revision of heteroblemma gen. nov. dissochaeteae ... · 230 blumea – volume 58 / 3, 2013 be, too....

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Blumea 58, 2013: 229–240www.ingentaconnect.com/content/nhn/blumea http://dx.doi.org/10.3767/000651913X674945RESEARCH ARTICLE

INTRODUCTION

Blume(1849)distinguishedanumberofsectionsinMedinilla Gaudich.(Melastomataceae),whichhavenotbeenacceptedbylaterauthors,e.g.Cogniaux(1891)andBakhuizenvandenBrinkJr(1943,1946,1947).However,averypeculiaroneissect.Hetero blemma Blumewith (then) as the only species M. alternifolia Blume.Stapf(1895a,b)alreadynotedthatthelatter is “a well-characterised group … differs from the other species of Medinilla very strikingly in habit; a difference which is brought about by the long and slender branches, which climb by means of aerial roots, and by the alternate, generally long-petioled, and often large leaves, the transverse venation is more marked than in most Medinillae”.Curiously, in between the illustrations for his new species belonging to Heteroblemma(t.2411,2417)hedescribedthegenus Hederella Stapfwith4species(nowCatanthera F.Muell.)(t.2415,2416)butdidnotnote thesimilarityordifferencesbetween this and Heteroblemma.Asurveyoftheinfra-familiarclassificationwasgivenbyRen-ner(1993).SheincludedCatanthera, Kendrickia Hook.f.,andMedinilla in the Miconieae DC. with38generaandc.2200species.Later(Clausing&Renner2001a)theDissochaeteae Triana were recognised as distinct with 380 species in 9 gen-era among which Catanthera, Kendrickia and Medinillasect.Heteroblemma.Medinilla has opposite or whorled leaves, which usually are moreorlessisophyllous.However,inHeteroblemma the leaves aresostronglyanisophyllousthattheyappeartobealternate.Apparently only in very young shoots one minute member of thepairisstillpresent(e.g.inSF 32755 (Corner), Veldkamp 7902),butgenerallyinolderonesthishastotallydisappearedor has become invisible because of the overgrowth by mosses andhumus.Onlyrarelysomeleavesareopposite,showingtheplesiomorphicstate.Over the years other specieswith such leaves have beendescribed.Theyallshareacuriouswoodystele.InMedinilla it is terete in transverse section, but in Heteroblemma (and

KendrickiafromSriLankawithisophyllous,oppositeleaves;seeBremer1988)itismoreorlessstellatelylobed(seeFig.1). InCatanthera (incl.PhyllapophysisMansf.) the lobes are further brokenupintolobedbodies.ThusCatanthera, Heteroblemma, and Kendrickia are united by a complex wood anatomical syn-apomorphythatappearstorelatetotheirgrowthform(Claus-ing&Renner2001a).Thisremarkable‘anomalous’anatomyisdiscussedbyVanVliet(1981)andsummarisedbyClausing&Renner(1992a:55–56).Inbrief,afteraclosedcylinderofsecondary xylem has been formed in young stems, the cam-bium starts to produce parenchymatous tissue at usually four equidistantpositions(nodoubtduetothedecussateleaves).Continuing production of xylem between these meristematic patches results in a radiate, sometimes cloverleaf-shaped xylem.Heteroblemma never has coroniform hairs, while these are usual in Catanthera.Kendrickiaisglabrous.InHeteroblemma the sessile fascicles of flowers are directly placed on opposite tubercles on the stem, while in Catanthera the inflorescences are axillary to cauliflorous, and often umbel-lateandpedunculate.Growththusseemstobemonopodial,which appears to be exceptional, for in Medinilla it apparently issympodial.InKendrickia the flowers are solitary or in umbel-shaped few-flowered terminal cymes, growth here is therefore alsosympodial.The 8 stamens of Heteroblemma and Kendrickia are equal in shape,size,anddevelopment,while inCatanthera they are alternatingly4longand4short,thelatteroftenstaminodial.The extra-ovarian pockets extend nearly to the base of the ovary, while in Medinilla they are said to reach at most to its middle (Bakhuizen vandenBrink Jr 1943, 1946, 1947: 41,keylead7b).Although especially Kendrickia is morphologically and phyto-geographically quite different from the other two, the three form a monophyletic clade when their ndhF sequences are analysed (Clausing&Renner2001a).The combination of these features clearly sets Heteroblemma apart, and one might imagine a line from Medinillas.s.toKen-drickia through Heteroblemma to Catanthera. As these are consideredtobedistinctgenera,sect.Heteroblemma should

Revision of Heteroblemma gen. nov. (Dissochaeteae – Melastomataceae) from Malesia and VietnamR.Cámara-Leret1,J.W.A.Ridder-Numan1,J.F.Veldkamp1

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

Key words

HeteroblemmaMalesiaMedinillaMelastomataceaetaxonomyVietnam

Abstract A taxonomic revision is presented of the new genus Heteroblemma(Dissochaeteae – Melastomataceae),formerly a section of MedinillawhichoccursinMalesiaandVietnamwith14species,3new,and11newcombina-tions.Descriptions,illustrations,akey,andanindextocollectorsareprovided.

Published on18October2013

230 Blumea – Volume 58 / 3, 2013

be,too.ThenameBlumegavetothesectionappearstobeindicativeofthis,also.Hegavenoexplanationforitsmean-ing, but clearly it is derived from the Greek words hetero and blemma:‘differentlooking’.BakhuizenvandenBrinkJr(1943,1946,1947)inhisrevisionoftheIndonesianMelastomataceae based on material present inLeiden(L)andUtrecht(U,nowinLeiden)casuallymentionedthe section (p.177),butdidnotmaintainit,asheregardedthecharactersastooinconstant.HeincludedM. alternifolia and twonewspeciesclearlyrelatedtoit insect.Heteromedinilla Bakh.f.,whichnameisthereforesuperfluous.In1977JRN,inacourseontaxonomicbotanysupervisedbyJFV,andBakhuizenvandenBrinkJrpreparedamanuscriptbasedontheLeidenholdings.Duetodutieselsewhereitre-maineduneditedandunpublished.SincethensomespeciesfromBorneowere publishedbyRegalado (1990).The firstauthor,RCL,aspartoftherequirementsofaMaster’sDegreeoftheLeidenUniversity,hasnowcompletedtherevision.ThegenusascircumscribedhereoccursinVietnam(3spp.),PeninsularMalaysia(1),Sumatras.l.(3),Borneo(11),Celebes(1),andNewGuinea(1).CuriousistheabsenceinThailandandCambodia, Java (erroneouslymentioned byRegalado1990:16),theLesserSundaIslands,thePhilippines,andtheMoluccas.Labelsindicatetheplantstobeepiphytes,onceasanepilith.Several species havebeennoted to grownear rivers or inperiodically flooded areas and may be at least facultative rheophytes(e.g.H. capillipes: Zainudin 4611).Theseareplantsthatoccurinriverbedswhichexperienceflashfloods(banjir)and have special adaptations to such an event: extensive root systems,tenaciousbranches,streamlinedleaves,etc.Inthiscase the numerous vegetative rootlets with which the plants are attached to their hosts or rocks can be expected to prevent the plantstobesweptaway.NonewerementionedbyVanSteenisinhisstudiesregardingrheophytes(1981,1987).Mostspeciesare rarely collected, either because they are indeed rare and local, or because they grow way up in the canopy, and so are only occasionally seen and within reach, or grow in dangerous placesalongriverbeds.

As is usual in the family, petals, and stamens are easily lost in the drying and mounting processes, so specimens often only haveundevelopedhypanthia.Fortunately,contrarytothesitu-ation in Catanthera, the leaves appear to be quite diagnostic andusefulinthedelimitationofthetaxa.However,whenthereare only a few collections it is not always clear whether two different leaf forms belong to the same taxon or represent two distinctones(seethenotesunderH. capillipes and H. loratum).

MATERIAL AND METHODS

HerbariumcollectionsfromAAU,BM,C,E,K,L,M,SAR,andSINGwereexamined.Because flower parts elongate after opening, for the measure-mentofthelengthofpetals,filaments,anthers,andstylematurebut still closed or nearly open buds were studied after briefly simmeringinwater.

Heteroblemma (Blume)Cámara-Leret,Ridd.-Num.&Veldk.,stat. nov.

MedinillaGaudich.sect.HeteroblemmaBlume,Mus.Bot.1(1849)19.—Type: Medinilla alternifolia Blume[≡Heteroblemma alternifolium (Blume)Cámara-Leret,Ridd.-Num.&Veldk.].

Woody, branchingmonopodially, climbingwith numerousadventitious roots.Hairsnevercoroniform.Branches terete, sometimes grooved, often pubescent when young; the older ones, in transverse section, with a lobed stele and a round marrowchannel.Leaves long-petiolate, apparently alternate, rarely some opposite, blades 3–9-plinerved, with or without a marginalvein,secondaryvenationscalariform(rarelyabsent),moreorlesselevatedonthelowersurface.Flowers in opposite, cymose bundles, 1–many together depending on the age of thestem,ramifloroustocauliflorousontubercles,4-merous.Hypanthium broadly to narrowly campanulate, calyx inconspicu-ous or conspicuous, sepals sometimes represented by short teeth.Petalsbroadlyfalciformtotriangular,somewhatfleshy.Stamens8,equalinshapeandsize;plectrumtriangular,rarelyrectangular, somewhat wrinkled, auriculate, shortly stalked to sessile, passing into the ventral ridge along the middle of the anther.Ovary 4-locular, obconical, extra-ovarian chambers 8, reachingnearlytothebaseofthehypanthium.Fruits glabrous tohirsute,oftenfilledwithajelly-likesubstance,calyxremnantspersistentorabsentinfruit. Distribution—14speciesinMalesia(PeninsularMalaysia,Sumatras.l.,Borneo,Celebes,NewGuinea)andVietnam.

MORPHOLOgy

Habit IncontrasttoMedinilla, in Heteroblemma all species are woody rootclimbers.Rootclimbinghasevolved in9genera in thePalaeotropicsandin9or10intheNeotropics(Clausing&Ren-ner2001a:46).Adventitiousrootsprovideafirmholdonsup-porting trunks and branches but also on rocky boulders along streams.Otherthansupport,adventitiousrootssimultaneouslyallow for the up-take of the stem run-off, which contains many nutrientsleachedfromthecanopy(Nadkarni1981).Apparently,in age the roots may die off and the stems become free from thehosts,fieldlabelsthenreportthespecimensasvinesorlianas,sometimesofconsiderablelengthe.g.c.25minH. coronatum.Mostspeciesarefoundgrowingontreesonprimaryorsecond-ary dipterocarp forests, on swamp forests, some appear to be restrictedto limestonehills(H. serpens), inmontaneforestsfromc.1500to2000m(H. bisetosum),andpresumablyas

Fig. 1 Heteroblemma alternifolium(Blume)Cámara-Leret,Ridd.-Num.&Veldk.Transversesectionthroughstele(Achmad 355,L).—Scalebar=1mm.

231R.Cámara-Leretetal.:RevisionofHeteroblemma

facultative rheophytes.The occurrence in generally closedundisturbed forests may be due to the fact that their exposed livingclimbingrootsaredraughtsensitive(Clausing&Renner2001a:57).Medinilla radicans Blume and M. rubrifronsRegaladoarealsorootclimbers,butClausing&Renner(2001a:t.3)regardedthem as parallel developments in Medinilla.

VestitureThe Melastomataceae presents the greatest diversity of ves-titure in the angiosperms (Wurdack 1986).Conventionally,trichome characters have been used in species delimitation in the Melastomataceae because of their diversity and constancy (Gleason1939).However,onyoungstemsorinnovationstheindument when present may be quite variable in expression and it may change with maturity and also may vary considerably between individuals or populations, yet many species display a remarkable constancy in trichome size, posture, pattern,anddistribution.Remarkable isH. bisetosa which has both apparently stipitate stellate hairs on the innovations and sessile stellate hairs on the hypanthia, these types being extremely rareforthegenus.Termsusedincharacterisingthevestitureare used here in the following sense: hirsute: coveredwithlong,ratherstifftrichomes. hirsutulous: minutelyhirsute. pilose: with soft, shaggy trichomes, they appear shorter

andsparserthanhirsute. stellate: trichomesstar-shaped,sessileorstipitate.

Innovations and branchletsWithtimetheinnovationsincreaseindiameterandthematurestemsorbranchletsdevelopthedistinctivelobedwoodystele.Youngbranchletsappearpubescent(hirsute)onlyinonespe-cies(H. barbatum).The bark of the branchlets is either smooth, roughly striate, or coveredbywartyelements(heretermedpustulate).Branchletscanattainadiameterofat least15mm.Yetthismaybeanartifact, as old and thicker stems perhaps have become leaf- and flowerless and so are not collected, just as in the youngest innovationswhereanisophyllymighthavebeenobserved.Onlyonecollectormentioned thedirectionof climbing: sini-strorse for H. barbatum from New Guinea (BW 8404 (Vink)).Thisdirection agrees with our observation in a collection from Borneo (S. 12139 (Ashton))ofH. decurrens, which suggests that the directionofclimbingmightbeuniformatthegenericlevel.

PetiolesA generic character seems to be that the leaves are long-petiolate and the contour of the petioles proved to be useful in the distinction of a group of species, where they are narrowly winged: H. coronatum, H. decurrens, and H. flagellatum. Ves-titureofthepetiolesvariesbetweenhirsutulous(H. barbatum, H. coronatum, H. flagellatum, H. serpens),pilose(H. formanii),orgenerallyglabrous.

LeavesThe Melastomataceae appears to be the largest family of flow-ering plants characterised by acrodromal or campylodromous pinnatevenation(Clausing&Renner2001b:486,492).The path followed by the primary nerves from the base of the leaf to the apex is an important qualitative character.Thusleaves can be either campylodromous or basal- or suprabasal acrodromous.Campylodromousnervation(H. cordatum, H. serpens) is usedhere in the senseofHickey (1973): “several primaryveins[nerves]originatingat,orcloseto,asinglepoint

and running in strongly developed, recurved arches before converging towards the leaf apex”. Incontrast,acrodromousnervation differs in the arches which are not recurved at the base.Acrodromousnervationcanbefurthersubdividedbythepositionatwhichthenervesoriginate.Thusinbasalacrodro-mous nervation, the acrodromous nerves originate at or very close(afewmm)tothebaseoftheblade,whileinsuprabasalacrodromous nervation, they originate several cm above the baseoftheblade(H. coronatum and H. loratum).Thischarac-terisindicatedherebythedistanceofthefirstpairofnervesfromthebaseoftheblade.Anothercharacterusingvenationis the distance between the most distal pair of nerves and the precedingpair.The number of nerves, type of venation, presence of secondary nervesandothercharactersfoundintheleaves(seebelow)areveryusefulfordistinguishingspeciesinMelastomes.Thebladesmayhaveaconspicuoussubmarginalvein(lesswidethanthesecondarynerves)butthiscanbeinconspicuoustoapparentlyabsent(H. bisetosum, H. sandakanense).The shape of the base of the blade is also important as a vegeta-tive character and the terminology followed here is that as found inHickey(1973).Itiscordateintwospecies:H. cordatum and H. serpens.Itiscuneate(marginsstraightornearlyso,forminga‘basalwedge’oflessthan90°)inH. barbatum.Anacuteleafbase(theverybaseacute,marginscurvedandnotstraight)is themost common case (H. alternifolium, H. bise tosum, H. capillipes, H. clemensiae, H. ke mulense, and H. sandaka-nense).Attenuateleaves(inH. formanii)arecharacterisedbythe margins progressively converging to the base and slightly extendingdownwardalongthepetiole.Long-attenuateleaves(H. loratum) differ from thepreviousone in theabruptarchtheyformatthepointoforiginofthefirstpairofnerves,theirmargins converging to the leaf base already very high up in theleaf.Theleafbaseisdecurrentwhenitsmarginextendsdownwardalongthepetioleatalowangletoit(H. coronatum, H. decurrens, H. flagellatum).Secondaryvenationisgenerallyprominent(exceptinH. bisetosum).Theangleofdivergenceofthesecondariesismeas-ured between the branch and the continuation of the source veinabovethepointofbranching(seeHickey1973).Itisacute(65–80°)inallcasesbutforH. sandakanense where the sec-ondariesdivergeatarightangle.

Domatia WithinMelastomataceae myrmecodomatia are known to occur inc.10genera,e.g.Clidemia D.Don,Maieta Aubl.,Miconia Ruiz&Pav.,Myrmidone Mart., Tococa Aubl.,andTopobeaAubl. (Jacobs1968,Cabrera&Jaffe1994).OfallthecollectionsstudiedhereonlyH. barbatumonce(outoffour)hadwhatappeartobeacaridomatiaontheleafunderside(BW 8404 (Vink)). They appear as adjacent leaf pouches on eachnervepairandtheirsizeisgreaterinthedistalmostpairofnervesthanintheproximalones.Inthefirstproximalpairofnervesdomatiaappearabsent(orifpresenttheymustbeverymuchreduced).Nearly 20 % of the myrmecophytes known worldwide belong to the family Melastomataceae(Benson1985,Huxley1986)butit seems ants have never been associated with Heteroblemma andthesizeofthepouches(afewmm2)seemstobetoosmallforthem,anyway.Asmitesaresmallandsoftbodiedtheyareusually lost when plants are pressed and dried and therefore are rarely observed in the domatia of herbarium specimens (Almeda2001),butinthefieldinCentralAmericatheyhavebeen collected in the domatia of Melastomataceae (Almeda1989,Pemberton&Turner1989).

232 Blumea – Volume 58 / 3, 2013

Inflorescences and pedicelsFlowers are crowded in dense or lax clusters on the old wood or onleafaxils.Theyarebornsinglyorrarelyincymes,subtendedbylanceolateandglabrous0.5–2.5mmbracts,theseattimesarepapillose.Thepedicelsareslenderandvaryinlength:intheflowerstheyrangefrom5–20mm.Infruittheyareusuallylonger,from5–40mm.Inflowerandfruittheycanbeeitherglabrousorpubescentinvariousdegrees(seevestiture).

Hypanthium and sepals–Fig.2The hypanthium is elliptic or urceolate in unopened buds and during anthesis its rim may become truncate whereby it may becomecampanulate.HypanthiaarehirsuteinH. barbatum, H. capillipes, and H. serpens.Theirlengthismeasuredfromthetorusdownwardstotheattachmenttothepedicel.Theterm‘torus’isusedhereinthesenseofGleason(1939)fortheringof vascular tissue at the apex of the hypanthium upon which thepetalsandstamenareinserted.Thesepalscanvaryfrom0.5–2mminlengthandinseveralspeciestheyareminutelytoconspicuouslytooth-shaped.Teetharebroadlytriangulartoovate,thoselessthan0.5mmwereconsideredtobe‘minute’;thoselongerthan0.5mmwerecalled‘conspicuous’.Theteethare mostly glabrous but in certain species they can be hirsute (H. capillipes)orprolongedbythepresenceofaslender1–3mm long hair (H. barbatum, H. coronatum, H. serpens).

PetalsPetalsuponexpansionareorientatedinaspreadingwhorl.InHeteroblemma as in most Melastomataceaetheyarefugacious.Theirnumber(4)andshapeisconstant:theyareovate,acute,andwithaclawedbase.Outsidetheyaregenerallycoveredbyminutereddish(i.s.)papillaewhichcanbebestobservedaftersimmering the flowers.Notesonpetalcolourhavenotbeenrecordedforallspecies(theyaremissinginH. capillipes, H. clemensiae, H. coronatum, H. kemulense, H. sandakanense). When present, they are usually noted aswhite or various

shadesofpink(whitishpinkish,palered),orpurple.Extremelyrare for this alliance is the report of yellow in H. decurrens by the collectors(S. 22818, Jugah ak. Kudi and Chew Wee-Lek 346),for Catanthera royeniiM.P.Nayar(Van Royen & Sleumer 7405),and for Phyllapophysis schlechteriMansf.(Kalkman s.n.).

AndroeciumIn bud theanthers lie in extra-ovarianpockets, their dorsalsides facing in towards the ovary and their ventral sides facing out.Whenthefloweropens,theanthersarepulledfromthepockets, become more or less erect and slightly twist into a row atthelowerregionoftheflower.Thestylecurvesupwardsandthestigmaispositionedabovethisrow.SeeFig.2.AnthesisishereusedinthesenseofWurdack(1953:352)asthetimeatwhichthiserectionoccurs.Atanthesistheirpositionchangesfromthebeinginflexedinbudtoerect.Filaments are taxonomically trivial, they are equal or slightly un-equal, ligular, broader at the their base than at the apex, dorso-ventrally flattened, striate, and glabrous, the basal margins are onlyslightlyexpanded.Theirlengthvariesfrom1.5–3.5mm.Because in the drying and mounting process petals and anthers are often lost in the Melastomataceae they are unknown for H. capillipes, H. clemensiae, H. coronatum, H. kemulense, and H. sandakanense.Itmaybededucedfromtheotherspeciesthat they, too, are diplostemonous, isomorphic, glabrous, taper from the bases of the thecae to the single apical pore, and present a slightly to well-developed connective that appears intheformoftwoslightlyelevatedridges.Fromthenumberofthe calyx teeth, extra-ovarian pockets, and the number of ovary loculesitisinferredthattheyarealso4-merous.The dorsal spur of the anthers, where known, is conspicuous and usually lanceolate but the lateral appendages that protrude to the sides are usually straight and minute, usually less than 0.5mm,butreaching1mminH. barbatum, H. bisetosum, and H. serpens.

Fig. 2Examplesofsomehypanthia.a.Heteroblemma barbatum (BW 8404 (Vink));b.H. capillipes(S. 36279(Ilias Paie));c.H. serpens(Ridley July 1893);d.H. coronatum (SAR 25080 (Banyeng ak Nudong));e.H. clemensiae (Clemens 3829); f.H. kemulense (Endert 4330);g.H. loratum (Haviland 1682); h.H. flagellatum(Haviland 68);i.Heteroblemma alternifolium(Soedarsono 378).—Scalebar=1mm.

cbd

a

gh

i

e

f


The exception seems to be that in H. barbatum the dorsal spur isrectangularandthelateralappendagesarerecurved.Pollination is likely by insects like bees and bumblebees, hover-flies,andthelike.Thepositionoftheanthersandtheapicalporessuggestpollinationby‘buzzing’,asiswell-knownfromtheflowersofe.g.Solanum.Forthistheyhavetoalightontheflower.

GynoeciumThe ovary is 4-locular, concrescent with the hypanthium, and with8extra-ovarianchambers.InMedinilla the depth of these does not reach beyond the middle of the ovary, in contrast in Heteroblemmatheyreachnearlytothebase.The style in bud ranges from 5–8 mm, its colour is white, pink, orreddishpurple.Generally,itiscoveredbyminutereddish(i.s.)papilsfromthebaseupwards.Atanthesisitisusuallyexsertedandcontinuestoincreaseinlengthevenafterthecorolladrops.The stigma is always punctiform, and only for two species the colourhasbeenrecorded:white.

FruitInthepalaeotropicalDissochaeteae there has been an inde-pendentevolutionofberriesfromcapsules(Clausing&Renner2001b). Fruits inHeteroblemma are hard berries (sclerifiedpericarp)withpersistentplacentas,theirinsideisusuallyfilledwithjelly.Catantherahassoftberrieswithaweaklysclerifiedpericarp, while in Kendrickia in contrast has a fleshy capsule that opens by four longitudinal cracks (Clausing&Renner2001a).Generally,thesurfaceissmooth,rarelyrugose-wrinkled (H. kemulense) andoccasionally it appears ribbed (H. bar-batum)andwiththickerwalls.Seedsarenumerous,comma-shaped with a papillate testa, up to 1 mm long, bright yellow toorange.The way of dispersal is unknown, but at least in the instances wherejellywasobservedinthefruitsitislikelytobebybirds.Clausingetal.(2000)mentionedthepresenceof‘woodyber-ries’, ‘fleshy capsules’,whichare terms in contradiction, forthefruitsherearenotberries(fleshy,indehiscent,mesocarpwell-developed, the seeds immersed in it, no cavities present, nodeveloped septs), nor capsules (dry, dehiscent, several-seeded,pericarpsplittingopen,thevalvesremainingattached).Curiously, although this seems to be a fairly common type of fruitinthetropics,thereisnogeneralterm.

DistributionMostofthespeciesareMalesianwithacentreofdiversityinBorneo(11species,8endemic),followedbyVietnam(3,1en- demic),Sumatra(3),PeninsularMalaysia(2),Celebes(1)andtheislandofNewGuinea(1endemic).OftheBorneoendemics,five(sofar)areexclusivetoSabahandSarawak(H. capillipes, H. coronatum, H. flagellatum, H. loratum, H. sandakanense)andthreearerestrictedtoKalimantan(H. cordatum, H. formanii, H. kemulense).Themostwidespreadspecies(andalsothebestcollectedone)is H. alternifolium distributedfromVietnam,theMalaysianPenin-sula(butsofarnotinbetween),Sumatra,Borneo,andCelebes.InVietnamAveryanovetal.(2003)statedthatH. alternifolium is very common in closed evergreen tropical monsoon sub-montanebroadleavedforestsbetween600–1400mandalsoin riparian communities between600–1000m.The collec-tionsmadebyAveryanovetal.in2001–2002havenotbeenstudied but we believe they may belong to any of the three species found inVietnam(H. alternifolium, H. bisetosum, or H. clemensiae).Fivespeciesarerepresentedbyonlyonecollection(H. clemen-siae, H. cordatum, H. coronatum, H. kemulense, H. sandakan-

ense).Ofthe9remainingspeciesallbutone(H. alternifolium: 107collections)havelessthan15collections(H. decurrens 13; H. loratum 11; H. serpens 11; H. capillipes7;H. formanii 7; H. bisetosum 5; H. flagellatum 5; H. barbatum 4).

KEy TO THE SPECIES

1.Matureleafbladesunderneathpubescent. . . . . . . . . . . 21.Matureleafbladesunderneathglabrous . . . . . . . . . . . . . 72. Leafbladesbasecordate,primaryvenation9-plinerved,cam-

pylodromous, distalmost primaries 5 mm away from the precedingpair.Hypanthium3mmlong. . . . . . . . . . . . . . . . . 3

2. Leafbladesbasecuneate,attenuateordecurrent,primaryvenation3–7-plinerved,basalorsuprabasalacrodromous,distalmostprimaries10–65mmawayfromtheprecedingpair.Hypanthium3.5–4mmlong . . . . . . . . . . . . . . . . . . 4

3. Innovationsandleafbladesunderneathhirsutulous.Petiolesand pedicels glabrous.Hypanthium glabrous, teethmi- nuteorabsent,0–0.2mmlong,glabrous.Antherslateralap- pendages0.2mmlong.Fruitglabrous,calyxremnantsin- conspicuous or absent in fruit, pedicels in fruit glabrous . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.H. cordatum

3. Innovations,leafbladesunderneath,pedicels,hypanthiumhirsute.Petioleshirsutulous.Hypanthiumteethconspicuous,0.5–1mmlong,crownedwithaslenderhair.Antherslateralappendages0.5mmlong.Fruitglabrescent,calyxremnantsconspicuous in fruit, pedicels in fruit glabrescent . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14. H. serpens

4. Innovationsglabrous.Petioleswinged. Leaf bladesbasedecurrent.Calyxteeth0.25–1mmlong. . . . . . . . . . . . . 5

4. Innovations hirsute or hirsutulous. Petioles terete. Leafbladesbasecuneateorattenuate.Calyxteeth0–1.5mmlong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Leafbladeslinear-lanceolate,5–6cmwide,5–6timesaslong as wide, margin entire, underneath hirsutulous, primary venation3-plinerved,firstpairofprimaries50mmawayfromtheleafbase.Calyxteethconspicuous,1mmlong.Fruit5–6by 5 mm, calyx remnants conspicuous in fruit . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. H. coronatum

5. Leafbladesellipticoroblong,11.5–16.5cmwide,1.7–2.1times as long as wide, margin laxly serrulate, underneath hirsute,primaryvenation7-plinerved,firstpairofprimarynerves5–15mmawayfromthebase.Calyxteethminute,0.2mmlong.Fruit7–8by7–8mm,calyxremnantsincon-spicuous or absent in fruit . . . . . . . . . . . . 9. H. flagellatum

6. Innovationshirsute,branchletssmooth.Petioles3–6mmwide,hirsutulous.Leafblades27.5–42.5cmlong,basecu-neate, margin glabrous, underneath hirsutulous, distalmost primaries35–65mmawayfromtheprecedingpair.Bracts2.5mmlong.Pedicelsoftheflowers5mmlong,hirsute.Hy-panthiumhirsute,teethconspicuous,1.5mmlong,crownedwithaslenderhair.Antherslateralappendagesrecurved, 1mmlong,dorsalspurrectangular.Fruit9–10by8–9mm,hirsute, calyx remnants conspicuous in fruit, pedicels in fruit 5mmlong,hirsute.—NewGuinea . . . . . . 2. H. barbatum

6. Innovationshirsutulous.Branchletsstriate.Petioles2mmwide,pilose.Leafblades13.5–23.5cmlong,baseattenu-ate, margin ciliolate, underneath pilose, distalmost primaries 10–30mmaway from the preceding pair. Bracts 1–1.5mmlong.Pedicelsof theflowers10–20mmlong,piloseorglabrous.Hypanthiumpiloseorglabrous,teethabsent.Antherslateralappendagesnotrecurved,0.2–0.4mmlong,dorsalspurtriangular-lanceolate.Fruit5–7by5–6.5mm,glabrous, calyx remnants inconspicuous or absent in fruit, pedicelsinfruit15–30mmlong,glabrous.—Borneo . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. H. formanii

234 Blumea – Volume 58 / 3, 2013

7. Leafbladesbaselong-attenuateordecurrent . . . . . . . . 8 7. Leafbladesbaseacute.........................98. Innovationsglabrescent.Petioleswinged.Leafblades9–17

cmwide,2–3.4timesaslongaswide,1.5–2.8timesaslong as the petiole, base decurrent, margin laxly serrulate, ciliolate,apexacuminate,primaryvenation5–7-plinerved,basalacrodromous,firstpairofprimaries5–15mmawayfromtheleafbase,tertiarynervesconspicuous.Calyxteethabsent.Anthers lateral appendages0.2mm long.Calyxremnants inconspicuous or absent in fruit 8. H. decurrens

8. Innovations hirsutulous. Petioles terete. Leaf blades3.5–6.5cmwide,4.6–6.6timesaslongaswide,3–5.6times as long as the petiole, base long-attenuate, margin entire, glabrous, apex acute, primary venation 3-plinerved, suprabasal acrodromous, first pair of primaries 35–110mm away from the leaf base, tertiary nerves faintly visible orinconspicuous.Calyxteethminute,0.2–0.4mmlong.Antherslateralappendages0.3mmlong.Calyxremnantsconspicuous in fruit . . . . . . . . . . . . . . . . . . 12. H. loratum

9. Submarginal vein inconspicuous, tertiary nerves faintlyvisible or inconspicuous. . . . . . . . . . . . . . . . . . . . . . . . 10

9. Submarginalveinconspicuous,tertiarynervesconspicu-ous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Innovationshirsutewithstellatehairs.Branchletsstriate.Petioles3.5–6.5cm long,1–1.5mmwide.Leafblades8–14cmlong,1.9–2.6timesaslongaswide,secondarynervesinconspicuous.Pedicelsoftheflowerswithsessilestellatehairs.Hypanthiumwithsessilestellatehairs.Fruit7mmlong,pedicelsinfruit15mmlong 3. H. bisetosum

10. Innovationsglabrous.Branchletspustulate.Petioles7–12.5cmlong,3–5mmwide.Leafblades23.5–26.5cmlong,2.9–3.8timesaslongaswide,secondarynervesconspi-cuous.Pedicelsoftheflowersglabrous.Hypanthiumgla-brous.Fruit6mmlong,pedicelsinfruit25–30mmlong . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. H. sandakanense

11. Innovationseitherhirsute,glabrescentorglabrous.Leafbladesapexacuminate.Fruit glabrous, pedicels in fruitglabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

11. Innovations hirsutulous. Leaf blades apex acute. Fruithirsute, pedicels in fruit hirsutulous.....4. H. capillipes

12. Innovationsglabrescentorglabrous.Primaryvenation5–9- plinerved.Calyx remnants inconspicuous or absent infruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

12. Innovationshirsute,primaryvenation7-plinerved.Calyxremnants conspicuous in fruit . . . . . . . 5. H. clemensiae

13. Fruitarisinginfasciclesfromoldwoodorleafaxils,7–9by6–7mm,smooth.Innovationsglabrescent,branchletspustulate or striate . . . . . . . . . . . . . . . . 1. H. alternifolium

13. Fruitarisingsinglyfromleafaxils,c.11by10mm,rugoseorslightlyribbed.Innovationsglabrous,branchletssmooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. H. kemulense

1. Heteroblemma alternifolium(Blume)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.—Fig.1,2I

Medinilla alternifoliaBlume,Mus.Bot.1(1849)19.—Lectotype:Korthals s.n. ‘Indrapura’ (holo L, sh. no. 908.129-1597), partly designated byBakhuizenvandenBrinkJr(1943,1946,1947:177,‘Blume s.n.’),sheetnumberselectedhere.

Medinilla scandensKing(1900)60.—Lectotype:King’s Collector 1814 (holoCAL;BM),designatedhere.

Innovationsglabrescent.Branchlets striate to pustulate, gla-brous.Petiolesterete,3.5–29cmby2–5mm,glabrous(rarelypilose).Leaves elliptictolanceolate,9–32by6–16.5cm,1.1–4times as long as wide, 1–4 times as long as the petioles, base acute, margin entire or laxly serrulate, glabrous or ciliolate,

apexacuminate,pergamentaceous(i.s.),underneathglabrous,5–9-plinerved,basalacrodromous,firstpairofprimaries2–10mm away from the base, distalmost primaries 1–100 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, angle of divergence from mid-veinacutewide,tertiarynervesconspicuous.Bracts 1–2 mm long.Flowers born on fascicles on old wood, pedicels 5–20 mm long, glabrous (rarely pilose).Mature floral buds 8–10 by3–4mm.Hypanthium urceolate to campanulate, 3–4 mm long,glabrous(rarelypilose).Sepals0.5–1.5mmlong,teethabsentorminute,0–0.5mm,apexglabrous.Petals5–6mmlong,white,pink,orviolet,outsidefinelypapillose.Filaments 1.5–3mm long.Anthers 4–5 mm long, lateral appendages not recurved,0.25–0.4(–2)mm long,dorsal spur triangulartolanceolate,1–1.5mmlong.Style5–7mmlong,glabroustopapillose.Fruit7–9by6–7mm,glabrous(rarelypilose),calyxteeth inconspicuous or absent; pedicels in fruit 5–30 mm long, glabrous(rarelyhirsutulous);insidefilledwithjelly. Distribution—Vietnam(ThuaThiên-Huê),PeninsularMalay-sia(Johor,Pahang,Perak,Terengganu),Sumatra(widespread),Borneo(widespread),Celebes(UpperSopuValley). Habitat—Primarymixeddipterocarp forest, secondary for-est, marshy forest, fresh water swamp forest, near water, along river side, on sandy clay, loam, sandstone, sometimes ultra-mafic,0–1100maltitude. Collector’snotes—Succulenttwiningclimberwithroots,be-cominglianaceous,vine,upto15mhigh.Innovationspurplish,bronze.Stems,twigsgreybrown,brownish,leavesapparentlyon short shoots, growth in length apparently by other stems withmoredistant leaves(Stevens et al. 521).Young leavesopposite,veryanisophyllous,olderonesapparentlyalternate.Petiolesthick-fleshy-juicy,palegreen,purplishreddish.Leavesmid-green above, beneath reddish purple becoming pale green, deeplyribbed,nervesreddish,bullatebetweenthenerves.In-florescencescauliflorous,deeppinkishred.Pedunclesgreen,red.Pedicelsyellowishgreen.Hypanthiumpink,red,reddishgreen,yellowishgreen.Petalswhite,pinkish(especiallyatthetips),pinklilac,violet.Filamentswhite,antherscreamyyellow,yellow,appendageundulatelymargined.Styleswhite.Stigmayellowish,brightred.Fruitsberries,dirtypalegreenish,yellow-ish green, pale yellow, yellow, lemon yellow, greenish orange, orange,reddishorange,brightred.

Notes—Thebestcollectedspeciesofthegenus,itpresentsgreat variability in the morphology of the leaves and the reported colourofthefruits,wherebythe‘fruits’clearlyarehypanthiafromwhichthepetalsandstamenshaddroppedoff.FromCelebesthereisonlyonecollection(Van Balgooy 3531) and it has hirsutulous pedicels but otherwise it falls within the variabilityofthespecies.FromVietnamthereisonlyonecol-lection(Poilane 29857)foundintheThuaThiên-Huêarea,withthe longest lateral appendages of all H. alternifolium collections (upto2mm).ItdiffersfromH. cordatum in the leaf with an acute base, basal acrodromousvenation,andtheglabrousunderside.107collectionswereseen.

2. Heteroblemma barbatum (Bakh.f.)Cámara-Leret,Ridd.- Num.&Veldk.,comb. nov.—Fig.2a

Medinilla barbata Bakh.f.,Contr.Melastom.(1943)176.≡RecueilTrav.Bot.Néerl.40(1946)176.≡Meded.Bot.Mus.Herb.RijksUniv.Utrecht91(1947)176.—Type:Lam 663 (holoL;BO).

Medinilla jappenensis Ohwi,ined.—Voucher:Aet & Idjan 825 (BO,L).

Innovationshirsute(setaeupto5mmlong).Branchlets smooth, hirsute(setaeupto8mmlong).Petiolesterete,5–14.5by3–6mm,hirsutulous.Leavesoblongtolanceolate,27.5–42.5by6.5–13cm,2.8–3.9timesaslongaswide,2.2–3.5timesas


long as the petioles, base cuneate, margin entire, glabrous, apexshortlyacuminate,texturepergamentaceous(i.s.),under-neathhirsutulous(especiallyontheprimaries),5–7-plinerved,basalacrodromous,firstpairofprimaries3mmawayfrombase,distalmostprimaries35–65mmawayfromtheprecedingpair,submarginal vein conspicuous, secondary nerves conspicuous, 30–35 pairs, angle of divergence from midvein acute wide, tertiary nerves conspicuous.Bracts 2.5mm long.Pedicels 5mmlong,hirsute.Mature floral buds urceolate to campanu-late,8mmlong.Hypanthiumurceolatetocampanulate,3.5mmlong,yellowishbrown,hirsute,hairsupto5mmlong.Sepals 0.5mm long, teeth conspicuous, 1.5mm long, crownedbya2mmlonghair.Petals5mmlong,pink,outsidepapillose.Filaments2–3mmlong.Anthers 3 mm long, lateral append-agesrecurved,1mmlong,dorsalspurrectangular,1mmlong.Fruit 9–10 by 8–9 mm, light green when young, later on white, surface ribbed, hirsute, calyx teeth persistent; pedicels in fruit 5mmlong,hirsute;insidefilledwithjelly,seedslessthan0.5mmlong,yellow(i.s.). Distribution—IrianJaya(Biak,Jayapura(Hollandia)),Jap-penBiak,Mamberamo),PapuaNewGuinea(WProv.). Habitat—Mixedrainforest,soilpeaty,claywithgravel,60– 850maltitude. Collector’s notes—Sinistrorse root climber.Hypanthiumyellowishbrown.Petalspink.Fruit palegreenwhenyoung,laterwhite.

Notes—BW 8404 (Vink) is the only collection out of a total of fourwhichhaswhatapparentlyareacaridomatiaonitsleaves.These small pouches are a few mm2 in area and are located adjacenttothemidvein.4collectionswereseen.

3. Heteroblemma bisetosum (Bakh.f.)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.

Medinilla bisetosa Bakh.f.,Contr.Melastom. (1943)177.≡RecueilTrav.Bot.Néerl.40(1946)177.≡Meded.Bot.Mus.Herb.RijksUniv.Utrecht91(1947)177.—Type:Forbes 2342 (holoL;BM,BO).

Medinilla alternifolia auct.nonBlume:Baker,J.Bot.62,Suppl.1(1924)40.

Innovationshirsute,hairsstipitatestellate.Branchlets striate, glabrous.Petiolesterete,3.5–6.5cmby1–1.5mm,glabrous.Leaveselliptictooblong,8–14by4–7cm,1.9–2.6timesaslongaswide,1.7–2.8timesaslongasthepetioles,baseacute,margin entire, glabrous, apex acuminate, texture somewhat leathery,thin(i.s.),underneathglabrous,5-plinerved,basalac-rodromous,firstpairofprimaries1–5mmawayfromthebase,distalmost primaries 5–20 mm away from the preceding pair, submarginal vein inconspicuous, secondary nerves inconspicu-ous.Bracts1mmlong.Flowers born in dense fascicles on old branchlets, pedicels 5 mm long, with sessile stellate hairs or glabrous.Hypanthium urceolate to campanulate, 3 mm long, withsessilestellatehairstoglabrous.Sepals 1 mm long, teeth minute, less than0.5mm,glabrous.Petals pink.Filaments 2mmlong.Anthers 3–4 mm, lateral appendages not recurved, 0.5–1mmlong,dorsalspurtriangular,0.5–1.5mmlong.Fruit 7 by 5mm, surface smooth, glabrous, calyx teeth absent;pedicelsinfruit15mm,glabrous;insidefilledwithjelly,seedsgreaterthan0.5mmbutlessthan1mm,yellow(i.s.). Distribution—Sumatra (Wcoast)andVietnam(LaoCai,LamDong,NinhThuanandKontumprovinces). Habitat—NotesonhabitatandecologyareonlyavailablefromVietnamcollections.Inclosedprimarybroadleavedmoun-tainforest,between1500–2000m,common. Collector’snotes—Epiphyticcreepingvineupto15mlong,cauliflorous.Petalspalepinktopink.

Notes—BakhuizenvandenBrinkJrerroneouslycitedBün- nemeijer 5662 asthetype.ThisisthetypeofM. buenne meijeri

Bakh.f.(1943,1946,1947:192).TheForbescollectioniscor-rectlycitedinhislistofcollectorsandthespecimenitselfinLisalsosolabelledbyBakhuizenvandenBrinkJrhimself.Thismix-up of type citations is not covered by the Code, but we have regarded it as a bibliographic error that can be corrected, similartoArt.33.5.In1943onewasnotrequiredtodesignateatype,anyway(Art.8).InexplicableisthedisjunctdistributionbetweenVietnamandSumatra.PreviouslyonlyknownfromthetypeinSumatra,ithasalsobeenfoundinVietnam.Althoughwetried,thematerialis indistinguishable except for the stipitate stellate hairs on the innovations and the sessile and stellate hairs in the younger flower pedicels and buds, unique for Heteroblemma.5collectionswereseen.

4. Heteroblemma capillipes (Regalado)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.—Fig.2b

Medinilla capillipes Regalado,Blumea35(1990)61,t.14.—Type:S. 36279 (Ilias Paie)(holoK;KEP,L,MO,SAN,SAR).

Innovations hirsutulous.Branchletsglabrous.Petioles terete, 6.5–13cmby2–3mm,glabrousorsetose(red).Leaves ellip-tictooblong,14–24by6.5–11.5cm,1.6–2.8timesaslongaswide, 1.5–2.4(–3.5) timesas longas thepetioles, baseacute, margin entire, ciliolate, apex acute, pergamentaceous (i.s.),underneathglabrous,5-plinerved,basalacrodromous,firstpairofprimaries2mmaway from thebase,distalmostprimaries 5 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, 20–25 pairs, angle of divergence from midvein acute wide, tertiary nervesconspicuous.Bracts1.5–2mmlong.Flowers born on old wood or axillary on young stems, pedicels 15–20 mm long, hirsutulous.Hypanthium urceolate to campanulate, 3–4 mm long,hirsute,hairsyellowish.Sepals0.5mmlong,teethmin-ute,lessthan0.5mmlong,hirsute.Petalspalepink.Fruit6–8by5–7mm,brightyellow,orangeorreddishorange,smooth,hirsute(rarelyglabrescent),hairsbrown,calyxteethpersistent;pedicelsinfruit25–40mmlong,green,hirsutulous;insidefilledwithjelly,seedslessthan0.5mmlong,yelloworange(i.s.). Distribution—EndemicofSarawak(3rd,7th,and9thDiv.). Habitat—Alongstreambanksinforestsonyellowsandysoil, on mossy boulders, in seasonally flooded riverine forest on pale yellow clay, as a rheophyte in lowland dipterocarp forest, andonsubmontanemossyforest,60–820maltitude. Collector’snotes—Creeper,climber,orrheophyte,orter-restrial,withadventiveroots,upto2.1m.Stemdarkbrown,brownhairy.Leavesslightlypapery,abovegreen,beneathpalegreen.Peduncledarkred.Pedicelspalegreen.Youngcalyxpinkwithlongstraw-colouredhairs.Petalspalepink.Fruitbrightyellow, orange yellow to orange, red, with yellowish orange to brownbristles,pedicelgreentoyellowishgreen.

Notes—S. 37559(Chai et al.)differsinitsshorterpetioles,leaves being linear-lanceolate, decurrent base, 3-plinerved, but showsthedistinctivehirsutefruits.WeareunsureofS. 66556 (Yahud Hj Wat et al.) from the 5th Div.ofSarawak. Itdiffersinitslanceolatetolinear-lanceolateleaves, its decurrent base, venation which is 3-plinerved and itsglabrousfruits.7collectionswereseen.

5. Heteroblemma clemensiae Cámara-Leret,sp. nov.—Fig.2e

Innovationeshirsutae,foliilaminabasiacuta,apiceacuminato,7-plinervatainfra glabra, vena submarginali conspicua, nervis tertiariis conspicuis, fructus glabri, pedicellis glabris, calycis reliquiae conspicuae.—Type:Clemens 3829(holoK;L,notfound).

236 Blumea – Volume 58 / 3, 2013

Fig. 3 Heteroblemma coronatumCámara-Leret.a.Habit; b. firstpairofprimarynerves;c.infructescences(all:SAR 25080(Banyeng ak Nudong)).—Scalebars:a,c=1cm;b=1mm.

c

b

a


Innovationshirsute.Branchletsstriate,glabrous.Petioles terete, 1–6.5cmby3mm,glabrous.Leaves oblong to lanceolate, 13–23by5–7cm,2.5–3.3timesaslongaswide,3.5–15timesas long as the petioles, base acute to obtuse, margin entire, glabrous,apexacuminate,pergamentaceous(i.s.),underneathglabrous,7-plinerved,basalacrodromous,firstpairofprimaries3–5mmawayfromthebase,distalmostprimaries4–5.5cmaway from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, 15–20 pairs, angle of diver-genceacutewide,tertiarynervesconspicuous.Bracts 2 mm long.Pedicels up to 20 cm long, glabrous.Flowers born in densefasciclesonoldwood.Budsc.12by7mm.Hypanthium campanulate,c.8by7mm,veryfinelyred-papillose.Petals c.5by5mm,red-purple(i.s.).Filamentsc.2mmlong;antherc.1.2mmlong,dorsalspuradaxiallydirected,c.0.5mmlong,lateralappendagestriangular,c.1mmlong.Stylec.7mmlong, whitepapilloseinlowerpart.Fruits8.5–10by7–9mm,smooth,glabrous,calyxteethpersistent,c.2mmlong;seedslessthan0.5mm,yellow(i.s.). Distribution—EndemicofVietnam,Tourane(nowDaNang). Habitat—Onlargeforesttree. Collector’snotes—Liana.Flowersinfasciclescauline,neartheground.

Note—Onlyknownfromthetype.Thespecimenwithflow-ersinLusedbyJRNforher1977descriptioncouldnotbefoundagain.

6. Heteroblemma cordatum Cámara-Leret, sp. nov.

Innovationeshirsutullae,petioliglabri,foliilaminabasicordatainfrahirsutulla9-plinerva nervatione campylodroma nervis primariis a pare precedenti 5 mm distanti, pedicelli glabri, hypanthium 3 mm longum glabrum dentibus nullis velminutis0–0.2mmlongisglabris,antheraeappendiceslaterales0.2mmlongae, fructus glabri calycis reliquiae nullae vel inconspicuae pedicellis glabris.—Type:Burley, Tukirin et al. 3155 (holoL;A,BO).

Innovationshirsutulous.Branchletspustulate,glabrous,c.10– 15mmdiam.Petiolesterete,c.16cmby3–4mmwide,glabrous.Leaves 15.5cmwide,basecordate,marginlaxlyser-rulate, ciliolate, pergamentaceous, underneath hirsutulous alongreticulations,9-plinerved,campylodromous,firstpairofprimaries 0–5 mm away from base, distalmost primaries 5 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, angle of divergence acute wide,tertiarynervesconspicuous.Bracts2mmlong.Flowers born on dense fascicles on old wood, pedicels 10 mm long, glabrous.Mature floral buds7–8by3.5mm.Hypanthium ur-ceolatetocampanulate,3mmlong,glabrous.Sepals0.5–1mmlong,teethabsentorminute,0–0.25mmlong,glabrous.Petals6–7mmlong,white,outsidepapillose.Filaments 2 mm long.Anthers5mmlong,lateralappendagesnotrecurved,0.25mmlong,dorsalspurtriangulartolanceolate.Style6mmlong.Fruit6by5mm,smooth,glabrous,calyxteethinconspicuousor absent; pedicels in fruit 15–35 mm long, glabrous; seeds lessthan0.5mm,yellow(i.s.). Distribution—EndemicofWKalimantan:NEofPontianak,G.Bentuangarea. Habitat—Mixeddipterocarpforestonsandstoneridge,oc-casional,400maltitude. Collector’snotes—Climber,25mtall.Petalswhite.

Notes—AlthoughsimilartoH. serpens by its cordate leaf base and the 9 primary nerves, it is different in its glabrous flower and fruit pedicels and hypanthium, while its calyx teeth areminuteorabsentandglabrous.ItdiffersfromH. alterni-folium in its cordate leaf base and campylodromous venation andabaxiallyitishirsutulousalongthereticulationsbelow.Onlyknownfromthetype.

7. Heteroblemma coronatum Cámara-Leret,sp. nov.—Fig.2b, 3

Innovationesglabrae,petiolialati,foliilaminalinearilanceolata5–6cmlata5–6-plolongaquamlatabasidecurrentiinfrahirsutulla3-plinervavenationesuprabasaliter acrodroma nervorum parimariorum pare prima 50 mm laminae basidistanti,hypanthium3.5–4mmlongumdentibusconspicuis1mmlongis,fructus5–6mmlongi5mmlaticalycisreliquiaeconspicuae.—Type:SAR 25080(Banyeng ak Nudong) (holoL,sh.107500;A,K,L,SAN,SAR,SING).

Innovations glabrous.Branchlets absent.Petioles narrowly winged,6–16cmby3mm,hirsutulous.Leaves linear-lanceo-late,28–33by5–6cm,5–6timesaslongaswide,2–4timesas long as the petioles, base decurrent, margin entire, ciliolate, pergamentaceous(i.s.),underneathhirsutulous,3-plinerved,suprabasalacrodromous,firstpairofprimaries50mmawayfrom the base, distalmost primaries 50 mm away from the precedingpair(thesubmarginalvein),submarginalveincon-spicuous, secondary nerves conspicuous, 20–25 pairs, angle ofdivergenceacutewide,tertiarynervesconspicuous.Bracts 1mmlong.Flowersabsent,bornon fasciclesonoldwood.Hypanthiumurceolatetocampanulate,4mmlong,glabrous.Sepals1mmlong,teethconspicuous,1mmlong.Fruit5–6by5 mm, glabrous, red, calyx teeth persistent, crowning the fruit conspicuously, prolonged by a caducous slender hair, pedicels infruit25–35mmlong,glabrous;insidefilledwithjelly. Distribution-–EndemicofSarawak:4thDiv.,LambirNationalPark. Habitat—Nearbyriver. Collector’snotes—Climbing3mhighinatree.Fruitsred.

Notes—Namedafteritsdistinctivecrownedfruits,adornedbyveryconspicuouscalyxremnants.Onlyknownfromthetype.

8. Heteroblemma decurrens (Cogn.)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.

Medinilla decurrens Cogn. inA.DC.,Mon.Phan. 7 (1891) 591.—Type:Beccari PB 4016(holoFI;BR).

Innovationsglabrescent.Branchlets striate, glabrous, up to 10 mmdiam.Petioles winged,5–18.5cmby2–4mm,glabrous(rarelypilose).Leaveselliptic to lanceolate,10–31by9–17cm,2–3.4timesaslongaswide,1.5–2.8timesaslongasthepetioles, base decurrent, margin laxly serrulate, ciliolate, apex acuminate,pergamentaceous(i.s.),underneathglabrous(rarelypilose),5–7-plinerved,basalacrodromous,firstpairofprimaries5–15 mm away from the base, distalmost primaries 10–30 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, 25 pairs, angle of divergence frommidveinacutewide,tertiarynervesconspicuous.Bracts 0.5–1.5mmlong.Flowers born in dense fascicles on old wood, pedicels10–15mmlong,palered,glabrous.Mature floral buds 8–9by3–4mm.Hypanthium urceolate to campanulate, 3 mm long,orangeorpalered,glabrous(rarelypilose).Sepals0.5mmlong,teethabsent,glabrous.Petals6–7mmlong,yelloworpink,outsidefinelypapillose.Filaments2–3.5mmlong.Anthers 4–5mmlong,creamy,lateralappendagesnotrecurved,0.25mmlong,dorsalspurtriangulartolanceolate,1–2mmlong.Style5–6mmlong,creamyorwhite,stigmawhite.Fruit5–7by4–6mm,paleyellowtoorange,smooth,glabrous,calyxteeth inconspicuous or absent; pedicels in fruit 15–40 mm long,green,glabrous; insidefilledwith jelly,seedsc.1mmlong,yellow(i.s.). Distribution—Malaysia:Sarawak,Brunei(TemburongDistr.:KualaBelalong). Habitat—Clayrichalluviumalongstreams,riverbank,inoccasionallyfloodedriverineforest,25–950maltitude. Collector’s notes—Climber, reportedly up to 10mhigh,stemnaked except for the top 1.5m, greyish,mid-brown,

238 Blumea – Volume 58 / 3, 2013

chocolate brown.Petiole proximally tinged brown. Leaveswhenyoungpinkish(green),laterabovedeepgreen,beneathpaler,drillpurple.Flowerscauliflorous.Pedunclespalegreen,green.Pedicelpalered;hypanthiumpalered,orange.Petalspink; pale yellow to yellow flowers as noted for S. 22818(Jugah ak. Kudi )andChew Wee-Lek 346 are extremely exceptional inthealliance.Filamentswhite,antherscream.Stylecreamy,white;stigmawhite.Fruitspaleyellow,orange;pedicelgreen,pale orange

Note—13collectionswereseen.

9. Heteroblemma flagellatum (Stapf)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.—Fig.2h

Medinilla flagellataStapfinHook.f.,Icon.Pl.(1895)t.2411.—Type:Havi-land 68(holoK).

Innovations glabrous.Branchlets striate, glabrous, 6–7mmdiam.Petiolesnarrowlywinged,5.5–19cmby2–4mm,hirsu-tulous.Leaves elliptictooblong,21–28by(8.5–)11.5–16.5cm,1.7–2.1timesaslongaswide,1.9–4.4timesaslongasthepetioles, base decurrent, margin laxly serrulate, ciliolate, apex acuminate,pergamentaceous(i.s.),underneathpalergreen,hirsute,hairspurplishbrown,7-plinerved,basalacrodromous,firstpairofprimaries5–15mmawayfromthebase,distalmostprimaries 10–30 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, 20–25 pairs, angleofdivergenceacutewide,tertiarynervesconspicuous.Bracts1–1.5mmlong.Flowers born in fascicles on old wood, pedicels15–20mm,pinkishred,glabrous.Mature floral buds 9–10by3mm,purplishordarkpink.Hypanthium urceolate tocampanulate,3.5mmlong,pinkishred,glabrous.Sepals 1mmlong,teethminute,lessthan0.5mm,glabrous.Petals 6–7mmlong,paletodarkpink,outsidepapillose.Filaments 2mmlong,white.Anthers 4 mm long, yellow, lateral append-agesnotrecurved,0.2mmlong,dorsalspurtriangulartolan-ceolate.Style6mm,whiteatthebase,reddishpurpleupwards,stigmawhite.Fruit7–8by7–8mm,orange,smooth,glabrous,calyx teeth absent; pedicels in fruit 20–40 mm long, pinkish red,glabrous;seedsupto0.5mmlong,yellow(i.s.). Distribution—EndemicofSarawak(1stand7thDiv.). Habitat—Hilldipterocarpforest,nearriver,limestone,darkclaysoil,150–365maltitude. Collector’snotes—Rootclimber,upto8mhigh,flowers1–2mabovetheground.Petiolessucculent,darkpurplish.Leavesabove green to dark green, beneath pale green with a purplish brownindumentonthenerves.Pedicelpinkishred,darkpink.Flowerbudsdarkpink,purplish.Hypanthiumpinkishred,darkpink.Petalspalepink,pinkishwhite.Filamentswhite.Stamensarrangedinagroupsothattheyareparallel.Filamentswhite.Anthersyellow,apexanddorsalappendageslightlypurplish.Topofovarypink.Stylewhiteatbase,upwardsreddishpurple,stigmawhite.Berriesorange,tasteless.


10. Heteroblemma formanii (Regalado)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.

Medinilla formanii Regalado,Blumea35(1990)56.—Type:Forman 521 (holoK;BO,L,SING).

Innovationshirsutulous.Branchletsstriate,glabrous,c.6mmdiam.Petiolesterete,5–9cmby2mm,pilose.Leaves oblong tolanceolate,13.5–23.5by4.5–9cm,2.1–3.6timesaslongaswide,1.7–4.3timesaslongasthepetioles,baseattenuate,marginentire,ciliolate,apexacuminate,pergamentaceous(i.s.), underneathpilose(onprimaries),5–7-plinerved,basalacro-dromous,firstpairofnervesbranching5–10mmawayfrom

the base, distalmost primaries 10–30 mm away from the pre-ceding pair, submarginal vein conspicuous, secondary nerves conspicuous, c. 25pairs, angleof divergence frommidveinacutewide, tertiary nerves conspicuous.Bracts 1–1.5mm.Flowers born in fascicles on old wood, pedicels 10–20 mm long,glabrousorpilose.Mature floral buds8–10by3–4mm.Hypanthiumurceolatetocampanulate,3.5mmlong,glabrousorpilose.Sepals0.5mmlong,teethabsent.Petals5–7mmlong,purple,outsidepapillose.Filaments1.5–2(–3)mmlong.Anthers 4–5 mm long, lateral appendages not recurved, up to 0.5mm,dorsalspurtriangulartolanceolate,1–1.5mmlong.Style5–8mmlong.Fruit5–7by5–6.5mm,orangetored,smooth, glabrous, calyx teeth inconspicuous or absent; pedicels infruit15–30mm,glabrous. Distribution—EndemicofCKalimantan(UpperKatinganRiver),EKalimantan(CKutei,Tabang,BelajangRiver;Balik-papan). Habitat—Dryobalanops or Dipterocarpus mixed forest near river, primary swampy forest, on sandy yellow loam, 0–100 m altitude. Collector’snotes—Epiphytic root climber, lianescent,uptoc.15mhigh,upwardswithlateralbraches.Planthigherupwithlateralbranches.Leavespalegreenonbothsides,nervesbrown.Flower(pale)purple.Petalswhite,paledirtypinkish,palepinklilac.Anthersyellowish,yellow.Fruitgreenwhitewhenyoung,turningmid-orangeorred.

Notes—WeareunsureofthecollectionsAfriastini 15, Am-briansyah 604, Ambriansyah & Arbainsyah 1969, Mogea & De Wilde 4210 & 4250.TheydifferfromH. formanii by the leaves withanacutebase,glabrousunderneath,andare(5–)7–9-pli-nerved.7collectionswereseen.

11. Heteroblemma kemulense (Regalado)Cámara-Leret, Ridd.-Num.&Veldk.,comb. nov.—Fig.2f

Medinilla kemulensis Regalado,Blumea35(1990)57,t.13.—Type:Endert 4330(holoK;BO,L).

Innovationsglabrous.Branchletssmooth,glabrous,c.8mmdiam.Petiolesterete,(7.5–)12–17.5cmby3–5mm,glabrous.Leaves elliptic, (13.5–)19–23by (9.5–)14–15 cm, 1.4–1.5timesaslongaswide,1.3–1.8timesaslongasthepetioles,base acute, margin laxly serrulate towards the apex, ciliolate, apexacuminate,pergamentaceous(i.s.),underneathglabrous,9-plinerved,basalacrodromous,firstpairofprimaries2–5mmaway from the base, distalmost primaries 5 mm away from the preceding pair, submarginal vein conspicuous, secondary nerves conspicuous, c. 25pairs, angleof divergenceacutewide, tertiarynervesconspicuous.Bracts notseen.Flowers bornsingly in the leafaxils,pedicels10mmlong,glabrous.Mature floral budsabsent.Fruitc.11by10mm,green(unripe),rugose or slightly ribbed, glabrous, calyx teeth inconspicuous orabsent;pedicelsinfruitc.20mm,glabrous;insidefilledwithjelly,seeds0.7mmlong. Distribution—EndemicofKalimantan,WKutai, nearMtKemul. Habitat—Primaryforest,1600maltitude. Collector’snotes—Climber.Fritgreen.

Note—Onlyknownfromthetype.

12. Heteroblemma loratum (Stapf)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.—Fig.2g

Medinilla lorata StapfinHook.f.,Icon.Pl.(1895)t.2417.—Type:Haviland 1785(holoK,barcodesK000277066,-067).

Innovationshirsutulous.Branchletsstriate,glabrous,c.7mmdiam.Petiolesterete,3.5–13cmby2–4mm,glabrous.Leaves


lanceolatetolinear-lanceolate,22.5–33by3.5–6.5cm,4.6–6.6timesaslongaswide,3–5.6timesaslongasthepetioles,baselong-attenuate, margin entire, glabrous, apex acute, leathery (i.s.),underneathglabrous,3-plinerved(rarely5-),suprabasalacrodromous,firstpairofprimaries35–110mmawayfromthebase, distalmost primaries 30–100 mm away from the submar-ginal vein, submarginal vein conspicuous, secondary nerves conspicuous, 20–30 pairs, angle of divergence acute wide, tertiarynerves faintlyvisible to inconspicuous.Bracts 1 mm long.Flowers born in fascicles on old wood or axillary on young stems,pedicels10–15mmlong,glabrous.Mature floral buds 9–11by3mm.Hypanthium urceolate to campanulate, 3 mm long,glabrous.Sepals0.5mmlong,teethminute,0.25–0.4mmlong,glabrous.Petals6–7mmlong,pink,outsidefinelyred-papillose.Filaments3mm long.Anthers5–6mm long,lateral appendages not recurved, 0.3mm long, dorsal spurtriangulartolanceolate,1mmlong.Style6–7mmlong.Fruit 6by5–6mm,green(unripe),smooth,glabrous,calyxteethpersistent; pedicels in fruit 20–30 mm long, glabrous; inside filledwithjelly,seedsupto1mm,yellow(i.s.). Distribution–-EndemicofSarawak(1stand4thDiv.;Serian,Bt.Gaharu) Habitat—Primary forestonyellowclay,near river,up to660maltitude. Collector’snotes—Climber,upto3m.Flowersscatteredonmiddle(older)leafbearingpart,pinkishinbud.Stigmaviolet.Immaturefruitpalegreen,green.

Notes—WearenotsurewhetherChurch et al. 348, 409, and Hansen 1230areconspecificorperhapsrepresentaproperspecies.They differ by having 5–7-plinerved leaves, basalacrodromous venation, acute leaf bases and their distribution, andarefoundinWKalimantan.Collectorshavenotedforthis:Climber, 3.5m long. Leaves above glossy green, beneathpale green, venation very prominent, brownish purple, sunken above; secondary nerves parallel,widely spaced.Pedicelsgreen.Hypanthiumpalegreen.Petalswhite,whitewithpinktinge,palepink.Filamentsyellownearapex.Stylepurple.Fruitorange.11collectionswereseen.

13. Heteroblemma sandakanense (Regalado)Cámara-Leret,Ridd.-Num.&Veldk.,comb. nov.

Medinilla sandakanensis Regalado,Blumea35(1990)63, f.15.—Type:SAN 20602(Patrick Ping Sam)(holoK;L,SAN).

Innovationsglabrous.Branchletspustulate,glabrous,c.8mmdiam.Petiolesterete,7–12.5cmby3–5mm,glabrous.Leaves oblongtolanceolate,23.5–26.5by7–8.5cm,2.9–3.8timesaslongaswide,2.1–3.8timesaslongasthepetioles,baseacute,margin entire, glabrous, apex acuminate, pergamentaceous (i.s.),underneathglabrous,5-plinerved,basalacrodromous,firstpairofprimaries2–3mmawayfromthebase,distalmostprimaries10–20(–50)mmawayfromtheprecedingpair,sub-marginal vein inconspicuous, secondary nerves conspicuous, 20–25 pairs, at a right angle of divergence from the midvein, tertiarynerves faintlyvisible to inconspicuous.Bracts 1 mm long.Flowers unknown, born in fascicles on old wood, pedicels glabrous.Hypanthium urceolate to campanulate, glabrous.Fruit6by5mm,yellow,smooth,glabrous,calyxteethabsent;pedicelsinfruit25–30mmlong,glabrous. Distribution—EndemicofSabah(SepilokFR). Habitat—Primaryforestnearstream,atlowaltitude. Collector’snotes—Climber.Fruityellow.

Note—Onlyknowfromthetype.Jaheri 18(BO),fromKali-mantan possibly belongs here, but it has shorter pedicels and bristlyfruits(Regalado1990:65).

14. Heteroblemma serpens (Stapf)Cámara-Leret,Ridd.-Num. &Veldk.,comb. nov.—Fig.2c

Medinilla serpensStapf,Icon.Pl.(1895a)t.2411.—Type:Haviland’s col-lector 551(holoK,barcodesK000277072,-073;SING).

Innovationshirsute.Branchletspustulate,glabrous,7–8mmdiamandupto10minlength.Petiolesterete,9–31.5cmby2–6mm,hirsutulous.Leavesbroadlyelliptictoelliptic,13–27by12–23cm,0.9–1.5timesaslongaswide,0.7–1.8timesas long as the petioles, base cordate, margin laxly serrulate, ciliolate,apexacuminate,pergamentaceous(i.s.),underneathhirsute, 9-plinerved, campylodromous, first pair of primaries0–5 mm away from the base, distalmost primaries 5 mm away from the preceding pair, submarginal vein conspicuous, sec-ondary nerves conspicuous, 20–25 pairs, angle of divergence frommidveinacutewide,tertiarynervesconspicuous.Bracts 1–2mmlong.Flowers born in dense fascicles, on old wood oraxillaryonyoungstems,pedicels10–15mmlong,hirsute.Mature floral buds9–10by3–4mm.Hypanthium urceolate to campanulate,3mmlong,pinkishgreen,hirsute.Sepals0.5–1mmlong,teethconspicuous,0.5–1mmlong,crownedwitha2–3mmhair.Petals6–7mmlong,palepinkorpalered,outsidepapillose.Filaments 2–3mm long,white.Anthers 4–5 mm long,yellow,lateralappendagesnotrecurved,0.5mmlong,dorsalspur1mmlong,triangulartolanceolate.Style6–8mmlong,darkpink.Fruit5–6by5mm,orangetobrown,smooth,glabrescent, calyx teeth persistent; pedicels in fruit 15–30 mm long,pinkish,glabrescent. Distribution—Sumatra(RiauArch:TudjuhIsland),Sarawak(1stDiv.),Sabah(TambunanDistr.). Habitat—Ontreesorrocks,limestonehills,onlyonerefer-encetoaltitude:460maltitude. Collector’snotes—Creepingherb(!),climbertoaheightof 7m,orcreeper.Cauliflorousfromgroundleveltotop.Leavesbeneath pinkish. Pedicels pink, hairy.Hypanthium pinkishgreen,hairy.Petalspalered,palepink,whitishpink.Filamentswhite,anthersyellow.Stylepink,darkpink.Fruitsbrown,or-ange;pediclespinkish.


Acknowledgements(especiallybyRCL)Dr.MaxM.J.vanBalgooycontri-butedwithhisexpertiseinthegenericidentificationoftheincomingMelasto-mataceae.Prof.PieterBaaswasverykindinpointingoutsomekeyliteratureinregardstovestitureandplantanatomy.Dr.WimVinkwashelpfulwithmydoubtsonthesubjectofdomatia.Dr.PaulKeßleroncollectingmethodsandtheirinfluenceonherbariummaterial.Dr.MarcoRoosintheadministrativeissuesconcerningthethesis.Dr.FritsA.C.B.Ademaforhishelpinsearch-inglocalities.Ms.AnitaV.M.WalsmitSachs-Jansenwasextremelykindinpreparing the illustrationsand initiatingme inbotanical illustration. In theearlystagesoftherevisionvaluableadvicewasgivenbythelateDr.ReinC.BakhuizenvandenBrinkf.

Weallarealsogratefultothedirectorsandcuratorsofthefollowingherbariawho on very short notice were able to send on loan their valuable material or sendhighresolutionimages:AAU,BM,C,E,FI,K,M,andSING.

JFVvisitedSARin2012andthanksitsstafffortheirgenerousassistance.

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INDEx TO COLLECTIONS

Heteroblemmaalt = H. alternifoliumbar = H. barbatumbis = H. bisetosumcap = H. capillipescle = H. clemensiae

crd = H. cordatumcro = H. coronatumdec = H. decurrensfla = H. flagellatumfor = H. formanii

kem = H. kemulenselor = H. loratumsan = H. sandakanenseser = H. serpens

(ST) = syntype(T) = type(V) = voucher

Achmad355:alt–Aet&Idjan825(V):bar–Afriastini15:for–Ambriansyah604:for–Ambriansyah&Arbainsyah1969:for–Atkinsetal.563:alt.

Beaman7080:alt–BeccariPB2000:alt;PB4016(T):dec–Boyceetal.425:dec–Brooks&Hewitts.n.September1908:lor–BRUN491(Ashton):alt;3106(Pukol):alt;17519(Ahmadetal.):alt;17926(AriffinKalatetal.):alt–Burley,Tukirinetal.469:alt;1186:alt;3155(T):crd–Butt8288:alt–BW8404(Vink):bar.

ChewWL346:dec;1005:alt–ChewWLetal.44:alt–Churchetal.1955:alt–Clemens3829(T):cle;21595:alt;26059:alt;31321:alt;31522:alt;32389:alt–Coodeetal.6406:alt–Curtis3294:alt.

DeWilde&DeWilde-Duyfjes19399:alt;19897:alt;19986:alt.Elmer21594:alt–Endert4330(T):kem.Forbes2342(T):bis–Forman521(T):for.Geh&Samsuri905:alt.Hallierf.520:alt;2833:alt;3314:alt–Hansen26:alt;65:alt;615:fla;876:fla;962:fla–Haviland68(T):fla;701:ser;1682:lor;1785(T):lor;3133:ser;3388:lor;3633:ser–Haviland’scollector551(T):ser–Haviland&Hose3632:alt.

Iboet82:ant.Jeremy14232:alt–Johns7509:alt.Keith5246:alt–KiewKBH47:alt–King’sCollector1814 (ST):alt–Kohawa&Hotta1006:alt;1592:alt–Kostermans10446:for.

Lam663(T):bar.Mogea4210:for;4250:for–MS.1182(Shah&Sidek):alt;1415(Shah):alt;3579(Shah&Samsuri):alt;5722(Sandsetal.):alt–Murataetal.B-3030:alt;B-3118:alt.

NGF42780(Hentyetal.):bar–Nielsenetal.1054:alt–Nooteboom1124:alt.Ohai3718:alt.Parris24/85:alt–Poilane29857:alt.Ridleys.n.1890:ser;July1893:ser;2933:alt;14104:alt.

S.2810(Anderson):alt;12139(Ashton):dec;12912(Anderson):ser;16662(Paie&Ashton):lor;22818(Jugah):dec;24380(Jugah):dec;25141(Ander-son):ser;27474(Erwin&Paul):fla;27728(Smith):cap;28337(Anderson&Paie):alt;33273(Tong&Banyeng):dec;33990(Chai):alt;34673(Chai):alt;35651(Ilias&Azahari):alt;36279(IliasPaie)(T):cap;36954(Martin&O.):alt;37559(Chaietal.):cap;38613(LeeB.):ser;40260(George):dec;41034(Paie):alt;41909(LeeB.):ser;42846(Georgeetal.):alt;45296(DygAwa&Paie):lor;45358(LeeB.):dec;46900(DygAwa&YiiP.C):cap;46996(Awa&Paie):ser;48370(YiiPuanChing):alt;49466(Mohtaretal.):alt;49694(Mohtar&OthmanIs.):alt;50062(DygAwa&B.Lee):dec;51408(Mohtar):dec;53648(Ching):alt;54780(LeeB.):dec;56872(Ching):dec;62921(Runietal.):alt;66556(YahudHjWatetal.):cap;90301(Kamarudinetal.):alt;91003(Kiewetal.):alt;95792(Juliaetal.):cap–SAN20602(PatrickPingSam)(T):san;34104(Aban):alt;68354(Talip&Terimiji):alt;82506(Cockburn):alt;84854(Cockburn):alt;85133(Stone):alt;90679(Patricketal.):alt;94499(AbanG.):alt;104265(Krispi-nus):alt;119044(Donggop&Gambio):alt;128277(Krispinus):alt;128361(Sumbing):alt–SAR25080(BanyengakNudong)(T):cro–Scortechini80:alt;86(ST):alt;88:alt;130(NOT150;ST):alt–SF6200(Sinclair):alt;10393(Henderson):alt;14827(BodenKloss):alt;30127(Corner):alt;32048(Kiah):alt;32456(Corner):alt;32755(Corner):alt;36301(Spare):alt;38804(SinclairetKiah):alt;40448(Sinclairetal.):alt–Soedarsono214:alt;378:alt–Stevensetal.521:alt.

Torquebiauetal.463:alt;3598:alt.VanBalgooy3531:alt–VanValkenburgetal.1086:alt–Veldkamp7902:alt–VH640(Averyanovetal.):bis;2911(Averyanovetal.):bis;3112(Averyanovetal.):bis;3493(Averyanovetal.):bis.

WongKM668:alt.YalinSurunda58:alt.Zainudin4611:cap.

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revision of heteroblemma gen. nov. dissochaeteae ... · 230 blumea – volume 58 / 3, 2013 be, too....

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