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Preview Three new species of frogs and a new tadpole from eastern Thailand

Three New Species of Frogs and a New Tadpole from Eastern Thailand Bryan L. Stuart/ Yodchaiy Chuaynkern,^ Tanya Chan-ard,^ and Robert F. Inger* Abstract We describe three new species of frogs from eastern Thailand based on old and new material. These represent a species of Megophrys from Chantaburi and Sa Kaeo Provinces, a species ofOdorrana from Loei Province, and a species ofFejervarya from Ubon Ratchatani Province. Tadpoles are assigned to the new species of Megophrys and Fejervarya and to a recently described species of Rhacophorus from eastern Thailand using molecular identification. Introduction descriptions of tadpoles (Heyer, 1971a,c), and the description of Odorrana indeprensa (Bain & The frog fauna of eastern Thailand has Stuart, 2005). Most recently, Matsui and Panha received sporadic attention. Smith and Kloss (2006) described Rhacophorusjarujini from Ka- (1915) were among the earliest workers and lasin and Roi Et Provinces. reported on a frog collection from coastal In a review on the amphibians of Thailand, Chantaburi Province and neighboring offshore Chan-ard (2003) provided accounts and illustra- islands. Taylor's (1962) monograph on the tions of three unnamed species from eastern amphibians of Thailand was based in part on Thailand, one each of Megophrys Kuhl and van frogs collected from Loei, Nong Khai, Ubon Hasselt, 1822; Odorrana Fei et al. (1991 "1990"); Ratchatani (as "Ubon"), and Chon Buri Pro- and Fejervarya Bolkay, 1915. The new species of vinces. Inger (1970) described Paafasciculispina, Megophrys was first collected more than two andWassersugetai. (1981)described thetadpole decades ago but has since remained unnamed. ofThelodermastellatum Taylor, 1962, both using The new species of Odorrana was originally material from Chantaburi Province. Fieldwork reported as Rana livida (Blyth, 1856) by Taylor in Nakhon Ratchasima Province (primarily at (1962) and has since been hidden under that Sakaerat Experimental Research Station) and name in natural history collections. In 2004, we adjacent Nakhon Nayok Province in 1969-1970 conducted fieldwork and obtained additional resulted in publications on frog calls (Heyer, material of these three undescribed species as 1971b), community ecology of tadpoles (Heyer, well as an unusually colored tadpole belonging 1973, 1974) and adults (Inger & Colwell, 1977), to the genus Rhacophorus Kuhl and van Hasselt, 1822. Tadpoles are assigned to adults of the unnamed Megophrys and Fejervarya and to the ' Field Museum, Department ofZoology, Division ofAmphibians and Reptiles, 1400 South Lake Shore recently described Rhacophorus jarujini Matsui Drive, Chicago, IL 60605-2496, U.S.A. and Panha, 2006, using molecular identification. " National Science Museum, Thailand Natural Herein, we describe the new species ofOdorrana, History Museum, Technopolis, Klong 5, Klong Fejervarya, and Megophrysand the tadpole ofR. Luang, Patumthani 12120, Thailand. jarujini. FIELDIANA: ZOOLOGY, N.S., NO. Ill, DECEMBER 13, 2006, PP. 1-19 NatFiiogn.al1.ParMk;ap3 =ofPlhoucalSirtiieTsarnefWeirlrdeldifteoSiannctthueartye;xt.4 1==PhPuhPuhaLuNaanmgtiWpildNloinf-ehuSnantcitnugarAyr;ea2; 5==PhPuhuKrJaodnuge-Nnag Yoi National Park; 6 = Pang Si Da National Park; 7 = Khao Soi Dao Wildlife Sanctuary. Fieldwork was conducted at locaHties 1, 5, and 6 in this study. Materials and Methods These three protected areas are situated in the uplands at the northwestern, southeastern, and Fieldwork southwestern margins, respectively, of the Khorat Basin in eastern Thailand. In September 2004, two of us (YC, BLS) Phu Luang Wildlife Sanctuary was sampled at conducted fieldwork under the auspices of the 850-1460 m elevation in hill evergreen, bamboo Thailand Natural History Museum at Phu mixed with evergreen, and rhododendron heath m Luang Wildlife Sanctuary in Loei Province, forests. The last was encountered at 1460 Phu Jong-Na Yoi National Park in Ubon elevation on the summit of Kok Nok Kraba, Ratchatani Province, and Pang Si Da National where weather conditions were cool and foggy Park in Sa Kaeo Province, Thailand (Fig. 1). duringourvisit. PhuJong-NaYoiNational Park FIELDIANA: ZOOLOGY was sampled at 160^00 m elevation in decidu- proximal edge of inner metatarsal tubercle to ous dipterocarp forest with grassy understory tip offourth toe. (often with large areas of exposed igneous bedrock), hill evergreen forest, and disturbed DNA Extraction and Sequencing evergreen forest. Pang Si Da National Park was m DNA sampled at 90-600 elevation in hill evergreen Total genomic was extracted from forest, evergreen mixed with bamboo forest, adults and tadpoles using PureGene Animal DNA disturbed evergreen forest, disturbed vegetation Tissue Isolation Protocol (Centra Systems, along abandoned roads, and around buildings. Inc.). A fragment of mitochondrial DNA that Specimens were caught in the field by hand, encodes part of the 16S ribosomal RNA gene preserved in 10% buffered formalin, and later (16S) was amplified from extractions of Me- transferred to 70% ethanol. Tissue samples were gophrys and Fejervarya by the polymerase chain 2ta0k%enDfMrSomOfrsoaglst-bsaytuprraetseedrvsitnograpgieecbeusffoefrlbiveefrorien rfoerac3f5iocnyc(lPesCRu;si9ng4tChe4p5sr,im6er0sCL-301s6,SR7a2nCaII1Imi(5n')- the specimen was fixed in formalin. Tissue GAGTTATTCAAATTAGGCACAGC-3') and samples were taken from tadpoles by preserving H-16SRanaIII (5'-CATGGGGTCTTCTCGT- one or two representatives from a tadpole lot in CTTAT-3'). A fragment of mitochondrial 20% DMSO salt-saturated storage buffer. DNA that encodes part of the cytochrome c Specimens are housed in the Field Museum of oxidase subunit III gene, the complete tRNA Natural History (fmnh) and the Thailand glycine, the complete NADH dehydrogenase Natural History Museum (thnhm). Most speci- subunit 3 gene, and part of the tRNA arginine mens are cross-cataloged at both institutions, in (ND3) was amplified from extractions of Rha- which case voucher numbers are reported as cophorus by PCR (94C 45s, 49 C 30s, 72"C FMNH/THNHM. The locatious of holotypes are 1 min) for 35 cycles using the primer pair L- explicitly stated. The locations ofparatypes and COXIII (5'-CCGCATGATACTGACACTT-3') additional material can be obtained from the and Arg-HND3III (5'-AACTGTCTTTTTTG- Field Museum ofNatural History. GACTAGC-3'). PCR products were electro- phoresed in a 1% low-melt agarose TALE gel Morphology stained with ethidium bromide and visualized under ultraviolet light. The bands containing Comparative material was examined in the DNA were excised, and agarose was digested holdings of fmnh, thnhm, California Academy from bands using GELase (Epicentre Technolo- aofndScUineincveesrs(ictays)o,fRCoaylailfoOrnntiaariMousMeuusmeuomf(Vreormt)e,- dgiierse)c.tiPonCsRbypdriordeuctctdsoubwleer-estsreaqnudecnyccelde sienqubeontch- brate Zoology (mvz). Data for Megophrys ing using Big Dye version 3 chemistry (Perkin jingdongensis were taken from Fei et al. (1983), Elmer) and the amplifying primers. Cycle se- for M. pachyproctus from Huang et al. (1998), quencing products were precipitated with etha- and for syntype males of Odorrana graminea nol, 3 M sodium acetate, and 125 mM EDTA from Boulenger (1920). and sequenced with a 3730 DNA Analyzer t=heMeneaasruersetm0e.n1tmsmw.erAebHbmrDaedLveia=wtiitohnsdiuaslecdalairpeerSsVtLo (SAeBqIu)e.ncSheeqruevnecressiowner4e.1edi(tGeedneacnoddeasl)ignaenddwidteh- snout-vent length; head lenHgtDhWfro=m posited in GenBank (accession numbers mtiapxiofmusmnouhteadto wriedatrh;ofSNtThe =jawss;nout length DQ860092-DQ860097). from tip of snout to the anterior corner of the etyhee;eEyeYbaEll;=ldOiaDme=ternaorfrotwheesetxppooisnetd opforitnitoenroor-f Species Accounts TMP = bital distanceT;EY = horizontal diameter of Family Megophryidae tympanum; tympanum-eye distance Genus Megophrys Kuhl and van Hasselt, 1822 cforronmeraontfetrhieoreyee;dgSeHoKf =tysmhpaannkulmengttoh;pTosGteHrio=r Megophrys lekagulisp. nov. thigh length, from vent to outer edge of knee; Megophrys sp. Chan-ard, 2003: 86. HND = hand length, from base ofpalm to tip of HoLOTYPE FMNH 213946 (field tag DLD Finger III; and FTL = foot length, from 996.1), deposited at fmnh, adult female (Fig. 2), STUART ET AL.: THREE NEW SPECIES OF FROGS FROM THAILAND Fig. 2. The adult female holotype (fmnh 213946) of Megophrys lekaguli sp. now in preservative. Clockwise from upper left: palmar view ofhand; plantar view offoot: lateral view ofhead; dorsal view; ventral view. collected in Khao Soi Dao Wildlife Sanctuary. Da National Park, Muang Sa Kaeo District, Chantaburi Province. Thailand. 600-700 m ele- Sa Kaeo Province, Thailand, 14 07'39.6"N vation,on26September 1979byDoyleDamman. 102 15'33.1"E, 600 m elevation, on 23-24 Sep- P.AR.ATYPES FMNH 213947 (onc adult male). tember2004 by Yodchaiy Chuaynkern, Bryan L. FMNH 213948 (one adult female), same data as Stuart, Chatchay Chuechat, and Sunchai Mak- holotype. fmnh 191456 (one adult male), fmnh chai. FMNH 265958/thnhm 05321. fmnh 265959/ 191457 (one adult female), collected in Pong thnhm 05322 (two adult males), collected in Nam Ron. Khao Soi Dao Tai Mountain, Huay Km 29 Stream. Pang Si Da National Park, Chantaburi Province. Thailand. 1000 m eleva- Muang Sa Kaeo District, Sa Kaeo Province, tion, on 28 November 1971. by Sukhum Pong- Thailand, 14 06'20.7"N 102 15'41.6"E, 600 m sapipatana. thnhm 1101 (one adult male). elevation, on26-27 September2004byYodchaiy thnhm 1034 (one adult female), collected in Chuaynkern, Bryan L. Stuart, Chatchay Chue- Khao Sabab Mountain, Namtok Phliu National chat. and Sunchai Makchai (Fig. 3). Park, Khlung District, Chantaburi Province, Other Material fmnh266341/thnhm05391, Km Thailand, on 13 July-01 August. 2001, by tadpoles (Figs. 4-5), collected in Huay 29 Narongrit Sukprakarn. fmnh 265955/thnhm Stream, Pang Si Da National Park, Muang Sa 05318, fmnh 265956/thnhm 05319, fmnh Kaeo District, Sa Kaeo Province, Thailand, 265957/thnhm 05320 (three adult males), collect- 14 06'20.7"N 102 15'41.6"E, 600 m elevation, on ed in Huay Kong Mou Now Stream, Pang Si 21 September 2004 by Yodchaiy Chuaynkern, FIELDIANA: ZOOLOGY jnr.ohfiqacispniSe:anslril ddge of dia Mil il doRsri a lores SS mthhofi body ataam ljw|ii insUt^ 61 Ac c^c to sfcia of tcnponl icgiow. ivaKnae locih oa two shiA tobodfaxiSp( tocachoiha; Tlpsoffi IV<n<I<III;l HKtMy tahudcs^an m Tipsoftoesniaa Toe loaeeWrthwaaAToeV: toes with laiKmim ofwdn Toe fieeofwcbfai^] s|i.BOS',ailac. of Toe IV; Bryaa L. Stmit, Oalchav^ rhrhat, nd lalEfLV, bat ao sabarticBlv tJuwhs: Soncliai Makciai, fmnh 2663^fiii9iBii QS397 (SKltaatedcpoaonlveDsiXFs,turcigcotl.SBdSDeadaKiNaatitBHiOaoanPayrloKvFitoneck^e..MMTnohaaaifilNsaonSndar. wtHohacknaagklgmhofolmfeaoTDdodeeantIa;pea^oa aa^cstoAebodljF; 14=0r39.(rN IQT15'33.1"E.600mekvatiQa.cm 49% 23 Sqptanbcr 20O4 hf Yodchaiy Chaaynkem, topofkad Bkyan L. Stmt, ChaichajF n>hii aod back wiA SnndiaiMakdiaL A DiAcaMKB medium-siipd species of 3e~ ^qphjj haTine males with S\X 56.6-66.6. females with SVL 71Jt-9C0^ slniq^ jMujeOi^e SDOol; m> <lrinul pragectiOD on lulimn; wcaUy visililepalpehrritnlicwfc';vomometecdiontwo diatrii^BStnusvcnetothebod^axis;wide dnk irticalbarbdoweve: ooi^colMiedannrIBp Stripe: narrow, cilriiMil, icttLal fine of c^cMdaikblownwitiha^^dtareainils< <fetiDcttym|MwnmorethanhalfofcyedumiHk relative finger kqeOs IV < II< I < ID: toes of th^ webbedonlyatbaseandhdMngdenn pectoralBaad' DEsaumoN of IfcunYtE lUbihB lal^aad, faasBr ateiy stocky: had sli^hlly widerthan long, aot : ifA^^doserto1 wider than trunk: snout obtuselv pointed in Cma^ ofhokrcpe: S\X 71JK; doisal view, stnm^ pwyrtine beyond lower HDL343:HDW36*S!^a S^EYE7.5:lOD Fk>.4. Lateral<iem^ofi^potefraaattHUiawMMaSl)efMi4if*jI Mi^ai^a^ STUARTETAL: THREE NE^'SPECIESOF FROGS FROMTHAILAND Fig. 5. Mouthpartsoftadpole(fmnh266341/thnhm05391)o{Megophryslekagulisp. nov. Scalebar = 1 mm. 8.2; TMP 4.6; TEY 5.4: SHK 35.5; TGH 35.3; Descriptionof Tadpole Body slender, oval, HND 19.5; FTL 32.6. flattened above; tail about as deep as body, Color of Holotype in Preservative Dor- dorsal fin arisingbehind origin oftail, maximum sum brown to grayish brown; dark brown band depth near mid-length, tapering gradually to between eyes; narrow, external, vertical face of narrow, rounded tip; tail 2.1-2.2 times body upper eyelid dark brown with a light area in its length, tail depth 20-23% of tail length. Maxi- center; dark hourglass-shaped marking on back; mum body width 50-54% of body length, body side of head light brown, with a wide, vertical, depth 40-42% ofbody length. Eyes dorsolateral, dark bar below eye, a narrow dark bar at lip pointing laterally. Nares open, dorsolateral, anterior to eye, and a narrow dark bar at lip slightly closer to eyes than to oral fuimel. rim posterior to eye, all separated by light areas on not raised; intemarial wider than interorbital. the margin of the upper lip; tympanum and Spiracleclosertoeyethantoendofbody,midway adjacent area dark brown; laterally body light up side, tube not free of body wall. Anal tube brown with dark spots ventrolaterally; throat ending at margin ofventral fin, opening medial. and chest brown with irregular darker markings, Oral disk terminal, lips expanded and directed becoming lighter posteriorly, creamy-white near upwardly into typical Megophrys funnel; trans- groin; limbs with narrow dark brown crossbars; verse width of expanded funnel 75% of body lower half of posterior surface of thigh brown, length. with a dark brown area beginning at vent and Colorinpreservativeofbodylight grayabove, extending distally; ventral surface of thigh and sides andventerwith small black spots; proximal ventral surface ofshank dusky brown; posterior halfofcaudal muscle with two or three irregular surface of shank with three large black spots; dark streaks, distal half dusky, mid-dorsal edge ventral surface of tarsus and foot uniform ofmuscle light; fins distinctly pigmented only in purplish brown. distal portions. Color of Paratype Male fmnh 265955 Body lengths (stage = mm): 25 = 9.0, 10.4; 37 IN Life Dorsum light grayish brown with dark = 12.r. 12.9; 38 = 13.8; 42 = 14.2. brown markings; upper eyelid, supratympanic Variation Measurements ofparatypes sum- fold, and flank with yellowish wash; upper lip, marized in Table 1. Chantaburi males (SVL inguinal region, anterior and posteriorsurface of 63.6-66.6. mean S.D. 64.9 1.5. N = 3) are thigh, and posterior surface of shank with largerthan Sa Kaeo males (SVL 56.6-58.8, mean salmon wash; dorsal surface of fingers and toes S.D. 57.8 0.9. N = 5). Males with dark orange with black spots; ventral surface ofhand brown nuptial pad covering most of the dorsal and foot purplish gray; throat and chest dark surface ofFinger I and a smaller, round area on gray with salmon wash; posterior part of belly the dorsal surface ofFinger IL Four males have pinkish white; pectoral and femoral glands small, dark asperities on the upperand lower lip. creamy-white; forelimb, knee, posterior surface concentrated anteriorly. Ova in preservative mm ofshank, and dorsal surface oftarsus with black approximately 2 diameter, uniformly light spots; eye orange with black reticulations and yellow, without pigmented hemisphere. Dorsum black, vertical pupil. in preservative light brown to dark gray. In Molecular Results A tadpole (fmnh preservative, the dark hourglass-shaped marking 266341/thnhm 05391) and an adult male para- on the back and the dark band between the eyes type (fmnh 265959/thnhm 05322) are identical in is more conspicuous in some paratypes than in a 704 bp fragment of 16S. the holotype. The dark brown band is an FIELDIANA: ZOOLOGY Table 1. Measurements (mm) ofMegophrys lekagulisp. nov. Abbreviations defined in the text. Measurement Fig. 6. The adult male holotype (fmnh 265925/thnhm 05341) of Odorrana aureola sp. nov. in preservative. Clockwise from upper left: palmar view of hand; plantar view of foot; lateral view of head; dorsal view; ventral view. boulders and rock outcrops above a swift, rocky Megophrys as more closely related to Ophryo- stream with waterfalls in hilly evergreen mixed phryne than to Xenophrys. However, their with bamboo forest and on leaf litter next to analysis was based only on a single species of a large tree root network 20 m from a swift, Megophrys and Xenophrys, and we feel that this rocky stream in disturbed vegetation next to an issue remains unresolved. We therefore leave the abandoned road through hilly evergreen forest. new species in the genus Megophrys, pending Tadpoles were collected at night (2000-2015 h) a phylogenetic analysis with improved taxonom- swimming at the water surface in a 3-m-wide ic sampling. swift, rocky stream in hilly evergreen mixed with bamboo forest and in a shallow stream pool with Family Ranidae silt substrate in disturbed vegetation next to an Genus Odorrana Fei et al. (1991 "1990") AalblanfidvoenoefdthreoaSdatKhareoougahduhlitllmyaelveesrgwreereencaflolriensgt.. Odorrana aureola sp. nov. Remarks The two species that most closely Rana livida (part) Taylor, 1962: 468; Inger and resemble the new species, M. auralensis and M. Chan-ard, 1997: 65. major, were placed in the genus Xenophrys Rana sp. Chan-ard, 2003: 138. Giinther, 1864 by KhMonsue and Thirakhupt Odorrana cf livida Stuart, Inger and Voris, 2006: (2001; M. major as lateralis) and Frost fig. 2. (2004) without supporting argument. Frost et Holotype fmnh 265925/thnhm 05341 (field HKV al. (2006) argued for recognition of Xenophrys tag 66220), deposited atthnhm, adult male because their phylogenetic analysis recovered (Fig. 6), collected on a large boulder on the bank FIELDIANA: ZOOLOGY of Toe V; posterior surface of forearm, ventral halfoffiank, anterior surface ofthigh, posterior surface ofthigh, posterior surface ofshank, and dorsal surface of foot black with distinct, vermiform, bright yellow (creamy-white in pre- servative) markings; upper lip stripe yellow (creamy-white in preservative); males with gular pouches; and unpigmented eggs. Description of Holotype Habitus moder- ately slender; head narrow, longer than wide; snout obtusely pointed in dorsal view, projecting beyond lower jaw, round in profile, slightly depressed; nostril lateral, slightly closer to tip of snout than eye; canthus distinct, slightly constricted behind nostrils; lores concave and Fig. 7. An adult female paratype (fmnh 265919/ oblique; eye diameter 81% snout length; in- THNHM 05346) of Odonana aureola sp. nov. in life. terorbital distance less than width of upper eyelid; pineal body visible; distinct, round of Sung Mak Hing Stream above Tad Loei tympanum, 64% the eye diameter, not depressed Waterfall in hilly evergreen forest in Phu Luang relative to skin oftemporal region, tympanic rim Wildlife Sanctuary, Phu Rua District, Loei elevated relative to tympanum; vomerine teeth Province, Thmailand, 1715'32.4"N 10L'30'22.8"E, on two slightly oblique ridges, closer to each 1100-1300 elevation, on 03 September 2004 by otherthan tochoanae; tongue notched; vocal sac Yodchaiy Chuaynkern, Bryan L. Stuart, opening at corner ofmouth; sac-like gularpouch Chatchay Chuechat, and Sunchai Makchai. at corner ofthroat. Paratypes FMNH 265924/thnhm 05351, fmnh Tips ofall four fingers expanded with circum- 265926/thnhm 05352 (two females), fmnh marginal grooves; width ofFinger III disc about 265923/thnhm 05350, fmnh 265927/thnhm 2 times the width of phalanx, 42% the diameter 05353 (two immature females), same data as oftympanum; relative finger lengths II < I < IV holotype. fmnh 265919/thnhm 05346 (Fig. 7), < III; ventral callous pad on Fingers II, III, and FMNH 265920/thnhm 05347, fmnh 265921/thnhm IV from distal edge of proximal subarticular 05348, fmnh 265922/thnhm 05349 (four fe- tubercle to base ofdisc; movable fiap ofskin on males), collected at Kok Nok Kraba, Phu Luang preaxial side of Fingers II and III; one sub- Wildlife Sanctuary, Phu Rua District, Loei articular tubercle on Fingers I and II, two Province, Thailand, 1716'48.7"N 10r3r07.5"E, subarticular tubercles on Fingers III and IV; m 1460 elevation, by Yodchaiy Chuaynkern, one supernumerary tubercle proximal to proxi- Bryan L. Stuart, Chatchay Chuechat, and mal subarticular tubercle on Fingers II, III, and Sunchai Makchai. fmnh 264541, thnhm 05721 IV; two palmar tubercles at base of III and IV, (two males), fmnh 264542, thnhm 05720 (two large, oval, in contact; velvety nuptial pad on females), collected near Lon Moei in Phu Luang Finger I, covering the dorsal surface to the level Wildlife Sanctuary, Phu Rua District, Loei of the distal edge of the subarticular tubercle, m Province, Thailand, 1400-1500 elevation, in covering themedial surfaceto baseoffingerdisc; September-October 1998 by Tanya Chan-ard. forearm robust. FMNH 172378-80 (three males), collected at Phu Tips of toes expanded, width of Toe IV disc Kading (=Phu Kradueng National Park), Loei slightly less than width ofFinger III disc; Toe III Province, Thailand on 14 March 1958 by Edward shorter than Toe V; all toes fully webbed to base H. Taylor. ofdiscs, notch in webbing between Toes IV and Diagnosis A species of Odonana having all Vextends to second subarticulartubercleon Toe digittips expanded withcircummarginal grooves; IV; movableflapofskin on preaxial sideofToe I first finger longer than second; smooth skin; from subarticular tubercle to base of disc, on males with SVL 63.6-67.4, females with SVL postaxial side ofToe V from slightly proximal of 87.1-96.5; males sometimes with white spinules proximal subarticular tubercle to base of disc; on temporal region, tympanic rim, rictal gland, elongate, oval innermetatarsal tubercle; noouter upper and lower lip near gape, and postaxial side metatarsal tubercle. STUART ET AL.: THREE NEW SPECIES OF FROGS FROM THAILAND Table 2. Measurements (mm) ofOdorrana aureola sp. nov. Abbreviations defined in the text. Measurement Adult males (holotype and paratypes) Adult females (paratypes) Range; mean t S.D. (N = 6) Range; mean S.D. (N = 7) SVL 63.6-67.4 65.3 1.6 87.1-96.5; 92.4 3.1 HDL 22.9-25.5 24.3 1.1 31.7-33.8; 32.6 0.8 HDW 21.0-22.9 22.1 0.8 29.3-31.8; 30.8 1.0 SNT 10.2-11.3 10.6 0.4 14.0-16.1; 15.2 0.7 EYE 7.9-9.0 8.4 0.4 10.0-11.4; 10.3 0.5 lOD 4.1-5.0 4.8 0.4 7.5-9.5; 8.6 0.6 TMP 5.3-5.5 5.4 0.1 5.8-6.7; 6.2 0.3 TEY 1.6-2.0 1.9 0.2 2.2-3.4; 3.0 0.4 SHK 40.0^3.7 41.3 1.4 55.5-61.6; 59.5 2.4 TGH 35.0^0.0 37.6 2.0 50.7-56.0; 53.7 2.2 HND 17.4-18.7 18.4 0.5 22.6-27.0; 25.8 1.6 FTL 33.9-36.5 35.5 1.0 47.0-53.6; 51.4 2.5 Range; median (N - 6) Range; median (N = 7) HDL:HDW 1.04-1.21; 1.08 1.02-1.09; 1.07 SNT:HDL 0.42-0.46; 0.44 0.43-0.48; 0.47 TMP:EYE 0.61-0.70; 0.65 0.57-0.63; 0.60 EYE:SNT 0.75-0.87; 0.77 0.62-0.75; 0.66 SHK:SVL 0.61-0.66; 0.63 0.60-0.67; 0.65 Skin smoothonall surfacesexceptgranularon creamy-white; upper lip stripe creamy-white; venteroposterior surface of thigh; few small rictal gland creamy-white; toe webbing brown. tubercles in temporal and tympanic regions; no Variation Measurements ofparatypes sum- humeral gland; weak supratympanic fold from marized in Table 2. SVL ofadult males 66-77% rear ofeye to rear oftympanum, no dorsolateral SVL ofadult females. Ova in preservative appro- mm fold; single,elongaterictalglandcontinuouswith ximately2 diameter, uniformlycreamy-white, upper lip, extending to axilla; scattered, white without pigmented hemisphere. Some specimens spinules on temporal region, tympanic rim, rictal with Toe IV webbed only to distal subarticular gland, upper and lower lip near gape, and tubercle, with narrow extension to base of disc. postaxial side ofToe V. Somespecimenswithdividedrictalgland. Females Measurements (mm) of holotype: SVL 65.4; with interorbital distance greater than width of HDL25.5; HDW 21.0; SNT 10.6; EYE 8.6; lOD upper eyelid. Females without robust forearm. 5.0; TMP 5.5; TEY 1.9; SHK 40.3; TGH 37.8; Females without whitish spinules or with few HND 18.3; FTL 34.9. whitish spinules only on loreal region and Colorof Holotypein Life Dorsum uniform tympanic region. Two males without whitish green; side of head to dorsal half of flank dark spinules. Females in life with coppery-brown brown; posterior surface offorearm, ventral half dorsum, three females with small bright green of flank, anterior surface of thigh, posterior spotsonanteriorhalfofdorsum,fivefemaleswith surface ofthigh, posterior surface ofshank, and three to ten round black spots on dorsum. dorsal surface of foot black with distinct, Comparisons Six other species of Odorrana vermiform, bright yellow markings; upper sur- have the combination of first finger longer than faces oflimbs brownish olive with brown cross- the second, all digit tips expanded with circum- bars; upper lip stripe yellow, lower lip black; marginal grooves, smooth skin on dorsum, light- nuptial pad gray, rictal gland yellow; throat with coloredupperlipstripe,maleswithgularpouches, brownish wash, gular pouch black, belly white; males without humeral glands, and females with toe webbing purplish brown. unpigmented eggs: Odorrana banaorum (Bain, Color of Holotype in Preservative Dor- Lathrop, Murphy, Orlov and Ho, 2003), O. sum bluish gray; markings on posterior surface chloronota (Giinther, 1875), O. graminea (Bou- offorearm, ventral halfofflank, anteriorsurface lenger, 1899), O. indeprensa(Bain&Stuart,2005), of thigh, posterior surface of thigh, posterior O. livida (Blyth, 1856), and O. morafkai (Bain, surface of shank, and dorsal surface of foot Lathrop, Murphy, Orlov and Ho, 2003). Odor- 10 FIELDIANA: ZOOLOGY

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