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Blumea 64, 2019: 108–114www.ingentaconnect.com/ontent/nhn/blumea https://doi.org/10.3767/blumea.2019.64.02.02RESEARCH ARTICLE

INTRODUCTION

Duringfieldtrips inThailandin2010,a limestonehill inLopBuriprovince,centralThailand,wasvisitedbythefirstauthor,whereashrubbyfiginhabitingalimestonecrackwasfound.Morphologically, it shows several diagnostic characters in a combinationunknownsofar.Youngtwigsarebrownpubescentto tomentose tovillous.The leafbladesareovate tonearlygloboseandabaxiallycoveredwithbrownfloccosehairs.Thefigsarepedunculate,solitaryorinpairsintheleafaxils(orjustbelowtheleaves).Specimensofthisplantwerecollectedandplacedinherbariumcollectionsbutremainedunidentified.In2011,Bergetal.(2011)identifiedabrownfloccosespecimen,Pooma et al. 3820, as Ficus talbotii King, but added a note: “moreattentionisneeded”.Later,in2013,thefirstauthorvisitedtheNationalHistoryMuseum(BM)inLondontostudyFicus L.specimensandhefoundasamplecollectedbyA.Marcanin1924fromalimestonehillintheprovinceRatchaburi(A. Marcan 143).ThespecimenisverysimilartothefigfoundinLopBuriand to Pooma et al. 3820.

MATERIAL AND METHODS

MacromorphologyIn2015thefirstauthorbegantostudyFicus subsection Cono-sycea (Miq.)Corner.ThemorphologyofPooma et al. 3820 andthesamplescollectedinLopBuriwerecarefullystudiedtogether with and in comparison to specimens of F. talbotii using classicaltaxonomicaltechniques.

Leaf anatomyTheleafanatomicalproceduresfollowedareconsistentwiththeonesused inChantarasuwanetal. (2014).Sixsampleswere selected, three from the new species, F. pongumphaii and three from F. talbotii.Dried leaves were rehydrated and cross sections were then madeusingaReichert slidemicrotome.Sixteen transversesectionsincludingmargin,midribandpetiolewerecollected.Free hand paradermal sections were taken from the adaxial andabaxialleafsurfaces.Halfofthesectionsandparadermalsectionswerebleachedandstainedwithsafranin/haematoxylin.AllsectionsweredehydratedandmountedinEuparal.Cuticularmacerations were made by placing a leaf sample in a 1 : 1 mix-tureofhydrogenperoxide(30%)andaceticacid(99–100%)at60°Covernight.Thecuticlewascleanedthefollowingdayandthesectionswerethenplacedinamixtureof0.5%SudanIVin70%alcoholat40°Cfor2–3handmountedinglycerinejelly.Theslideswereobservedfor11microscopiccharacterssimilarto the ones used as diagnostic features by Chantarasuwan et al.(2014):Indumentum,cuticle,epidermalcells,mesophyll,thestomatalcomplex,lithocysts,silicifiedcells,crystals,petioleandmidrib vascularization, veins and bundle sheaths, and scleren-chymafibresandsclerifiedgroundtissue.Descriptionsofbothspecies were made and they were compared for consistent differencesincharacterstates.

RESULTS AND DISCUSSION

ThenewspeciesdiffersinseveralcharacterstatesfromF. tal-botii (seeTable1 formorphologicaldifferencesandTable2for leafanatomicaldifferences).Thenewspeciesisashrubthatnormallygrowsupto3mtall,whileF. talbotii is a tree of upto22mhigh(Table1).Theperidermoftheleafytwigsispersistent in the new species but flaking off in F. talbotii.Thefigpedunclesare2–3mmlonginthenewspeciesandabsentto up to 2 mm long in F. talbotii.Thenewspecieshasobovatesyconia with outside a normally brown floccose indumentum and inside internal hairs are present, F. talbotii has subglobose (toobovate),normallyglabrousorrarelypuberuloussyconia,

Ficus pongumphaii (Moraceae), a new species from Thailand, compared with the ambiguous species F. talbotii B.Chantarasuwan1,S.Sungkaew2,3,K.Pruesapan4,P.Baas5,P.C.vanWelzen5,6

1 ThailandNaturalHistoryMuseum,NationalScienceMuseum,Klong5,KlongLuang,PathumThani,Thailand.

2 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok,Thailand;correspondingauthore-mail:fforsws@ku.ac.th.

3 Center forAdvancedStudies inTropicalNaturalResources,KasetsartUniversity,Bangkok,Thailand.

4 PlantVarietiesProtectionDivision,DepartmentofAgriculture,Bangkok,Thailand.

5 NaturalisBiodiversityCenter,SectionBiodiversityDiscovery,P.O.Box9517,2300RALeiden,TheNetherlands.

6 InstituteofBiologyLeiden,LeidenUniversity,P.O.Box9505,2300RALeiden,TheNetherlands.

Key words

Ficusleaf anatomyMoraceaenew species

Abstract A deciduous shrub previously included in Ficus talbotii for many years, is now regarded as a new spe-cies, Ficus pongumphaii.ItismorphologicallydistinctfromF. talbotii with as typical characters the densely brown pubescenttotomentoseorvillousonleafytwig;theelliptic,suborbiculartoobovateleafbladesthatarebrowntomentellousontheuppersurfaceandbrownfloccosetomentosetovillousunderneath;thepedunculatefigsareobovate,brownfloccoseorvillousoutsideandhaveinternalhairs.Theleafanatomyshowsamultipleepidermisonbothsurfaces;enlargedlithocystsonbothsidesofthelamina,whicharemoreabundantadaxiallyandwithveryfewabaxially.Thespecies,endemictoThailand,isnamedafterthegreatThaidendrologist,AssociateProfessorSomnuekPongumphai.

Published on 10 May 2019

109B.Chantarasuwanetal.:Ficus pongumphaiifromThailand

Fig. 1 Ficus pongumphaiiChantaras.&Sungkaew.a.Twigwithleavesandfigs;b.figs;c.figinlongitudinalsection;d.staminateflower;e.stamen;f.sessilepistillateflower;g.pedicelledpistillateflower(B. Chantarasuwan 180910-4,THNHM).—Drawing:PajareeInthachup,2015.

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whichlackinternalhairs.ThenewspeciesresemblesF. calci-cola Corner, treated as a form within the F. talbotii complex by Bergetal.(2011),butdiffersinfoliarindumentum,figspeduncle,andindumentumonreceptacle(syconia).Theleafanatomyyieldssupportingevidenceforthedistinctionbetween F. pongumphaii and F. talbotii, with the presence/absenceofglandularhairswithdiscoid,4-cellularheads(Fig.3c2) and the relative frequency of lithocysts on upper andlowerleafsurfaces(Fig.4)asthemostimportantdistinguishingfeatures.However,thesedifferences,andtheotherdifferenti-atingcharacters(Table2)havetobetestedinmorematerial,becausetheycanbedependentonthe(unknown)ageofthepersistentleaves.Morphologically and anatomically the new species, F. pongum-phaii, is clearly distinct from F. talbotii with which it was con-fused.Table1and2 show that thenewspecies iseasy toidentifybasedalreadyonhabitandindumentum.Theleafanatomy(amultipleepidermisandthepresenceoflithocystsonbothsidesofthelamina)supportsthepositionofboth species in subsection Conocyea(seeChantarasuwanetal.2014,forsubsectionaldivisions).

TAXONOMY

Ficus pongumphaiiChantaras.&Sungkaew,sp. nov.—Fig.1,2,3a2,b2,c2,d2,4a2,b2

Shrub,atleastupto3mhigh.Leafytwigbrownpubescenttotomentosetovillous.Leaflaminaelliptictosuborbiculartoobovate,uppersurfacewhitishorbrowntomentellous,lowersurfacebrownfloccosetomentosetovillous.Figpedunculate,basalbractspersistent,strigose.Receptacleobovate,brownfloccoseorvillous.Synconialinternalhairspresent.Epidermismulti-layers.Enlarged lithocysts abundant adaxially and only few abaxially.—Type:B. Chantarasuwan 180910-4 (holoTHNHM; iso L),Thailand, LopBuri,Thawung,WatKhaoSamorkhorn,18Sept.2010.

Deciduousshrub,upto3mtall,intermittentgrowthnotpromi-nent.Leafy twig1.5–2.5mmthick,brownpubescenttotomen-tose to villous, peridermpersistent.Leaf spirally arranged, laminaelliptictosuborbiculartoobovate,2.7–6.5by1.9–4.8cm, apex apiculate, the acumen blunt, base cuneate, upper surface whitish or brown tomentellous, lower surface brown floccosetomentosetovillous,lateralveins5–7pairs,usuallybranching(furcatedawayfrommargin),basalpairsendingup

to2/5–1/2thelengthofthelamina,usuallybranching.Petiole 0.8–1 cm long, brown tomentose, drying brown, epidermispersistent.Stipule0.4–0.6cmlong,persistent,brownstrigoseortomentose,epidermisofbudscalepersistent.Figs in axils ofleavesorjustbelowtheleaves,solitaryorpairs,peduncle2–3mm long, tomentose; basal bracts 3, 1–1.5mm long,brownstrigose,persistent.Receptacleobovate,0.6–0.7cmdiam when dry, brown floccose or villous, apex convex, ostiole c.2mmdiam, upper ostiolar bracts glabrous; internal hairspresent.Staminate flowers dispersed, sessile to pedicellate, tepals2(or3),ovate,obovateorspatulate,free,red-brown.Pistillate flowers sessile to pedicellate, ovary red-brown, tepals 3,ovate,obovateorspatulate,free,red-brown. Distribution&Habitat—Aspeciesseeminglyendemic toThailand,occurringonlimestonehillsata30–600melevation.

Specimens examined.Thailand, P. Palee 278(L),ChiangMai,SanhGahmPang,2May1995; A. Marcan 1437(BM),Ratchaburi,15July1924; R. Pooma, K. Phattahirankanok, S. Sirimongkol, M. Poopath 3820(BKF,L),Lopburi,ThaWung,WatKhaoSamorkhorn,20Apr.2004; P. Phonsena, D. Chusithong, N. Loetsombunsuk 5718(BK,BKF,L),Lopburi,ThaWung,WatKhaoSamork-horn,21Dec.2007;B. Chantarasuwan 180910-4(L,THNHM),Lopburi,ThaWung,WatKhaoSamorkhorn,18Aug.2010;B. Chantarasuwan 081212-2 (L),Lopburi,ThaWung,WatKhaoSamorkhorn,8Dec.2012;Put 2394(BK),Saraburi,HinLap,18Aug.1929;Nai Noe 130 (BK),MuakLek,KaoMakKok,17July1925;C.F. van Beusekom & T. Smitinand 2036(BKF,L),ChonBuri,Sriracha,SriChangisland,7Nov.1969.

Ficus talbotii King —Fig.3a1,b1,c1,d1,4a1,b1

Ficus talbotiiKing(1887)51,t.63;Talbot(1911)511,t.521;Corner(1965)19;(1977)139,t.14;C.C.Berg(2007)24;C.C.Bergetal.(2011)647.—Type:W.A. Talbot 1100(lectoCAL),India,N.KanaraDistrict.

Ficus pierreiGagnep.(1927)93;(1928)763.—Type:Pierre n° 1676(holoP),Cambodia,SonrongTong,montibusKéréev,Apr.1870.

Ficus calcicolaCorner(1960)392;Kochummen(1978)142;C.C.Berg&Corner(2005)636.—Type:M. Nur SFN 34388(holoSING;isoL),MalayPeninsula,Selangor,Kaching,BatuTakun.

Tree,upto22mtall,deciduous,intermittentgrowthnotpromi-nent.Leafy twig1.5–2mmthick,glabrousorwhitishpuberulousorbrownsubtomentose,peridermflakingoff.Leaves spirally arrangedtosubdistichous,laminaovatetoelliptic,4.5–9by1.5–3.5cm,apexacuminate,theacumenblunt,baseroundedto cuneate, upper surface glabrous or minutely puberulous on midrib and main veins, lower surface glabrous to puberulous orsubtomentoseonmidribandmainveins,lateralveins6–7pairs, furcated away from margin, basal pairs ending up to 1/5–1/3 the lengthof the lamina,usuallybranching.Petiole 1–1.6cmlong,glabrousorminutelyandsparselypuberulous,dryingblackish,epidermispersistent.Stipule0.4–0.5cmlong, caducous,puberulous,brownstrigoseorsubtomentose.Figs inaxillary,solitaryorinpairs,sessile(orpeduncleupto2mm

Characters F. pongumphaii F. talbotii

Habit Shrub,normallyup Tree,upto22mtall to3mtall

Indumentumofleafy Brownpubescentto Glabrousorwhitishtwig tomentose to villous puberulous

Peridermofleafytwig Persistent Flakingoff

Petiolelength 0.8–1cm 1–1.6cm

Endingofbasallateral Basalpairsupto Basalpairsupto1/5–1/3veins 2/5–1/2thelengthof thelengthofthelamina the lamina

Foliar indumentum Upper surface brown Upper surface glabrous tomentellous Lowersurfacebrown Lowersurfaceglabrousor tomentose minutely puberulous on midrib and main veins

Figpeduncle Peduncle2–3mmlong Absent(–2mmlong)

Receptacle(Syconia) Obovate,brownfloccose Subglobose(toobovate), orvillous glabrous(orpuberulous)

Synconialinternalhairs Present Absent

Tepalsofstaminate 2(or3) 3flowers

Table 1 Comparison of some morphological differences between F. pong-umphaiiChantaras.&SungkaewandF. talbotiiKing.

Characters F. pongumphaii F. talbotii

Glandularhairs Hairswithellipsoid- Onlyhairswithellipsoid- capitate 1- or 2-celled capitate 1- or 2-celled heads and discoid-capitate heads hairs with 4-celled heads (Fig.3c2))

Cristarquecells Absent Present

Silicifiedcellgroups Presentinepidermisand Presentinmesophyllonly mesophyll

Enlarged lithocysts More abundant adaxially Abundant on both sides

Sclerificationofsub- Absent Abaxiallyofmidribepidermal ground tissue

Table 2 Comparison of leaf anatomical characters between F. pongumphaii Chantaras.&SungkaewandF. talbotiiKing.

111B.Chantarasuwanetal.:Ficus pongumphaiifromThailand

Fig. 2PhotosofliveFicus pongumphaiiChantaras.&Sungkaew.a.Habitinnaturalhabitat;b–d.twigswithleavesandfigs;e.fig;f.figinlongitudinalsec-tion.—PhotosbyBhanumasChantarasuwan.

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Fig. 3 Cross sections and cuticular macerations of the leaf laminas of Ficus talbotiiKing(leftcolumn)andFicus pongumphaiiChantaras.&Sungkaew(rightcolumn).a1.LeafmarginwithoutindumentumofF. talbotii;a2.leafmarginwithindumentumofF. pongumphaii;b1.laminaofF. talbotii showing multi-layered epidermisonbothsidesandenlargedlithocystsonbothsurfaces;b2.laminaofF. pongumphaii showing multi-layered epidermis on both sides and enlarged lithocystsadaxially;c1.abaxialcuticularmacerationofF. talbotiishowingfewhairs(blackarrows);c2.abaxialcuticularmacerationofF. pongumphaii showing manyhairs;d1.petioleofF. talbotiiwithfewhairs;d2.petioleofF. pongumphaiiwithanabundanthairs(a1,b1,d1:M. Nur SFN 34388,L;a2,b2,c2:R. Pooma et al. 3820,L;c1:M.F. Newman et al. 1148,L;d2:B. Chantarasuwan 180910-4,THNHM).—PhotosbyBhanumasChantarasuwan.

113B.Chantarasuwanetal.:Ficus pongumphaiifromThailand

long);basalbracts3,2–3mmlong,brownpuberulous,per-sistent.Receptaclesubglobose(toobovate),0.6–0.8cmdiamwhen dry, glabrous or minutely puberulous, apex convex, ostiole c.2mmdiam,upperostiolarbractsglabrous; internalhairsabsentorfew.Staminate flowers dispersed, sessile to pedicel-late,tepals3,ovate,free,red-brown.Pistillate flowers sessile topedicellate,ovarywhitewithreddot, tepals3(–4),ovate,obovateorspatulate,free,red-brown. Distribution&Habitat—Distributed inSri Lanka, India,Myanmar,China(Yunnan),Laos,Vietnam,Cambodia,ThailandandMalayPeninsula.Foundinmixeddeciduous,evergreenand dry evergreen forests, and on limestone hills, up to 1 100 m elevation.

Specimens examined.Cambodia, L. Pierre 1676 (P),SonrongTong,mon- tibusKéréev,Apr.1870.–india, K.M. Matthew RHT25340(L),Tamilnadu,Na-makkal,Kollihill,Salem,21Dec.1979.–malay Peninsula, M. Nur SFN34388 (L,SING),Selangor,Kaching,BatuTakun,3Nov.1937;T.C. Whitmore FRI 12162 (L),Selangor,AnakTakun,31July1968;T.C. Whitmore FRI 15633 (L),Selangor,BatuCaves,20Sept.1970.–myanmar, N. Tanaka, T. Suga-wara, S. Sakai, K. Aoki, A. Tanaka, H. Miwa, Than Than Aye, Khin Myo Htwe 021824 (L),Mandalay,Peik-Chin-Myaung,E96°37'12"N22°05'32",13Jan.2002.–Thailand, Put 4024(BK,L),Lampang,MaungNgao,17July1929;A.F.G. Kerr 19843(BK);PrachinBuri,KabinBuri,10Nov.1930; M.F. New-man, T. Boonthavikoon, C. Hemrat, D.J. Middleton 1148 (L),Chumphon,KawWeing,11Jan.1927;A.F.G. Kerr 11376(BK),PrachuapKiriKhan,Pranburi,SamRoiYot,30June2000.

LEAF ANATOMY

Ficus pongumphaiiChantaras.&Sungkaew — Fig.3a2,b2,c2, d2, 4a2, b2

Material studied.B. Chantarasuwan 180910-4;R. Pooma, K. Phattahi-rankanok, S. Sirimongkol, M. Poopath 3820, P. Palee 278(seeaboveforlocalities).

Surfaceview—Indumentumpresentabaxiallyandadaxially,consisting of glandular ellipsoid-capitate hairs with 1- or 2-celled heads, discoid-capitate glandular hairs with 4-celled heads and simpleseptateandnon-septatehairs.Cuticlesmooth.Anticli-nalwallsstraightonbothsurfaces.Radiatingepidermalcellsaroundlithocysts5–8onbothsurfaces.Stomataactinocytictoanomocytic,20–30µmlongand17–25µmwide;giantstomata28–38µmlongand25–30µmwide. Transversesection—Cuticlelessthan2µmthickabovethelamina,abovemidrib2–3µmthickandmarginally2.5–3µmthick.Epidermismulti-layeredonbothsides,cellsinouterlayersmallerthanintheinnerlayer.Stomataslightlysunken,onlyoutercuticular ledgespresent.Enlarged lithocystsabundantadaxially, fewabaxially.Mesophylldorsiventral; silicifiedcellgroups present in mesophyll and epidermis especially near the stomataoftheabaxialepidermis.Palisade2-layered.Midribwithtwoopposingarcssurroundedbyfibrecaps.Petiolewithacylinderofseparatebundles,withoutafibrecap(orrarelywithasmallfibrecap);peripheralgroundtissuenotsclerified.Pith bundles present inmidrib and petiole.Veins vertically

Fig. 4 Free hand paradermal leaf surfaces of Ficus talbotiiKing(leftcolumn)andFicus pongumphaiiChantaras.&Sungkaew(rightcolumn).a1.Abaxialparadermal leaf surface of F. talbotiishowingradiatingepidermalcellsaroundlithocysts(blackarrows);a2.abaxialparadermalleafsurfaceofF. pongumphaii showingafewradiatingepidermalcellsaroundlithocysts(blackarrows)andabundanthairs;b1.adaxialparadermalleafsurfaceofF. talbotii showing radiating epidermalcellsaroundlithocysts(blackarrows)andnoindumentum;b2.adaxialparadermalleafsurfaceofF. pongumphaii showing radiating epidermal cells aroundlithocysts(blackarrows)(a1,b1:M.F. Newman et al. 1148,L;a2,b2:P. Palee 278, L).—PhotosbyBhanumasChantarasuwan.

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transcurrent;minor veins embedded inmesophyll.Marginalsclerenchymastrandsabsent.Drusespresent inmesophyll,ground tissue parenchyma and phloem parenchyma of midrib andpetiole,fewinthebundlesheathsaroundtheveins;pris-matic crystals absent or extremely rare in the parenchyma of midribandpetiole.

Ficus talbotiiKing—Fig.3a1,b1,c1,d1,4a1,b1

Material studied. M.F. Newman, T. Boonthavikoon, C. Hemrat, D.J. Middleton 1148;M. Nur SFN34388; T.C. Whitmore (KEP) FRI 15633(seeaboveforlocalities).

Surfaceview—Indumentumpresent,consistingofellipsoid-capitate glandular hairs with 1- or 2-celled heads and simple septateandnon-septatehairsabundantonthepetiole.Cuticlesmooth.Anticlinalwallsstraightonbothsurfaces.Radiatingepidermalcellsaroundlithocysts5–8onbothsurfaces.Sto-mataactinocytictoanomocytic,25–28µmlongand17–25µmwide;giantstomata30–38µmlongand25–30µmwide. Transversesection—Cuticle2–4µmthickabovethelamina, c.4µmabovemidribandmarginally5–8µmthick.Epidermismulti-layered on both sides, cells in outer layer smaller than in the inner layer. Stomata levelwith epidermis, inner andoutercuticularledgespresent.Enlargedlithocystspresentincomparablefrequenciesonbothsides.Mesophylldorsiventral;silicifiedcellgroupspresent,especiallyinmesophyll.Palisade2-layered.Midribwithtwoopposingarcssurroundedbyfibrecaps;subepidermalgroundtissuesclerifiedabaxially.Petiolewithacylinderofseparatebundleswithfibrecap;peripheralgroundtissuenotsclerified.Pithbundlespresentinmidribandpetiole.Veinsvertically transcurrent;minorveinsembeddedinmesophyll.Marginalsclerenchymastrandsabsent.Drusespresent in mesophyll, the bundle sheaths around the veins, ground tissue parenchyma and phloem parenchyma of midrib andpetiole;prismaticcrystals(partlyincristarquecells)presentin periphery of the bundle sheaths above and below the veins andintheparenchymaofmidribandpetiole.

AcknowledgementsThefirstauthorwishestothanktheNationalScienceandTechnologyDevelopmentAgency(NSTDA)fortheresearchgrantundertheprojectcodeSCH-NR2015-792.WearegratefultotheherbariaBK,BKF,BM,CAL,K,L,PandTHNHMforcomparingherbariumspecimensandusingspecimendatafromtheirdatabases.SpecialthanksforassistanceduringfieldworktothestaffofPhuKaeBotanicalGardensandThamPetThamThongForestPark,toDr.SomranSuddeeofBKFfortaxonomicdiscussionsandmanythankstoMr.CholawitThongcharoenchaikitandstaffoftheThailandNationalScienceMuseumfortheirhelpinvariousways.

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