Studies on Schismatoglottideae (Araceae) of Borneo VII: Schottarum and Bakoa, Two New Genera from Sarawak, Malaysian Borneo
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Botanical Studies (2008) 49: 393-404. TAXONOMY Studies on Schismatoglottideae (Araceae) of Borneo VII: Schottarum and Bakoa, two new genera from Sarawak, Malaysian Borneo Peter C. BOYCE1,* and Sin Yeng WONG2 1Malesiana Tropicals, Level 5, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia 2Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300 Samarahan, Sarawak, Malaysia (ReceivedNovember27,2007;AcceptedMay30,2008) ABSTRACT. SchottarumP.C.Boyce&S.Y.WongandBakoa P.C.Boyce&S.Y.Wongaredescribedas newgenerafromSarawak,eachwithonespecies:Schottarum sarikeense (Bogner&M.Hotta)P.C.Boyce& S.Y.WongbaseduponSchismatoglottis sarikeensis (Bogner&M.Hotta)Bogner&A.HayandBakoa lucens (Bogner) P.C.Boyce&S.Y.WongbaseduponPiptospatha lucens (Bogner)Bogner&A.Hay.Bothspecies wereformerlyplacedinnow-defunctHottarum(=Piptospatha).Akeytothegeneraandprinciplesubgeneric divisionsofTribeSchismatoglottideaeinBorneoispresented.Bothspeciesareillustrated. Keyword:Araceae;Borneo;Rheophytic;Schismatoglottideae. INTRODUCTION chanicsandantherfunction,morphologiesthat,byplotting ontorecentmolecularphylogeneticresults,tobepublished AstreatedinThe Genera of Araceae(Mayoetal., elsewhere(Wong,inprep.),arenowknowntobeofcon- 1997)theSchismatoglottideaecomprisedsevengenera siderable significance in the tribe, has confirmed the long- (Schismatoglottis Zoll.&Moritzi,Piptospatha N.E.Br, heldsuspicionthatneitherspeciesfitsunreservedlyinto Hottarum Bogner&Nicolson,Bucephalandra Schott, anypre-existinggenusandthatbotharebestaccommodat- Phymatarum M.Hotta,Aridarum RidleyandHeteroari- edinnew,separate,genera.Thesenovelgeneraarehereby darum M.Hotta)withgenericboundariesbasedonthe describedandthenecessarynewcombinationsmade. presenceorabsenceofmorphologiessuchasconstricted spathes,motilestaminodes,thecaewithhorn-likeor Schottarum P.C.Boyce&S.Y.Wong, gen.nov. needle-likestructures,placentationandseedmicropylar Herbarheophytica,foliorumpetiolusinvaginumsu- appendages. prainpertemliberamtriangularispersistentiproductus. Themostrecentspecies-levelrevisionofthetribe(Hay Pedunculussemierectusvelpatensveldeclinatus.Flores andYuzammi,2000;BognerandHay,2000)although unisexualesnudi.inflorescentiafeminaintotospatham recognizingnumerousnovelspeciesreducedthenumber adnate.Floresmasculifertilesadapiceacicularispost- ofgeneratofivebysynonymizingHetroaridarumwith florescentia feminiis producens. Ovula pleura, orthotropa Aridarum(BognerandHay,200)andHottarumwith adbasimloculiinserta.Spadicisquamparssuperiorpistil- Piptospatha(BognerandHay,2000;HayandYuzammi, lodiisinstructa.Spathaetubusinfructiferoruminaquilatera 2000). infundibuliformis,persistens. IndismantlingHottarumandthesubsequenttransfer- Typus:Schottarumsarikeense (Bogner&M.Hotta) raloftheconstituentspeciesintoPiptospatha (most)and P.C.Boyce&S.Y.Wong,comb.nov. Schismatoglottis (one)placementproblemsarosewithtwo Smallrheophyticherbs.Stem usuallycondensed(very species:H. sarikeense Bogner&M.HottaandH. lucens rarelyelongatedandformingadecumbenttoweakly Bogner. Due to their unique inflorescences and infructes- creepingrhizome).Leaves severaltogether;petiole, cences,especiallyspathesenescenceandfruitdispersal sheathingonlyatextremebaseandthencewithacoria- mechanics,placementinSchismatoglottis (H. sarikeense) ceouspersistentligularportion;bladeverynarrowlyellip- andPiptospatha (H. lucens) isatbestweaklysupported tic, thinly but somewhat stiffly coriaceous, primary lateral andoursubsequentobservationsofplantsinthewildand veins extremely fine abaxially (barely differentiated from incultivation,especiallynotingspathesenescenceme- secondary venation in dry material; flush with but darker thansurroundingtissueinfreshstate);secondaryvenation *Correspondingauthor:E-mail:[email protected]. faintly prominent adaxially, fine and dense; tertiary vena- 394 Botanical Studies, Vol. 49, 2008 tionobscure.Inflorescence solitarypershootbutplants emergingfromthethecae(onepertheca)onlyattheonset usuallybearingseveralinflorescencesinsequencefrom ofmaleanthesis.Suchastructureemerginginthisman- separatebutdenselyaggregatedshoots;pedunclelong, nerisuniqueinthefamily-inallotherspeciessuchthecae stifflyarching-spreading.Spathe weaklynodding;lower structures are present from well before the inflorescence spathenarrowlyovoid,slightlydown-curved,weakly opens and are not topped with flap of any sort. Although differentiatedfromthelimbbyaconstrictionbutlower yettobeconfirmedbydirectobservationwespeculate and upper spathe differing in texture: lower spathe stiffly thattheseflap-likestructuresareassociatedwithpollen coriaceous,spathelimbsomewhatsoftlycoriaceous;limb dispersalandthatapollendropletformsonthesurfaceof marcescentfromthemarginsinwardsanddownwardsand the flap. The very slender nature of these needle-like struc- then,withtheportionclosesttotheabscissionlayerstill turesrecallsthoseofPhymatarumalthoughinthatgenus fresh,thespathelimbshedding,ovate-lanceolate,spathe thestructuresarepresentwellbeforetheonsetoffemale slightly inflated and gaping during female anthesis, then anthesis;furtherthethecaeofPhymatarumarenotably spreadingduringmaleanthesis,narrowedintoabeaked verrucate(uniquelysointhetribe)whilethoseofSchotta- tipthroughout.Spadix subcylindric;dorsal(inrelationto rumaresmooth.Schottarumsharesbasalplacentationwith spathe)sideoffemalezoneadnatetothespathe;pistils PhymatarumbuttheovulesandseedsofSchottarumlack subglobose;stigmasessile,discoid;ovulesorthotropous, thecharacteristicmicropylarappendageofPhymatarum. micropyleshortlybeakedbutnotextendedintoanap- BasalplacentationalsooccursinPiptospathabutseeds pendage;placentationbasal;interpistillarstaminodes ofthatgenusalsohaveapronouncedmicropylarappend- absent from among the pistils, confined to a row along the age;PiptospathadiffersfromPhymataruminhavingtrun- spathe/spadixadnationandoccasionallyascatteredindi- catestamenslackinganeedle-likeprocess. vidualamongstthetwolowermostrowsofpistils,sterile ThespadixwithdistalpistillodesinSchottarumis intersticeconfinedtoabout2irregularwhorlsofsterile uniqueforthetribe;inallotherSchismatoglottideaeanthe stamensatthebaseofthemalezone;malezonerather terminalpartofthespadix,andanappendix,ifpresent,is narrowerthanthefemalezone;stamenscrowded,partially comprisedofstaminodes.Weascribethedistalorgansas to completely connate into groups of 2-3, truncate and flat- pistillodesbasedonobservationsoflivingplantswhere toppedatfemaleanthesiswithatransversetoobliquehya- thehighlydistinctivepinkpistilsareclearlyhomologous lineridgeendinginanobliquedisk,butatanthesiseach withthestructuresformingtheterminalpartofthespadix thecaerectingthisstructure intoaneedle-likeprojection arequitedifferenttothestipitate-clavatewhitestaminodes terminatingwithaweaklypeltateovate-triangularflap; associatedwiththepistilsandtheintersticebetweenthe occasionallyascatteredpistillodeamongthestamens;ap- male and female flower zones. pendixabsenttobullet-shaped,whenpresentcomprised onpistillodes,taperingandnarrowlyobtuse;distal-most Schottarumspathelimbsenescencemechanicsisunu- pistillodesoftenunitedtothetopintocurvedorsinuous sual,althoughnotunique,inthetribebythespathelimb groups.Fruiting spathe unequallyfunnel-form,themar- marcescentfromthemarginsinwardsanddownwards ginsobliquelydeclinedtowardstheconvolution;fruiting andthen,withtheportionclosesttotheabscissionlayer pedunclearching/declinatewithlowerspathemouthheld stillfresh,thespatheisshed.Similar(butprobablynot laterallyorslightlydownwardswiththeconvolutionven- homologous)marginalmarcescenceoccursintheshoot- tralinrespecttothepeduncle;berrygibbous-cylindricto architecturallyquitedifferentSchismatoglottis tecturata ellipsoid-oblong,,withratherfewseeds;seedellipsoid,1-2 (Schott)Engl.withonlythemarginmarcescentandlater mmlong,0.25-0.3mmdiam.,verypalebrown,minutely sheddingwhilethegreaterportionofthespatheispersist- scabrid,micropylebeakedbutlackingamicropylarap- ent,turninggreenandremainingmore-or-lessclosedat pendage. the orifice and then shedding by abscission at the insertion ofthepedunclebysplittingandrecurvingbasipetallyat Distribution.Malesia: endemictoSarawakinSarikei maturationofthefruits. andSriAmanDistricts;atbothlocalitiesitisscatteredand rare. ThespatheduringfruitingofSchottarumispecu- liarinthatwhileitisalmostcertainlyasplash-cup,the Habitat. Old secondary and fragments of primary low- marginsofthepersistentlowersmarginsdonotforma landriparianevergreenmoistforestonshales.Schottarum levelrimbutinsteadareobliquelydeclinedtowardsthe isrheophyticonverticalclay-loamriverbanks.55-80m convolutionssuchthatwespeculatethatthefruits/seeds asl. areejectedforwardsandawayfromthefrontofthecup Notes.Theuniquecombinationofmorphologiesdis- ratherthanupwardsandoutasisknowntobetheinstance playedbySchottarum issmooththecaewithahyaline inorthodoxsplash-cupdispersers,e.g.,Aridarum.This ridgethatbecomeserectintoaneedle-likeprojectionat viewisfurtherreinforcedbythefruitingpedunclebeing theonsetofmaleanthesis,anunconstrictedspathe,aspa- arching/declinatethuspresentingthelowerspatheopen- dixfrequentlywithdistalpistillodesandseedslackinga inglaterallyordownwardsratherthenthepedunclebeing micropylarappendageandcarriedonabasalplacenta. erectandthelowerspathehelderectasisthesituationin Schottarumismostremarkablefortheneedle-likestruc- Bucephalandra,AridarumandthePiptospatha elongata tures,tippedwithaweaklypeltateovate-triangularflap, Group—all orthodox splash-cup dispersers. BOYCE and WONG — Studies on Schismatoglottideae (Araceae) of Borneo 395 Etymology. The generic name honours theAus- out,duringlateanthesislimbmarcescentfromthemargins