2014 three new ginger species (zingiberaceae) from laos

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Three New Ginger Species

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© 2014 Naturalis Biodiversity Center

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Blumea 59, 2014: 106–112www.ingentaconnect.com/content/nhn/blumea http://dx.doi.org/10.3767/000651914X685221RESEARCH ARTICLE

INTRODUCTION

Cambodia, Laos and Vietnam comprise one of the least known hotspots of the family Zingiberaceae, the latest comprehensive studybeingoveracenturyold(Gagnepain1908).OnlyAmo-mumL.hasbeenstudiedindepthsincethen,resultinginthedescriptionofelevennewspecies(Lamxay&Newman2012).There have been, by contrast, numerous recent descriptions of newspecies(e.g.Leong-Škorničková&Lý2010,Lýetal.2010,Nguyen&Leong-Škorničková2012,Leong-Škorničková&Trần2013,Leong-Škorničkováetal.2013,2014,Leong-Škorničková&Lưu2014)andanewgenus,Newmania(Leong-Škorničkováetal.2011),fromtheIndochinesefloristicregion(sensuTakhta-jan1986),suggestingthatalargeamountofgingerbiodiversityisyettobedocumented.DuringourextensiveexplorationsoftheIndochineseregion,the following three ginger species have been found to be new, andaredescribedandillustratedbelow.Theoriginaldescrip-tions and types of all the species in their respective genera/subgenera have been consulted, especially those from adjacent territories.While two of these three species are currently known only from their type localities, we believe that detailed descriptions ac-companied by colour plates will aid local botanists and students inidentificationandbringmorereportsoftheirdistributioninthenearfuture.The descriptions were made from full-grown, flowering, living material.InlinewithgoodpracticeoncollectingZingiberaceae, the type collections include spirit material and the descriptions are accompanied by detailed photographic documentation.ConservationassessmentsfollowthecriteriaoutlinedbyIUCNversion3.1.(2012).

TAXONOMIC TREATMENT

Curcuma corniculata Škorničk.,sp. nov. —Fig.1

Similar to Curcuma glans K.Larsen&Mood,butdiffers in theredcorollalobes(vswhiteorwhitewithareddishtinge),brightorange,diamond-shapedlabellumwithhorn-liketipscurvingoutwards(vsobovatewhitelabellumwithgoldenyellowmedianbandandapex)andbrightorangelateralstaminodeswithdarkpurplepatchatthebase(vswhitelateralstaminodeswithpurplebaseandgoldenyellowapex).—Type:Jana Leong-Škorničková, Trần Hṹu Đăng, Otakar Šída, Udone Souvannakhoummane & Kittisack Phoutthavong JLS-1752(holotypeSING(incl.spirit);isotypeE,HNL,P),Laos,Louang-phrabang,BanLongLaoMai,866masl,N19°45'16.6"E102°02'56.7",20June2012.

Etymology. Thespecificepithet ‘corniculata’ isderivedfromLatinandmeanssmallhorns,inreferencetothehorn-liketipsofthelabellum.

Herbto1mtall. Rhizomeovoid,c.1.5–3.5by0.8–1.5cm,withthinbranches(c.3–6mmdiam),brownexternally,lightyellowinternally,slightlyaromatic;root tuberselliptic,3–5.5cmlong,light brown externally, pure white internally with translucent whitecentre,distanced5–15cmfromrhizome. Leafy shoot with5–9leaveswhenflowering;pseudostemto35cmlong,green, composedof leaf sheaths; leafless sheaths perhaps two,decayedatanthesis;leaf sheathsgreen,puberulent;ligule 5–11 mm long, bilobed, hyaline, greenish white, translucent, puberulent;petiole10–35cmlong(petioleoffirstleafshortest,innermostleaveslonger),canaliculate,green,shortlysparselyhairy;laminabroadlyelliptictoelliptic-ovate,to47by22cm,prominently plicate, adaxially bright green, sparsely shortly hairyalongmainveins,abaxiallylightergreen,glabrous;midribglabrous,green;basecordate;apexacute,c.1–1.5cmlong,puberulent. Inflorescence lateral,many-flowered;peduncle 4–15 cm long, to 1 cm diam, cream-white, with up to 5 sheath-ingbracts(mostlydecayed,1–8cmlong);spike7–11cmlong, c.4–6cmdiaminthemiddle,withoutcoma;fertile bracts20–30perinflorescence,bract3–6by1.2–3.3cm(largeratthebaseoftheinflorescence),ovatetotrullate,smallerandovateattheapex, whitish at base to light green at apex with various degrees ofredtingethroughoutthebract(resultinginlightredtodeepred bracts, colour varyingwithin population), both surfacesshortlypubescent,connateinthelower1/2to1/4;enclosingcincinnuswith3flowersatthebaseoftheinflorescence,1–2

Three new ginger species (Zingiberaceae) from LaosJ.Leong-Škorničková1,O.Šída 2,S.Bouamanivong3,K.Souvannakhoummane 4, K.Phathavong 4

Key words

CurcumagingersLaosnew speciesvulnerableZingiberZingiberaceae

1 TheHerbarium,SingaporeBotanicGardens,NationalParksBoard,1ClunyRoad,259569,Singapore;

correspondingauthore-mail:jana_skornickova@nparks.gov.sg.2 DepartmentofBotany,NationalMuseuminPrague,Cirkusová1740,Praha9–HorníPočernice19000,CzechRepublic.

3 BiotechnologyandEcologyInstitute,MinistryofScienceandTechnology,Vientienne,LaoP.D.R.

4 PhaTadKeBotanicalGarden,BanWatThat,P.O.Box959,06000LuangPrabang,LaoP.D.R.

Abstract Three new Zingiberaceae species from Laos, Curcuma corniculata and C. flammea(Curcumasubg.Ecomata),andZingiber jiewhoei(Zingibersect.Zingiber),aredescribedandillustratedhere.

Published on10October2014

107J.Leong-Škorničkováetal.:GingerspeciesfromLaos

Fig. 1 Curcuma corniculataŠkorničk.a.Habit;b.inflorescence;c.seeds;d.ligules;e.rhizome;f.flowerdissection,withdetailofstamenfromsideandfrontview(allfromtype:JLS-1752).—Scalebars:c,f(inset)=1cm;e,f=2cm.—PhotosbyJanaLeong-Škorničková.

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flowersatthetop;bracteoles much reduced, one per flower, ovate,boat-shaped,c.1–2.5mmbyc.1mm(outeroneslarger,inneronesgraduallysmaller),hyaline,translucentwhite,gla-brous.Flowers6.5–8cm,muchexsertedfromthebracts;calyx to21mmlong,3-toothed,unilaterallysplittoc.6–8mm,trans-lucent white or with slight red tinge at upper part, sparsely hairy orglabrous;floral tubec.4–5.5cmlong,externallywhiteatthebase, with red tinge at the apex, densely puberulent, internally white, sparsely puberulent with dorsally placed groove hold-ingthestyle;dorsal corolla lobe 22–27by7–9mm,triangularovate, concave, with sides rolled inwards, glabrous, red out-side,lightredtowhiteinside,apexmucronate,mucroc.2mm, withafewshorthairs;lateral corolla lobes21–24by7–8mm,triangular ovate with sides incurved, concave, glabrous, red outside,lightredtowhiteinside;labellum23–25by16–20mm,diamond-shaped,marginsrecurved,apexbifidwithanincisionup to 14 mm long and strongly recurved horn-like lobes, base of the labellum light orange with two orange lines in the centre and often with slight purple tinge laterally, middle and apex of the labellum light to deep orange at the margin, with deep orangebandrunningthroughthecentre(medianband);lateral staminodesc.22–24by8–10mm,slightlyrhomboidwithsidesflexingoutwards,brightorangewith3–4mmatbasetingeddarkpurple(sometimesmissing),sparse,shortglandularhairspresentonbothsurfaces. Stamen18–20mm;filament8–9mmlong,whitewithpurpletingetowardsbase,2.5–3mmbroadatbase,1.5mmbroadatapex(thepointofattachmenttotheconnective),coveredwithdense,veryshort,glandularhairs;an-therc.13–15mmlong,spurred,connectivelightyellowtoverylightorange,denselycoveredwithshortglandularhairs;anther spurs c.3mm long, triangularwithsharp tips,pointingout-wards;anther crestthin,rounded,c.1.5mmlongandc.1mm broadatbase,orange;anther thecae 9–11 mm long, dehiscing alongtheirwholelength.Epigynous glands2,c.4–6mmlong,0.3–0.5mmdiam,cream-white.Style thin, white, glabrous, held ingrooveindorsalsideoffloraltube;stigmac.1mmlong,0.8mmwide,white,ostioleciliate,facingupwards.Ovary 2–3by2–2.5mm,trilocular,denselypuberulent,hairsc.0.2mmlong.Fruitaglobosetrilocularcapsule,c.1cmdiam,white,puberu-lous;seedsirregularlyobovoid,4–7mmlong,lightbrown,shiny,enclosedintranslucentwhite,laciniatearil.Descriptionbasedon living and spirit material of JLS-1752. Distribution—EndemictoLaos,currentlyknownonlyfromthetypelocality. Habitat&Ecology—Growinginbamboovegetationonlime- stoneoutcrops,at800–900masl. Phenology—Floweringinearlyrainyseason,endMaytoJuly, flowers last a single day, open in the morning and closing inlateafternoon.FruitingJunetoJuly(August?). Conservation status — Specimen records exist so far only fromthetypelocality.Localpeoplewhowereinterviewedandshown images of this species, reported that it occurs in vari-ousareasadjacenttoLouangphrabang.Thenumberandsizeofthepopulationsare,however,notyetexactlyknown.MostCurcumaspeciesinsubg.Ecomata, especially those allied to C. bicolorMood&K.Larsen,wherethisspeciesbelongs,tendtobefairlyrestrictedintheirdistribution.Accordingly,weestimatethat the extent of occurrence is less than 20 000 km2.Thetypepopulation is not in a protected area and is used for cultivation ofmaize(Zea maysL.),groundnuts(Arachis hypogaeaL.)andother crops, therefore continuing decline in area, extent and qualityofhabitat canbe foreseen.CategoryVulnerableVUB1ab(iii) is,withcaution,proposedhere.FurtherexplorationoftheregionisneededasthisspeciesmaybeEndangeredorCriticallyEndangeredsoon.

Notes — Curcuma corniculata belongs to the recently es-tablishedsubg.Ecomata(Záveskáetal.2012).Thetwomost

closely related species in this subgenus are C. glans and C. bicolor.Itmaybefairlydifficulttodeterminedriedherbariumspecimens but there is no confusion when living, flowering material is available (seeMood& Larsen 2001) or whenthere are appropriate notes or photographs accompanying specimens.Also,ifaspiritspecimenispresent,confusionisnotlikely.Amongthesespecies,C. corniculata can be readily recognisedbyitsbifidlabellum,withthetwostronglyreflexedhorn-likelobes.

Curcuma flammea Škorničk.,sp. nov.—Fig.2

Similar to Curcuma bicolorMood&K.Larsen,butdiffers in inflorescencewithlongerpedunclesto13cmlong(vsalmostsessileinflorescencewithpedunclec.1.5cmlong),bractswhite,pinktodarkred(vslightgreenbractswithmoreorlessreddishtingeandprominentveins),brightorangelabellumwithvariousdegreeofredornamentation(vsmostlyyellow-orangelabellumwith reddish to purplemargins).—Type:Jana Leong-Škorničková, Trần Hṹu Đăng, Somsanith Bouamanivong, Khantheo Santhammavong, Udone Souvannakhoummane, Kittisack Phoutthavong JLS-1777 (holotypeSING(incl.spirit); isotypeE,HNL,P),Laos,VientianeProv.,VangViengDist.,BanPhouDinDeng,467masl,N18°57'43.2"E102°26'09.9",23June2012.

Etymology. ThespecificepithetisLatinforfieryorflaming.Itisgivenonaccount of the bright orange-red colours of the flowers and bracts, which resembleflamesontheforestfloorwheninfullbloom.

Herbto70cmtall.Rhizomeovoid,c.1–1.5by0.8–1.2cm,withthinbranchesc.3–6mmdiam,brownexternally,lightyellowinternally,slightlyaromaticwithbittersmell;root tubers elliptic, c.3.5by2cm,lightbrownexternally,purewhiteinternallywithtranslucentwhite centre, distanced5–15 cm from rhizome.Leafy shoot to90cm longwith1–5 leaveswhen flowering;pseudostemto30cmlong,composedofleafsheaths;leafless sheathsdecayedatanthesis;leaf sheaths white green or with pink tinge at base turning green towards the apex, densely puberulent;ligule3–5mmlong,bilobed,lobesround,hyaline,greenishwhite,translucent,hairy;petiole6–27cmlong(petioleof first leaf shortest, innermost leaves longer), canaliculate,green,denselypuberulent;lamina elliptic, broadly elliptic to el-liptic-ovate,to36by17cm,prominentlyplicate,adaxiallybrightgreen, shortly hairy along main veins, abaxially lighter green, shortlypuberulent;midribglabrous,green;basecordate;apexacute,c.5–10mmlong.Inflorescence central(oftenbreakingthroughthepseudostem)orrarelylateral(bothcanoccuronthesameplant),many-flowered;peduncle3–13cmlong,to7mmdiam,cream-white;spike4–9cmlong,c.3cmdiaminthemiddle,withoutcoma;fertile bracts7–24perinflorescence,3–6by1.2–3.3cm(largeratthebaseoftheinflorescence),ovate to trullate, smaller and ovate at the apex, whitish to light green at base with various degree of red tinge throughout the bract(resultinginverylightpink,lightredtodeepredbracts;the colour of the bracts is consistent within a spike, but varies withinapopulation),verysparselypuberulentoutside,glabrousinside,connateinthelower1/2to1/3;enclosingcincinniwith3flowersatthebaseoftheinflorescence,1–2flowersatthetop;bracteoles strongly reduced, one per flower, ovate, boat-shaped,c.1–2byc.1mm(outeroneslarger,inneronesgra- dually smaller or totally absent), hyaline, translucentwhite,glabrous.Flowers6.5–7.5cm,muchexsertedfromthebracts;calyxto22mmlong,3-toothed,unilaterallysplitc.8–10mm,translucent white with more or less red tinge especially distally, sparselypuberulent;floral tubec.4–5cmlong,externallywhiteat the base, with rich red tinge at the apex, densely puberulent, internally white to very light orange, puberulent with dorsally placedgrooveholdingthestyle;dorsal corolla lobe 22–25 by 9–11 mm, triangular ovate, concave, with sides rolled inwards, glabrous,redoutside,lightredinside,apexmucronate,mucroc.2mm;lateral corolla lobes20–21by8–9mm,triangularovatewith sides rolled inwards, concave, glabrous, red on outside,

109J.Leong-Škorničkováetal.:GingerspeciesfromLaos

Fig. 2 Curcuma flammea Škorničk.a,b.Habit;c,d.inflorescences;e.ligules;f–h.flower(sideview,frontview,topview);i.detailofstamen(side,backandfrontview);j.flowerdissection(allfromtype:JLS-1777).—Scalebars:i=1cm;j=2cm.—PhotosbyJanaLeong-Škorničková.

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rhizomes),internallypurple,smellingstronglyoflemongrass,scalesbroadlytriangular,soondecaying. Leafy shootto2.5mlong,arching,withupto42leaves,baseswollento3cmdiam,pink-red,basal1/2to1/3ofpseudostemleafless,sheathedbynumerous(c.13) leaflesssheaths; leafless sheaths dark greenwith faint, dark red-brown tinge towards base; leaf sheathsgreen,glabrousatbase,pubescenttowardstheligule;ligule to 9 mm long, shallowly bilobed, green, pubescent when young,glabrescentinoldpseudostems;petiole reduced to a pulvinus(to5mmlong);laminato30×8cm(graduallysmallertowards apex of leafy shoot), narrowly ovate (lanceolate),gradually tapering to attenuate apex, base rounded, adaxially glossy green, glabrous, abaxially lighter green and sparsely villose. Inflorescence radical, on erect peduncle to 16 cm longsheathedby4–8sterilebracts;spike9–18by4–5cm,fusiform,withupto50bracts;bractsobovate,to4.7by2.7cm(smallertowardsapexofinflorescence),apexbroadlyacute,externally bright red, whitish towards base, internally whitish except red margins, glabrous and glossy both sides, enclosing asingleflower;bracteole to 4 cm long, 15 mm broad at widest point, narrowly ovate, translucent white with apex tinged red, glabrous throughout. Flowers 7–9cm long;calyx c.21mmlong, translucent white, glabrous, with three inconspicuous, reddishteethatapex,withunilateralincisionc.10mm;floral tubec.42mmlong,whitetowardsbase,cream-yellowtowardsapex,externallyglabrous,internallyshortlypuberulousatapex;dorsal corolla lobe narrowly ovate-triangular, to 40 mm long, c.11mmbroadatwidestpoint,semi-translucentlightyellow,glabrous throughout, apex mucronate, mucro < 1mm;lateral corolla lobesnarrowlyovate-triangular,to35mmlong,5–6mmbroad at widest point, semi-translucent light yellow, glabrous throughout, apexacute; labellum c. 30mm long, c. 16mmwide(c.28mmincludingthestaminodes),ovate,cream-white;lateral staminodes16–20mmlong,c.7mmbroad,moreorlesselliptic,connatetothelabellumbybasal2/3,cream-whitewithdensebrightpurple-violetblotches.Stamen25mmlong;filament reduced to < 1mm;antherc.25mmlong(crestnotstraightened),c.4mmbroad,connectivetissuecream-whitewithpinktinge,glabrous;anther crestc.20mmlong(straight-ened),wrappedaroundstigma,white,glabrous;anther thecae c.14mmlong,dehiscingalongtheirentirelength.Epigynous glands2,c.5mmlong,c.0.75mmdiam,ochraceous.Style white,glabrous;stigma2–3mmlong,slightlythickerthanstyle,white,ostiolefacingdownwards(towardsthelabellum),ciliate.Ovary4.5by3mm,white,glabrous,trilocularwithcentralpla-centation. Fruit (youngfruit,perhapsimmature)ovoidorslightlytrigonalcapsule,2.3by1.1cm,white,glabrous;seedsc.8by3mm,darkmaroon,enclosedintranslucentwhite,irregularlylaciniatearil.DescriptionbasedonlivingandspiritmaterialofLeong-Škorničková et al. JLS-1807. Distribution — Currentlyknownonlyfromthetypelocality. Habitat&Ecology— Growingnearstreamsinlowlandde-ciduousforest. Phenology— Flowering in the early rainy season, June to July,flowersopenintheafternoon,around3pmandclosethenextmorning.FruitingoccursattheendofJunetoAugust. Conservation status — Specimen records exist so far only fromthetypelocality.Accordingly,weconcludethattheextentof occurrence is less than 20 000 km2.Thetypelocality isapopular tourist destination and the type population is not in a protected area, continuing decline in area, extent and quality of habitatcanbeforeseen.CategoryVulnerableVUB1a,b(iii)isthereforeproposedhere.FurtherexplorationoftheareaaroundVang Vieng is needed to understand the situation better and to amendtheconservationstatus.

Notes — By its fusiform inflorescences and bright red bracts, Z. jiewhoei resembles Z. peninsulare and Z. newmanii Theilade

lightredoninside;labellum21–25mmlong,7–9mmbroadatbasal1/3,thendiamond-shapedindistal2/3,c.16–21mmatwidestpoint,apexbifidwithanincisionto8mmlong,baseofthe labellum light orange with dark red lines, middle and apical part of the labellum orange at apex and margin, with dark red tinge towards the lower part, with deep orange band marked with dark red line on the outside running through the centre (medianband);lateral staminodes20–22by10–13mm,rhom-boid, bright red in basal half, orange at apex, glabrous outside, with short glandular hairs present on the middle part facing the centreoftheflower.Stamen16–17mmlong;filament7–8mmlong, orange with red tinge, 4 mm broad at base, 2 mm broad at apex(thepointofattachmenttotheconnective),coveredwithglandularhairs;antherc.14mmlong,spurred,connectivelightorange,denselycoveredwithshortglandularhairs;anther spurs 3–4mmlong,narrowlytriangularwithsharptipspointingout-wards;anther crestthick,rounded,c.1.5mmlongandc.2mm broadat base, orange;anther thecae 9–10 mm, dehiscing alongtheirentirelength.Epigynous glands2,c.5–6mmlong,c.0.6mmdiam,lightyellowatbase,darkeratapex.Style thin, white,glabrous,placedinagrooveindorsalsideoffloraltube;stigmac.1.5mmlong,1.5mmwide,white,ostioleciliate,facingupwards.Ovary2–3by2–2.5mm,trilocular,denselyhairy,hairsc.0.2mmlong.Fruits notseen.Descriptionbasedonliving and spirit material of Leong-Škorničková et al. JLS-1777. Distribution—Currentlyknownonlyfromthetypelocality. Habitat&Ecology— Growingonsteep limestoneslopescoveredwithprimarydipterocarpforestat400–500masl. Phenology— Floweringintheearlyrainyseason,endMayto July, flowers last a single day, open in the morning and clos-inginlateafternoon.FruitingJunetoJuly(August?). Conservation status — Specimen records exist so far only fromthetypelocality.MostCurcumaspeciesinsubg.Ecomata, especially from the alliance of C. bicolor, where this species belongs, tend to be fairly restricted in distribution.Accord-ingly, we estimate that the extent of occurrence is less than 20 000 km2.While the habitat of this species is unsuitablefor fast agricultural exploitation, the fact that Vang Vieng is a popular tourist destination and the type population is not in a protected area, continuing decline in area, extent and quality ofhabitatcanbeforeseen.CategoryVulnerableVUB1ab(iii)isthereforeproposedhere.FurtherexplorationoftheareaaroundVang Vieng is needed to understand the situation better and amendtheconservationstatus.

Notes — Curcuma flammeaalsobelongstosubg.Ecomata.Morphologically, themostsimilarspeciesareC. bicolor and C. rhombaMood&K.Larsen.Preliminarymolecularanalysesofsubg.Ecomata(Záveskáetal.unpubl.)suggestthatC. flam-mea is closer to C. bicolor than to C. rhomba.

Zingiber jiewhoei Škorničk. sp. nov.—Fig.3

Similar to Zingiber peninsulare Theilade, but differs in cream-white label-lum and cream-white staminodes with dense, bright, purple-violet blotches (vslabellumandstaminodesdarkpurplewithwhitespots).—Type:Jana Leong-Škorničková, Trần Hṹu Đăng, Somsanith Bouamanivong, Khantheo Santhammavong, Udone Souvannakhoummane, Kittisack Phoutthavong JLS-1807 (holotypeSING (incl. spirit); isotypeE,HNL), Laos,VientianeProv.,VangViengDist., KaengNyuiwaterfall, 370masl,N18°57'21.5"E102°29'45.5",25June2012.

Etymology. WenamethisbeautifulspeciesinhonourofMr.TanJiewHoe,PresidentoftheSingaporeGardeningSociety,forhiscontinuouscontribu-tionstowardsgingerresearchatSingaporeBotanicGardensandgeneroussupportofplanttaxonomyinLaos.

Rhizomatous,clump-formingherb. Rhizome 1–3cmdiam,branch- ed,1–3cmbetweenleafyshootsarisingfromthesamerhi-zome,externallypinkish(youngrhizomes)tolightbrown(older

111J.Leong-Škorničkováetal.:GingerspeciesfromLaos

Fig. 3 Zingiber jiewhoei Škorničk.a,b.Habit;c.detailofpulvinusandligule;d.internalcolourofrhizome;e.inflorescence;f.flower(sideview);g.flower(frontview);h.seeds;i.youngfruitinbract;j.stamen(frontandsideview);k.fertilebractandflowerwithbracteole(sideview);l.dissectedflower(allfromtype: JLS-1807).—Scalebars:h,j=1cm;k,l=2cm.—PhotosbyJanaLeong-Škorničková.

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112 Blumea – Volume 59 / 2, 2014

but it can be readily recognised by its cream-white labellum and cream-white staminodes with dense bright purple-violet blotches(vslabellumandstaminodeswithdensedark-maroonblotches).

AcknowledgementsWethankthecuratorsofBK,BKF,E,HN,K,P,SING,andVNMforlettingusexaminespecimensintheircare,theAsianZingi-beraceaeInformationCentreatSingaporeBotanicGardens,andtheZingi-beraceaeResourceCentreattheRoyalBotanicGardenEdinburgh(http://elmer.rbge.org.uk/ZRC/)forprovidingprotologuesandrelatedreferences.WearegratefultoRikGadella,DirectorofPhaTadKeBotanicGarden,andDr.SourioudongSundara,DirectorBiotechnologyandEcology Institute,MinistryofScienceandTechnology,fortheirsolidsupportduringourfieldworkinLaos.WealsothankTrầnHṹuĐăng,SomdiOudomsackandLeeVuefortheirassistanceinthefield.FundingbytheNationalParksBoard(Singapore),andbytheCzechScienceFoundation(GAČR14-13541S)isgratefullyac-knowledged.FurtherfundingwaskindlyprovidedbytheMinistryofCultureof theCzechRepublic (DKRVO2013/10,NationalMuseum, 00023272).The two reviewers are acknowledged for their constructive comments and languageimprovementsofthismanuscript.

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