Russian Journal of Herpetology

Vol. 14, No. 1, 2007, pp. 65 72

A NEW SPECIES OF Theloderma (ANURA: RHACOPHORIDAE) FROM SOUTHERN THAILAND AND PENINSULAR MALAYSIA David S. McLeod1* and Norhayati Ahmad2
Submited September 19, 2006. A new rhacophorid frog tentatively assigned to the genus Theloderma is described on the basis of five specimens collected from southern Thailand and peninsular Malaysia. The new species is most similar to Theloderma asperum in external morphology and coloration, and differs from this and all other congeners by body size, characteristics of skin tuberculation, digital characters, and coloration. This species is the fourth Theloderma described from Malaysia and the third from Thailand. Keywords: New species, peninsular Malaysia, Rhacophoridae, Thailand, Theloderma.

INTRODUCTION Theloderma is a genus of rhacophorid frogs that has rough, tuberculate skin, and Y-shaped terminal phalanges (Liem, 1970; Manthey and Grossmann, 1997). Subsequent to the original description of Theloderma by Tschudi (Tschudi, 1838), members now recognized in this genus were placed in the genera Rhacophorus and Philautus until Taylor (1962) resurrected Theloderma and expanded its content to include those rhacophorids having a distinct tympanum and small numbers of large eggs (8) laid above water-filled cavities in trees. Liem (1970) recognized Theloderma based on the presence of numerous calcified warts on the dorsum, two slips of the m. extensor digitorum communis longus in the foot, and the reproductive behavior reported by Taylor (1962). In their re-analysis of Liem (1970) and Channing (1989), Wilkinson and Drewes (2000) found no synapomorphy for Theloderma. The monophyly of Theloderma remains in question until a thorough re-assessment of the relationships among rhacophorid genera is completed. Therefore, the generic allocation of the new species described herein is tentative. Currently, 13 species of Theloderma are distributed from Sri Lanka and northeastern India to Myanmar and
Division of Herpetology, Natural History Museum and Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas, 66045-7651, USA; E-mail: dsmcleod@ku.edu. 2 Faculty of Science and Technology, 43600 Bangi, University Kebangsaan Malaysia, Malaysia; Email: noryati@pkrisc.cc.ukm.my. * Author to whom correspondence should be addressed.
1

southern China, through Indochina to Malaysia and Sumatra (Manthey and Grossmann, 1997; Inger et al., 1999; Iskandar and Colijin, 2000; Frost, 2004; Orlov et al., 2006; Ohler et al., 2002). A recent summary of the known species and their current distributions is provided by Orlov et al. (2006). Five species of Theloderma occur in Thailand and Malaysia (T. asperum, T. gordoni, T. horridum, T. stellatum, T. leporosum); of these, only three (T. asperum, T. horridum, T. leporosum) occur on the Thai-Malay Peninsula (Berry, 1975; Chan-Ard, 2003). In 1997, Norsham Yakob (Forest Research Institute Malaysia) collected a distinctive, white rhacophorid frog at a field site in peninsular Malaysia. The authors independently collected similar specimens during the course of their respective field programs in Thailand and Malaysia. The specimens that we collected have Yshaped terminal phalanges and tuberculate skin typical of Theloderma (sensu Liem, 1970; Manthey and Grossmann, 1997), but are distinguished from other members of this genus by a combination of several features of the adult morphology. Owing to these morphological distinctions, the new species is described herein. MATERIAL AND METHODS Frogs were collected opportunistically in the field by hand. The Thai and Taman Negara specimens were euthanized in the field by immersion in dilute chlorobutanol, fixed in 10% buffered formaldehyde, and preserved in 70% ethanol. Tissue samples were taken prior to fixation and stored in 95% ethanol. The Taman Nega-

1026-2296/2007/1401-0065 2007 Folium Publishing Company

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David S. McLeod and Norhayati Ahmad MBE, mandible-back of eye distance; MFE, mandiblefront of eye distance; MN, mandible-nostril distance; NS, nostril-snout distance; PAL, palm length; SVL, snout-vent length; TAD, tympanum-back of eye distance; TBL, shank (tibial) length; TND, tympanum-nostril distance; TPD, tympanum-front of eye distance; TYE, tympanum diameter; UAW, upper arm length; UEW, upper eyelid width; NM, no measurement taken; FLL, forelimb length (PAL + LAL + UAW); HLL, hind limb length (FOL + TBL + FEL). Digital webbing formulae follows that of Savage and Heyer (1997). Digits of the hand are numbered IIV based on homology (Alberch and Gale, 1985). Radiographs were made of specimens to examine osteological characters. Sex was determined by examination of gonads, vocal slits, and the presence of nuptial pads in males. The paratype UKM MZ 1245 was cleared and double stained using a modified protocol based on (Taylor and Van Dyke, 1985). All illustrations were prepared by DSM with the aid of a binocular dissecting microscope and camera lucida. Specimens were deposited in the Raffles Museum of Biodiversity Research (ZRC), Forest Research Institute Malaysia (FRIM), University Kebangsaan Malaysia Museum Zoology (UKM MZ), and the National Science Museum of Thailand (THNSM). Comparative materials were also examined from the holdings of the Field Museum of Natural History (FMNH). SYSTEMATICS Theloderma licin sp. nov. (Figs. 1 4; Table 1) Holotype. FRIM 0967, adult male, collected at the Taman Negara Resort, Kuala Tahan Taman Negara (034012 N 1024434 E) Pahang State, Peninsular Malaysia, 82 m elevation, on 15 Oct. 2003 by Dennis Yong. Paratypes. ZRC.1.11400 (formerly, UKM MZ 2401), adult female, collected at Ulu Muda Forest Reserve (5500 N 100550 E), Kedah State, Peninsular Malaysia, 385 m elevation, on 25 Oct. 2001 by Norhayati Ahmad. ZRC.1.10535, adult male, collected at Lakum Forest Reserve (33712 N 102022 E) Pahang State, Peninsular Malaysia, 72 m elevation, on 17 March 2003 by Lim Ching Chiz. UKM MZ 1245, adult male, collected at the Sungai Lasor base camp (06037 N 1005727 E), Ulu Muda Forest Reserve, Kedah State, Peninsular Malaysia, 193 m elevation, on 25 March 2003 by Manohar Mariapan. THNSM 01468, gravid adult female, obtained in secondary forest at Khao Lu-

5.0 mm

5.0 mm

Fig. 1. Dorsal (a) and lateral (b) views of head of Theloderma licin sp. nov. holotype (FRIM 0967).

ra specimen was kept alive in captivity for 1 week prior to euthanasia. Three specimens were placed directly into 70% ethanol after euthanasia. Measurements were made on preserved specimens with digital calipers to the nearest 0.01 mm and rounded to 0.1 mm. Measurements of the holotype of Theloderma leporosum were made from high-resolution scaled digital photographs provided by the National Museum of Natural History (Leiden, Netherlands). Abbreviations and measurements used follow those of ManamendraArachchi and Pethiyagoda (2005): DL, length of disc of third finger; DW, width of disc of third finger; ED, eye diameter; EN, eye to nostril distance; ES, eye to snout distance; FEL, thigh (femur) length; FOL, foot length; HL, head length; HW, head width; IN, internarial distance; IO, interorbital width; LAL, lower arm length;

A New Species of Theloderma

67

a b

1.0 mm 0.5 mm

1.0 mm

1.0 mm

Fig. 2. Hands and feet of Theloderma licin sp. nov. (a) Terminal phalanx, male, UKM MZ 1245; (b) dorsal view of hand, holotype FRIM 0967; (c) palmar view of hand, THNSM 01468; (d) plantar view of foot, THNSM 01468.

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David S. McLeod and Norhayati Ahmad dorsum and lateral parts of body and limbs with numerous fine, pearly-tipped tubercles; ventral skin weakly granular; (3) pupil horizontal; (4) tympanum distinctly visible, free of tubercles, 58 89% of ED; (5) supratympanic fold reaching angle of jaws, tubercles absent on fold; (6) canthus rostralis prominent, transverse sectional profile rounded; loreal region slightly concave to flat; (7) fingers with lateral dermal fringes and basal webbing (II222III23IV2-32V) (8) three oval metacarpal tubercles, inner (thenar) and median large, outer small; (9) toes fully webbed (I11 2II12III112IV211V); (10) prominent oval inner metatarsal tubercle; no outer tubercle apparent; (11) terminal phalanges distinctly Y-shaped; (12) digital tips dilated, wider than long, 45 80% of TYE; (13) vomerine teeth absent; (14) choanae oval; (15) tongue acutely triangular, narrow at attachment, lacking medial papillae; (16) males with exposed nuptial pad on Digit II, vocal slits present; (17) in life, dorsum pale white-brown with dark brown spots in inguinal and axial regions; dark brown bar extending posteriorly from snout to supratympanic fold; ventral surface of body and limbs white with brown reticulations; chin brown with small white spots; iris red. Description of the holotype. Habitus moderately slender; head width about equal to head length; snout obtusely pointed in dorsal view (Fig. 1a), rounded in ventral view, projecting slightly beyond lower jaw in lateral view (Fig. 1b); nostril oriented dorsolaterally, closer to tip of snout than to eye; internarial space dorsally concave, narrower than interorbital distance (52 71% IO); canthus rostralis prominent and rounded in section; loreal region slightly concave to flat; pupil round; interorbital space dorsally convex; tympanum distinct, round, free of dermal tubercles, smaller than eye (58 89%

ang National Park (84231 N 994149 E) Nakhon Si Thamarat, Thailand, approximately 400 m elevation, on 17 July 2003 by David S. McLeod. Etymology. The specific name licin is derived from the Malay word for smooth in reference to the skin texture of this peculiar species of the genus Theloderma. Diagnosis. Theloderma licin is distinguished from members the genus Philautus by the presence of Yshaped terminal phalanges (simple or weakly-bifurcate in Philautus), and from members of the genus Nictixalus by the absence of co-ossification of head skin to the skull (co-ossified in Nictixalus) (Liem, 1970). Theloderma licin is characterized by the following combination of characters: (1) SVL of adult males 28.0 30.1 mm, females 27.5 31.2 mm; (2) skin of

Fig. 3. Holotype of Theloderma licin sp. nov. (FRIM 0967). Photo by Norsham Yaakob.

Fig. 4. Paratype of Theloderma licin sp. nov. from Thailand (THNSM 01468) illustrating individual variation in color pattern between pre-capture (a) and post-capture (b) events. Photo b by J. Murphy.

A New Species of Theloderma ED); supratympanic fold present, extending almost to insertion of arm; vomerine teeth absent; tongue acutely subtriangular, narrow at point of attachment, deeply notched behind; vocal slits at corners of mouth (beyond edge of tongue); choanae oval, parallel with maxilla, widely separated from each other. Fingers with Y-shaped terminal phalanges (Fig. 2a); discs of fingers semicircular, wider than long, bearing circummarginal grooves (Fig. 2c); ventral surface of each disc with distally oriented, ridged sub triangular pad; relative finger lengths: II < III < V < IV; inner metacarpal tubercle large, oval; median metacarpal tubercle large, round; outer metacarpal tubercle small, oval; subarticular tubercles prominent, round, one large tubercle on Fingers II V, one small proximal tubercle on Fingers III IV; well developed white nuptial pad on dorsal and lateral portions of Finger II; lateral dermal fringes on Fingers III IV, webbing formula: II2 2III23IV22V (Fig. 2b).
TABLE 1. Measurements of Theloderma licin Character DBE DFE DL DW ED EN ES FEL FOL HL HW IN IO LAL MBE MFE MN NS PAL SVL TAD TL TND TPD TYE UAW UEW FRIM 0967 Holotype Male 9.0 6.0 1.3 1.4 3.5 3.3 4.5 15.4 19.5 11.2 11.7 2.5 4.3 6.0 5.0 7.8 10.2 1.4 7.9 30.1 0.6 15.6 6.7 4.0 3.1 5.9 1.9 ZRC.1.11400 Paratype Female Gravid 9.4 5.7 1.4 1.8 3.8 3.4 4.0 14.3 19.7 10.3 10.8 2.1 3.4 5.5 4.2 7.3 9.1 1.3 7.6 27.7 1.0 15.3 6.4 4.2 2.2 5.0 2.0 ZRC.1.10535 Paratype Female Gravid 9.4 6.3 1.0 1.3 3.3 3.0 4.0 14.2 19.8 10.2 10.1 2.4 3.3 6.7 4.9 7.9 9.9 1.4 8.2 30.4 0.8 15.8 6.9 4.4 2.4 5.2 2.1 UKM MZ 1245 Paratype Male (C&S) NM NM NM NM 3.8 2.8 3.9 14.6 12.6 8.7 9.3 1.7 3.3 NM NM NM NM NM 7.3 28.0 NM 15.6 NM 2.8 NM NM NM THNSM 01468 Paratype Female Gravid 9.6 6.6 1.57 2.24 3.5 3.3 4.48 16.8 23.1 12.2 11.7 2.8 4.0 7.3 5.5 8.6 11.1 1.4 10.0 31.2 0.9 17.2 7.6 4.4 2.9 5.6 2.4 Mean S.D. 9.4 0.2 6.1 0.4 1.3 0.2 1.7 0.4 3.6 0.2 3.2 0.2 4.2 0.3 15.1 1.1 18.9 3.8 10.5 1.3 10.7 1.0 2.3 0.4 3.6 0.4 6.4 0.8 4.9 0.6 7.9 0.5 10.1 0.8 1.4 0.0 8.2 1.1 29.5 1.6 0.8 0.2 15.9 0.8 6.9 0.5 4.0 0.7 2.6 0.4 5.4 0.4 2.1 0.2

69 Hind limb more than twice length of forelimb (FLL = 39.3% HLL). Toes with Y-shaped terminal phalanges (Fig. 2a); discs of toes semicircular, wider than long, bearing circummarginal grooves; elevated ridges present on discs as in fingers, smaller than discs of fingers; relative toe lengths: I < II < III < V < IV; webbing formula: I112III12IV11V (Fig. 2d); inner metatarsal prominent, oval; no outer metatarsal tubercle; subarticular tubercles prominent, round. In life, dorsum with many small, pearl-tipped tubercles; larger tubercles on sides, above cloaca and on upper surface of limbs; skin on head smooth, not co-ossified with skull; chin smooth, lacking tubercles; palpebrum smooth; ventral surface weakly granular; tuberculate texture lost in preservative; humeral gland absent. Color in life. Iris red; dorsal color variable, pale whitish brown to light brown; inguinal blotches dark brown, partly outlined with white; dark brown bar extends from snout, covers tympanum and continues later-

Note. See text for abbreviations. NM, not measured.

70 ally to anterior of inguinal blotches; throat brown, flecked with small white spots; venter of trunk, arms, and legs white with brown reticulation; faint transverse streaks of brown crossing dorsal surface of femur, tibia, and tarsus, when leg is adpressed and viewed from above, these streaks and the inguinal blotch form a continuous line. It is possible that the white coloration of the dorsum matches the white crusotose lichens, common on trees in the areas where the frogs were collected. Color in preservative. In ethanol, dorsal coloration changed to light gray-brown. Dark brown coloration and white markings on lateral and ventral aspects remain as in life. Variation. Like other rhacophorids (e.g., Polypedates leucomystax), Theloderma licin can vary the pigmentation of portions of its dermis in response to stress, and presumably also to diel period and microhabitat. We noted that the region of variation occurs dorsally from the tip of snout, continuing laterally above the loreal region and upper eyelid to the back, but not reaching the cloaca. The dorsum of the Thai individual (THNSM 01468) was almost entirely white when collected, but changed to brown after the frog was handled, photographed, and maintained in captivity for 2 days (Fig. 4a, b). Similar color change was observed in the holotype (FRIM 0967), which was kept in captivity for a week. The coloration of this species in ethanol seems to depend on its color before euthanasia. The body of paratype ZRC.1.11400 was paler than the holotype FRIM 0976 and paratype UKM MZ 1245, both of which were brown prior to euthanasia. In ethanol, the body colorations of these individuals remained pale white and brown, respectively. In preservative, the reticulated ventral pattern is distinct, and the inguinal spot remains brown with a prominent white outline. In two gravid females, eggs varied in size within ovaries. Large, pigmented, well-developed eggs were 1.0 1.6 mm (n = 10, x = 1.1 0.19) in diameter, whereas small, white, less-developed eggs were 0.4 0.8 mm (n = 10, x = 0.6 0.12) in diameter. Distribution and ecology. Additional fieldwork is necessary to confirm the full distribution of this species. However, on the basis of the known localities, it seems likely that Theloderma licin may occur throughout the Malay Peninsula and perhaps as far north as the Isthmus of Kra in Thailand. The holotype (FRIM 0976) was collected on Crinum Lily (Crinum asiaticum) leaf. This individual was heard before being seen, because of its cryptic coloration. Based on observations by the collector (Dennis Yong), this species was reported to be common within the gardens of the resort area in Taman Negara. Yong

David S. McLeod and Norhayati Ahmad also reported that this species was found calling from within a sewage tank. Paratype ZRC.1.11400 was collected from a fern leaf (15 cm wide) on a steep cliff (almost 90) near a logging road. The Thai paratype (THNSM 01468) was found during the afternoon, approximately 40 cm above ground, resting on a leaf in vegetation alongside a walking trail in a secondary tropical evergreen forest. Paratype ZRC.1.10535 was collected at a temporary forest pool. Paratype UKM MZ 1245 was found inside a bathroom at the Sungai Lasor base camp. Comparisons. Orlov et al. (2006) identified three species-groups based on SVL: small (28 35 mm), medium (40 45 mm), and large (48 75 mm). Theloderma licin is referred to the small group, and therefore is easily separated from members of the other two groups: T. horridum, T. moloch, T. phrynoderma, T. ryabovi, T. nagalandensis, T. bicolor, T. corticale, T. gordoni, T. kwangsiense, and T. leporosum (Orlov et al., 2006). Theloderma licin also can be separated from the Sri Lankan endemic T. schmardana by SVL [23.0 25.6 mm (Dutta and Manamendra-Arachchi, 1996)]. We compared Theloderma licin to six phenotypically similar and/or geographically proximate congeners from Thailand, Malaysia, Laos PDR, Cambodia, and Vietnam (Tables 2 and 3). Theloderma licin is morphologically similar to T. asperum. Both species have large patches of white skin on the dorsum and red eyes, but T. licin differs from T. asperum in having dorsal surfaces of the trunk and limbs being nearly completely white, and digits with basal webbing (free in asperum). In life, T. licin has nearly smooth skin with fine pearly-tipped tubercles, whereas its congeners have dorsal and lateral surfaces that are heavily tuberculated with small to large pearly tipped tubercles. Remarks. Base pairs of a portion of the 12S and 16S mtDNA gene regions from the Taman Negara, Pahang, and Thai specimens were sequenced and revealed that T. licin is closely related to the morphologically similar T. asperum (G. Wogan, unpublished data). Some of diagnostic characters of Theloderma proposed by Liem (1970) seem to be unstable, and it is evident that a complete phylogenetic revision of Theloderma and the phenotypically similar genera of Nictixalus and Philautus is needed. Liem (1970), remarked that the skin of the head is characteristically co-ossified to the skull in the genus Theloderma. None of the species examined in this study had this condition. Additionally, Liem (1970) used the presence of two slips of m. extensor communis longus in the foot as a defining character for Theloderma; however, Wilkinson and Drewes (2000) mention that this character state is also shared by

A New Species of Theloderma Boophis tephraeomystax and Mantidactylus luteus. Taylor (1962) stated that no supratympanic fold was present in T. asperum. In the specimens of T. asperum we examined, it was evident that a fold was present, albeit indistinct. Additionally, in other congeners (e.g., T. gordoni), tubercles obscured the supratympanic fold, but the presence of the fold was apparent, whereas in others (e.g., T. horridum), tubercles surrounded the tympanum, and showed no evidence of a line from eye to arm. In the original description of the genus, Tschudi (1838) described the tympanum of the T. leporosum as hidden. Taylor (1962) and Berry (1975) considered the frogs of

71 this genus to have distinct tympana. The apparent discrepancy may be explained by the presence of tubercles on the tympanum of some species (Table 2), although they do not appear to be present on T. leporosum. Taylor (1962) also commented that Theloderma appeared to be tree-hole breeders, and while true, at least T. stellatum has been observed to deposit eggs on structures above small permanent man-made ponds on the floor of a dry evergreen forest in Thailand (DSM, personal observation).
Acknowledgments. We thank our respective institutions, KU and UKM, for supporting the research and preparation of

TABLE 2. Distribution of Selected Diagnostic Characters in Selected Members of the genus Theloderma Character Vomerine teeth Dorsal asperites Metacarpal tubercles Webbing of hand Webbing of foot T. licin T. asperum T. corticale present present, large 3: thenar + 2 palmar (not bifid) None I11II1 2III1 2IV21V present green-brown T. gordoni present present, large 2: thenar and bifid palmar None I22II1 2III1 3IV31V present brown T. horridum absent present, large 2: thenar and bifid palmar II11III1 2IV11V I11II1 1III1 1IV11V absent brown T. leporosum present present, large 2: thenar and bifid palmar None I12II1 2III1 3IV21V present brown T. stellatum absent present, small 2: thenar and bifid palmar II22III1 3IV22V I12II1 2III1 2IV21V absent brown with white markings present (tuberculate) present

Chin asperites Dorsal color in life Supratympanic fold Tympanum with tubercles

absent absent present, minute present, small 3: thenar + 3: thenar + 2 palmar 2 palmar (not bifid) (not bifid) II22III2 None 3IV32V I12II1 II2II1 2III12IV2 2III1 1V 2IV21V absent absent white to whitish white with brown gray-brown patch present present absent absent

absent present

present (tuberculate) present

absent present

absent absent

Note. See Table 3 for morphometric proportions.

TABLE 3. Morphological Proportions (in %) in Adults of the Theloderma Species Examined Character SVL, mm HL/HW DL/DW HL/SVL FEL/SVL TYE/ED FEL/TBL FLL/HLL ES/HL DW/TYE FEL + TBL/ SVL IN/IO IO/UEW T. licin n=5 27.6 31.1 93.5 105.0 70.0 92.1 31.1 39.3 46.6 54.0 57.6 89.3 89.9 98.7 36.7 40.3 36.6 44.8 45.5 79.6 98.5 109.2 51.5 70.6 161.7 228.5 T. asperum n=2 21.1 29.7 95.7 106.5 77.2 87.8 37.3 38.5 41.5 49.9 66.1 71.8 78.5 85.5 37.9 39.2 41.9 44.9 50.8 63.9 94.4 108.1 57.0 62.2 128.2 144.9 T. corticale n=2 29.7 34.4 100.0 113.3 60.9 79.8 39.3 42.8 43.3 58.0 78.1 91.0 89.8 105.2 37.4 43.1 37.8 42.9 61.7 88.6 91.5 113.1 58.7 74.7 111.9 137.7 T. gordoni n=4 43.9 49.2 101.3 108.9 66.1 109.8 40.4 45.4 44.6 51.7 72.8 92.9 88.7 102.4 41.6 44.0 38.8 62.5 59.7 80.4 94.8 106.2 54.3 71.2 123.9 158.9 T. horridum n=4 37.1 48.7 114.1 123.0 53.1 68.4 41.3 46.6 46.8 53.8 54.7 78.2 87.0 95.7 38.5 41.3 42.6 41.0 93.7 146.1 95.6 111.5 69.3 79.6 82.3 116.5 T. leporosum n=1 62.6 110.0 60.2 43.1 45.2 70.7 80.2 33.5 32.9 90.1 101.5 56.7 121.6 T. stellatum n=7 66.6 68.9 105.1 109.2 65.6 71.4 42.5 43.6 46.7 50.1 57.1 88.1 91.0 96.9 37.4 42.8 40.3 40.6 84.4 94.2 98.0 101.8 66.3 69.7 119.4 124.4

Note. See text for character abbreviations and Table 1 for intraspecific comparisons of each variable.

72
this paper. DSM thanks Mr. Tanya Chan-Ard, Dr. D. Karns and Mr. J. Murphy for the opportunity to work at Khao Luang during the July 2003 survey. Norsham Yaakob (FRIM) was instrumental in the organization and development of this manuscript. NA thanks the Forestry Department of Peninsular Malaysia and the Forestry Department of Kedah for their help and support in all her research in Kedah. We thank Kelvin Lim for examination of comparative material under his care. H. K. Voris, R. F. Inger, and A. Resetar at FMNH helped with specimen examination and loan of materials. We are indebted to our mutual colleague, G. Wogan (California Academy of Sciences); through her interactions with NY and DSM she realized that we seemed to have collected the same new taxon. Her introductions prompted the development of this collaborative work. Rafe Brown, Juan M. Guayasamin, Edgar Lehr, Linda Trueb, and Lee Grismer provided constructive reviews of early drafts of this manuscript. Funding was provided, in part by the University of Kansas department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Research Centers Panorama fund.

David S. McLeod and Norhayati Ahmad

Ohler A., Swan S. R., and Daltry J. C. (2002), A recent survey of the amphibian fauna of the Cardamon Mountains, Southwest Cambodia with descriptions of three new species, The Raffles Bull. Zool., 50(2), 465 481. Orlov N. L., Dutta S. K., Ghate H. V., and Kent Y. (2006), New Species of Theloderma from Kon Kum Province (Vietnam) and Nagaland State (India) [Anura: Rhacophoridae], Russ. J. Herpetol., 13(2), 135 154. Savage J. M. and Heyer W. R. (1997), Digital Webbing Formulae for Anurans: A refinement, Herpetol. Rev., 28(3), 131. Taylor E. H. (1962), The amphibian fauna of Thailand, Kansas Univ. Sci. Bull., 43, 265 599 Taylor W. R. and Van Dyke G. C. (1985), Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study, Cybium, 9, 107 121. Tschudi J. J. V. (1838), Classification der Batrachier, mit Bercksichtigung der fossilen Thiere dieser Abtheilung der Reptilien, Mem. Soc. Sci. Nat. Neuchtel, 2, 1 99. Wilkinson J. A. and Drewes R. C. (2000), Character assessment, genus level boundaries, and phylogenetic analyses of the family Rhacophoridae: A review and present day status, Contemp. Herpetol., 2, 1 18.

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APPENDIX I. Specimens Examined

Theloderma asperum. Cambodia: Mondolkiri Province: FMNH 262787, 261896; Lao PDR: Khammouan Province: FMNH 255379; Vientiane Province: FMNH 261897; Malaysia: Pahang, Frasers Hills: ZRC.1.8043, ZRC.1.9321. Theloderma corticale. Vietnam: Vinh-Phu Province: FMNH 254285, 254286. Theloderma leprosum. Indonesia: Sumatra: RMNH 1754. Theloderma licin. Malaysia: Kedah State: ZRC.1.11400, UKM MZ 1245; Pahang State: FRIM 0967, ZRC.1.10535; Thailand: Nakhon Si Thamarat: THNSM 01458. Theloderma gordoni. Thailand: Chiang Mai Province: FMNH 172248; Vietnam: Gai-Lai Province: FMNH 253615, 2536156; Vinh-Phu Province: FMNH 254287. Theloderma horridum. Malaysia: Bukit Timah Nature Reserve: ZRC.1.3220; Selangor: FMNH 186600, 186602; Tioman Island: ZRC.1.9661; Sabah: FMNH 240962; Singapore: ZRC.1.3278. Theloderma stellatum. Cambodia: Mondolkiri Province: FMNH 262786; Thailand: Chanthaburi Province: FMNH 172249, 211831; Nakhon Ratchasima Province: FMNH 183711; Nakhon Nayok Province: FMNH 183715; Vietnam: Akhe District: Gai-Lai Province: FMNH 253622 24, FMNH 253617.