Taxonomic Status of the Population of Rhacophorus From Sumbar

8/13/2019 Taxonomic Status of the Population of Rhacophorus From Sumbar

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New Rhacophorus from Sumatra

All articles available online at http://www.salamandra-journal.com 2011 Deutsche Gesellscha fr Herpetologie und Terrarienkunde e.V. (DGHT), Rheinbach, Germany

SALAMANDRA 47(3) 133143 20 August 2011 ISSN 00363375

Taxonomic status of the population of

Rhacophorus angulirostris A, 1927 (Anura: Rhacophoridae)

from Sumatera Barat (West Sumatra) and its description as a new species

J. M D

Institut fr Integrierte Naturwissenschaen, Abteilung Biologie, AG Zoologie, Universitt Koblenz-Landau, Universittsstr. 1,56070 Koblenz, Germany

e-mail: [email protected]

Manuscript received: 26 November 2010

Abstract.e taxonomic status of the population of Rhacophorus angulirostrisfrom West Sumatra is reassessed. is recordwas based on five specimens that had been collected in Padang, West Sumatra, more than a century ago, are stored in thecollection of the Naturhistorisches Museum Wien, Austria, and were assigned to R. angulirostrisfrom Borneo by I(). Reexamination revealed that the specimens from Sumatra share only few diagnostic characters with R. angulirostrisfrom Borneo. Bornean and Sumatran specimens differ in the presence of a dermal projection at the tibio-tarsal articula-tion, male body size, snout shape, shape of the canthus rostralis and the supratympanic fold, extent of webbing between thefingers, size of the thenar tubercle, position of the nostrils, internarial distance, the eye-to-nostril distance, size of eye andtympanum, and size and position of the dentigerous processes of the vomers. e specimens from Padang were comparedto all other Southeast Asian members of the genus and found to represent a distinct species, herein described as new, thatappears to be not even closely related to R. angulirostris. Because the only record of R. angulirostrisfrom Sumatra was basedon these specimens, the geographic distribution of R. angulirostrisis again restricted to northeastern Borneo. An identifica-tion key for the Rhacophorusspecies from Sumatra is provided.

Key words. Amphibia, Rhacophorus pseudacutirostrissp. n., identification key, Indonesia, Borneo, endemism.

Introduction

Species of the genus Rhacophorus K H, are distributed from India to China and southern Ja-pan and southeastwards to the Greater Sunda Islands andthe Philippines. e genus currently contains species(F , C G ).

Fourteen species of Rhacophorus have been reportedfrom Sumatra: R. achantharrhena H, P S, ; R. angulirostris A, ; R. appendicula-tus (G, ); R. barisani H, P S, ; R. bifasciatus K, ; R. catami-

tusH, P S, ; R. cyanopuncta-tus M & S, ; R. margaritifer (S,); R. modestus B, ; R. nigropalmatusB, ; R. pardalis G, ; R. poecilono-tus B, ; R. prominanus S, ; and R.reinwardtii (S, ). Recently, R. norhayatiaeC G, has been described from south-ern ailand and Peninsular Malaysia. e species wasprovisionally included in the herpetofauna of Sumatra byC G () for closely matching the descrip-tion of Rhacophorus reinwardti var. lateralis which hadbeen described by W () from Laut Tador, BatuBahra, Sumatra, and formerly been considered a synonym

of R. reinwardtii.It is currently unclear, to which speciesthe Sumatran population belongs that has been referred toas R. reinwardtiior if even both species occur on Sumatra.

erefore, both R. reinwardtii and R. norhayatiae are in-cluded in the identification key to the Sumatran species ofRhacophorusprovided below.

Rhacophorus angulirostris was originally described asRhacophorus acutirostris by M () fromKina Balu [= Gunung Kinabalu, Sabah, Malaysia, north-eastern Borneo]. Subsequently, the species, which wasoriginally described as Ixalus acutirostrisby P ()and which is now in the genus Philautus, was transferredto the genus Rhacophorus, rendering R. acutirostrisM-, a junior homonym. erefore, Ms() original name was replaced with Rhacophorus an-

gulirostrisby A ().Rhacophorus angulirostrishas been recorded only oncefrom Sumatra by I () who referred specimensof a Rhacophorus species collected at Padang, West Su-matra, and deposited in the collection of the Naturhis-torisches Museum Wien, Austria (NHW), to the Borneanspecies R. angulirostris(using the old name R. acutirostrisM, ) based on the morphological charactershe had studied. In the course of an ongoing study on therhacophorids of the Sunda region, I examined the speci-mens of the genus Rhacophorus from Sumatra depositedin the collection of the NHW and noticed many morpho-logical differences between the specimens from Padang as-

signed to R. angulirostris by I () and specimens ofR. angulirostris from Borneo. e former represent a spe-cies that appears to be not even closely related to R. an-


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gulirostris and furthermore does not match any of the de-scribed species from Southeast Asia. erefore, I formallydescribe it herein as new to science. As a consequence, thegeographic distribution of R. angulirostris is restricted tonortheastern Borneo.

Material and methods

Type specimens are stored in the collection of the Naturhis-torisches Museum Wien, Austria (NHW). I took the fol-lowing measurements with a digital calliper (to the near-est . mm): snoutvent length (SVL, from tip of snoutto vent); tibiofibula length (TFL, measured with both theknee and tibio-tarsal articulation flexed); knee to knee dis-tance (KNE, distance between knees with thighs being heldlaterally at right angles to the body); thigh length (THL,half the knee to knee distance); tarsus + foot length (TarL;distance from tibio-tarsal articulation to tip of fourth toe);foot length (FOT, distance from proximal end of innermetatarsal tubercle to tip of fourth toe); total leg length(LEG, distance from vent to tip of fourth toe, measuredwith leg fully extended at a right angle to the body); lengthof fingers (FL, FL, FL, FL, distance from proximal endof thenar tubercle to tip of first, second, third, and fourthfinger, respectively); forearm + hand length (ARM, dis-tance from elbow to tip of third finger); elbow to elbowdistance (ELB, measured with upper arms held laterally atright angles to body and elbows flexed); head width (HW,distance between angles of jaw); head length (HL, distancefrom angle of jaw to tip of snout); horizontal eye diameter(ED); horizontal tympanum diameter (TD); upper eyelidwidth (EW); interorbital distance (IO, shortest distance be-

tween upper eyelids); eye to nostril distance (EN, distancebetween anterior margin of eye and centre of nostril); nos-tril to snout distance (NS, distance between centre of nos-tril and tip of snout); snout length (SL, distance betweenanterior margin of eye and tip of snout); internarial dis-tance (NN, distance between centres of nostrils). e web-bing formulae are given as proposed by M D- ().

For comparisons, I examined museum specimens ofseveral species of Rhacophorus (see Appendix). Museumabbreviations are as follows: e Natural History Museum(British Museum [Natural History]), London, United King-dom (BMNH); e Field Museum (Field Museum of Natu-

ral History), Chicago, Illinois, USA (FMNH); Naturhisto-risches Museum Basel, Switzerland (NHMB); Naturhisto-risches Museum Wien, Austria (NHW); NaturhistorischesMuseum der Burgergemeinde Bern, Switzerland (NMBE);Museum und Forschungsinstitut Senckenberg, Frankfurtam Main, Germany (SMF); Sabah Parks Zoological Mu-seum, Kinabalu Park Headquarters, Ranau District, Sabah,Malaysia (SP); Zoological Museum of the Department ofBiology, Universiti Brunei Darussalam, Bandar Seri Bega-wan, Brunei Darussalam (UBD); Museum fr Naturkunde,Leibniz-Institut fr Evolutions- und Biodiversittsfor-schung an der Humboldt-Universitt zu Berlin, Germany(ZMB). JMD field numbers refer to specimens in my work-

ing collection, which eventually will be deposited in thecollection of the NMBE. To ensure that no available nameis referable to the new species, I also examined the types of

all species that are currently regarded as synonyms of spe-cies occurring on Sumatra and/or Java, i.e., Rhacophorusbarbouri A, , R. javanusB, , R. phyllo-pygusW, , and R. pulchellus W, .

Additional information on characters used to compareSoutheast Asian species of Rhacophorus and for working

out the identification key to the Sumatran species was takenfrom: B (, ), W (), K- (), A (), I et al. (), O et al.(), O et al. (), H et al. (), I I (), W et al. (), M P- (), O (), D G (), O- et al. (, ), O (), and C G- (). Where respective data was available, compari-sons were made for males and females separately.

Rhacophorus pseudacutirostris sp. n.(Figs. )

Rhacophorus acutirostris (nec Rhacophorus acutirostris M,, archeonym replaced by Rhacophorus angulirostris A, ):I (, partim).

Rhacophorus angulirostris (nec Rhacophorus angulirostris A, ):M G (, partim); H et al. (, par-tim); M et al. (, partim); D (, partim); T- et al. (, partim).

Holotype: NHW :, adult male, from Padang, Sumat-era Barat (West Sumatra), Indonesia, collected between and by Consul J S.

Paratypes: NHW :, NHW :, two adult females,

NHW:, :, two adult males, same data as holo-type.

Diagnosis: A species of the genus Rhacophorus, distin-guishable from its congeners by the combination of thefollowing characters: () small to medium size (SVL ofadult males .. mm, adult females .. mm);() snout obtusely pointed in both dorsal view and profile;() head wider than trunk and wider than long; () canthusrostralis distinct; () nostrils closer to tip of snout than toeye, separated from each other by distance subequal to dis-tance between eye and nostril; () eyes moderately large;() eye diameter greater than eye-to-nostril distance; ()

interorbital distance wider than upper eyelid and greaterthan internarial distance; () tympanum diameter half theeye diameter; () dentigerous processes of vomers aboutthree times as long as broad and separated from each otherby distance slightly less than their length; () dorsal faces,ventral side of limbs, chin, throat, and chest region smooth;() supratympanic fold thick and conspicuous, not con-cealing tympanum; () low transverse ridge present insupracloacal region; () dermal flaps absent on postax-ial edges of forearm and tarsus; () large conical tuberclepresent at tibio-tarsal articulation; () fingers webbed forabout one-third of their lengths, toes for three-fourth; ()nuptial pads absent in males.

Description of holotype: Measurements are given in Table. Body moderately slender, widest at temporal region, ta-


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pering to groin (Fig. ); head large (HL/SVL .), widerthan trunk and wider than long (HW/HL .); snout long-er than eye diameter (SL/ED .); snout projecting beyondmandible, obtusely pointed in both dorsal view and profile(Fig. ); canthus rostralis distinct, sharp, slightly curvedin profile, concave in dorsal view; loreal region sloping,

moderately concave; nostrils directed dorsolaterally, situ-ated in low protuberances, closer to tip of snout than to eye(EN/NS .), separated from each other by distance sub-equal to distance between eye and nostril (NN/EN .);eyes directed anterolaterally, protruding, moderately large(ED/HL .); pupil horizontal; eye diameter greater thaneye-to-nostril distance (ED/EN .); interorbital distancewider than upper eyelid (IO/EW .) and greater than in-ternarial distance (IO/NN .); tympanum visible, sep-arated from orbit by distance equal to half its diameter;tympanic diameter subequal to half eye diameter (TD/ED .); tympanic annulus visible; skin not co-ossifiedto forehead; upper jaw with dentition; teeth on premax-illary larger than those on maxillary; choanae located farlaterally, at margins of roof of mouth; dentigerous proc-esses of vomers strongly developed, beginning at antero-medial edge of choanae, directed posteromedially, aboutthree times as long as broad, bearing small teeth, separatedfrom each other by distance slightly less than their lengths;tongue moderately broad, bifurcated at distal end, and freefor about half its length; median lingual process absent;median subgular vocal sac present; vocal slits on both sidesnear base of tongue.

Dorsal surfaces, ventral side of limbs, chin, throat, andchest region smooth; abdomen coarsely granular; supra-tympanic fold thick and conspicuous, extending from pos-terior edge of upper eyelid to scapular region, not conceal-

NHW 16301:5 16301:2 16301:4 16301:1 16301:3

Status holotype paratype paratype paratype paratype

Sex male male male female female

SVL 39.6 39.9 37.0 52.3 51.8

HW 14.1 14.9 13.6 18.6 17.8

HL 12.1 12.4 11.7 14.9 14.6

ED 5.0 4.7 4.7 6.4 5.7

TD 2.4 2.5 2.4 2.8 2.6

EW 3.3 3.4 3.1 4.5 4.2

IO 4.5 4.7 4.3 5.0 5.1

EN 3.8 3.7 3.6 4.3 4.4

NS 3.1 3.2 3.0 3.9 3.8

SL 6.6 6.6 6.3 7.9 7.5

NN 3.8 3.9 3.8 4.8 4.4

TFL 21.5 21.4 20.0 26.0 25.7

KNE 38.1 35.9 34.2 48.4 48.5

THL 19.0 18.0 17.1 24.2 24.3

TarL 29.2 28.5 25.8 37.6 37.0

FOT 17.8 17.6 16.2 23.9 24.0

LEG 65.3 63.5 60.3 83.9 83.8

1FL 6.4 6.2 6.3 8.6 9.0

2FL 8.2 8.5 8.4 11.2 11.3

3FL 13.1 12.8 12.3 17.0 17.3

4FL 11.2 11.2 10.6 14.7 14.9

ARM 20.1 19.5 19.3 27.2 27.4

ELB 25.5 27.1 24.6 36.0 36.1

Table 1. Morphological measurements of the type series ofRhacophorus pseudacutirostrissp. n. (in mm). For abbreviations,see Material and methods.

Figure 1. Dorsal (le) and ventral views (right) of the male holotype of Rhacophorus pseudacutirostrissp. n. (NHW 16301:5). Scalebar = 5 mm.


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ing tympanum; low transverse ridge in supracloacal re-gion, bearing row of small tubercles; few small tuberclesin infraanal region and on ventral sides of thighs; small,indistinct white tubercles along postaxial edge of forearm;large conical tubercle at tibio-tarsal articulation.

Forelimbs moderately slender; hand longer than fore-

arm (FL/ARM .); tips of fingers enlarged into broadoval discs, each with circummarginal groove; disc of Fin-ger III wider than tympanum diameter; relative lengths offingers: I < II < IV < III; subarticular tubercles rounded,well developed, numbering one on Fingers I and II, twoon Fingers III and IV; distal subarticular tubercles on Fin-gers III and IV much larger than proximal ones; webbingformula of the hand: I..II-III..IV (Fig. );thenar tubercle oval, comparatively small, about one-thirdof base of Finger I; inner and outer palmar tubercles oval,small; narrow dermal ridge on postaxial edge of Finger IVbetween base of disc and outer palmar tubercle; metacar-pals with several supernumerary small tubercles; nuptialpads or asperities absent.

Hindlimbs slender, moderately long (LEG/SVL .);distance between knees considerably larger than betweenelbows (KNE/ELB .); tibio-tarsal articulation reachingto tip of snout when legs are adpressed to body; tibiofibulamoderately long (TFL/SVL .), longer than thigh (TFL/THL .); heels overlapping each other considerably whenknees are flexed and thighs are held laterally at right an-

gles to the body; foot shorter than tibiofibula (FOT/TFL.); relative lengths of toes: I < II < III < V < IV; discsof toes smaller than those of fingers; subarticular tuberclesnumbering one on Toes I and II, two on Toes III and V,and three on Toe IV; pedal webbing formula: I..II+.III..IV.+V (Fig. );inner metatarsal tubercle

oval, low and small (length . mm), no outer one; few su-pernumerary small tubercles on metatarsals; dermal ridgeon postaxial edge of Toe V from disc to base of metatarsus.

In preservative, dorsum greyish with darker markings.Markings irregular on dorsum, as broad crosslines on dor-sal faces of limbs. Ventral side of body cream-coloured, oflimbs darker, more yellowish. Flanks and postaxial lateralface of thigh speckled with light, irregularly shaped blotch-es. Tubercles on ventral sides of thighs, along postaxialedge of forearm, and in supraanal and infraanal regionwhitish. Colour of iris bluish grey. No information is avail-able regarding the colouration in life.

Variation:Measurements of the paratypes are given in Ta-ble . e male paratypes are very similar to the holotypein measurements and proportions. ey likewise have me-dian subgular vocal sacs and lack nuptial pads and asperi-ties. Variation of webbing in males is summarized by I..II-III.(+.)IV for the hand and I.(.-)II+(..)III(+.)(..)IV(+.)+V for thefoot. Females are larger than males, have slightly shorter

Figure 2. Lateral (le) and dorsal views (right) of the head of the male holotype of Rhacophorus pseudacutirostris sp. n.(top, NHW16301:5) and a topotypic male Rhacophorus angulirostris(bottom, ZMB 61674) (not to the same scale).


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heads with an HL/SVL of . (vs. ..), and havecomparatively shorter tibiofibulae with a TFL/SVL of .(vs. . in males), TFL/THL .. (vs. ..), andTFL/FOT .. (vs. ..). Variation of webbing infemales is summarized by I..II.III.+IV forthe hand and I+-II++III+.IV++V for the foot.

Distribution: So far, the new species is only known froma single locality, which was given as Padang, Sumatra byJ S, the collector of the type series (Fig. ).e specimens were probably not collected in the city itselfbut more likely in the forests in the vicinity of Padang. No

further specimens have been collected or reported fromSumatra since the collection of the type series more than years ago.

Etymology: e species epithet is composed ofpseud-, theLatinized form of the Greek prefix -, meaning false,and the Latin words acutus, meaning sharpened, and ros-trum, meaning snout; in allusion to the former confusionof the new species with Rhacophorus acutirostris M-, (name replaced with R. angulirostris A,) from Borneo. As common name, I suggest SumatranSharp-nosed Tree Frog.

Comparisons:Rhacophorus angulirostris and R. penanorumD, , differ from R. pseudacutirostris (charac-ters in parentheses) in the following characters: e malesare smaller with an SVL of .. mm in R. angulirostrisand .. mm in R. penanorum(vs. .. mm); thetibio-tarsal articulation lacks a dermal projection (large,pointed dermal projection present at tibio-tarsal articula-tion); the snout is sharply pointed and projects consider-ably beyond the lower jaw, sloping forward from nostrilthan back to mouth in profile (obtusely pointed, projectingless; Fig. ); the canthus rostralis is more pronounced; thewebbing between the fingers is more developed (Fig. ),formula being III,(.)III(.)(.)IV in R.

angulirostrisand I..II.III(-

)IV in R. pen-anorum (vs. I..II[.-]III.[+.]IV); thenostrils are situated about half way between eye and tip of

snout with EN/NS .. in male R. angulirostris (Fig.), .. in female R. angulirostris, and .. inmale R. penanorum(females of R. penanorumunknown)(vs. situated closer to the tip of snout than to eye with EN/NS .. in males [Fig. ] and .. in females); theinternarial distance is relatively larger with NN/EN .. in R. angulirostrisand .. in R. penanorum (vs... in males, .. in females); the eye-to-nostrildistance is relatively smaller in relation to the eye diameterwith ED/EN .. in male R. angulirostrisand .. in male R. penanorum(vs. ..; Fig. ); the eyeis relatively larger with ED/HL .. in male R. an-

gulirostrisand .. in male R. penanorum(vs. ..; Fig. ); the tympanum is relatively smaller with TD/ED .. in male R. angulirostrisand .. in R.penanorum(vs. .. in males; Fig. ); the supratym-panic fold conceals the tympanum for one-fih on the up-per edge (not concealing tympanum; Fig. ); the dentiger-ous processes of the vomers are shorter and separated fromeach other by a distance equal to twice their length in R.angulirostrisand four-thirds their length in R. penanorum(processes longer, separated by distance equal to less thanthe length of an individual process); the thenar tubercle ismuch larger, being almost the size of the base of Finger I(Fig. ; vs. smaller, about one-third of base of Finger I, Fig.

). Additionally, R. angulirostrisdiffers by a relatively small-er interorbital distance in males with IO/EW .. (vs...) and a relatively larger such distance in femaleswith IO/EW .. (vs. ..). R. penanorum has aslightly shorter tibia with TFL/SVL .. in males (vs..) and a slightly wider head with HW/HL .. (vs...).

In the following congeners the webbing reaches thedisc on the postaxial side of the third finger (f), on pre-axial and postaxial sides of the fourth toe (t) or both (f &t), and these species are thus readily distinguished fromRhacophorus pseudacutirostris (webbing reaching at mostto proximal edge of distal subarticular tubercle on postax-

ial side of third finger, preaxial side of fourth finger, andpreaxial and postaxial sides of the fourth toe): Rhacopho-rus annamensisS, (f & t); R. dennysiB,

Figure 3. Volar view of right hand (le) and plantar view of lefoot (right) of the holotype of Rhacophorus pseudacutirostris sp.n.(NHW 16301:5). Scale bar = 2 mm.

Figure 4. Volar view of hand of the male holotype of Rhacophoruspenanorum(le, ZMB 70718) and a male Rhacophorus anguliros-tris sp. n. (right, SP 2913) (not to the same scale).


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(t); R. dulitensisB, (t); R. exechopygusI, O D, (f & t); R. fasciatusB-, (f & t); R. feaeB, (f & t); R.georgiiR, (f & t); R. harrissoni I H, (f & t); R. jarujiniM P, (t); R. kioO & D, (f & t); R. maximusG,

(f & t); R. nigropalmatus(f & t); R. norhayatiae(f & t);R. pardalis(including its synonym R. pulchellus; f & t); R.poecilonotus(t); R. prominanus(t); R. reinwardtii(f & t); R.rhodopusL H, (f & t); R. robinsoniiB, (f & t); R. rufipes (f & t).

Webbing between the fingers is confined to the bases inR. dugritei(D, ), R. everettiB, , R.hungfuensisL H, , and R. kajauD, .

e following Southeast Asian species differ from thenew species in having conspicuous dermal flaps or fring-es along the postaxial edge of the forearm (F), the tarsus(T), or both (F & T): Rhacophorus achantharrhena (F &T); R. appendiculatus(including R. phyllopygus; F & T); R.baluensisI, (F & T); R. barisani(F & T); R. bi-fasciatus (F & T); R. bipunctatusA, (F & T); R. cal-caneusS, (T); Rhacophorus chuyangsinensisO-, N H, (F & T); R. edentulusM, (F & T); R. hoanglienensisO, L, M H, (F); R. margaritifer(including R. javanusand R.barbouri; F & T); R. monticolaB, (F & T);R. turpesS, (F & T).

e remaining Southeast Asian Rhacophorus spe-cies differ in the following characters from the new spe-cies (characters given in parentheses). Rhacophorus be-lalongensisD G, , R. bimaculatus(P-, ), R. gadingensisD H, , and R. gauni(I, ) are somewhat smaller species (SVL of males


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of characters that show little or no intraspecific variation(but sometimes are sexually dimorphic) and are thereforewell suited to distinguish species of the genus Rhacopho-rus, including the presence/absence of dermal flaps or ap-pendages on the limbs, snout shape, extent of webbing,and others more. Rhacophorus pseudacutirostrisand R. an-

gulirostrisin fact do share some of these characters. eyboth lack dermal flaps or fringes on the postaxial edges offorearm and tarsus, have a smooth dorsal skin, and a simi-lar extent of foot webbing. ese characters, however, arealso shared by a number of other species. Two characters,male body size and the pointed snout, are similar in thetwo species, but show constant differences on closer ex-amination. On the other hand, R. pseudacutirostrisdiffersfrom R. angulirostrisin a number of diagnostic characters,among them some that were already used by W ()(who considered R. angulirostrisone of many subspeciesof a wide-ranging R. schlegelii) to differentiate R. anguli-rostris from other subspecies of R. schlegelii (which werelater re-elevated to species rank), i.e., the presence of der-mal appendages on the heel and the size and relative posi-tion of vomerine odontophores. ese characters were alsomentioned by I () in his description of R. angu-lirostris. erefore, it is unclear why I () chose toassign the Padang specimens to R. angulirostris. Rhacopho-rus pseudacutirostrisdiffers in a number of other diagnos-tic characters from R. angulirostris(see above). In fact, R.angulirostris has more diagnostic characters in commonwith its sister species R. penanorum and also with someother congeners like R. rufipesand R. cyanopunctatus, allof which have a sharply pointed snout and lack dermal ap-pendages on the heel, by which characters they differ fromR. pseudacutirostris.

Furthermore, the distribution range of the Sumatranendemic R. pseudacutirostris, which is known only fromPadang, is widely separated from those of the Bornean en-demics R. angulirostris and R. penanorum, the former of

which has been recorded so far from Gunung Kinabalu,Gunung Trusmadi and the Crocker Range in Sabah andthe latter is known only from Gunung Mulu in Sarawak(D , D et al. ; Fig. ).

To validate that no available name is referable to thepopulation from Padang, West Sumatra, I examined the

types of all species that are currently regarded as synonymsof Rhacophorusspecies occurring on Sumatra and/or Java.In the process, I was able to confirm the status as syno-nyms of R. barbouri (type locality: West Java; junior syno-nym of R. margaritifer), R. javanus (type locality: VulkanTjisurupan, West Java; junior synonym of R. margaritifer),R. phyllopygus (type locality: Indragiri, Sumatra; juniorsynonym of R. appendiculatus), and R. pulchellus (type lo-cality: Djapura, Indragiri, Sumatra; junior synonym of R.pardalis). All these taxa are clearly distinguishable from R.pseudacutirostris(see above). I furthermore examined thetype of Rhacophorus depressus A, , a species origi-nally described from West Java and currently regarded asincertae sedis by F (). e type of this species isa male measuring . mm in SVL. Its toes are completelywebbed, but there is no webbing between the fingers. etips of fingers and toes are enlarged into broad discs. Con-trary to character states observed in species of the genusRhacophorus,metatarsal tubercles are absent, and the dor-sal face of Finger I bears a nuptial pad covered with manylarge spines. is species is probably not at all a member ofthe Rhacophoridae but more likely belongs to the Hylidae,possibly Litoria. Because members of the hylid family donot occur autochthonally on Java, the type specimen mightnot even originate from this island.

Although the description of R. pseudacutirostris pre-sented here is based on five specimens only and bioacous-

tic and molecular data are unfortunately lacking, detailedmorphological comparisons revealed that the availablespecimens exhibit a unique combination of characters thatdistinguish the species from all other known Southeast

Figure 5. Point locality distribution map of Rhacophorus pseudacutirostris sp. n., R. angulirostris, and R. penanorum.


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Asian members of the genus. erefore, it seems justifiedto regard the population from Padang as a distinct evolu-tionary lineage, now named Rhacophoruspseudacutiros-tris.

Key to the Sumatran species of Rhacophorus

1 Fingers IIIV and all toes webbed to the base of the disc ..................... ...................... ....... 2

1* Fingers IIIV and toes not or not entirely webbed to the disc ................... ...................... ...............5

2 Finger I webbed to the disc, webbing black proximally, yellow distally .............. R. nigropalmatus

2* Finger I not webbed to disc, colour of webbing different .................... ...................... ................3

3 Dorsum green, webbing black with bluemarkings ......................................................................... 4

3* Dorsum brown, grey, or reddish, never green, webbing red .................... ...................... ........ R. pardalis

4 Dorsum with small, black spots; lateral faces of thigh and tibia yellow ..................... . R. reinwardtii

4* Dorsum without black spots; lateral faces of thigh and tibia black with blue markings ..................... ...................... ..... R. norhayatiae

5 Dermal flaps along the postaxial edges of forearm and tarsus .................... ...................... ............. 6

5* Dermal flaps absent .................... ...................... .......... 12

6 Dorsum green in life; webbing between

Toes IIIV red ...................... ...................... ................... 76* Dorsum not green; webbing between Toes IIIV not red ..................... ...................... .............. 8

7 Toes fully webbed; SVL of males > 50 mm; snout pointed ................... ..................... R. prominanus

7* Toes not fully webbed; SVL of males 4047 mm; snout acuminate ................... R. achantharrhena

8 SVL of adult males < 38 mm ..................... ................. 9

8* SVL of adult males > 40 mm ...................... .............. 10

9 Dorsum tuberculate; broad, wavy skin flaps along forearm; webbing reaching to distal subarticular tubercles on Finger IV and

Toe IV .............................................. R. appendiculatus

9* Dorsum shagreened; skin flap along forearmweakly expressed or reduced to a line of tu-bercles; webbing not reaching distal subar-

ticular tubercles ...................... ... R. catamitus(partim)

10 Snout sharply pointed ..................... ............ R. barisani

10* Snout rounded or acuminate .................... ................ 11

11 Tibio-tarsal articulation extending beyond tip of snout when the leg is adpressed forward ...................... ...................... ...... R. margaritifer

11* Tibio-tarsal articulation reaching a point

between eye and tip of snout when the leg is adpressed forward ...................... .. R. bifasciatus

12 Snout pointed ...................... ...................... .................. 13

12* Snout rounded or subacuminate ...................... ........ 14

13 Webbing on Finger III does not reach dis-tal subarticular tubercle; dermal projectionpresent at tibio-tarsal articulation; SVL of

females > 50 mm .................... .... R. pseudacutirostris

13* Finger III webbed beyond distal subarticular tubercle; dermal projection at tibio-tarsal articulation absent; SVL of females < 45 mm ........................................................... R. cyanopunctatus

14 SVL of adult males > 40 mm ...................... ............. 15

14* SVL of adult males < 38 mm ..................................................... R. catamitus (partim)

15 Toes fully webbed; dermal calcar at tibio- tarsal articulation present .................. R. poecilonotus

15* Toes not fully webbed; dermal calcar absent ..................... ..................... ................ R. modestus

Acknowledgements

I would like to thank H G (NHW) for the loan ofthe type specimens. For providing working space and permis-sion to examine comparative material under their care or forloans of comparative material, I am grateful to B C(BMNH), H V and A R (FMNH), ST. H and B B (NMBE), T. UG (UBD), G K (SMF), M Land P Y (SP), and M-O R, FT, R G, and D L (ZMB). R- W (NHMB), M D (SenckenbergGesellscha fr Naturforschung), and F T (ZMB)

sent detailed photographs of the types of Rhacophorus phyllopy-gus and R. pulchellus, R. javanus, and R. barbouri, respectively.F T also sent literature. J K and A- O made helpful comments on an earlier version ofthe manuscript. My visit to the Natural History Museum (Lon-don) was funded by the Synthesys Project (http://www.synthesys.info), which is financed by European Community Research In-frastructure Action under the FP Structuring the European Re-search Area programme (GB-TAF-).

References

A, E. (): Zur Systematik der asiatischen Arten der Frosch-gattung Rhacophorus. Sitzungsberichte der Gesellschanaturforschender Freunde zu Berlin, : .

B, O. (): Neue Reptilien und Batrachier aus West-Java. Zoologischer Anzeiger, (): .

B, G. A. (): Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum. Second Edi-tion. Taylor and Francis, London. xvi + pp., plates.

B, G. A. (): Reptiles and batrachians collected inKorinchi, West Sumatra, by Messrs. H. C. Robinson and C.Boden Kloss. Journal of the Federated Malay States Muse-ums, : .

C K. O. & L. L. Grismer (): Re-assessment of the Rein-wardts Gliding Frog, Rhacophorus reinwardtii (Schlegel )(Anura: Rhacophoridae) in southern ailand and Peninsular

Malaysia and its re-description as a new species. Zootaxa,: .


9/11

141


D, J. M. (): A new treefrog (Anura: Rhacophoridae:Rhacophorus) from Gunung Mulu, Borneo. Salamandra, :.

D, J. M. & T. U. Grafe (): A new treefrog of the genusRhacophorus (Anura: Rhacophoridae) from Brunei Darussal-am (Borneo). Salamandra, : .

D, J. M., P. Y M. B. L (): Advertisementcalls of Rhacophorus angulirostrisAhl, and Rhacophoruseveretti macroscelis Boulenger, from Gunung Kinabalu,Sabah, East Malaysia (Borneo). Herpetozoa, : .

F, D. R. (): Amphibian species of the world: an onlinereference. Version . ( January ). American Museumof Natural History, New York, USA. Electronic database ac-cessible at http://research.amnh.org/herpetology/amphibia/index.php (..).

H, M. B., A. J. P E. N. S (): New andpoorly known parachuting frogs (Rhacophoridae: Rhacopho-rus) from Sumatra and Java. Herpetological Monographs,: .

I, R. F. (): e systematics and zoogeography of the Am-phibia of Borneo. Fieldiana: Zoology, : .

I, R. F. D. T. I (): A collection of amphibi-ans from West Sumatra, with description of a new species of

Megophrys(Amphibia: Anura). e Raffles Bulletin of Zoo-logy, : .

I, R. F., N. O I. D (): Frogs of Vietnam:a report on new collections. Fieldiana: Zoology, New Series,: iiiiv + .

M, R., U. M, G. V, P. H J. K- (): Amphibians & Reptiles of Mount Kinabalu(North Borneo). Koeltz Scientific Books, Knigstein, pp.

M, U. W. G (): Amphibien & ReptilienSdostasiens. Natur und Tier - Verlag, Mnster, pp.

M, M. S. P (): A new species of Rhacophorus

from eastern ailand (Anura: Rhacophoridae). ZoologicalScience, : .

M, M. F. (): Recherches sur la faune herptologiquedes iles de Borno et de Palawan. Nouvelles archives duMusum dHistoire Naturelle, Srie , : .

M, C. W. W. E. D (): A new species of Hylafrom Cerro Colorado, and other treefrog records and geo-graphical notes from western Panama. American MuseumNovitates, : .

O, A. (): Rhacophorus burmanus (Andersson, ) the valid nomen for Rhacophorus taronensis Smith, andRhacophorus gongshanensis Yang & Su, . Herpetozoa, :.

O, A., O. M, S. S S. G (): Am-

phibian biodiversity of Hoang Lien Nature Reserve (Lao CaiProvince, northern Vietnam) with description of two new spe-cies. Herpetozoa, : .

O, N. L. (): Description of a new species of Rhacopho-rus genus (Amphibia: Anura: Rhacophoridae) from Kon ChaRang Area (Gia Lai Province, Vietnam). Russian Journal ofHerpetology, : .

O, N. L., S. G, T. P K. P- ( ): A new species of Rhacophorusgenus (Am-phibia: Anura: Rhacophoridae: Rhacophorinae) from Kham-mouan Province, Lao PDR. Russian Journal of Herpetology,: .

O, N. L., A. L, R. W. M H T. C. ():Frogs of the familiy Rhacophoridae (Anura: Amphibia) in

the northern Hoang Lien Mountains (Mount Fan Si Pan, SaPa District, Lao Cai Province), Vietnam. Russian Journal ofHerpetology, : .

O, N. L., N. S. N H T. C. (): Description ofa new species and new records of Rhacophorus genus (Am-phibia: Anura: Rhacophoridae) with the review of amphibi-ans and reptiles diversity of Ghu Yang Sin National Park (DacLac Province, Vietnam). Russian Journal of Herpetology, :.

P, W. (): Herpetologische Notizen. Monatsberichte

der Kniglich Preussischen Akademie der Wissenschaen zuBerlin. Aus dem Jahre : .

T, A., P. D A. O (): Note on a collectionof amphibians and reptiles from western Sumatra (Indone-sia), with the description of a new species of the genus Bufo. Zootaxa, : .

K, P. N. (): e Amphibia of the Indo-AustralianArchipelago. E. J. Brill Ltd., Leiden, xii + pp.

W, F. (): Reptilien und Batrachier aus Sumatra, gesam-melt von Herrn Gustav Schneider jr. im Jahre . Zoo-logische Jahrbcher. Abteilung fr Systematik, Geographieund Biologie der iere, : , plates .

W, S. () Revision der Untergattung Rhacophorus (aus-

schlielich der Madagaskar-Formen). e Bulletin of theRaffles Museum, : .

W, J. A., T. T, K. S. L A. K. S (): Anew species of Rhacophorus (Anura: Rhacophoridae) fromMyanmar (Burma). Proceedings of the California Academyof Science, : .

Appendix

Comparative material examined

Rhacophorus angulirostris MALAYSIA: Sabah: GunungTrusmadi (SP , , , , , ); Sungai

Silau-Silau, Headquarters, Kinabalu National Park (ZMB, , ).

Rhacophorus appendiculatus INDONESIA: Indragiri, Su-matra (NHMB [holotype of Rhacophorus phyllopygusWerner, ]). MALAYSIA: Sabah: Taman Bukit Tawau,Tawau District (SP , , ); LinumunsutLake, Maliau, (SP ); Lower Segama, Lahad Datu Dis-trict (SP , , ). Sarawak: GunungMulu National Park, Miri Division (NMBE ). PHILLIPINES: Culion, Calamian Group (SMF); Northeast Mindanao (SMF , ); no locality(ZMB , ).

Rhacophorus baluensis MALAYSIA: Sabah: Headquar-ters, Taman Kinabalu, Ranau District (SP , ); Restau-rant Bayu, near Kinabalu Park, Ranau District (SP ).

Rhacophorus belalongensis BRUNEI DARUSSALAM:Kuala Belalong Field Studies Centre, Temburong District(ZMB [holotype], , [paratypes]; UBDGK -, -, -, -, -, -, G-, -,-, -, -, -, -, -, - [paratypes]).

Rhacophorus cf. bipunctatus INDIA: Travancore, Ker-ala State (ZMB ). MYANMAR: Birma (ZMB ,).

Rhacophorus bimaculatus PHILLIPINES: Agusan Riv-

er, Mindanao (ZMB [syntype]); Dapitan, Mindanao(SMF ).


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Rhacophorus cyanopunctatus BRUNEI DARUSSAL-AM: Kuala Belalong Field Studies Centre, TemburongDistrict (UBD GK-). MALAYSIA: Sabah: East CoastResidency, Kinabatangan District (FMNH ); Sun-gai Tawau, Tawau Hills Park, Tawau District (FMNH). Sarawak: Gunung Mulu National Park, Miri Di-

vision (NMBE ; ZMB ); Nanga Teka-lit Camp, Kapit Division (FMNH , ,, , , ); Labang camp on SungeiSeran, Bintulu Division (FMNH , ); Tubaucamp on Sungei Pesu, Bintulu Division (FMNH ,, , , , ). SINGAPORE:no locality (FMNH ). THAILAND: Khao Sok Na-tional Park, Surat ani (ZMB [holotype]).

Rhacophorus dennysi CHINA: Northern GuangdongProvince (ZMB ); Gao-fung Provinz, Kuangtun[= Guangdong Province] (ZMB ); Pingshiang,Jiangxi Province (ZMB ); no locality (ZMB ,, ).

Rhacophorus depressus INDONESIA: West Java (ZMB [holotype]).

Rhacophorus dugritei CHINA: Batang, Sichuan Province(ZMB , , [syntypes of Rhacopho-rus pleurostictus batangensisVogt, ]).

Rhacophorus dulitensis BRUNEI DARUSSALAM: BatuApoi, Temburong District (UBD ). MALAYSIA: Sabah:Marak Parak, Kota Merudu District (SP , ); Hutan Simpan, Mandamui, Pitas Kudat (SP , ); Taman Bukit-bukit Tawau, Tawau District (SP, ). Sarawak: Gunung Mulu National Park, Miri Di-vision (NMBE ).

Rhacophorus edentulus INDONESIA: Northern Sulawesi(ZMB ).

Rhacophorus everetti MALAYSIA: Sabah: Marai Parai (SP, ); Sayap, Kota Belud District (SP , , ,, ); Headquarters, Kinabalu Park, Ranau District(SP , , , , , , , ,, ); Trusmadi (SP , ). Sarawak: Gu-nung Mulu National Park, Miri Division (NMBE ).

Rhacophorus fasciatus MALAYSIA: Sarawak: Akar Riv-er (BMNH ... = ... [syntypes]); Gu-nung Mulu National Park, Miri Division (NMBE ,; ZMB ); Mount Penrissen (BMNH ...= ... [holotype of Rhacophorus shelfordiBouleng-er, ]).

Rhacophorus gadingensis MALAYSIA: Sarawak: GunungGading National Park, Lundu Division (JMD ); KubahNational Park (JMD ).

Rhacophorus gauni MALAYSIA: Sabah: Sungai Kil-ampun, Purulon Camp, Crocker Range National Park(FMNH ; SP , ); Mendolong Camp, SipitangDistrict (FMNH , , , , ,, ; SP , ); Poring Station, Mt. Kina-balu Park, Ranau District (FMNH ; SP , );

Tawau Hills Park, Tawau District (FMNH , ,; SP , , ); Marak Parak, Kota

Marudu District (FMNH ); Rangkam Kimanis, Pan-tod Besar, Tambunan District (FMNH ); DanumValley Field Centre, Lahad Datu District (FMNH ,, , , , , , ,, , , , , , ,, , , , , , );

Sungai Agathis, Maliau Basin (SP ). Sarawak: Men-giong River, Nanga Tekalit, Kapit Division (FMNH ,, , , , [paratypes], FMNH, , , , , ,, ); Gunung Mulu National Park, MiriDivision (ZMB , NMBE ).

Rhacophorus georgii INDONESIA: Tanke Solokko, Me-kongga Mountains, Southeast Sulawesi (ZMB ).

Rhacophorus harrissoni BRUNEI DARUSALAM: BatuApoi, Temburong District (UBD ). MALAYSIA: Sabah:Lower Segama, Lahad Datu District (SP );Maliau Basin (SP ). Sarawak: Gunung MuluNational Park, Miri Division (NMBE ).

Rhacophorus kajau MALAYSIA: Sarawak: Camp I, m,Gunung Mulu National Park, Miri Division (BMNH. [holotype], . [paratypes]; NMBE); Kubah National Park (JMD ).

Rhacophorus margaritifer INDONESIA: Penglengan,West-Java (SMF ); West Java (ZMB [holotypeof Rhacophorus barbouriAhl, ]); Vulkan Tjisurupan,West-Java (SMF [holotype of Rhacophorus javanusBoettger, ]).

Rhacophorus maximus INDIA: Khassja [= Khasi Hills,Assam] (ZMB ); North India (ZMB ).

Rhacophorus monticola INDONESIA: Southern Sulawesi(SMF ).

Rhacophorus nigropalmatus BRUNEI DARUSALAM:Batu Apoi, Temburong District (UBD ). INDONESIA:Palembang, Sumatra (NMBE ). MALAY-SIA: Sabah: Sungai Stuebing, Trusmadi, Tambunan Dis-trict (SP ); Tawau Hills Park, Tawau District (SP );primary forest (SP ).

Rhacophorus orlovi VIETNAM: Ky Anh-Ke Go, HaTinh Province (ZMB , [paratypes]); KannackTown, Buon Luoi village, Ankhe District, Gia-Lai Province(FMNH ).

Rhacophorus pardalis BRUNEI DARUSALAM: KualaBelalong Field Studies Centre, Temburong District (UBDGK-); without locality (UBD ). INDONESIA: Djapu-ra, Indragiri, Sumatra (NHMB [holotype of Rhacoph-orus pulchellusWerner, ]). MALAYSIA: Sabah: TamanBukit Tawau, Tawau District (SP , ); Danum Val-ley Field Centre, Lahad Datu District (SP ); Pulau TigaNational Park (SP , , ); Kg. Tipasu,Napong , Ranau District (SP ); Mongkopo, RanauDistrict (SP ); Sungai Kokoguan, Marak Parak, KotaMarudu District (SP , ); Mendulong, Sipitang Dis-trict (SP , ); Maliau Basin (SP );Sungai Rompon, Trusmadi (SP ); Pulau Jembongan

(SP ); Lower Segama, Lahad Datu District (SP ,, , , ); Hutan Simpan, Mendamai,


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Pitas Kudat (SP ); no locality (SP , ,); PDC Lembak Inbak, Telupid (SP ); Ma-langkap Tomis, Kinabalu Park, Kota Belud District (SP); Sg. Kimanis, Kg. Kindosodon, TambunanDistrict (SP ). Sarawak: Bako National Park, Kuch-ing Division (NMBE , ); Batang Ai Nation-

al Park, Sri Aman Division (NMBE ); Gu-nung Mulu National Park, Miri Division (NMBE );Kubah National Park, Kuching Division (NMBE ).PHILLIPINES: Palawan (SMF ); Claveria, NorthernLuzon (SMF ).

Rhacophorus penanorum MALAYSIA: Sarawak: GunungMulu National Park, Miri Division (ZMB [holo-type], , [paratopotypes]).

Rhacophorus prominanus MALAYSIA: Pahang: GunungBrinchang, Cameron Highlands (ZMB ); Bukit Fra-ser (ZMB , ).

Rhacophorus reinwardtii INDONESIA: Java (NMBE

, ). MALAYSIA: Sarawak: Batang Ai Na-tional Park, Sri Aman Division (NMBE ).Borneo (SMF [two specimens]).

Rhacophorus rufipes BRUNEI DARUSALAM: Kuala Be-lalong Field Studies Centre, Temburong District (UBDGK-). MALAYSIA: Sarawak: Gunung Mulu NationalPark, Miri Division (NMBE ).

Documents

Taxonomic Status of the Population of Rhacophorus From Sumbar