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Blumea63,2018:150–156www.ingentaconnect.com/content/nhn/blumea https://doi.org/10.3767/blumea.2018.63.02.12RESEARCH ARTICLE

INTRODUCTION

At the end of a mission to research NepenthesL.inIndonesianNewGuineainJune2017(Cheeketal.2018)anexcursionwasmadetoGunungBotak.GunungBotak,twohourssoutheastofManokwari by road, is a popular destination for local touristic daytrips.ItcomprisesfourridgesthatdescendgraduallyintotheseafromthenorthcoastofNewGuinea.TheridgesofGunungBotak are remarkable, in a country blanketed with evergreen forest, for being treeless, presumably owing to the reputedly nickeliferous,phytotoxicsubstrate.Duringthemission,seedswerecollectedfortheIndonesianSeedBank(Cheek2017a).Amongthespeciesofwhichseedwas collected at Gunung Botak was a thicket-forming road-sideshrub.Theherbariumvoucherspecimen,Cheek 18779, isidentifiedinthispaperasanewspeciesofSeringiaJ.Gay(Byttneriaceae or Malvaceae-Byttnerioideae). Searching theNaturalis specimen website resulted in what appeared to be a secondspecimenofthesametaxoncollectedafewyearsear-lier,alsoinIndonesianNewGuinea,c.135kmfromthefirst,insavannah: Takeuchi et al. 15719 (misidentifiedasKeraudrenia corollata(Steetz)Druce.ThisidentificationwasconfirmedwhenthespecimenarrivedonloanfromLtoK.Seringia consists of one species in Madagascar and 20 in Australia(Wilkins&Whitlock2016).Thisinvestigationinitiallyappeared tobe the first evidenced, published recordof thegenusfromSEAsia.PreviousrecordsofthegenusfromNewGuinea have been unsubstantiated by specimens (Wilkins&Whitlock2016).However,areviewerpointedtoanearlierrecord,overlookedbyWilkins&Whitlock(2016)andallotherauthors, of what is almost certainly the same species, also at Gunung Botak, that of Kanehira & Hatusima 12963, 13205 (Merrill&Perry1949,discussedbelow).

Traditionally,Seringia was regarded as monotypic and close to the genus KeraudreniaJ.Gay.Wilkins&Whitlock(2016)unitedthe latter with the former on morphological grounds after a care-fulanddetailedmonographicrevision.Bothgenerahadbeenincluded within tribe Lasiopetaleae of Sterculiaceae in Malvales.Phylogenetic analysis of the four families of coreMalvales (Malvaceae, Bombacaceae, Tiliaceae and Sterculiaceae)byBaumetal.(1998)andbyBayeretal.(1999)showedthatwhilethe monophyly of traditional Malvaceae and Bombacaceae (excludingDurionaceae)wasunchallenged,Grewioideae of former Tiliaceae are more closely related to Byttnerieae of former Sterculiaceae than to the rest of the former Tiliaceae.Thesolutionistorecogniseeitherninesubfamiliesinasuper-MalvaceaeasproposedbyBayeretal.(1999)andmaintainedinBayer&Kubitzki(2003),ortorecognisenine,notfourfamiliesin the core Malvales(Baumetal.1998,Cheek2007b,2017b,Cheek&Dorr2007,Cheeketal.inpress).Underthesecondoption, preferred here, Seringia is placed in the Byttneriaceae, sister to Sparrmanniaceae(Cheek2007c).Byttneriaceae is a pantropicalfamilyofc.650speciesinc.24generabestknownfor the genus TheobromaL.fromwhichchocolateisproduced(Cheek2007a).TheByttneriaceae have been divided into three tribes since the19thcentury:Hermannieae, Byttnerieae and Lasiopetaleae (Cheek2007a).ThefirsttwoarebothcentredintropicalAfricaandAmerica,thelastinAustralia.Wilkins&Whitlock(2016)give a key to the nine genera of Lasiopetaleae.Allarerestrictedto Australia, apart from Maxwellia Baill.(NewCaledonia)andboth CommersoniaJ.R.Forst.&G.Forst.,andSeringia which have outliers inMadagascar and (Commersonia) SEAsiaandthePacific.Cheek 18779 clearly falls in Seringia since it lacksadistinctepicalyx,hasstylesfree,inconspicuoushighlyreducedpetals,staminalfilamentsprominentandyellowwithshort simplehairs on theabaxial surface, andantherswithverticalslitdehiscence.WiththekeytospeciesofSeringiainWilkins&Whitlock(2016),Cheek 18779 keys out to S. corollata Steetzsincetheinnersurface of the calyx lacks glandular hairs and instead hasdense stellate hairs which are appressed and fan-shaped, and

Seringia (Byttneriaceae / Malvaceae-Byttnerioideae) new to Southeast Asia and S. botak endangered in Indonesian New Guinea grassland and savannahM.Cheek1,J.Wanma2,M.Jitmau3,M.Jebb4

1 Science,RoyalBotanicGardens,Kew,Richmond,Surrey,TW93AE,UnitedKingdom:correspondingauthore-mail:m.cheek@kew.org.

2 FacultyofForestry,UniversityofPapua,Manokwari,PapuaBarat,Indo-nesia.

3 HerbariumManokwariense,UniversitasPapua,Manokwari98314,WestPapua,Indonesia.

4 NationalBotanicGardens,Glasnevin,Dublin8,RepublicofIreland.

Key words

BomberaiGunung BotaknickeliferousseedbankTangguhLNGProject

Abstract Seringia botakCheeksp.nov.,basedonacollectionmadebytheauthorsin2017,isdescribedfrommetalliferousgrasslandandsavannahhabitatsinIndonesianNewGuinea.Initiallythisappearedtobethefirstverifiedrecord of SeringiafromSEAsiaandtoextendtherangeofthegenusc.1500kmfurthernorththanthepreviouslyrecordednorthernlimitinNorthernTerritory,Australia.Furtherresearch,however,showedtwopreviousspecimensofthetaxonfromNewGuinea,whichhadbeenoverlooked.TheextinctionriskofSeringia botak is assessed as EndangeredusingtheIUCN2016standard.

Published on 1 October 2018

151M.Cheeketal.:Seringia in Southeast Asia

because glandular hairs are concealed below the stellates on theabaxialsurfaceofthecalyx,onthestemsandontheupperleafsurface.Indeed,Takeuchi et al. 15719 had been previously identifiedasKeraudrenia corollata (Steetz)Druce(≡S. corol-lata).BecauseCheek 18779 and Takeuchi et al. 15719 do have conspicuouslongglandularhairsalongthemarginsofthecalyxand leaf-blade, they do have resemblances to S. adenolasia. Yet, Cheek 18779 and Takeuchi et al. 15719 do not match ref-erence material of S. corollata or S. adenolasia, differing from thatspecies(andallothersofthegenus)inhavinguniformly1-floweredinflorescences,not2–5ormore-flowered.ThenewtaxonalsodiffersfromS. corollata in having a dense band of long-stalkedglandularhairsonthemarginofbothcalyxandleaf-blade, and in the densely orange-brown tomentose indu-mentumoftheyoungstems,clothedinmassive(2mmdiam)robuststellatehairsexceedinginsizethoserecordedbyWilkins&Whitlock(2016)inotherspeciesofthegenus.Accordingly,itis here described as new to science as S. botak.New plant species are still steadily being described from IndonesianNewGuinea.Recent examples areSchuiteman&DeVogel(2011),Schuiteman(2013,2015),Juswaraetal.(2016),Hambalietal.(2017),Schuiteman&Wanma(2017),Cheeketal.(2018).Thenumberof floweringplant species known to science isdisputed, [with published figures varyingwidely dependingonthemethodofestimation(NicLughadhaetal.2017),]butareasonableestimateis369000(NicLughadhaetal.2016),while the number of species described as new to science each year has been fairly constant in the 21st century, regularly exceeding2000(RBGKew,2016).Approximately5%ofknownplantspecieshavebeenassessedfortheirglobalextinctionriskfollowingtheIUCN(2012)‘goldstandard’ categories and criteria,with the results beingpu-blishedontheIUCNRedList.Thisnumberrisesto21–26%whenevidence-basedassessmentsbeyondthoseonIUCNsgloballistareconsidered,and30–44%oftheseassessmentsconsider the speciesassessed to be threatened (Bachmanetal.2018).Newlydiscoveredspeciessuchasthatreportedin this paper, are increasingly likely to be threatened, since widespreadspeciestendtobediscoveredsooner.[Althoughtherearenotableexceptionstothisrule(e.g.,DeOliveiraetal.2013).]Generallyspeaking,it isthemorelocalised,rarerspeciesthatremainundiscovered.Thismakesitallthemoreurgent to discover, document and protect such species before theybecomeextinct.

MATERIALS AND METHODS

Nomenclatural changes were made according to the Code (McNeilletal.2012).NamesofspeciesandauthorsfollowIPNI(continuouslyupdated).HerbariummaterialwasexaminedwithaLeicaWildM8dissectingbinocularmicroscopefittedwithaneyepiecegraticulemeasuringinunitsof0.025mmatmaximummagnification.ThedrawingwasmadewiththesameequipmentwithaLeica308700 camera lucidaattachment.Specimenswere inspected from the following herbaria: BM, BO, K, L, MANandP.ThoseforLwerefirstaccessedviatheNaturalisbioportalwebsite:http://bioportal.naturalis.nl.TheformatofthedescriptionlargelyfollowsthoseinWilkins&Whitlock(2016)modifiedtoBlumeaformat.Specimenscitedhavebeenseenunlessindicated‘notseen’.Theconservationassessmentfol-lowstheIUCN(2012)standard.HerbariumcodesfollowIndexHerbariorum(Thiers,continuouslyupdated).

RESULTS

Seringia botak Cheek, sp. nov.—Fig.1,2

Differs from all other known species of SeringiaJ.Gayinhavinguniformly1-floweredinflorescences(not2–many-flowered).Differingfromthosespe-cieswithappressed,fan-shapedstellatehairsontheadaxialsurfaceofthecalyx(S. adenolasiaF.Muell.,S. corollata SteetzandS. hookeriana (Walp.)F.Muell.)inthedenselong-stalkedglandularhairsatmarginsofcalyxandleaf-blade,andinthelarge(2mmdiam)brown,densestellatehairsonthestems. — Type:Cheek 18779(holoMAN;isoBO,K,L),NewGuinea,Indo-nesia,WestPapuaProvince,SEofManokwari,ManokwariRegency,RansikiDistrict,GunungBotak,S1°38'44.69"E134°05'27.54",fl.,fr.25June2017.

Etymology. Named as a noun in apposition for the type locality, Gunung Botak(meaningBaldMountain)wheretheauthorsdiscoveredthisspeciesandwhichisapossiblefutureImportantPlantArea(Darbyshireetal.2017).

Thicket-forming shrub, probably spreading by rhizomatousgrowth,1.5–2mtall.Stems stiff, branching only sparingly, and onlyat1mormoreabovetheground.Stemserect,terete1–1.5cmwideatbase,withside-branchesascendingatc.45°fromthevertical,arisingatintervalsof10–30cmfromtheprincipalaxis,internodes1.2–1.8(–2)cmlong,0.15–0.2mmwide;stemapicesheldhorizontallyornodding,indumentumdensebrightsessilestellateorange-browntomentose,hairs10–40-armed,2 mm long, arms stiff, erect, copper-coloured, concealing sparse stalkedtranslucentcapitateglandularhairs0.25–0.3mmlong,mixedwithwhiteerectarmedstellatehairs0.3–0.5mmlong.Stipulesearlycaducous,verynarrowlytriangularatbase,apexfiliform,4–5by0.5mmwideatbase,0.25mmwideatapexmembranous, pink-brown, indumentum as stem, but shorter andsparser.Leaves alternate, distinctly distichous, in two ranks oneachsideofthehorizontalorascendingstems,declinate,held30–80°fromthehorizontal.Petioleteretec.4mmlong,indumentumasstem.Blade discolorous, the upper surface dark green, rugose, lower surface bright white tomentose, narrowly oblong-lanceolate,5–9by0.9–1.8cm,baserounded,abruptlyandslightlycordate,sinus1by1mm;marginflat,entire;apexacute,withoutapiculus.Adaxial surface with primary to tertiary nervesdeeplyimpressed,20–30%coveredinwhitestellatehairsof2sizeclasses:1)sparse8-armedhairs0.5mmdiam;2)minute3–5-armederecthairs0.1mmdiam,glandularhairsabsent;marginsdenselyglandularhairy,hairsmulticellular,0.25mmlong,translucent,headblack.Abaxial surface with second-arynerves8or9oneachsideofthemidrib,barelydetectable,nerves with sparsely scattered large stellate hairs, white with tancentres,12–14stiffarms,spreadingc.1.5mmdiam,mixedwith black-headed glandular hairs, the rest of the surface be-tween the nerves completely covered in bright white arachnoid stellatehairs0.25–0.3mmdiam,10–15-armed,thearmsfine,sinuous,entanglingwitheachother;glandularhairsabsentorextremelyinconspicuous.Inflorescence 1-flowered, pendulous, heldbeneaththehorizontalyoungstems,leaf-opposed,oftentwo inflorescences per stem at successive nodes, opening in succession;22–25mmlong,indumentumamixtureoffourhairtypes:1)largestellatehairs,c.10-armed,armsrobusterect1.5mmlong,moderatelydense;2)glandularhairsaboutaslongas previous, heads pale brown or black, stem translucent, with 6–10redtransversewalls;3)small,stellate,whitehairs,armsfine,sinuous,0.25mmtall;4)shortglandularhairs0.1–0.25mmlong,headswhite.Bracts(2or)3,caducous,pink-brownor red, inserted close to each other, in an uneven whorl, nar-rowlylanceolate-oblong,membranous6–7(–8)by1–1.5(–2)mm,apexacute,baserounded,indumentumasinflorescenceaxis;bractsinserted:1)6–8mmfrombase;2)c.15–17mmfrombase;3)17.5–19mmfrombase.Calyx spreading, purple, 7–8mmlong;tube1/8–1/9oftotalcalyxlength;lobesovate,longerthanwidec.7.5by3.5–4mm,marginentire,apexacute.Abaxial surface with moderately dense, clavate, white-headed

152 Blumea–Volume63/2,2018

Fig.1 Seringia botakCheek.a.Habit,floweringandfruitingstem;b.leafmargindetail,fromabaxialsurface;c.trichomes,adaxialleafsurface;d.largestel-latehair,ingroundofdensesmallstellatehairs,leafabaxialsurface;e.floweratanthesis,sideview(fromphoto);f.flower,fromabove(driedspecimen);g.androeciumandgynoecium,petalinforeground;h.detailofstaminalfilament;i.detailofstylesandstigmas;j.indumentumofsepalmargin;k.trichometypes,sepaladaxialsurface;l.fruit(hydrated)inclined,frombelow;m.fruit,dried,fromabove(fieldphoto);n.trichomes,fruitingcalyxabaxialsurface;o.setafromfruitwing.—Scalebars:a=5cm;b,d,g,i– j=1mm;c,h,k,n–o=0.5mm;e–f=5mm;l–m=1cm.—DrawnbyAndrewBrownfromphotosandspecimens of Cheek 18779(K).

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153M.Cheeketal.:Seringia in Southeast Asia

glandularstalkedhairs,0.1–0.25mmlong,over-toppedandconcealed by 80% cover ofwhite stellate hairswith c. 12fineerect arms.Adaxial surface with midrib raised in distal quarter,surfaceentirelylackingglandularhairs(orifpresenthighlyinconspicuous),midlineoflobeswithappressedsilveryfan-likestellatehair0.5–0.75by0.1mm,6-or7-armed,thearms all robust, parallel and touching, directed towards lobe apex,covering60%of thesurface;marginal thirdsofcalyxlobeswith stellate hairswith fine, sinuous, entangled armsc.0.1mm long,coveringc.50%of thesurface,calyx tubeproximaltoandroeciumglabrousinaring0.75–1mmwide;marginofacalyxentire,denselyglandularhairy,hairs0.3–0.5mmlong,translucent,multicellular,headsblack,elliptic,c.0.1mmlong.Petals5,patent,canaliculatetrullate,0.75–1by0.3mm,theapextriangular,yellow,thetwolateraltrianglesdarkred,slightlyinflexed.Androecium and gynoeciumerect(inthependulousflower).Staminaltubeabsentor0.2mmlong.Sta-minodesabsent,rarelypresent,thenligulate,1.25by0.2mm.Filamentsyellow, redatapex, ligulate,1.75–2by0.75mm,outersurfaceconvex,minutelydenselypuberulentwithsim-ple hairs 0.05mm long.Anthers yellow, glabrous, 2-celled, oblong,1–1.1by0.5mmtouching(cohering?)laterally,pollenyellow.Gynoeciumovary5freecarpels,outersurfacedenselycoveredintranslucent,erecttostellatehairsc.1mmlong,thearmsexsertedbeyondthestaminalfilamentsby0.75–1mm;ovules6percarpel;stylesasmanyascarpels,2–2.5mmlong,glabrous, appressed together and appearing as one when live, nottwisted,stigmastruncate,coherent.Fruit pendulous dur-ing development, the inflorescenceaxis slightly accrescent,27–29mm long.Atmaturity infructescence erect, fruit andcalyxhelderect, above thestems,dull brown,matt; sepalsbecoming concave in fruit, dull dark purple accrescent, doubling indimensions,14–15by8.5–9mm,glossywitharaisedre-ticulumofnerves,papery,enfoldingthedevelopinggreenfruit.Fruitsubglobose,11by15mm;outersurfacewithnumerous

sessilewhite,stellatehairswith10–14erectarms1mmlong,glandularhairsabsent.Carpelwingsdevelopedalonglinesofdehiscence,3–4mmwide.Carpelsfreefromeachotherapartfromthebasal1/4–1/3,carpelwall0.25–3mmthick,woody,each2-or3-seeded.Setae numerous, densest along the wing apices,1.75by0.25mm,alsoonthesidesofthewingsand,moresparsely,betweenthewings.Seeds glossy black, straight, ellipsoid,3.75by2by1.75mm,arilbrightwhite,dome-likeovermicropyle,1by1mm;chalazahorseshoe-shaped,epidermalcells longitudinal, seed-coatcartilaginous.Embryogreen,c.2mmlong,cotyledonsequal,orbicular,flat,0.9–1mmdiam,radiclecylindric,c.1.1mm long,0.25mmdiam.Seedlings,juvenileleavesandchromosomenumberunknown. Phenology—Flowering:June,andprobablysomemonthspreviouslyandsubsequently(fruitsseenatallstagesofdevel-opment,andflowerbudsseen). Distribution&Ecology—Brokenmineralsubstrateinnatur-allyopen,areas(presumednickeliferous)inanotherwiselow- landevergreenforestarea(Botak)or“lowlandsavannah…growing under Melaleuca … substrate unconsolidated qua-ternaryalluvialandlittoraldeposits…rainfall3000mmp.a.” (Bomberai:Anon2005).EndemictoIndonesianNewGuinea,nearsea-level,atcoast.

Additional specimens.IndonesIa, New Guinea, Takeuchi, Sambas, Ma-turbongs 15719(BOnotseen,CANBnotseen,Knotseen,L4152226,MANnotseen),BomberaiPeninsula,TangguhSurveyareaEofBPSaenggabasecamp,nearS02°27'26"E133°06'38",fl.13Feb.2002;Kanehira & Hatusima 12963, 13205 (FUnotseen:seenotebelow),Waren,60kmSofManokwari,Mar.1940.

Conservation — Seringia botak has so far only been re-cordedattwolocations.TheBomberaisiteispartofthelicenceareaof theTangguhLiquifiedNaturalGas (LNG) industrialcomplexownedbyBP.Herethespecieswasdiscoveredduringa two week botanical survey that was part of an Environmental

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Fig. 2 Seringia botakCheek.a.Habit,floweringandfruitingstemswithMJebbandJWanmaforscale;b.floweringstemshowingleavesin2-ranks;c.close-upofflower;d.close-upoftheerectdryfruitshowingglossyblackseeds(Cheek 18779,K).—PhotosbyM.Cheek.

154 Blumea–Volume63/2,2018

ImpactAssessment(Anon2005).Inthisstudythespecieswas(asKeraudrenia corollata)oneof18‘NoteworthyFloraSpecies’listed(Anon2005:64,tableA5.1)whichincludedfivetoeightspeciesidentifiedasnewtoscience,amongwhichtwoofthespecies later described were: Glochidion daviesii W.N.Takeuchi(Euphorbiaceae)andScaevola burnettiiW.N.Takeuchi(Good-eniaceae) (Takeuchi 2003).The last specieswas indicatedas occurring in savannah, as was the Seringia. 123haofthishabitatweredue tobe clearedaspart of a total of 642halosttotheindustrialinstallation(Anon2005).However,Anon(2005:31)concludesthat“Allhabitatsandvegetation typesare widespread in the region, and no species of flora or fauna areconfinedtotheprojectproperty.Accordingly, theProjectwill not have significant impacts on the biodiversity of theregion”.Concerningthesavannah,areasarestatedtooccurtothewest,outsidetheprojectarea(confirmedbyviewingonGoogleEarth7March2018),whilethepredominantvegeta-tionislowlandrainforest.Whilethesavannahinthisareawasdescribed as species-poor, and not flagged up as a conserva-tionpriority,Takeuchi(2006)laterstatedthatthepresenceofendemic species such as the Scaevola suggested that the Melaleuca savannah in this area was not anthropogenic but natural.Seringia botak appears to be an additional endemic elementof thissavannahformation, theextent,compositionandconservationvalueofwhichareunknown.Clearly, at the Bomberai site the species may already have beenlostsubsequenttotheImpactstudyduetotheclearanceof savannah for the construction of the industrial area or for the accommodationareanearby.ThelabeldatasuggeststhatonlyasingleplantorpatchmayhavebeenseenbyTakeuchi.Itisnotknownwhetherthesubstrateatthissiteisnickeliferousornot.At the type location, Gunung Botak, Seringia botak was ob-served by the authors at a single site, where it occupied an area of6–8m2.Itverylikelyoccupiesasecondsiteclosebyalso(Merrill&Perry1949,seebelow).Owingtothedistinctivedensegrowth and showy purple flowers this is a conspicuous species thatisnoteasytooverlook.Yet,IndonesianNewGuinearemainsincompletely surveyed for plants and this species may well have othersitesinnaturallyopensavannahorgrasslandareas.However,usingtheprecautionaryprincipleandtheevidenceavailable,asrecommendedbyIUCN(2012),wehereassessSeringia botakasEndangeredB1+2ab(iii)sincetwolocationsareknown,withatotalofthreesitesbeingprobable.Therefore,the area of occupancy is 12 km2 (using thepreferred IUCNcell-sizeof4km2),whiletheextentofoccurrenceisbelowthecriterionB threshold.Threats areevident at both locations:theTangguh-Bomberai location is in the licenceareaof anexpandingindustrialcomplex(seeabove)whiletheGunungBotak location is threatened by road-widening work that was in progressduringourvisit.Theplantsseentherewereallcloseto the road,andat the footofasteepcliff.PartofGunungBotakisalsothreatenedbyquarryingformaterialsforcementmanufacture(Makuba2015).Thereisalsoariskthattheareamight be open-cast mined since the rock is considered to con-tainnickel(Utteridgepers.comm.toCheek2017).Currently,Gunung Botak is unprotected, but it may be worth recognising itasanImportantPlantAreaifitfulfilstherevisedcriteriaofDarbyshireetal.(2017).Ex situ conservation measures are in place for S. botak since several hundred seeds are now stored intheIndonesianSeedBank.

Ecology TheecologyoftheNewGuineanSeringia botak is similar to that of the Australian species in that all favour open habitats, avoiding forest.However,theNewGuineanspeciesgrowsinhigherrain-fallareas(3000mmp.a.)thanthoseinAustralia(wheremost

speciesgrowinareaswithrainfallof100–1200mmp.a.).ThenickeliferoussubstrateatGunungBotak(detailedsubstrateisunknownfortheTangguh-Bomberaisite)doesnotmatchthoseof Australian species which predominantly grow on sandstone orferraliticsubstrates(Wilkins&Whitlock2016).However,theMalagasy species, S. macrantha(Baill.)C.F.Wilkins&Whitlock,growsinultramafic(metalliferous)substrates(Wilkins&Whit-lock2016).WithinSEAsia,metalliferous-ultramaficsubstratesoftenhostnarrowlyendemicspeciesuniquetothoseoutcrops,e.g.,inthegenusNepenthes (Cheek2015,Cheek&Jebb2000,2001,2013a,b,c,Jebb&Cheek1997).Similarly,specificityto nickeliferous substrate may restrict the range of S. botak.

AffinitiesSeringia botakhasaffinitieswiththemajorityofeasternandnorthern(ratherthanwestern)AustralianspeciesofSeringia havingspreadingcalycesatanthesis(notreflexed),andwith(1–)5petalspresent(notnone,orrarely,1),leafapicesacute(nottruncateorretuse)andreddepressed-capitateglandsab-sent(vspresent).AmongthoseeasternandnorthernAustralianspecies S. botak is closest morphologically to three species in particular,sharing twosynapomorphies:1) thedistalpartoftheotherwiseyellowstaminalfilamentarered;2)theadaxialsurfaceofthecalyxhaspeculiarappressed,fan-shapedstellatehairs.Thesespeciesare:S. adenolasia, S. hookeriana and S. corollata.Oftheseitseemsclosesttothelast,whilehavingsimilarities with S. adenolasia.Seringia botak and S. corollata both differ from the other two species in lacking the prominent, long-stalked,glandulartrichomesthatextendbeyondthestel-late hairs on stems, upper surface of the leaves and the outer surfaceofthecalyx.However,S. botak does have long-stalked glandularhairsonthemarginsofbothcalyxandleaf-blade,andon the lower leaf-blade midrib, and so has more resemblance to S. adenolasia in respect of long-stalked glandular indumen-tum than does S. corollata. Thedistichousleafarrangementof S. botak also resembles S. adenolasia.Seringia corollata and S. botak also differ from S. adenolasia and S. hookeriana inhavingcalyxlobesandleaveswithanacuteorobtuse,notanacuminateapex.

Seringia in New Guinea Wilkins&Whitlock(2016)statethatthefirstreportofKerau-drenia (=Seringia)inNewGuineaappearstobethatofWillis(1919:605),followedbyGreen(1935:488),VanSteenis(1987:23)andHarden(1990:308).“However,nospecimensofthesegenerahavebeenlocatedininternationalherbaria”(Wilkins&Whitlock2016).Currently, the main generic checklist for New Guinea is that ofHöft(1992:34,whichincludesKeraudrenia).ThischecklistwasmainlybasedonacardindexattheLAEherbarium(LAE,PapuaNewGuinea)(Frodinpers.comm.toCheek,Oct.2017).ChecksatLAEbyTiberiusJimbo(pers.comm.toCheek.Oct.2017)showedthattheonlyspecimensatLAEofSeringia(in-cluding Keraudrenia)derivefromAustralia,notNewGuinea,indicatingthatthiswasthelikelysourceofthiserror.Thepresentpaperwasthusthoughttobethefirstverified,evidence-basedpublished record of Seringia (includingKeraudrenia) inNewGuinea, and inSEAsia.However, ananonymous reviewerof this paper brought to attention an earlier record based on Kanehira & Hatusima 12963, 13205 (FU) (Merrill & Perry1949).Collectedwithin4kmofthetypelocality,and,aswithTakeuchi 15719, identified asKeraudrenia corollata, this is almostcertainlythefirstrecordofSeringia from New Guinea thatwenowknowof.However,sincetheherbariumofKanehira&HatusimaisatKyushuUniversity,Japan(FU)(Thiersetal.continuouslyupdated),whichdoesnot loanspecimensand

155M.Cheeketal.:Seringia in Southeast Asia

doesnothaveadigitalherbarium,itisnotpossibletoconfirmabsolutely the identificationwithoutvisitingJapan.EnquirieshavebeenmadetoherbariaintheUSAwhereitisconceivablethat a duplicate may have been deposited, but so far, without asuccessfulresult.

A proto-epicalyx in Seringia?Anepicalyx is a calyx-like structure of bracts (‘epicalycularbracts’) that subtends the true calyx of a flower.Within therelated Malvaceaes.str.itisknowninmanygenera,suchasPavoniaCav.andUrenaL.Theoriginoftheepicalyxappearsto be linked with the reduction of the number of flowers in a dichasialcymoseinflorescencefromseveraltoone,followed(inMalvaceaes.str.)byadoublingofthenumberofbracts.Seringia botakisuniqueinSeringia in having a constantly 1-flowered inflorescence which otherwise only occurs as an atypical state in S. corollata, or in S. macrantha of Madagascar, the sole non-Australianspeciesofthegenusprevioustothispaper.InSeringia botakalooseimperfectwhorlofthree±equalbracts(insertionseparatedfromeachotherbyseveralmm)occurswhich partly covers and protects the flower bud, but which, un-liketheepicalyxusualinmostgeneraofMalvaceaes.str.,isnota verticil and is caducous, falling soon, but sometimes persisting inthedevelopingfruit.Clearlythisstructuredoesnotqualifyasanepicalyxbutitmaybeconjecturedtobeaproto-epicalyx.

Dispersal and biogeographyNothing is recorded on the dispersal of Seringia (Wilkins&Whitlock2016).Weconjecturethatbirdsormoreprobablyantsmight disperse the seeds of S. botak, in view of the fleshy aril andhardsmoothseed-coat.Themature,dehisced,fruitsareheldcloseabovethehorizontalstemapices,presenting theglossyblackpartofthesmallseeds(c.1mm),butnotthearil,toviewfromabove.Theseedsareheldsecurelyinthefruitssothattheyarenotshedanddonotfalleasily.TherecanbelittledoubtthatSeringia botak shares a common ancestor with S. corollata, S. adenolasia and S. hookeriana in viewofthesynapomorphiesnotedunder‘affinities’andsincethetwofirstmentionedspeciesareotherwisethemostnorth-erlyofthegenus,bothoccurringinnorthernNorthernTerritoryofAustralia.ThegreatgeographicaldisjunctionbetweentheS. botak and its nearest neighbours is likely to be due to scarcity ofsuitableavailablehabitatbetween.Mostoftheinterveningdistance is either sea, or in New Guinea dense, high mountain or dense closed forest, none of which provide habitat for the species of Seringia.GunungBotakwithitssubstrate-inducednaturallyopenvegetationandtheTangguh-Bomberaisavan-naharea(despiterainfallofover3mp.a.),maybeamongthefew locations in New Guinea that Seringia might be capable ofcolonising.

Key to species of Seringia with fan-shaped stellate hairs on adaxial calyx and red-apexed staminal filamentsModifiedfromWilkins&Whitlock(2016)

1.Upperandlowersurfacesofcalyx,stemandbothsurfacesoftheleaves(exceptthemidrib)withoutlong,conspicuousglandularhairs(ifpresent,thenonlyonthemarginsofthecalyx and leaf-blade, or short and completely concealedbelowthestellatehairs) . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.Outersurfacesofcalyx,stemandbothsurfacesoftheleaveswithconspicuousglandularhairsextendingwellbeyondthestellate hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2.Glandularhairsabsent,orifpresentfewandallconcealedbelowthestellatehairsonthestemandleaf-blade;inflores-cence(1-or)2-or3-flowered,bractsgreen.—Australia.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. corollata

2.Glandularhairsdense,conspicuousandextendingbeyondthestellatehairsonmarginsofcalyxandmarginsof leafbladesandontheabaxialmidrib;inflorescence1-flowered,bractsred.—NewGuinea. . . . . . . . . . . . . . . . . S. botak

3.Matureleaf-bladeflat,minorveinsconspicuousbelow.Leafandcalyxlobeswithapexacute,apiculusabsent . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. adenolasia

3.Mature leaf-blade recurved,minor veins inconspicuousbelow.Leaf and calyx lobeswithapexacuminate (rarelyacute),mainlywithapiculus . . . . . . . . . . . . . S. hookeriana

AcknowledgementsDrCharlieHeatubunencouragedtheNepenthes re- searchvisittoIndonesia,whichresultedinthispaper.WethanktheIndone-sianNationalScientificauthoritiesforgrantingpermissionforourresearchinthatcountry(researchpermit165/FRP/E5/Dit.K1/V1/2017issued15June2017byRISTEKDIKTI).WealsothankDr.JoenistheKeeperofHerbariumBogoriense,LIPI,andDr.NurhaidahSinagaHead,ResearchandCommunityDevelopmentUnitUNIPA,ManokwariforsupportduringourstudyvisittoIndonesia.AboveallwethanktheexcellentFreddyPattiselanno,HeadoftheBiodiversityCentre,UNIPAourmainscientificcounterpart.HeexpeditedpermissionsandorganizedthefieldworkforourvisittoPapuaBarat. WethankTiberiusJimbooftheLAEherbariumforconfirmingthatnospecimensarerecorded of SeringiafromPapuaNewGuineaatthatherbarium;DavidFrodinandTimUtteridgeforgeneralinformationonNewGuinea.NicolienSolofListhankedforexpeditingtheloanoftheTakeuchispecimentoK.FieldworkforMCwasmadepossiblewithsupportfromtheplayersofPeople’sPostcodeLotteryandforMJbytheIrishMuseumsTrust.

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