Revision of Creochiton (Melastomataceae)

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Blumea 58, 2013: 217–227www.ingentaconnect.com/content/nhn/blumea http://dx.doi.org/10.3767/000651913X674134RESEARCH ARTICLE

INTRODUCTION

The genus Creochiton (Melastomataceae)was establishedbyBlume (1831a, b) for two species from Java previouslyincluded in Melastoma L.:C. bibracteatus Blume(1826) and C. pudi bundus Blume(1823).The latter isheredesignatedasthelectotype.ThenameisderivedfromtheGreek,kreaschitoon (κρεας χιτων), fleshy garment,which refers to theusually large and thickened bracteoles that cover the flower budbeforeanthesis.Notethatthegenderofchiton is male andallepithetshaveherebeencorrectedwherenecessary.Thegenusiseasilyrecognizedaswoodyclimbers(scramblingshrubs), creepers, or epiphytic shrubswith distinct fleshybracteolesenvelopingtheflowerbuds.Ithasbeenregardedby most authors as allied to Dissochaeta Blume.Thecircum-scriptionemployedhereisafterVeldkamp(1978)followedbyMaxwell(1980,1981,1983).There is a remark by Kadereit (néeClausing; 2006) thatCreochiton would partially belong to Dissochaeta but(inlitt.)she then thought that Creochiton anomalus (King&StapfexKing)Veldk., C. furfuraceus (M.P.Nayar)Veldk.andC. monticola(Ridl.)Veldk.wouldbelongtherebecauseoftheaxillaryplacentation, anthers with ventral appendages and extra-ovarialchambers.Naudin(1852:141,153)pointedoutsimilaritiestotheAmericansubtribe Pyxidanthinae (‘Pyxidantheae’) Naudin, but noted that perhaps it was better placed with the subtribe Dissochaetinae (‘Dissochaeteae’)Naudin(1851:67).Triana(1865,1871)el-evated this to the tribe Dissochaeteae (Naudin)Trianainwhichhewasfollowedbylaterauthors,e.g.Cogniaux(1891),Krasser(1893),Maxwell(1983).Baillon(1880:15)evenregardedCreo- chiton in a heterogeneous assembly of genera ranked as sec-tions of Dissochaeta but did not make the necessary combi-nations for the two Creochiton species.Becauseofthecloserelationships between the two genera both are included in the Dissochaeteae(Naudin)Triana,e.g.becauseoftheirhabitandtheshapeoftheirinflorescences(Triana1871,Cogniaux1891,Krasser1893,Maxwell1983).

Mansfeld(1925)putCreochiton into the capsular-fruited tribe Astronieae Triana on the basis of the basal placentation.BakhuizenvandenBrinkJr.(1943:306)agreedwiththisandalthough he knew that the fruits are berry-like, he created the subtribe Creochitoninae(Miq.)Bakh.f.becauseoftheleaf-likebracteolesthatclasptheflowerbuds,andthecuneateseeds.Thisdivisionisnotsupportedbywoodanatomy(VanVliet1981)andmacromorphology(Veldkamp1978)andCreochiton should be included in the Dissochaetinae.Enchosantherawas established byGuillaumin (1913: 341)for E.anomala (King&StapfexKing)GuillauminbasedonAnplectrum anomalum King&Stapf ex King.He decidedto create a separate new genus because it differed from AnplectrumA.Graybytheshapeandstructuresoftheeightheteromorphousanthers.HedistinguishedCreochiton by the homomorphousstamens.Veldkamp(1978) reducedEncho-santhera to Creochiton because the differences seemed only of specific value.Thestructureandnervationof the leaves(plinerved)and rather thickbracteolesweremoresimilar toCreochiton than to Dissochaeta.EisocreochitonwasdescribedbyQuisumbing&Merrill(1928:177) on thebasis ofE.bracteatus Quisumb.&Merr.Theysaid that it was similar to Creochiton in its vegetative and in-florescencescharacters.Thedifferenceswereinthenumberofstamensandthepresenceofextra-ovarialchambers.Al-though no fruits were available they placed their new genus in the capsular-fruited Oxysporeae Triana near BlastusLour.Judging from labels inBOOhwi in1945alreadyequated itwith Creochiton.Nayar(1970)revisedthegenusandcorrectlyassigned it to the baccate Dissochaeteae.Hedistinguishedthree species, E.bracteatus, E.furfuraceus, and E.monticola.Veldkamp(1978)andMaxwell(1980,1981,1983)notedthatthe characters supposedly differentiating between Creochiton and Eisocreochiton cannot separate them at the generic level andthattheymustbeunitedintoasinglegenus.Similartothesituation in Dissochaeta, the number of stamens and the pres-enceanddepthoftheextra-ovarialchambersareofspecificvalueonly,notgeneric.Withthisinmind,thethreespeciesof Eisocreochiton were transferred to Creochiton(Veldkamp1978).Furtado (1963)wasapparentlynotawareofEncho-santheraGuillauminanddescribedAnplectrella for the same species, Anplectrum anomalum.Beinghomotypic,thenameAnplectrellaissuperfluous.

Revision of Creochiton (Melastomataceae)A.Kartonegoro1,J.F.Veldkamp2

Key words

CreochitonMalesiaMelastomataceaenew combinationrevision

1 ResearchCenterforBiology,IndonesianInstituteofSciences(LIPI),Jl.Jakarta-BogorKM46,Cibinong,WestJava,Indonesia;

correspondingauthore-mail:[email protected] NaturalisBiodiversityCenter,P.O.Box9517,2300RALeiden,TheNether-lands;e-mail:[email protected].

Abstract A taxonomic revision of the Malesian genus Creochiton(Melastomataceae)ispresented.Twelvespeciesarerecognized,althoughoneisbasedontheoriginaldescriptiononly.Creochiton turbinatus (comb.nov.)israisedtospecificrankfromavarietyofC.ledermannii.Akeytothespecies,descriptions,ecologicalhabitats,distributions(withmaps),synonymy,taxonomicnotes,vernacularnamesandanindextocollectionsareprovided.

Published on 2October2013

218 Blumea – Volume 58 / 3, 2013

GENERAL MORPHOLOGY

HabitThe species of Creochitonarecreepers(scramblingshrubs)orepiphyticshrubs,2–40minheightorsometimeswoodyclimb-ers,e.g.thePhilippinespecies(C.bracteatus, C.dipterus and C.roseus).Inflorescencesareonlyseenonthintwigs,whichsuggest that the growth mode is sympodial as seems usual for the Dissochaeteae. This is most obvious in C. anomalus wheretheinflorescencesareterminal.Becauseofthisolderstems are rarely collected it cannot be ascertained how exactly theplantsclimb.For C. brevibracteatus Mansfeld reported that the stem is as thickasanarm.AspecimenofC. roseus(PNH 9924, Sulit)isdescribedas5–6mtalland1cmdiameter.Usually they have terete branchlets except for C.furfuraceus and C.monticola where they are obscurely triangular and be-cometeretewhenmature.Theyareneverflattened.Thenodesare usually thickened with or without a distinct interpetiolary ridge.Theindumentisvariablefromabsenttodenselystellatefurfuraceousorfloccose(C.bibracteatus and C.ledermannii).

LeavesThe leaves of Creochiton are more or less coriaceous when dry.Theshapeisquitevariable,usuallytheyareovate-ellipticoblongandonlyrarelysuborbicular.TheyarelargestinC. bibracteatus(6–12.5by3.2–7.5cm)andsmallestinC.monticola (2–3by1.4–1.5cm).Theapexusually isacuteorroundedwhileinthreespeciesitisretuse(C.anomalus, C.ledermannii and C. monticola).Thebaseismostlyroundedorcuneate,or sometimes subcordate in C.bibracteatus, and the margin al-waysentire.Similar to nearly allMelastomataceae genera, Creochiton has acrodromal venation with one or two pairs of lateralnerveswhichariseatorabovethebase.Thissubbasalor plinerved nervation is a character that usually distinguishes Creochiton from Dissochaeta. It is also found inBoerlagea Cogn.,Medinilla Gaudich.,Pachycentria Blume and Plethiandra Hook.f.,butthesearedifferentinhabit.Thesecondaryvena-tionisscalariform.Generally,theuppersideoftheleavesisglabrouswithsunkenmain nerves, while the pubescence of the lower sides is similar tothatofthebranchlets.

InflorescenceThe inflorescence of Creochiton is either a cymose pseudo-umbelwith3–5flowersorathyrsewithupto35flowers.Itismostly axillary except for C.anomalus whereitisterminal.Themainaxisisusuallytereteandsparselytodenselyfurfuraceous.Bracts are present at each node and there is a pair of bracteoles at the base of each pedicel which usually more or less cov-erstheflowerbud.Ingeneraltheyarecoriaceous,caducous,concave and usually glabrous, more rarely furfuraceous in- or outside.InC. bracteatus, C. brevibracteatus and C. dipterus they are thin and do not fully enclose the flower bud and are stillpresentatthetimeofanthesis.

FlowersThe flowers are tetramerous, similar to those of Dissochaeta. The calyx tube or hypanthium is usually campanulate or urceo-latewithatruncaterimorwith4wave-liketeeth.Ithasthesameindumentasthemainaxisandbranchlets.Thepetalsareovatetoelliptic.Thecoloursvarybetweenwhite,pinkorpurplish,but cannotbeused todistinguishspecies.Their length has here been measured as much as possible from budsabouttoopen.SimilartoDissochaeta, the characters of thestamensarealsoconsideredtohavespecifictaxonomic

value.Whenthereareonly4stamenstheanthersareusuallyequalandhomomorphic.Whenthereare8stamenstheymaybe either homomorphic or heteromorphic, in which case the oppositipetalousonesaresmaller.The anthers are elongate and glabrous and open distally with asinglepore.Theconnectiveusuallyhasadorsalappend-age(atriangular,hastateorsagittatecrestinalternipetalousanthersandspur-likeorligularinoppositipetalousones)andin C.anomalus and C.monticola there are two lateral appendages (ligularorfiliform).The ovary is from 1/2 to nearly 2/3 times as long as the hyp-anthium.Theapexisusuallyglabrous,butsometimesdenselypubescent. Similar to other genera inDissochaeteae it is concrescent with the hypanthium with or without septs forming extra-ovarial chambers in which the anthers are inserted before anthesis(Kartonegoro&Veldkamp2010).Thechambersvaryfrom absent to shallow to reaching to halfway or to the base oftheovary.

FruitsThe fruits of Creochiton as in all other Dissochaeteae are berry-like, globose or subglobose, colourful when mature, sometimes withfoursmallcalyxremnants.Theyarevariouslydescribedasberries,leatheryberries,orevenindehiscentcapsules.Trueberriesbydefinitionhaveawell-developedusuallyjuicyor fleshy mesocarp, with the seeds immersed in it and without cavitiesorsepts.Capsulesarebydefinitiondehiscent.Anan-cient term for fruits with a dry indehiscent pericarp with several hollow cells without contents except for the many seeds was carcerule.They are usually glabrous, but sometimes they are sparsely furfuraceous.InC.turbinatus there are distinct vertical ridges bywhichtheylookturbinate-shaped.Theseedsarenumerous,smoothandcurved.

DISTRIBUTION AND ECOLOGY

Creochiton is an endemic genus of the Malesian region ranging fromtheMalayPeninsulaandESumatra(C. anomalus:Johor,Perak;Indragiri,Bangka)totheCentralandNewBritainProv-incesofPapuaNewGuinea(C. novoguineensis).Thereisnoindicationforacentreofdiversityduetoamoreorlessequalnumberofspeciesineveryregion.Creochiton monticola and C.furfuraceusareendemictoBorneo;C.ledermannii, C.novoguineensis and C.turbinatustoNewGuinea;whileC.bractea-tus, C.dipterus and C.roseustothePhilippines.Creochiton pudibundus and C.bibracteatus were previously known only fromJavabutnowthefirsthasbeenfoundinCelebesandthesecondinCeram.Creochiton mainly grow as epiphytic shrubs or woody climb-ers with adventitious roots in rainforests that are not affected byseasonalclimate.PerhapsthisexplainsitsabsenceintheLesserSundaIslands.Somespeciesarefoundinmountainforests up to 2 000 m altitude like C.bibracteatus, C.monticola, C.novoguineensis and C.pudibundus.SeveralspeciessuchasC.anomalus, C.furfuraceus, C.leder-mannii and C.turbinatus are known from lowland primary forest orswampyforestbetween50–500m.Theplantsusuallygrowat the edge of primary or secondary forest, near river banks or indisturbedforestwithgapsinthecanopy.Thescarcenumberof specimens in the herbarium is probably due to the fact that theplantsgrowhighinthetrees.AccordingtoFurtado(1963)all specimens of A. anomalus in theMalayPeninsulawerecollectedfromloggedorfallentrees.

219A.Kartonegoro&J.F.Veldkamp:RevisionofCreochiton

TAXONOMIC TREATMENT

CreochitonCreochitonBlume(1831a)506;Naudin(1852)153;Miq.(1855)559;Tri-ana(1871)85;Cogn.(1891)604;Krasser(1893)179;Merr.(1905)32;(1923)191;Koord. (1912)693; (1923)212;Bakh.f. (1943)307; (1963)362.―Disso chaeta Blumesect.Creochiton (Blume)Baill.(1880)15,51.―Lectotype(designatedhere):Creochiton pudibundus (Blume)Blume.

EnchosantheraGuillaumin(1913)341.―Anplectrella Furtado(1963)106,nom. superfl.―Type:Enchosanthera anomala (King&Stapf exKing)Guillaumin [≡Anplectrella anomala (King&Stapf ex King) Furtado ≡Creochiton anomalus (King&Stapfex King)Veldk.].

Eisocreochiton Quisumb.&Merr.(1928)177;M.P.Nayar(1970)87.―Type:Eisocreochiton bracteatus Quisumb.&Merr. [≡Creochiton bracteatus (Quisumb.&Merr.)Veldk.].

Dissochaeta BlumefideKadereit(2006)3(p.p.).Diplectria auct.non (Blume)Rchb.:Kuntze(1891)246(p.p.).Melastoma auct.nonBurm.ex L.:Blume(1823)71;(1826)1071;DC.(1828) 148.

Woody creepers, climbers, scrambling or epiphytic shrubs;branchlets commonly terete or sometimes obscurely trian-gular, glabrous to densely covered with brown furfuraceous or floccosehairs, sometimeswith adventitious roots; nodesthickened or rarely flattened, with or without an interpetiolary ridge.Leaves with acrodromal venation, thin, subcoriaceous or coriaceous, ovate to oblong rarely suborbicular, apex acute or rounded to retuse, base cuneate to subcordate, margin entire; leavesglabrousabove,midribimmersed,glabroustodenselyfurfuraceous,glabroustodenselytomentosebelow;nervation plinerved with one or two main lateral nerves arising atorabovethebase,midribraisedbelow;secondarynervationpinnate,commonlyreticulate;petioleterete,dorsallygrooved,glabrous to densely furfuraceous. Inflorescences commonly axillary, either umbellate, 3–5-flowered or paniculate with up to35flowers,rarelyterminal(C.anomalus);mainaxisterete,sparselytodenselyfurfuraceous;bractsminutetoconspicu-ous,thinorcoriaceous;bracteolesdistinct,paired,commonlycoriaceous, concave, obovate or orbicular, caducous, glabrous or densely furfuraceous, enclosing the flower bud, rarely thin orminute; pedicel terete, glabrous to densely furfuraceous;flowertetramerous;hypanthiumcampanulateornearlyurceo-late,glabroustodenselyfurfuraceous,hairssimpleorstellate;calyxrimtruncateorwavywith4shortlobes;petalsovateorelliptic,glabrous,acutetip,white,pinktopurplish.Stamens4or8,equalorunequal,if8,4alternipetalouslarger,4oppositi-petaloussmaller;filamentsflat,straight;anthersbasifix,cylin-dric, elongate, straight or slightly curved at anthesis, opening withasingleterminalpore;connectivewithatriangular,hastateor sagittate crest in alternipetalous ones, triangular, spur-like or ligular in oppositipetalous ones, lateral appendages commonly absent,rarelypaired;ovaryglobose,0.5–0.75timesaslongas thehypanthium, glabrous, 4-locular, placentation axillaryorbasal;stigmacapitate;stylestraightorcurved,glabrous,slender;extra-ovarialchambersrarelydevelopedornarrowedat the top of the ovary, or extending to about half the length of theovary.Fruits berry-like, globose to subglobose, green orpurplishwhenmature,glabrous;stalkglabrousorsparselyfurfuraceous,seedsnumerous,cuneate,curved,flat-topped. Distribution―ThegenuswithtwelvespeciesisendemictotheMalesianregion(MalayPeninsula,Sumatra,Borneo,Java,Celebes,Moluccas,PhilippinesandNewGuinea),butisnotyetfoundintheLesserSundaIslands.

KEY TO THE SPECIES

1.Hypanthiumdenselyfurfuraceous . . . . . . . . . . . . . . . . . . 21.Hypanthiumglabrous, glabrescent or sparsely furfurace-

ous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Leaf underneath glabrescent (but formidrib).―Philip-pines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Leafunderneathdenselyfurfuraceousordenselytomen-tose.Bracteolesobovate,completelycoveringtheflowerbud,fleshy,denselyfurfuraceous.―Java,Moluccas. .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.C. bibracteatus

3. Bracteolespartiallycoveringtheflowerbud,thin,outsidedensely furfuraceous . . . . . . . . . . . . . . . . . 5.C. dipterus

3. Bracteoles completely covering the flower bud, fleshy,outside glabrous . . . . . . . . . . . . . . . . . . . . .11.C. roseus

4. Petioleglabroustosparselyfurfuraceous...........54. Petioledenselyfurfuraceousorfloccose. . . . . . . . . . . . 85. Petiole8–15mmlong.Leaf4.5–13cmlong,basecune-

atetorounded,apexacuminatetoacute.Inflorescencesaxillary.Bracteolesovatetoobovate,completelycoveringbud, fleshy, glabrousor densely furfuraceous.Stamensequal.―Borneo,Celebes,Java,NewGuinea . . . . . . . 6

5. Petiole2–6mmlong.Leaf1.5–4cmlong,baseacute,apex roundedor retuse. Inflorescences terminal.Brac-teoleslinear,partiallycoveringbud,thin,scaly.Stamensunequal.―MalayPeninsula,Sumatra . . 1.C. anomalus

6. Leafapexacute.Bracteolesobovate.Stamens8,lateralappendagesandextra-ovarialchambersabsent.―Cel-ebes,Java,NewGuinea........................7

6. Leafapexacuminate.Bracteolesovate.Stamens4,lateralappendagesandextra-ovarialchamberspresent.Petiolesparsely furfuraceous.Leafunderneathdensely furfura-ceous.Inflorescencesumbellate,few-flowered.Bracteolesdensely furfuraceous.Hypanthiumstellate furfuraceous.Fruitsglobose.―Borneo . . . . . . . . . . . 6.C. furfuraceus

7. Petiolesparselyfurfuraceous.Leafunderneathglabrous(butformidrib).Inflorescencespaniculate,many-flowered.Bracteoles densely furfuraceous.Hypanthium sparselystellatefurfuraceous.Fruitssubglobose.―NewGuinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.C. novoguineensis

7. Petioleglabrousorwithsomehairs.Leafunderneathpubes-cent.Inflorescencesumbellate,few-flowered.Bracteoles andhypanthiumglabrous.Fruitsglobose.―Celebes,Java . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10.C. pudibundus

8. Petiole8–20mmlong.Leaf3.5–7by1.8–4cm,under-neathglabroustoglabrescent(butformidrib) . . . . . . . . 9

8. Petiole2–6mmlong.Leafbasecuneatetorounded.Brac-teolescompletelycoveringtheflowerbud,4–8mmlong.Hypanthiumglabrous to sparsely stellate furfuraceous.Stamens8..................................10

9. Leafbasecuneate.Inflorescencesumbellate.Bracteolescompletelycoveringtheflowerbud,6–7mmlong.Hyp-anthiumsparselystellatefurfuraceous.Stamens4,lateralappendagespresent.―Philippines . . . 3.C. bracteatus

9. Leafbaserounded.Inflorescencespaniculate.Bracteolespartiallycoveringtheflowerbud,1mmlong.Hypanthiumglabrous.Stamens8,lateralappendagesabsent.―NewGuinea . . . . . . . . . . . . . . . . . . . . . . 4.C. brevibracteatus

10. Leaf 3.4–6 by 2.2–3.1 cm, underneath glabrescent todenselytomentose.Inflorescencesumbellate,few-flower-ed.Bracteolesorbicularorovate.Stamensequal,lateralappendagesabsent.―NewGuinea. . . . . . . . . . . . . 11

10. Leaf2–3by1.4–1.5cm,underneathglabrous(butformid-rib).Inflorescencespaniculate,many-flowered.Bracteolesobovate.Stamensunequal,lateralappendagespresent.Leafelliptictoobovate.Fruitssubglobose,smooth.―Bor-neo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.C. monticola

11. Leavesobovate,apexretuse,underneathdenselytomen-tose.Bracteolesinsidestellatefurfuraceous.Hypanthiumsparselystellatefurfuraceous.Fruitssubglobose,smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.C. ledermannii

220 Blumea – Volume 58 / 3, 2013

11. Leavesovate,apexacute,underneathglabrescent(butformidrib).Bracteolesinsideglabrous.Hypanthiumglabres-cent.Fruitsglobose,ridged . . . . . . . . . . 12.C. turbinatus

1. Creochiton anomalus (King&Stapfex King)Veldk.― Map 1

Creochiton anomalus (King&Stapfex King)Veldk.(1978)431;Veldk.&M.P.Nayar(1978)438.―Anplectrum anomalum King&Stapfex King(1900)58;Ridl.(1922)800.―Enchosanthera anomala (King&Stapfex King)Guillaumin(1913)341,f.1,2.―Anplectrella anomala (King&Stapfex King)Furtado(1963)106.―Lectotype(Veldkamp1978):King’s Collector 5779(holoCALn.v.;isoK,L,P,SING),MalayPeninsula,Perak,Gopeng.

Epiphytesorwoodyclimbers,10–40mhigh;branchletsterete,coveredbystellatebrownscales,glabrescent;nodesthickenedwith an interpetiolary ridge.Leaves coriaceous, obovate to elliptic,apexroundedtoretuse,baseacute;aboveglabrous,light green, sparsely stellate scaly on the midrib, below densely covered with stellate tomentose hairs, yellowish green, dark brownwhendry;1.5–4by1–3cm;nervationwith1–2pairsof lateralnervesfromabovethebase;petiole3–8cmlong,coveredwith stellate scales, 2–6mm long. Inflorescences terminal,paniculate,few-tomany-flowered;mainaxisangular,coveredwithstellatescales,furfuraceous;secondaryaxes,1–6mmlong;bractsminuteorlinear;bracteoleslinear,notenvel-opingtheflowerbuds,coveredwithbrownscales,2–3mm;pedicel covered with stellately furfuraceous hairs, 8–11 mm

Fig. 1 Creochiton bibracteatus (Blume)Blume.a.Habit;b.flower;c,d.stameninflowerbud(c=abaxial;d=adaxial);e.crosssectionfruit;f.maturefruit;g.flowerbudwithapairofbracteoles;h.bracteole(all:Backer 10623,BO).

cb d

a

g

h

e

f


long.Hypanthiumcampanulatetonearlyurceolate,4–5by2–3mm,stellatefurfuraceous,green,becomingbrownwhendry;calyxrimtruncate,furfuraceouswith4smallundulations0.5mmlong;petalselliptic,whiteorpink,4–5by2–2.5mm.Stamens 8,unequal,thealternipetalousones:filamentflat,3mmlong;antherelongate,yellow,glabrous,7mmlong;connectivewithatriangulartosagittate,crest2mmlong,purple; lateralap-pendagespaired,ligular,1–1.5mmlong;theoppositipetalousones:filament flat,3mmlong;antherelongate,5mmlong;connectivewithaligularcrest,1mmlong;lateralappendagesabsent;ovaryglobose,glabrous,3mmlong;styleglabrous,slender,straight,5–7mmlong;extra-ovarialchambers8,halfaslongastheovary.Fruits subglobose, glabrous to sparsely furfuraceous,greentoyellowishgreen,4–6by3–4mmdiam;seedsc.0.6mmlong. Distribution―PeninsularMalaysia(Johor,Perak)andSu-matra(Riau,Bangka). Ecology&Habitat―Lowlandorswampyprimaryrainforestbetween100–550maltitude.

Note―Creochiton anomalus differs from all species by theterminalpaniculateinflorescences.Ithaslinearbracteoleswhichpartiallycovertheflowerbud.Thespecieshas8hetero-morphous stamens that are similar to those of C. monticola from Borneo, but the oppositipetalous stamens are smaller and have nolateralappendages.SimilartoC.brevibracteatus but differs byitsshorterpetioleandleaves.

2. Creochiton bibracteatus (Blume)Blume―Fig.1,Map2

Creochiton bibracteatus (Blume)Blume(1831a)507;(1831b)248;Naudin(1852)154;Miq.(1855)560;Triana(1871)85,t.7,f.91;Cogn.(1891)605;Krasser(1893)178,f.77a;Koord.(1912)693;(1923)212;Bakh.f.(1943)308;(1963)362.―Melastoma bibracteatum Blume(1826)1071;DC. (1828) 148.—Diplectria bibracteata (Blume)Kuntze (1891) 246,nom.inval.prov.―Lectotype(designatedhere):Blume s.n. (holoL,sh.no.908.128–2131;iso908.129–921,-2141,-2146),Java.

Creochiton superbusNaudin(1852)153;Miq.(1855)560.―Type:Zollinger 3009(holoG;isoL,U),Java,MtSalak.

Epiphyteorclimber,upto10–26mheight;branchletsterete,denselybrownfurfuraceous;nodesthickenedbytheinterpetio-lary ridge.Leaves thin, ovate to suborbicular, apex acute to rounded, base rounded to subcordate, glabrous above but puberulous near base, midrib sunken, densely stellate furfura-ceoustofloccosebelow,6–12.5by3.2–7.5cm;nervationpli- nervedwith2pairsofmainnervesarisingabove thebase;petioledenselyfurfuraceous,10–27mmlong.Inflorescences axillary4–6cmlong,umbellatewith3–5flowers;mainaxisterete,upto4.5cmlong,denselyfurfuraceous;bractsoblong,glabrous,4mmlong;bracteolesobovate,coriaceous,concave,enveloping the flower buds, densely furfuraceous outside, gla-brousinside,violet,6–15by4–6mm;pediceldenselyfurfura-ceous,10–15mmlong;flowerbud10mmlong.Hypanthium campanulate,denselystellatefurfuraceous,3–5by2–3mm;calyxrimtruncatewith4smallundulations,0.5–1mmlong;petalsoblong,white,7–9by4mm.Stamens8,equal;filamentflat,4mmlong;antherelongate,4–6mmlong;connectivewithasagittatecrest1–1.5mmlong;lateralappendagesabsent;ovaryellipsoid,glabrous,3mmlong;styleglabrous,curvedwhenmature, 6–10mm long; extra-ovarial chambers shal-lowabout1/5aslongastheovaryorabsent.Fruits globose or subglobose, vertically depressed, glabrous, dark green to black,8–10by7mmdiam;stalkdenselyfurfuraceous;seedsc.1mmlong. Distribution―Java(west)andMoluccas(Ceram). Ecology&Habitat―Slopeofsecondaryforest,edgeoffor-estormossymountainforestbetween1100–1600maltitude. Vernacular name―Harendongareuy (Sundanese: usedfor about every climbing Melastomataceae).

Note―Creochiton bibracteatus can easily be distinguished from the other species by the densely furfuraceous petioles, rounded to subcordate leave base, large, ovate to suborbicular leavesupto12.5cmlong,belowfurfuraceoustofloccose,andthedenselystellatefurfuraceoushypanthium.Theaxillaryum-bellateinflorescencesand8equalstamensinthisspeciesaresimilar to C.pudibundus but it differs from the latter by being glabrous or with sparsely stellate hairs in most parts and by thecoriaceousleaves.Previouslythespecieswasknownonly

Map 1 DistributionofCreochiton anomalus.

Map 2 DistributionofCreochiton bibracteatus (●)andC.pudibundus (■).

222 Blumea – Volume 58 / 3, 2013

asepiphyticshrubsorclimbersinthewesternpartofJava,butin1917onecollectionwasmadeinMautipass,Ceram,thatisindistinguishablefromtheJavaspecimens(Rutten 1970, BO).

3. Creochiton bracteatus (Quisumb.&Merr.)Veldk.― Map 3

Creochiton bracteatus (Quisumb.&Merr.)Veldk.(1978)438.―Eisocreo-chiton bracteatus Quisumb.&Merr.(1928)177,pl.4;M.P.Nayar(1970)88(‘bracteata’).―Type:Ramos & Edaño BS 45610 (holoPNHn.v.,probablylost;isoBM,BO,K,NY,SING,US),Philippines,Luzon,NuevaVizcayaProvince,MtAlzapan.

Climbers;branchletsterete,sparselypuberulous,2mmdiam;nodeswithan interpetiolary ridge.Leaves thin, ovate, apex acute or acuminate, base cuneate, glabrous above, glabres-centbelowmidribdenselyfurfuraceous,3.5–6by1.8–3.6cm;nervation plinerved with two pairs of main nerves arising above the base; petiole densely stellate furfuraceous to floccose,10–20mmlong.Inflorescencesaxillary,thyrsoidupto4cm long,5-flowered;mainaxisangular,denselyfurfuraceous,2–4cm long;bractselliptic,sparsely furfuraceous,8–11by5–7mm; bracteoles ovate to suborbicular, glabrous to sparselyfurfuraceous, thin, 6–7 by 3–4mm, enveloping the flowerbuds;pediceldenselystellatefurfuraceous,7–8mmlong.Hy-panthiumcampanulate,sparselystellatefurfuraceous,3–4by2–3mm;calyxrimtruncatewith4shallowundulations,0.3mmlong;petalsovate,apexacute,baserounded,3–4by3mm. Stamens4,equal;filamentflat,c.3mmlong;anthersellipsoid,4–5mmlong;connectivewithatriangularcrest,0.75–1mmlong;lateralappendagepaired,0.8–1.5mmlong;ovaryglo-bose,glabrous,2–3mmlong;styleglabrous,c.6mmlong;extra-ovarialchambersdeep,halfaslongastheovary.Fruits andseedsnotseen. Distribution―Philippines(Luzon),onlyknownfromthetype. Ecology&Habitat―Climbingontreesonforestedslopesat1700maltitude.

Note―Creochiton bracteatus differs from all congeners by its densely furfuraceous to densely floccose petioles, 10–20 mmlong,leafbasecuneate,underneathglabrescent(exceptmidrib)andextra-ovarialchambershalfaslongastheovary.SimilartoC.furfuraceus byits4equalalternipetalousstamens,

but differs by having glabrescent branchlets and underside of the leaves while the latter has densely furfuraceous branch-letsandundersideoftheleaves.Itsbractsandbracteolesareverydistinctbeinglarge,thin,membranous,andveined.Thebracteoles are also not concave and do not fully enclose the flowerbudsasintheotherspecies.

4. Creochiton brevibracteatus Mansf.

Creochiton brevibracteatus Mansf.(1925)135.―Type:Ledermann 8660 (holoB†),NewGuinea,ESepik,Aprilfluβ(AprilRiver),Standlager(MainCamp).

Liana;stemasthickasanarm,innovationsterete,furfuraceous.Leavesovate,rarelybroadlyelliptic,5–7by3–4cm,5-nerved(excl.marginalnerves),baserounded,apexacute,darkgreen,above glabrous, below glabrous except for the prominent nerves,petiole8–15mm long, furfuraceous. Inflorescences paniculate,axillary,upto5cmlong,incl.the2cmlongpedun-cle,axessomewhatfurfuraceous;pedicels7mmlong,aboutglabrous;bracteoleshardly1mmlong.Hypanthium glabrous, 4mmlong,hardly4-dentate;petals4,ovate,shortlybeforeanthesis5mmlong,white.Stamens8,subequal,filaments3mmlong,anthers4mmlong,dorsalappendix1.5mmlong.Fruitsandseedsnotseen. Distribution―NewGuinea(ESepik). Ecology&Habitat―Primaryforestwithmanytreeferns,bamboos, small palms and Selaginella up to 1 m, at 100 m altitude.

Note―Nospecimenshavebeenseen,andthedescriptionisafterMansfeld(1925).Fromthisthespeciesseemtodifferfromallcongenersby:Petioles8–15mmlong,denselyfurfu-raceous.Leaves5–7by3–4cmwide,underneathglabrous(butformidrib).Inflorescencepaniculate.Bracteolespartiallycoveringtheflowerbud,1mmlong.Hypanthiumglabrous.ThetypewaslostduringtheSecondWorldWar.ThedescriptionagreesquitewellwiththePhilippineC. dipterus which differs mainly by the possibly more pubescent leaves, 3-flowered cymes, bracteoles 5–7 mm long, hypanthium densely stellate furfuraceous,rimtruncatewith40.3–0.5mmlonglobes.

5. Creochiton dipterus Elmer―Map3

Creochiton dipterus Elmer(1911)1192;(1939)3659;Merr.(1923)190.―Type: Elmer 9813 (holoPNHn.v.;isoE,MO),Philippines,Negros,NegrosOrientalProvince,Dumaguete(CuernosMts).

Climbers,upto25mhigh;branchletsterete,glabrescent tosparselybrownfurfuraceous;nodesthickenedwithinterpetio-lary ridge.Leaves coriaceous, ovate, apex acute to obtuse, baseroundedorsubcordate;glabrousabove,yellowishgreen,glabrescentbelow,midribdenselyfurfuraceous,4–7.5by2–4.5cm; nervation plinervedwith 1 pair ofmain nerves arisingabovethebase;petioledenselyfurfuraceous,5–17mmlong.Inflorescencesaxillary,a3-floweredcyme,1.5–2.5cmlong;mainaxisterete,denselyfurfuraceous,2cmlong;bractsovate,denselyfurfuraceous,veined,c.3mmlong;bracteolesovate,orbicular to oblong, thin, densely furfuraceous on both sides, veined, yellowish, half enveloping the flower buds, 5–7 by 2–5 mm;pediceldenselyfurfuraceous,8–9mmlong.Hypanthium campanulate, densely stellate furfuraceous, yellowish green, 3–4by2mm;calyxrimtruncatewith4smalldistinctlobes,0.3–0.5mmlong;petalselliptic,apexrounded,4–5by4mm,white.Stamens8,equal;filamentflat,cream,3–4mmlong;an-therelongate,pinkishorpurple,4–5mmlong;connectivewithasagittatecrest,lightcream,c.1mmlong;lateralappendages absent;ovary2–3mmlong,glabrous;styleglabrous,pink,9–11mm long;extra-ovarial chambersnotdeveloped.Fruits globose, glabrous,c.5by5mm,greenandturningredwhenripe;seeds0.5–0.7mmlong.

Map 3 DistributionofCreochiton bracteatus (▲),Creochiton dipterus (■)and Creochiton roseus (●).


Distribution―Philippines(Luzon,Mindanao,Mindoro,Ne-gros). Ecology&Habitat―Primaryordisturbedforestonridgeslope,between450–1800maltitude. Vernacularname―Buyon-Buyon(Visayan).

Note―Creochiton dipterus is distinguished by its coria-ceous leaves, underneath glabrescent (but formidrib), theumbellate 3-flowered inflorescence, thin, ovate, orbicular to oblong bracteoles which partially cover the flower bud, and the absenceofextra-ovarialchambers.ItresemblesC.ledermannii by the shape of the leaves and fruits but differs by its glabres-cent leaves, the single cyme and thin bracteoles, while the latter has densely tomentose leaves, umbellate inflorescences andcoriaceousbracteoles.Theshapeofstamensissimilarto C.novoguineensis but it differs by the shape and indument of theleaves.

6. Creochiton furfuraceus (M.P.Nayar)Veldk.―Map4

Creochiton furfuraceus (M.P.Nayar)Veldk. (1978) 438.―Eisocreochiton furfuraceus M.P.Nayar(1970)88,pl.1(‘furfuracea’).―Type:Jacobs 5293 (holoKn.v.;isoL;alsoinB,CANB,G,S,SAR,US,n.v.),Borneo,Sarawak,3rdDiv.,KapitDistrict,BelagaSubdistrict,RejangRiver.

Woodyclimbers;branchletsangular,becomingterete,denselypuberulouswithbrownhairs;nodesthickenedwithan inter-petiolaryridge.Leaves coriaceous, ovate to elliptic, apex acuminate,baserounded;glabrousabovewithstellatehairsonthemidrib,belowdenselystellatefurfuraceous,4.5–7by2.5–3cm;nervationplinervedwith1pairofmainnervesarisingabovethe base; petiole densely coveredwith brown puberuloushairs,8–11mmlong. Inflorescences axillary, thyrsoid, 3–10 cmlong,with10–35flowers;mainaxisterete,denselystel-latepuberulous,2–6cmlong;pediceldenselyfurfuraceous,4–5mm long;bracteolescoriaceous,ovate,concave,apexrounded,5–6by3–4mm,stellatefurfuraceousonbothsides,envelopingtheflowerbuds.Hypanthiumcampanulate,2.5by2mm,stellatefurfuraceous,palebrownish;calyxrimtruncate;petalsovate,apexacute,basetruncate,c.2.5by2mm,sor-didlypinkmargined,centrewhite.Stamens4,equal;filamentflat,white,1.5mmlong;antherselongate,curved,2mmlong,palegreen;connectivewithatriangularcrest,c.0.75mmlong,lateralappendagespaired,c.0.7mmlong;ovaryc.2mmlong,glabrous,4-ridgedatthetop;styleglabrous,pinkish,2–2.5mm

long;extra-ovarialchambersshallow,1/5aslongastheovary.Fruitsglobose,stellatefurfuraceous,5by5mmdiam;seeds0.7–0.8mmlong. Distribution―Borneo(Sarawak). Ecology&Habitat―Primaryforestonsandstonesubstra-tumhillsatbelow500maltitude.

Note―Creochiton furfuraceus can be distinguished by its dense furfuraceous indument in most parts, axillary thyrsoid inflorescences with 10–35 flowers, coriaceous bracteoles and 4equalstamens.ItissimilartoC.bracteatusbyhaving4equalstamens and the shape of the anthers but differs by a less dense indument on most parts, smaller inflorescences with only c.5flowers,largeandthinbracteolesanddeepextra-ovarialchambers.

7. Creochiton ledermannii Mansf.―Map5

Creochiton ledermannii Mansf.(1925)135.―Lectotype(designatedhere):Ledermann 12943 (holoB†;isoL),NewGuinea,Sepikgebiet,Felsspitze.

Creochiton ledermannii Mansf.var. turbinatus auct.nonJ.F.Maxwell:J.F.Maxwell(1981)323,quoadparatypi.

Epiphytesorclimbers,2–3mheight;branchletsterete,denselyfurfuraceoustofloccose;nodesthickenedwithaninterpetiolaryridge.Leaves coriaceous, obovate, apex retuse, base rounded tocuneate;glabrousabove,darkgreen,denselytomentosebe-low,midribdenselyfurfuraceous,glossyandpalergreen,3.4–4by2.4–3cm;nervationplinervedwith1pairofmainnervesarisingfromabovethebase;petioledenselyfurfuraceous,2–6mmlong.Inflorescencesaxillary,upto6cmlong,umbellatewith3flowers;mainaxisterete,denselyfurfuraceous,2.7cmlong;bractselliptic,glabrous,12by3mm;bracteolescoria-ceous, orbicular or ovate, concave, glabrous outside, densely furfuraceousinside,envelopingtheflowerbuds,whitish,4–8by6mm,whitish;pedicelglabrescenttosparselyfurfuraceous,7–18mmlong.Hypanthium urceolate, sparsely to densely stel-latefurfuraceous,3by2mm,teethtriangular,c.0.5mmlong;petalselliptic,3–4.2by2mm,red-purplish.Stamens8,equal;filamentflat,2–2.5mmlong;antherselongate,2–3mmlong;connectivewithatriangularcrest,1.5mmlong;lateralappend-agesabsent;ovaryglobose,1.5–1.8mmlong,glabrous;styleglabrous,c.5mmlong;extra-ovarialchambersshallow,c.1/5aslongastheovary.Fruits subglobose, glabrous to puberulous, greenishwhite,4–7by4mmdiam;seedsc.0.5mmlong. Distribution―NewGuinea. Ecology&Habitat―Rainforestonriverbank200–250maltitude. Vernacularname―Apunengapok(Waskuk).

Note―Characteristics ofC. ledermannii are the retuse apex leaf, axillary umbellate 3-flowered inflorescences, brac-teoles coriaceous and concave, hypanthium urceolate, calyx rimdistinctly truncate, and8 equal stamens. It differs from C.turbinatus by its dense indument and retuse leaf apex, while thelatterislesspubescentwithanacuteleafapex.Theshapeof the stamens is similar to C.pudibundus, but this differs by the sparsely puberulous leaves, glabrous petioles and bracteoles, absenceofextra-ovarialchambers,andthesubglobosefruits.The paratypes of C. turbinatus (Hoogland & Craven 10735, Docters van Leeuwen 10382)wereregardedtobelonghere.

8. Creochiton monticola (Ridl.)Veldk.―Map4

Creochiton monticola (Ridl.)Veldk.(1978)433;Veldk.&M.P.Nayar(1978)438.―Anplectrum monticola Ridl.(1946)31.―Eisocreochiton monticola (Ridl.)M.P.Nayar (1970)89,pl.2.―Type:Brooks 50 (holoKn.v.;isoBO),Borneo,Sarawak,MtBenKajan.

Creochiton kinabaluensis Heine(1953)214.―Lectotype(designatedhere):Clemens 32646 = 33951(holoM;isoAn.v., BM,BO,K,L),Borneo,MtKinabalu,MtNungkok.Map 4 DistributionofCreochiton furfuraceus (■)andC.monticola (●).

224 Blumea – Volume 58 / 3, 2013

Fig. 2 Creochiton pudibundus (Blume)Blume.a.Habit;b.flower;c.flowerbudwitapairbracteoles;d.maturestamen;e.crosssectionofflowerbud;f. fruit (a:Backer 35201; b–e:Bakhuizen van den Brink 5524; f:Backer 15169,allBO).

c

b

d

ae

f


Climbers,oftenepiphytic,upto30mtall;branchletsangularbecomingterete,furfuraceous.Leaves coriaceous, elliptic, apex acutetoretuse,basecuneate;glabrousonbothsides,2–4.5by1.4–2.6 cm;with 1–2pairs ofmain nervesarising fromabovethebase;petioleglabroustofurfuraceous,3–5mmlong.Inflorescencesaxillary,upto3.5cmlong,paniculate,4–10-flow-ered;mainaxisangular,stellatefurfuraceous,1.5–2.5cmlong;bractsovate,glabrous,veined,5–7by4mm;bracteolesthin,pale, obovate, glabrous to sparsely furfuraceous on both sides, veined,3–7by3.4mm,envelopingtheflowerbuds;pedicelfurfuraceous,2mmlong.Hypanthium campanulate, glabrous tosparselyfurfuraceous,green,pink,2–2.5by2mm;calyxrimtruncatewith4shallowundulations;petalsoblongtolanceolate,apexacute,c.4by2.5mm,creamtopinkorreddish.Stamens 8,unequal,thealternipetalousones:filamentsflat,2–2.5mmlong;antherelongate,red,c.2.5mmlong;connectivewithatriangular crest, 1–1.5mm long; lateralappendagespaired, c.0.5mmlong;theoppositipetalousones:filamentflat,1.5mmlong;anthersc.1.5mmlong,bent;connectivewithspur-likecrest,0.5mmlong; lateralappendages ligulate,c.0.75mmlongorabsent;ovaryglobose,glabrous,1–1.5mmlong;styleglabrous,c.5mmlong;extra-ovarialchambersshallow,about1/5aslongastheovary.Fruits subglobose, glabrous, whitish, yellowishgreen,orpurplewhenmature,2–5by1–5mmdiam;seeds0.75mmlong. Distribution―Borneo(Sabah,Sarawak). Ecology&Habitat―Mountainforestbetween820–2000m altitude.

Note―Creochiton monticola is easily recognized by its small leaves (<4.5 cm long), glabrous underneath, axillarypaniculateinflorescencesandobovatebracteoles.Morphologi-cally it cannot be mistaken for C.anomalus which has densely tomentose leaves, acute base, terminal inflorescences with thin bracteoles which partially cover the flower bud, and long extra-ovarialchambers(abouthalfaslongastheovary).Thestamens differ by the oppositipetalous ones having lateral ap-pendagesc.1–1.5mmlongwhiletheyare0.5–0.75mmlongorabsentinthelatter.

9. Creochiton novoguineensis (Bakerf.)Veldk.&M.P.Nayar―Map5

Creochiton novoguineensis (Bakerf.)Veldk.&M.P.Nayar(Veldkamp1978)438.―Dissochaeta novoguineensis Baker f. (1923)21;Mansf. (1925)114.―Lectotype(Veldkamp1978):Forbes 708 (holoBM;isoBM,E,K,L), NewGuinea,CentralDistrict,SogeriRegion,MtWori-Wori.

Creochiton divitiflorus Mansf.(1925)135.―Type:Ledermann 9576 (holoBn.v.;isoL),NewGuinea,EastSepikDistrict,Etappenberg.

Creochiton schlechteri Mansf.(1925)134.―Type:Schlechter 19234 (holoBn.v.,isoS),NewGuinea,KaiserWilhelmsland,Jaduna.

Epiphytesorwoodyclimbers,upto20mtall;branchletsterete,glabrous to furfuraceous, young ones more or less densely furfuraceous, up to 5mmdiam; nodes thickened.Leaves coriaceous,elliptictooblong,apexacute,basecuneate;gla-brous, dark green above, pale green below, midrib below brown furfuraceous,7.2–13by2.4–6cm;nervationwith1–2pairsofmainnervesarisingabovethebaseclosetothemargin;petiolesparselyfurfuraceous,10–13mmlong.Inflorescences axillary, umbellate topaniculate,up to14cm long,up to35 flowers;mainaxisterete,furfuraceous;bracteolescoriaceous,obovate,glabrous outside, sparsely stellate furfuraceous inside, white or yellowish,3–7by2–3mm,envelopingtheflowerbuds.Hypan-thium campanulate, light green, sparsely stellate furfuraceous, 3–5by2–3mm;calyxrimtruncate,withdistinctundulations,1mmlong;petalselliptic,apexacuminate,creamy,whiteorpinkish,c.9by6mm.Stamens8,equal;filamentflat,3–4mmlong;antherselongate,purplish,4–5mmlong;connectivewithasagittatecrest,0.5–1mmlong;lateralappendagesabsent;

ovaryglabrous,topconcave,2–3mmlong;styleglabrous,c.6 mmlong;extra-ovarialchambersnotdeveloped.Fruits subglo-bose,glabrous,5–6mmlong,darkredturningbluishpurpleorblack;seedsc.0.7mmlong. Distribution―NewGuinea. Ecology&Habitat―Ridgeforestbetween175–2000malti- tude. Vernacularname―Marendekan(Wagau).

Note―Creochiton novoguineensis differs from all conge-ners by: petioles 10–13 mm long, sparsely furfuraceous, leaves 7.2–13cmlong,basecuneate,underneathglabrous(butformidrib). Inflorescencepaniculate,many-flowered,bracteolescompletely covering the flower bud, 3–7 mm long, fleshy, and hypanthiumsparselystellatefurfuraceous.SimilartoC.leder-manniiintheshapeof8equalstamens,butdiffersintheshapeofleavesandinflorescences.Creochiton novoguineensis has oblong glabrous leaves and paniculate inflorescences, while C. ledermannii has obovate leaves underneath densely tomentoseandumbellateinflorescences.ContrarytotheotherspeciesinNewGuinea,C.novoguineensis has no developed extra-ovarialchambers.Theleavesofthisspeciessometimeshave basal lateral main nerves close to the margin of the leaf (Aet & Idjan 296).

10. Creochiton pudibundus(Blume)Blume―Fig.2,Map2

Creochiton pudibundus (Blume)Blume(1831a)506;(1831b)248;Naudin(1852)154;Miq.(1855)560;Triana(1871)85;Cogn.(1891)604;Krasser(1893)178, f.77b;Koord. (1912)693; (1923)212;Bakh.f. (1943)309;(1963)363.―Melastoma pudibundum Blume (1823)71; (1826)1071;DC.(1828)148.―Diplectria pudibunda (Blume)Kuntze(1891)246,nom.inval. prov.―Lectotype (designatedhere):Blume s.n. (holoL, sh. no.908.129–971;isoBO,Lsh.908.129–904,-911,-932,-961,-981,-991,-993),Java,G.Salak,Cicelebak.

Creochiton pudibundus (Blume)Blumevar.coriaceusNaudin(1852)154;Miq.(1855)560.―Type:Zollinger 3196 (holoGn.v.;isoL),Java.

Creochiton emarginatus Miq.(1855)561.―Type:Junghuhn s.n.(holoU,sh.000543;isoL,sh.908.129-952),Java,Pengalengan.

Epiphytes,upto1mhigh,orclimbers,c.30mhigh;branchletsterete,glabroustosparselyfurfuraceous,4.5–6.5mmdiam;nodes thickenedwith an interpetiolary ridge.Leaves coria-ceous, ovate to elliptic, apex acute, base rounded or cuneate, glabrous above, midrib sunken above, sparsely puberulous to furfuraceousbelow,7–10by3.7–5.8cm;nervationplinervedwith2pairsofmainnervesarisingfromabovethebase;petioleglabroustopuberulous,terete,8–15mmlong.Inflorescences axillary,umbellatewith4–7flowers,3–5cmlong;mainaxisterete,puberulous,1.5–3cmlong;bractsovate,denselystel-latehairy,2–3mmlong;bracteolesglabrous,obovate,cream,concave,envelopingtheflowerbud,7–10by4–5mm;pedicelsparselystellatefurfuraceous,5mmlong.Flowerbudc.11by10mm.Hypanthium campanulate, glabrous, rather angular, 2–4by2–3mm;calyxrimtruncatewith4undulations,c.0.5mmlong;petalsovate,violetorred,glabrous,6–13by6–7mm.Stamens8,equal;filamentflat,7mmlong;antherelon-gate,5–7mmlong;connectivewithatriangularcrest,0.5–1mmlong;lateralappendagesabsent;ovaryglobose,glabrous,c.3by2mm;styleglabrous,curvedwhenmature,c.10mmlong;extra-ovarial chambersnotdeveloped.Fruits globose, glabrous,white,3–5by4–6mm;stalkglabrous1.5–1.7cmlong;seeds0.7–0.75mmlong. Distribution―Java(west)andCelebes(southwest,south-east). Ecology&Habitat―Edgeofprimaryorsecondaryforest,edgeofriveroronamountainslopebetween1000–1400maltitude. Vernacularname―Caluncung,Harendongareuy(Sunda-nese for about any climbing Melastomataceae), Harendongareuyputih,Harendongareuymerah.

226 Blumea – Volume 58 / 3, 2013

Note―Creochiton pudibundus is easily recognized by its coriaceous ovate to elliptic glabrous leaves, underneath sparsely puberulous to densely furfuraceous, axillary umbel-lateinflorescences,coriaceous,concavebracteoles.Similarto C.bibracteatus in the axillary umbellate inflorescence, number and shape of the stamens, but differs in the shape and indu-ment of the leaves which in C.bibracteatus are thin, ovate and denselytomentose.Creochiton ledermannii differs by the small leaveswitharetuseapexandamoredenseindument.Appar-entlynotcollectedinJavaafterBakhuizen 5524(13June1922).

11. Creochiton roseus Merr.―Map3

Creochiton roseus Merr.(1905)32;(1923)191.―Type:Copeland 1297 (holo PNHn.v.,probablylost;isoK,NY,US),Philippines,Mindanao,DavaoDis- trict,Todaya.

Woodyclimbers,upto6mtall;branchletsterete,sparselyfurfu-raceous,distallyfloccose,2–3mmdiam;nodesthickenedwithaninterpetiolaryridge.Leaves thin to subcoriaceous, ovate to elliptic,apexacute,baseroundedtocuneate;aboveglabrous,below glabrescent,midrib densely furfuraceous, 6.2–9 by3.2–4.5cm;nervationplinervedwith1–2pairsofmainnervesarising fromabove the base; petiole densely furfuraceous,1.5–2cmlong.Inflorescences axillary, 2–3 cm long, umbellate topaniculate,3–5-flowered;mainaxisdenselyfurfuraceous,1–2.5 cm long; bracts not seen; bracteoles subcoriaceous,obovate to elliptic, veined, apex rounded, glabrous outside, densely stellate furfuraceous inside, enveloping the flower buds,6–12by4–6mm;pedicel0.5–1cmlong.Hypanthium campanulate or urceolate, densely stellate furfuraceous, 3–5 by2–3mm; calyx rim truncate; petals elliptic, apexobtusec.7by3–4mm,bluishpink.Stamens8,equal;filamentflat,greenish,4–4.5mmlong;antherselongate,yellow,4.5–5mmlong;connectivewithatriangularcrest,purple,c.1mmlong;lateralappendagesabsent;ovary3–4mmlong,glabrous,apexconcave;styleglabrous,greenish,8–9mmlong;extra-ovarialchambersnotdeveloped.Fruitsglobose,depressed,6–7by8–10mm,green,glabrous;seeds0.8–1mmlong. Distribution―Philippines(Mindanao). Ecology&Habitat―Primaryforestbetween800–1200maltitude. Vernacularname―Isag(Baguio).

Note―Creochiton roseus is distinguished by its floccose branchlets, 3–5-flowered inflorescences, densely stellate fur-furaceoushypanthiumwithacompletely truncatecalyx rim.Creochiton dipterus differsby the coriaceous leaves, and4smalldistinctteethontherimofthecalyx.AnotherPhilippinespecies, C.bracteatus,hasthinbracteolesandonly4stamens.

12. Creochiton turbinatus (J.F.Maxwell)Karton.,comb. & stat. nov. ―Map5

Creochiton ledermannii Mansf.var.turbinatus J.F.Maxwell,Gard.Bull.Sin-gapore33(1981)323, t.7,quoadtypus.―Type:BW 12260 (Schram)(holoL;isoBO;alsoinA,BRI,CANB,K,LAE,MANn.v.),NewGuinea,EofSorong,WarsamsonValley,Asbakin.

Epiphytes;branchletsterete,glabrescent,innovationsdenselyfurfuraceous; nodes thickenedwith an interpetiolary ridge.Leaves coriaceous, ovate, apex acute, base rounded, above glabrous, light green, below glabrescent midrib densely furfu-raceous,4–6by2.2–3.1cm;nervationplinervedwith1pairofmainnervesarisingfromabovethebase;petioledenselyfurfuraceous,3–5mm long. Inflorescences axillary, up to 5 cmlong,umbellate,2–5-flowered;mainaxisterete,denselyfurfuraceous;bractsovate,glabrescent,c.2by1mm;bracte-oles coriaceous, orbicular, concave, glabrous, enveloping the flowerbuds,yellowishgreen,5–7by3–4mm;pediceldenselyfurfuraceous,5–10mmlong.Hypanthium campanulate, gla-brescent,2–3by2mm,calyxrimtruncate;petalselliptic,2–3by2mm.Stamens8,equal;filamentflat,c.2mmlong;antherselongate,2–3mmlong;connectivewithatriangularcrest,1mm long;lateralappendagesabsent;ovaryglobose,1–1.4mmlong,glabrous;styleglabrous,c.4mmlong;extra-ovarialchambersshallow,c.1/5aslongastheovary.Fruits globose, glabrous, green,4–6by3–4mmdiam,turbinatewith8distinct lines;seedsc.0.5mmlong. Distribution―NewGuinea(Vogelkop). Ecology&Habitat―Primaryforestonhumidsandyclayin- undated in the wet season, riverbank, between 50–150 m altitude. Vernacularname―Apunengapok(Waskuk).

Note―Creochiton turbinatus differs from all congeners by: petioles3–5mmlong,leavesovate,2.2–3.1cmwide,baserounded,apexacute,underneathglabrescent(butformidrib).The inflorescences are umbellate with orbicular bracteoles which completely cover the flower bud, 5–7 mm long, fleshy, glabrous.It differs from C.ledermannii by having glabrescent leaveswithanacuteapex,andtheturbinatefruits.Theumbel-late inflorescences of C.turbinatusarenot3-,but5-flowered.ThisspeciesisonlyknownfromthetypecollectednearSoronginIndonesianNewGuinea.Theparatypes(Hoogland & Craven 10735, Docters van Leeuwen 10382)belongtoC. ledermannii, because the leaves are densely tomentose and have a retuse apex.

Map 5 DistributionofCreochiton ledermannii (●),Creochiton novoguineensis (■)andCreochiton turbinatus (▲).


Acknowledgements This study was based on the specimens available in BO,L,SINGandU(nowinL),thestaffofwhichismuchthankedfortheirassistance.VeryusefulwerethesitesontheinternetshowingspecimensprovidedbyE,L,MO,NY,SandUS.DrawingsweremadebyW.SantosoandA.Kusumawati.Dr.H.-J.Esser(M)providedinformationonthespecimensthere.MrT.-L.YaoismuchthankedforcheckingtheholdingsinKEP.Dr.G.Kadereit(néeClausing;MJG)kindlyprovidedinformationonherstudieson the Dissochaeteae.

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INDEX TO SPECIMENS CITED

Unnumberedandanonymouscollectionshavenotbeenincluded.

Creochitonano = anomalusbib = bibracteatusbra = bracteatusbre = brevibracteatusdip = dipterusfur = furfuraceusled = ledermanniimon = monticola

nov = novoguineensispud = pudibundusros = roseustur = turbinatus(LT) = lectotype(ST) = syntype(T) = type(n.v.) = cited in literature but not seen by us

Aet&Idjan296:nov.Backer5617:bib;10623:bib;15169:pud;21533:pud;25976:pud;35201:pud–BakhuizenvandenBrink5524:pud;7304:bib–Brooks50(T):mon–Bruggeman272:bib–BS8104(Merrill):ros;11640(Ramosn.v.):dip;24661(Amarillasn.v.):dip;38878(Ramos&Edaño):ros;45610(Ramos&Edaño)(T):bra–Burkill1792:ano–Buwalda6431:ano–BW12260(Schram;T):tur.

Carr12728:nov;15173:nov–Clemens1116:nov;1427:nov;6427:nov;28616:mon;28616A:mon;30389(An.v.):mon;32646=33951(LT):mon;34455(ST):mon–Coode&Ferguson7537:mon–Copeland1297(T):ros–Craven&Schodde1449:nov–Curran7280(n.v.):dip.

DeVriese40:bib;42:pud;61:bib;73:pud;74:pud;91:bib;94:pud–Doct-ersvanLeeuwen9915:nov;10382:led.

Elmer9813(Tn.v.):dip;10563:ros;14119:dip;16671:dip–Everett7280(n.v.):dip.

FB19069(Ramos&Edañon.v.):dip–Forbes708(LT):nov;809(ST):nov.Hassan&Kadim13:ano–Hoogland&Craven10735:led.Jacobs5293(T):fur.King’sCollector1619:pud;2637:pud;5779(T):ano–Koorders24251:pud;25873:bib;25907:bib.

LAE53969(Streimann):nov;54749(Stevens&Martin):nov–Ledermann8660(T):bre;9576:nov;12640(STn.v.):led;12943(LT):led.

NGF8144(Gray):nov;21596(Sayers):nov;26480(Frodin):nov;28989(Streimann&Katik):nov;47207(Johns):nov;48158(Foreman&Kumul):nov.

PNH2642(Sulit):dip;9924(Sulit):ros;41952(Mendoza):dip.Rutten1970:bib.S.14384(Othmanb.Haron):mon;16371(IliasPaie):mon;22128(SibatakLuang):mon;25865(IliasPaie):mon;29293(IliasPaie):mon;35814(Chai):fur;36332(IliasPaie):mon;38875(B.Lee):mon;40739(IliasPaie):mon;41199(IliasPaie):mon;41645(IliasPaie):mon;45870(Dyg.Awa&I.Paie):mon;46770(Dyg.Awa&P.I.Yii):mon;58254(Yii&AbuTalib):mon–SAN46777 (Mikil) (n.v.):mon;57814 (IbanGibot):mon;87181(Madani):mon–Sapei91:bib–Schlechter19234(Tn.v.):nov–SF30876(Corner):ano–SMHI1162(Ridsdale):dip.

Takeuchi5257:nov;6100:led.Uchida73:bib.VanRoyen&Sleumer6064:nov;6365:nov–VanSteenis1154:bib.Whitmore400:ano.Zollinger3009(T):bib;3196:pud.

Revision of Creochiton (Melastomataceae)

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