Link)Steud.(1840)799,insynonymy(‘coccineum’).―Jatropha pandurifolia Andrews var.coccinea(Link)Pax(1910)50,f.19b.―Jatropha integerrimaJacq.var.coccinea(Link)N.P.Balakr.(1980,publ.1982)176.―Type:Notindicated,Hort.Cels.(=GardenofJ.M.Cels).

Jatropha paucifloraC.WrightexGriseb.(1865)170;Müll.Arg.(1866)1095;Pax(1910)51.―Jatropha diversifoliaA.Rich.var.pauciflora (C.WrightexGriseb.)M.Gómez(1894)51.―Type:C. Wright 1954(holoGOET;isoBM,BREMs.n.,G-DC,GH,HAC[3],K,MO,NY,P,US[2x],YU),Cuba.

Jatropha moluensisSessé&Moc.(1894)224.―Type:Notindicated(MA?n.v.),[Cuba,]Havana.

Jatropha pandurifoliaAndrewsvar.latifoliaPax(1910)50,f.19a.―Jatropha integerrimaJacq.var. latifolia(Pax)N.P.Balakr.(1980,publ.1982)176. ―Lectotype(designatedhere):Pr. de la Sagra 595(holoW),Cuba,cul-tivatedingardensinHavana.

Jatropha glaucovirensPax(1910)51.―Type:A.H. Curtiss 458(holoB?lost;isoA,BM,G[2x],GH,HAC,K,L,M),[Cuba,]IsladePinos,NuevaGerona.

Shrubs to treelets, at least up to 8 m high, stem up to 10 cm diam,somewhatsucculent;floweringbranches2–3mmdiam,hairy,glabrescent.Outer barkdarkgrey,fairlysmooth;exudatewhite(onelabel).Indumentum of simple, pilose hairs, present onmost parts.Stipules triangular, 1.3–1.5by0.5–0.7mm,caducous,oftenwithbasal lobe, lattersometimesseparate.Leaves: petiole 0.8–7.5 cm long, diam c. 1mm, round intransverse section, but flat to slightly grooved above, espe-ciallyhairyabove,basallythickened,lightgreen;laminaovatetomostoftenobovate,5.2–12by3.4–7.5cm,1.3–2.6timeslonger than wide, papyraceous, basally rounded to somewhat emarginate,oftenwithtwoglandularextensions,marginentireexceptbasallyseveraltriangularmini-lobesendinginagland,apexacuminatetocuspate,darkgreenabove,usuallyhairyonvenation,dullgreenbelow,glabrous;venationbasally7-pal-mate,butmidribandnexttwoveinsstrongest,latterendingoverbladehalf,moreterminallypinnatewith5–12veinsperside.Inflorescences compounddichasiawithcentralflower(s)pistil-late,erect,subterminal,upto15cmlong;peduncle4.5–12.7cmlong,diamc.1.5mm;basalbractelliptic,upto8by1mm,withstipule-likebasalextensions,upperbractsusuallytriangular,c.3by1mm,becomingsmallerupwards,marginserratewith

glandontopoftheteeth,apexacute.Flowers: pedicels 5–8 mmlong,apically(andoftenalsobasally)anabscissionzone;calyxlobesbasallyconnate,lobestriangulartoovate,basallygreen,restdarkred,glabrous;petalsfree,obovate,darkred.Staminate flowersc.16mmdiam,lobesovate,2–3by1.5–2mm,margin entire, apex round; petals 9–24by5–10mm,curvedbackwards,apexslightlyemarginatetoround,basallyhairyinside,restglabrous;disclobes±square,c.0.5by0.5mm;stamens10,in2whorlsof5,glabrous,androphore4.5–5mmlong;freepartoffilaments2.3–4mmlong,anthersnar-rowlytriangular,2–2.5byc.0.7mm,basallyslightlydivaricate,dorsibasifixed, opening latrorse, connectivewith appendix.Pistillate flowersc.20mmdiam:lobes1.5–3.7by1.1–2mm,margin(entireto)serratewithglands,apexacute;petals10–13by5–6mm,apexemarginatetorounded,glabrous;disclobesrectangular, c. 1 by0.5mm;ovary ellipsoid, 2.5–5by2–3mm,glabrous,green,style0.8–1mmlong;stigmas4.2–5mmlong, red,bifid inupper2.4–2.5mm, flattenedand insomebroadened.Fruits oblong,c.1by1cm,dehiscingsepticidally,smooth,green;wallc.1mmthick;columellaT-shaped,c.9mm long.Seeds ellipsoid,c.8by3mm. Distribution―CaribbeanIslands(Cuba,DominicanRepu-blic,Haiti),introducedinMalesia(Java,Philippines,Celebes). Habitat&Ecology―Secondaryforest,alongtrailinlowlandmixedforest;soil:brownishclay,clay-loam.Altitude:sea-levelto4m.Flowering:January,February,April,May,August,De-cember. Uses―Ornamentalbecauseofrelativelargeorange-redflowersinredinflorescences.

Note―Thisspeciesisveryvariableintheshapeoftheleafblade.The formmostencountered isobovate tosomewhatpanduriform,butinthePhilippinesthebladesareovate.

4. Jatropha multifidaL.―Fig.1c,5,6;Map4

Jatropha multifidaL.(1753)1006;Miq.(1859)392;Müll.Arg.(1866)1089;Kurz(1877)403;Hook.f.(1887)383;Pax(1910)40,f.13;Merr.(1923)449;Gagnep.(1926)325;McVaugh(1945)277;Backer&Bakh.f.(1963)494;AiryShaw(1972)284;Radcl.-Sm.(1987)354;Grierson&D.G.Long(1987)790;Chantharaprasong&Welzen(2007)347,f.11C;LiBingtao&M.G.Gilbert(2008)269;Fern.Casas(2016)37,f.14-17,map4.―Manihot multifida (L.)Crantz(1766)167.―Adenoropium multifidum(L.) Pohl(1827)16.―Lectotype (designated byRadcliffe-Smith 1987): t. 173 (f. 213), oppositep.218inDillenius,HortusElthamensis(1732),Americameridio-nali(SAmerica).

Map 3DistributioninMalesiaofJatropha integerrima Jacq.


Fig. 5 Jatropha multifidaL.a.Terminalbranchlet;b.apexofleaflobe,abaxialsurface;c.staminatebud;d.youngstaminateflower;e.adultstaminateflower;f.pistillateflower;g.pistillatepetal(all:F. Melville 156,MA).—Drawing:JoséMaríaPizarroDomínguez,2006.

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Jatropha janiphaBlanco(nonL.,nonLour.)(1837)758;(1845)521;(1879)159, t.342;Merr. (1918)229.―Neotype(designatedhereafterMerrill1918):Merrill Species Blancoanae 625(holoL),Philippines,Luzon,Manila.

Shrubs,atleastupto2.5mtall,branchessomewhatsucculent;flowering branches5–9mmdiam.Outer bark greyish with green-brownlenticels.Indumentumabsent.Stipules dissected inmanyflagelliformpartsof15–20by0.1–0.2mm.Leaves palmatisect;petiole4–27cmlong,1–5mmdiam,roundbutflattenedtosomewhatgroovedabove,basallythickened;bladealmostcircularincircumference,14–34by12–30cm,basecordate; lobes (6–)9–11(–13),elliptic,4.5–17by0.7–8cm,margin entire but with 1 or more small, triangular, alternate side-lobes,slightlyconstrictedaboveside-lobes,apexgradu-allyacute;palmatelynerved,withalongmidribupto21pairsofnerves.Inflorescences subterminal, cymose, corymbiform, erect,upto30cmlong;peduncleupto26cmlong,2–3mmwide; rachis 4–21mm long; central flower pistillate, othersstaminate;bractsnarrowlytriangular,2–4by0.5–1mm,oftenfolded lengthwise, margin usually with a few side-lobe-like ini-tiations,apexacuminate,upwardsbecomingsmaller.Flowers c.6mmdiam,allpartsredororange;pedicels3–7mmlong;calyxc.3mmwideandlong,marginentire;petalsobovate,contort,apexemarginatetoobtuse.Staminate flowers:calyxlobesc.1by1–1.2mm,apexemarginatetorounded; petals 4–5.8by2.5–3mm;discglands±square,c.0.4by0.4mm;stamens8, 5 in outer and 3 in innerwhorl, free, filaments2.5–3mmlong,antherswithparallelthecae,2–2.5by0.4–0.7mm,basifixed,openingextrorse.Pistillate flowers: calyxlobestriangular,c.2by1mm,apexacute;petalsc.4by2.5mm;discglandspresent;ovaryovoid,styleshorttoabsent,stigmasshort and thick.Fruits irregularly shaped, shape dependent onnumberofdevelopedseeds,c.3by2cm,containing1–3

seeds, dehiscence septicidal, yellow;wall c. 0.5mm thick;columellanotseen.Seeds sub-ellipsoid,17.5–20by15–17by12–13.5mm. Distribution―SNorthAmerica,centralandNSouthAme-rica,introducedinMalesia(MalayPeninsula,Sumatra,Java,Philippines). Habitat&Ecology―Hill slopes, cultivatedwithinhumansettlements.Altitude: up to 700m.Flowering:March,April,May;fruiting:April. Vernacular names―MalayPeninsula:Hubiq (Semelai).Philippines (Merrill 1923):Apio (Visaya);Tubang-amerikáno(Bíkol);Maná(Spanish).Dutch:Koraalboom(coraltree). Uses(afterBurkill1935,Heyne1950)―Ornamentalplantofwhichtheyoungleavesandtuberscanbeeatenafterroasting;olderleavescanactaspurgative.Seedsaremedicinallyusedaspurgativeandcriminalpoisoningduetocatharticproperties;oilusedaslampoil.

5. Jatropha podagricaHook.―Fig.1d,7,8;Map4

Jatropha podagricaHook.(1848)t.4376;Müll.Arg.(1866)1093;Pax(1910)44;Merr.(1923)450;McVaugh(1945)277;Backer&Bakh.f.(1963)494;AiryShaw(1972)284;Radcl.-Sm.(1987)355;Chantharaprasong&Welzen(2007)348,f.11D;LiBingtao&M.G.Gilbert(2008)268;Fern.Casas(2016)49,f.18-22,map5.―Type(seeRadcliffe-Smith1987):Seemann s.n. (BMn.v.),Panama,SantaMarta.

Shrubs,upto70(–150)cmhigh,notorhardlybranching,ba-sal stem very thickened, succulent, flask-like, on top a more slenderapicalbranch(orloweronbasalpartside-branches);floweringbranchesc.1cmdiam.Barkgreyandgreen;waterytoslightlywhitesap.Indumentumabsent.Stipules dissected inslip-likelobesofc.4mmwide.Leaves:petiole16.5–35cmlong, c. 0.5 cmdiam, round,hollow;bladeovate, c. 15–28

Fig. 6 Jatropha multifidaL.,fruitsandseeds.a.Fruit,lateralview;b.fruit,apicalview;c.seed,ventralview;d.seed,dorsalview;e.seedlateralview(all:C.F. Baker 4,MA).—Drawing:JoséMaríaPizarroDomínguez,2006.

c

b

d

a

e


Fig. 7 Jatropha podagricaHook.a.Terminalbranchlet;b.abaxialleaftip;c.detailofinflorescence;d.staminateflowers;e.pistillateflowers;f.pistillatepetal;g.fruit(all:J. Lau 2024,MA).—Drawing:JoséMaríaPizarroDomínguez,2006.

cb

d ag

ef

e

d

d


Fig. 8 Jatropha podagricaHook.a.Habitofcompleteplant;b.branchlettip;c.maturefruit;d.seed,ventralview;e.seed,dorsalview;f.seedlateralview (a,b:D.A. Neill 217;c- f: W.D. Stevens 9142;allMA).—Drawing:JoséMaríaPizarroDomínguez,2006.

c

b

d

a

e f

b


by17–31cm,5-lobed, lobes lessthan1/3rdofblade,base4.5–7.5 cmpeltate,margin entire, dark green above, palelight greenishgreyishbelow; lobesobovate, apicesbroadlyacutetoacuminate;venationpalmate,slightlyraisedontop, 7veinsoriginatingfrombase,nerves4–10pairsperlobe.Inflorescences compoundcymes,corymbiform;peduncle34–43cm long, red, cymes with basally the central flowers pistillate, on the branches staminate flowers; rachis 3–40mmhigh;bractsovate,1–3by0.8–2mm,apexacuminate,becomingsmallerupwards.Flowers: pedicel with apically an abscission zone;sepalsandpetals red. Staminate flowers: pedicel 2–5 mmlong;calyx lobes1.4–1.5by0.8–1mm,apexrounded;petalsobovate,c.6.5by1.6mm,marginentire,apexrounded;disc glands thick, ± square, c. 0.3 by 0.3mm; stamens 8, 5inouterwhorl,3ininnerwhorl,filamentsfree,c.4mmlong,antherswithparallelthecae,c.2.5by0.8mm,basifixed,open-inglatro-extrorse,pollenorange.Pistillate flowerssubsessile; sepalsfree,triangular,c.1.5by1.5mm,apexobtuse;petalsearlycaducous,notseen;discglandslikestaminateflowers;ovaryellipsoid, c.5by3mm,stylenearlyabsent; stigmas:unreceptivebasalpartc.1mmlong,receptivepartc.1.2mmlong,almostcompletelysplit,U-shaped.Fruits flattened, de-pressedglobular,c.11by7mm,glabrous,openingsepticidallyandpartly loculicidally;wallc.0.3mmthick;columellac.10mmlong,apicallythickened,notT-shaped.Seeds ellipsoid, c.9 by4mm;carunclec.1.5by2mm. Distribution―CentralAmericaandCaribbeanIslands.In-troducedinMalesiaasornamental,occurringinthePhilippines,naturalizedintheCarolineIslands(KororIsland). Habitat&Ecology―OntheCaroline Islandscommon inthicketsaroundhouses,inthePhilippinesandThailandonlyplanted;granitebedrock.Altitude:sea-levelupto350m.Flow-ering:February–April,July,August,October;fruiting:February,March,July,August,October. Uses―Gardenornamental,potentialtoescapecultivation,notconsideredadangerousinvasiveintheCarolineIslands.Youngleavesandtuberscanbeeaten,butnotraw.

Acknowledgments The directors and keepers of the various herbaria mentioned under the types are thanked for information and/or pictures of the typespecimens.Thereviewersandeditorarethankedfortheirveryhelpfulsuggestions.AnitaWalsmitSachs,JoséMaríaPizarroDomínguezandJuanLuisCastilloGorroñomadethebeautifuldrawings.Jan-FritsVeldkampgaveadviceonnomenclature.

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IDENTIFICATION LIST

1 = J. curcasL. 2=J. gossypiifoliaL. 3=J. integerrimaJacq. 4=J. multifidaL. 5=J. podagricaHook.

Aseries231:1–Achmad1747:1–Anderson108:2;566:2–Arnoldo89a:2;98a:4;576:1;906:2;1951:2–Avé116:1.

Backer27085:2;36235:1;36330:3;36531:4;36734:5;36830:1;37625:1–Baker2279:2–BakhuizenvandenBrinkJr1660:2;2731:4–Baumann-Bodenheim12042:2–Beaman6288:1–Belshaw3372:2–Bhargavaetal.6455:1–Bloembergen3082:2;3083:1;3141:2;3287:2;3870:2;4634:1–BogorBotanicalGardenIX.A.8:4–Boldingh19B:2;516B:4;558B:2;570B:2;589B:2;771:1;1126B:2;1209:1;1360B:2;1639B:2;2235:4;2594:4;2595:1;2601:1;3106:1;3113B:2;4649:2;5129:1;6221:2;7166:2–Boom7169:2;38540:3–Bordenave139:2–Bouamanivong,Kien&Sinh143:1–BovienDjomb.58:2–Brantjes706710:2–Breteler3163:2;4352:2–Bullock879:1–Bünnemeijer8342:1–Burgers113:2;179:2–Buwalda5984:1;7152:2–Buysman185:1;306:2;307:4;406:1;1593:1–Buysman&Broadway2565:2.

Cancap6ExpeditionPh13:2;Ph143:2–Chayamaritetal.1858:2;1860:2–Chin3582:1–Chun&Tso43599:1–Coert74:1–Collins1513:2–Coode&Duffels6105:1–Cooray69111727R:2–Cuming687:1–Curry721:1–Curtiss434:2;451:1.

d’Alleizette6455:1–DangiDN3:1–Danser5528:1;6477:2–DeEscobar989:1–DeGranville&Poncy11706:2–DeVoogd755:2–P.A.W.J.deWildeCUWa138:2–DenOuter907:3–Dickason8644:2–Dorantesetal.896:1–Dorgelo699:3–Dubs1926:1–Dunbar339:2;351:2.

Elbert793:1;2034:1;2059:2;3829:2–Esser99-17:2–Evans1463:1–Eyma4059:1.

Farenholtz2009:1–Feuillet1569:2–Florschütz&Florschütz1035:1–ForestDepartment(Guyana)7075:2–Forster14791:2–Fosberg24768:2;25373:1;25769a:2;28922:2;32260:5;46217:2;60379:3–FuertesLorén474:4.

Gianno279:4–Gillespie&Gopaul1994:2–Grimes1042:1–Guillaumin&Baumann-Bodenheim11042:2.

Hahn,Tiwari&Ameer3800:2–Hanke592:2–Harini139:1–Harleyetal.21104:2–Harrison1671:2–Hartley10581:1–Hatschbach45828:2;60782:2;64656:2–Haught3710:2–Hekking1115:2–Henkel,Williams&Ameer2623:2–Henkel,Williams&James3394:2–Hidayat101:2;125:1–Hochreutiner2799:2–Hohenacker433:1–Hoogerwerf266:1–Hosaka2811:2–Hotta4985:1–How70514:1–Hu&But21137:3–Huq&Mia10498:2;10650:1.

Iwatsuki,Murata&Gutiérrez4:2.Jaag465:2;631:2;764:2–Jansen-Jacobs,TerWelle&James5355:2–Jayasuriya1248:2–Jensma14:1.

Kelloff&Roesel1143:2–Kern20276:2–Kessler707:2–King5516:2–KLseries2420:2–Koelz18964:1–Koorders25216:2;25245:1–Kooy807:2–Krukoff16498:1–Kvistetal.396:2.

LAEseries67813:1–Lagosa112:1–Lakshnakara900:1–Lanjouw626:2–Lazarides7137:1–Leefmans69:2–Leeuwenberg13194:3–Lei537:1;720:1–Lens801:2–Liao10172:3–Libman,Somsamut&XaiveuLAOS-441:1–Luna900:2.

Maas&Maas-vandeKamer360:2–Maasetal.5968:2–MacKee5487:2;26492:2;36508:1;43893:2–Madulidetal.7217:1–Maxwell71-199:2;88-524:2;89-778:1;91-356:5;91-381:2;93-851:2;98-457:1–Meijer9208:2;10170:2–Mejía&Zanoni9033:1–MerrillSpeciesBlancoanae10:1;SpeciesBlancoanae625:4–Metzner64:2;80:1–Mexia4477:2–Mocquerys825:2–Murata,Fukuoka&PhengklaiT-16564:2.

Nagata397:3–Nagelkerken168:2;304:2–Nair4469:4–Nee44109:2–NGFseries22177:2–Noerkas62:3–Nooteboom1340:1.

Ochoaetal.387:1.Paiva,Silveira&SousaT347:2–Pedersen5295:1–Pennington&DelaCruz10541:2–Pipoly11243:2–Pipoly&Ameer9068:2–Pires&Black793:1–Platenkamp15:1–PNHseries19215:1;20469:5;804691;91582:2;97900:1–Popta149/19:4;258:1–PPIseries1756:2;1778:1;1798:3;5515:1;9789:1;15216:3;18446:2;29136:1;29166:2;37358:2–Prawiroatmodjo&Maskuri1271:1–Proctor23506:2.

Raizada23765:2–Raynal-Roques15586:2;15816:2;15981:2–Reek-mans7729:5–Reinders&Torres40:1;60:2–RHTseries14178:2–Riley2:2;13:1–RobinsonPlantaeRumphiana333:1–Rugel380:2–Rutten428:1–Rutten-Pekelharing30:4;109:2;496:1.

S series 36436: 5 –Samuels 331: 1 –SANseries 134962: 1 –Santos4649:4–Sauvin380:1–Schimpff51:1–Setthi&Negi25758:2–SFNseries39309:2–Shimizu,Fukuoka&NalampoonT-7617:2–Sidiyasa1108:2–Sieber219:2;221:1;282:4;309:4;435:2–Singleton903:1–Sintenis4789:1–Skog,Feuillet&Rossman7539:2–Sloot&Reina118:1–H.H.Smith1472:2–Snellius-II10546:2;11150:2–Soejartoetal.6511:1–Soepadmo&Suhaimi254:2–Southavong,Kien&Sinh52:1–Splitgerber16:2;74:2;172:1;433:4–Stahl1075:4–Sterringa12445:2–Stoffers42:2;88:2;272:2;343:2;380:2;437:2;452:2;481:2;760:2;766:2;803:2;884:2;904:2;1053:2;1351:2;1453:2;1996:2;2058:2;2267:2;2527:2;2832:2;2897:2;3009:2;3430:2–Stone3997:2;4047:4;4060:1–Subbarao24269:1.

Trotz&Persaud9:4;29:2–TsugaruT-61643:1.UGP’cognosyGroup126:2–UPNGseries240:2.VanAndeletal.1279:1;1311:2;5361:2–VanBeusekom&VanBeusekom-Osinga1587:2;1588:1–VanBorssumWaalkes1705:1–VanGijzen48:1–VanGroll-Meyer52:1;66:4;132:2;133:2–VanHeel144:4–VanOoststroom12493:3–VanProosdij183:2;516:2–VanWelzen2003-6:4;2003-8:5;2003-10:1–Verheijen4716:2;5420:2–Versteeg445:1–Versteegh&DenOuter285:1–Vidal,Vidal&Niyomdham6087:1–VonEggers355:2–VonTürckheim2637:2;2773:2.

Waas&Tirvengadum814:2–Wagner139:2;440:2–Wallich7799E:1;7799F:1–Warnecke286:2;358:1;373:4–Watdahnahsahp81:3–Went1117:2–Wight2637:1–Winkler2187:2–Wittingthon78:1.

Yates1880:1;1910:4–Yuncker5557:1;15116:1;15637:1;15774:1.Zimmermann27:3;145:3;178:4–Zollinger73:4;481:1;3926:2.

A revision of Jatropha (Euphorbiaceae) in Malesia - Naturalis ...

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