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Lee Grismer, Evan S. H. Quah L., Syafiq, Muhamad Fatihah, Rujirawan, Attapol, Aowphol, Anchalee, Ahmad, Amirrudin B., Anuar, M.S. Shahrul (2023): Comments on the taxonomic status of Cyrtodactylus zebraicus Taylor, 1962 (Squamata: Gekkonidae) from Northern Peninsular Malaysia. Zootaxa 5318 (4): 489-503, DOI: 10.11646/zootaxa.5318.4.3, URL: http://dx.doi.org/10.11646/zootaxa.5318.4.3

Cyrtodactylus zebraicus Taylor, 1962 (Figs. 3–5; Table 2 & 3) Cyrtodactylus peguensis (in part): Das & Norsham 2007: 77; Grismer 2008: 30 Cyrtodactylys peguensis zebraicus: Ulber 1989: 4 & 5; Denzer & Manthey 1991: 314; Manthey & Grossmann 1997: 226; Das 2010: 213 Referred specimens. USMHC 2635, collected by Evan S.H. Quah and M.S. Shahrul Anuar on 7 November 2019 at approximately 21:00 h from Perlis State Park, Perlis, Malaysia (approximately 6.698222N, 100.191611E; 144 m elevation). USMHC 2639 bears the same collection data as USMHC 2635 but was collected on 9 November 2019. UMTZC 2413, collected by Muhamad Fatihah Syafiq, Baizul Hafsyam Badli-Sham and Amirrudin B. Ahmad on 5 October 2022 at 20:25 h from Bukit Ayer, Perlis, Malaysia (6.542000N, 100.168000E; 52 m elevation). Description of specimen USMHC 2635. Adult male with 62.0 mm SVL; head moderate in length (HL/SVL 0.29), wide (HW/HL 0.58), slightly flattened (HD/HL 0.40), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal region flattened, prefrontal region slightly concave, canthus rostralis rounded; snout short (ES/HL 0.40), rounded in rostral region, eye to snout distance slightly less than head depth; eye large (ED/HL 0.22), eyeball slightly protuberant, eye diameter less than the eye to ear distance, pupil vertical; ear opening elliptical, obliquely oriented, moderate in size (EL/HL 0.09); rostral large, subrectangular, height 1.5 mm, shorter than wide, 2.7 mm, medially divided by a dorsal furrow, reaching to approximately halfway down rostral height, bordered posteriorly by supranasals and internasal, laterally by first supralabials and nostrils; external nares at anterior angle of snout, directed lateroposteriorly, bordered anteriorly by rostral, dorsally by two large supranasals, posteriorly by two smaller postnasals, ventrally by first supralabial; internarial distance narrow; anterior pair of supranasals subrectangular, separated by one small internasal, bordered anteriorly by rostral, laterally by nostrils, posteriorly by posterior supranasal and three smaller scales; one internasal, subhexagonal, vertically arranged, slightly protruding rostral, bordered posteriorly by two small scales; 9/8 (right/left) supralabials extending to below midpoint of eye, 15/12 to below the posterior margin of the eyeball, subrectangular anteriorly, elliptical shape posteriorly; 6/7 infralabials extending to below midpoint of eye, 9/11 to below the posterior margin of the eyeball, larger than supralabials, tapering smoothly posteriorly; scales of frontonasal, prefrontal and lores, small, relatively raised, domed, slightly larger than granular scales on top of head and occiput; scales on occiput intermixed with scattered, slightly larger, more rounded, dome to subconical tubercles, more prominent tubercles between occiput and above ear opening; dorsal supraciliaries crenulated, not elongate or keeled; mental large, triangular, 2.0 mm in width, 1.7 mm in length, bordered laterally by first infralabials and posteriorly by large, right and left trapezoidal postmentals that contact medially for 50% of their length posterior to mental; one row of slightly enlarged chin shields extending posteriorly to sixth (right/left) infralabials; and gular and throat scales small, granular, grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales. Body relatively short, somewhat stout (AG/SVL 0.46), lacking ventrolateral folds; scales on dorsum small, mostly homogenous, granular, interspersed with larger, irregularly arranged, slightly prominent trihedral keeled tubercles; tubercles extending from occiput beyond to the base of the original portion of tail; tubercles on occiput, nape and anterior of body at level above shoulder smaller, subconical; those mid-dorsally and on the posterior section of the body larger, being more dense, slightly more prominently keeled, and more regularly arranged in sacral region and tail base; tubercles on flanks sparse; approximately 15 longitudinal rows of dorsal tubercles; approximately 22 paravertebral tubercles; 30 flat, imbricate, smooth ventral scales, those near midline larger than those laterally and dorsal scales; femoral are not enlarged; precloacal scales smooth, approximately twice the size of femoral scales; femoral pores absent; seven precloacal scales; six precloacal pores; four rows of enlarged postprecloacal scales; and precloacal groove or depression absent. Limbs moderately slender;forelimbs relatively short(FL/SVL 0.16); scales on dorsal surface domed to subconical, granular, slightly larger than those on body, interspersed with sparsely enlarged, subconical and trihedrally keeled tubercles; dorsal scales of wrist and palm flat, smooth, round, imbricate; ventral scales of palm flat, weakly rounded, slightly raised, not imbricate, smaller than those on body; 14/14 (right/left) total subdigital lamellae on fourth finger, 4/4 proximal subdigital lamellae rectangular with rounded to weakly rounded corners, broadly expanded proximal to joint inflection on fourth finger, 10/10 distal subdigital lamellae, slightly expanded immediately distal to joint, becoming gradually more expanded near the claw; digits well-developed, relatively long, inflected at basal interphalangeal joints; digits slightly narrower distal to inflections; no interdigital webbing; claw well-developed, relatively short, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.17); dorsal scales domed to subconical, granular, interspersed with enlarged subconical and trihedrally keeled tubercles, and anterior part of thigh covered by flat, slightly larger, imbricate scales; ventral scales of femora flat, smooth, imbricate, smaller than those on body; small postfemoral scales form an abrupt union with large, flat ventral scales of posteroventral margin of thigh; ventral scales of tibia flat, imbricate; dorsal scales of plantar surface relatively smooth, rounded, imbricate; ventral scales of plantar surface low flat, weakly rounded; 16/16 (right/left) total subdigital lamellae on fourth toe, 5/5 proximal subdigital lamellae, rectangular with rounded to weaky rounded corners, broadly expanded proximal to joint inflection on fourth toe, 11/11 distal subdigital lamellae, slightly expanded immediately distal to joint, becoming gradually more expanded near the claw; digits well-developed, relatively long, inflected at basal, interphalangeal joints; and claw well-developed, relatively short, claw base sheathed by a dorsal and ventral scale. Tail 41.4 mm in length, regenerated, shorter than SVL (TL/SVL 0.67), moderate in proportions, cylindrical, wide anteriorly, 5.3 mm in width at base, tapering to a tip, covered with small, flat, imbricating scales on the dorsal surface but slightly larger, flat, imbricating scales on ventral surface; dorsal scales of original portion of tail base granular, round, becoming larger, flatter, subimbricate posteriorly on regenerated portions of tail; those on tail base bearing trihedrally keeled tubercles forming paravertebral rows, no caudal furrow; base of tail forming hemipenial swelling; 2/2 (right/left) postcloacal tubercles on the enlarged smooth hemipenial swelling; and postcloacal tubercles approximately equal size. Coloration in life (Fig. 2). Dorsal ground color of head, body, and limbs light-brown; top of head bearing large, dark-brown blotches edged in yellow, forming a light-colored reticulated pattern; superciliary scales yellow; supralabials and infralabials off-white bearing darker markings; wide, discontinuous, dark-brown nuchal loop edged in yellow, extending from posterior margin of one orbit, across occiput and nape to posterior margin of the other orbit; five dark-brown paravertebral blotches on the dorsum starting from shoulder region on top of the forelimbs until the sacral band, all blotches edged in yellow but the second and third pair of paravertebral blotches not divided medially along the vertebral column; all yellow outlines of the paravertebral blotches extend along the flanks towards the venter to form a series of narrow, yellow bands along the flanks; sacral band between hindlimbs composed of paravertebral blotches; postsacral band composed of confluent blotches; regenerated tail dark-grey with lighter grey and orange-brown speckling; dorsal portion of forelimbs and hindlimbs bearing irregularly shaped yellow banding and reticulations; all ventral surfaces generally greyish white, immaculate, except for ventral surface of knee and elbow region which bear some dark mottling. Coloration in preservation (Fig. 2). The overall color pattern is similar to that in life except that it is faded. Ground color of head, body, and limbs light-brown; nuchal, body and sacral bands dark brown but slightly lighter than in life; the light-yellow reticulated pattern on the dorsal surfaces of the head, body and limbs faded to off-white; and all ventral surfaces light-beige. Variation (Fig. 2, 3 & 5). Morphometric, meristic and color pattern characters of the other referred specimens of C. zebraicus from Peninsular Malaysia are presented in Table 1. All the specimens are generally similar in most aspects of morphology with subtle variation in color pattern (Figs. 2 & 3). In USMHC 2639 the right 4TLU is damaged and approximately 2/3 of tail is original: bearing four longitudinal rows of dorsal tubercles, two transverse rows of dorsal tubercles extend from tail base to anterior margin of first light caudal band, 4.7 mm from tail base, approximately 8.3% of tail; a single enlarged median row of transverse scales on subcaudal region starting from second light caudal band on original portion of tail and ends at the start of the regenerated portion of the tail. The color pattern of USMHC 2639 is also slightly lighter both in life and preservation and the original portion of the tail bears light and dark caudal bands that completely encircle the tail. Similarly, the tail that is original in UMTZC 2413 bear light and dark caudal bands that completely encircle the tail. ......continued on the next page TABLE 1 ( Continued) Distribution in Peninsular Malaysia (Fig. 1). Cyrtodactylus zebraicus is currently only known from around the Nakawan Range in Perlis state but the species may range more widely in Peninsular Malaysia. However, given what is known about its habits in Thailand (Grismer et al. 2021), it is expected to be confined to only lowland areas in the extreme north at the Malaysian-Thai border, in the states of Kedah, Perak and Kelantan. The extralimital distribution of the species outside of Peninsular Malaysia is in peninsular Thailand, south of the Isthmus of Kra (Grismer et al. 2021). Natural history (Fig. 4). In Peninsular Malaysia, C. zebraicus is a habitat generalist and has been found in lowland, semi-deciduous forest (Sharma et al. 1996, 2001) associated with karstic areas along the Nakawan Range. Specimens USMHC 2635 and 2639 were both observed at night approximately 1m above the ground on the trunks of small trees, in an open area near the forest edge. It was raining the night USM 2635 was found and the gecko was perched on a dry section of the tree trunk and partially protected by low vegetation. In comparison, UMTZC 2413 was found at night crawling in a grassy area bordering the main road that cuts through the forested area. The weather was clear the night UMTZC 2413 was found but it had been raining for a few days beforehand. Conservation status. At present, there is only limited ecological knowledge for the species in Peninsular Malaysia. Anecdotal observations indicate that Peninsular Malaysian specimens are similar to Thai populations in habits, and it is a habitat generalist that can survive in habitats with moderate human disturbance (Grismer et al. 2021). In addition, the species occurs in protected areas managed by the Perlis Forestry Department such as Bukit Ayer. As such, we believe that the future of C. zebraicus in the country is secure and we recommend listing it as a species of Least Concern (LC). Comparison and comments on taxonomy. Cyrtodactylus zebraicus can be easily differentiated from all other species of Cyrtodactylus known in Peninsular Malaysia based on a combination of color pattern and scale characters. From the agamensis group in Peninsular Malaysia composed of C. jarakensis, C. metropolis, C. pantiensis, C. payacolus, C. semenanjungensis and C. tiomanensis (Grismer et al. 2021), C. zebraicus differs by having fewer paravertebral tubercles (22–27 vs. 32–40), fewer ventral scales (30–36 vs. 36–61), fewer subdigital lamellae on fourth toe (13–17 vs. 17–24) and the color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. dark speckling with absence of white reticulum) (Grismer 2011; Grismer et al. 2014a). From C. aurensis of the philipinicus group (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 95.0– 99.4 mm SVL), fewer ventral scales (30–36 vs. 45–51), fewer subdigital lamellae on fourth toe (13–17 vs. 18–23), precloacal groove (absent vs. present and deep), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. lightly colored reticulate pattern) and patterning on body (dense bold banding or blotched pattern on body, and bold light and dark banding on original tail vs. narrow light bands on body and tail) (Grismer 2011). From members of the pulchellus group consisting of C. astrum, C. australotitiwangsaensis, C. bintangrendah, C. bintangtinggi, C. dayangbuntingensis, C. evanquahi, C. hidupselamanya, C. jelawangensis, C. langkawiensis, C. lenggongensis, C. macrotuberculatus, C. pulchellus, C. sharkari, C. timur and C. trilatofasciatus (Grismer et al. 2021), C. zebraicus can be easily differentiated on the basis of lacking a deep precloacal groove, smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 99.0– 122.2 mm SVL), contiguous series of enlarged pore-bearing femoral and precloacal scales (absent vs. present), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. generally immaculate with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. 3–7 bold, dark bands on the body edged in white) (Grismer 2011; Quah et al. 2019; Wood et al. 2020). From members of the brevipalmatus group composed of C. cf. brevipalmatus and C. elok (Grismer et al. 2021), C. zebraicus differs by webbing at the base of digits (absent vs. present), caudal fringe of spiny tubercles along tail (absent vs. present), fewer ventral scales (30–36 vs. 36–47), fewer subdigital lamellae on fourth toe (13–17 vs. 19–21), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. mottled with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. color similar to tree bark with usually faint bands and mottling) (Grismer 2011; Grismer et al. 2022). From C. durio of the lateralis group (Grismer et al. 2021), C. zebraicus differs by its smaller adult length in both sexes (50.0– 65.7 mm SVL vs. 79.3 mm SVL in adult male), elongate, spinose tubercles in ventrolateral body fold and on ventrolateral margin of tail (absent vs. present), fewer ventral scales (30–36 vs. 59), fewer subdigital lamellae on fourth toe (13–17 vs. 22), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. irregular mottling) and patterning on body (dense bold banding or blotched pattern on body vs. squarish blotches on body) (Grismer 2011). From C. leegrismeri of the condorensis group (Grismer et al. 2021), C. zebraicus differs in its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 90.0–92.0 mm SVL), fewer subdigital lamellae on fourth toe (13–17 vs. 18–21), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. weak reticulum) and patterning on body (dense bold banding or blotched pattern on body, and bold light and dark banding on original tail vs. dark, indistinct symmetrical blotches on body and tail) (Grismer 2011). From members of the sworderi group consisting of C. guakanthanensis, C. gunungsenyumensis, C. quadrivirgatus, C. sworderi and C. tebuensis (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 71.0– 82.2 mm SVL), fewer paravertebral tubercles (22–27 vs. 32–40) and color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. mottled with absence of white reticulum) (Grismer 2011; Grismer et al. 2016). From members of the darmandvillei group composed of C. batucolus and C. seribuatensis (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 70.1–75.2 mm SVL), fewer ventral scales (30–36 vs. 28–42), fewer subdigital lamellae on fourth toe (13–17 vs. 17–22), contiguous series of enlarged pore-bearing femoral and precloacal scales (absent vs. present), color pattern on top of head (prominent blotched patterning outlined by white reticulum vs. dark speckling with absence of white reticulum) and patterning on body (dense bold banding or blotched pattern on body vs. dark blotches on body, bordered by diffuse, light colored bands or spots) (Grismer 2011). From populations of the C. consobrinus complex of the malayanus group (Grismer et al. 2021), C. zebraicus differs by its smaller adult size in both sexes (50.0– 65.7 mm SVL vs. 117.4–125.0 mm SVL), fewer ventral scales (30–36 vs. 58–65), fewer subdigital lamellae on fourth toe (13–17 vs. 23–28), enlarged pore-bearing femoral scales in males (absent vs. present), and patterning of body (dense bold banding or blotched pattern on body vs. narrow, relatively widely spaced lighter bands) (Grismer 2011).

Subdoluseps vietnamensis sp. nov. (Figs. 3 & 4) Holotype. ITBCZ 5842, adult male, collected from Ho Tram Area, Xuyen Moc District, Ba Ria-Vung Tau Province, Vietnam; coordinates 10°29’38.2” N, 107°27’40.0” E; elevation 11 m asl by Sang N. Nguyen and Vu D.H. Nguyen, on 21 March 2017 (Fig. 3). Paratypes. Two adult males ITBCZ 5843 and 5844, and one adult female ITBCZ 5856 collected by Sang N. Nguyen and Vu D.H. Nguyen from the same site as the holotype; two adult males, ITBCZ 7038 and 7046, collected by Manh V. Le on 04 June 2020 at Ho Tram Area, Xuyen Moc District, Vietnam, coordinates 10°28’59.8” N, 107°26’03.6” E, elevation 10 m asl; two adult females ITCBZ 7056 and 7059, and one juvenile ITBCZ 7070, collected by Manh V. Le on 04 June 2020 at La Gi Town, Binh Thuan Province, Vietnam, coordinates 10°42’38.8’’ N, 107°42’10.8” E, elevation 62 m asl (Fig. 4). Diagnosis. Subdoluseps vietnamensis sp. nov. is distinguished from its congeners by a combination of the following morphological characters: medium size in adults (SVL up to 48.7 mm); tail length approximately equal to SVL; 27–30 smooth midbody scale rows; dorsal scales not enlarged; 55–57 paravertebral scales; 34–37 axilla-groin scales; 55–62 ventral scale rows; 64–74 subcaudal scales; four supraoculars; frontoparietal single; prefrontals not in contact with one another; two loreal scales; seven supralabials, the fifth below the center of the eye; one anterior and two posterior enlarged temporal scales; ear opening with two lobules on the anterior margin; 9 or 10 smooth subdigital lamellae beneath finger III and 12–15 beneath toe IV; six enlarged precloacal scales; and four distinct black stripes on dorsum. Description of holotype. Adult male, SVL 46.5 mm; snout short and obtuse; lower eyelid scaly; body rather robust; tail regenerated. Head scales smooth; rostral convex, distinctly visible from above, broader than long (2.0 mm width, 1.0 mm height); a pair of supranasals forming short median suture, touching nasals and anterior loreals laterally, width equal to length (0.7 mm); prefrontals small, quadrangular, widely separated from each other, each touching both loreals; four supraoculars; frontal truncate anteriorly, longer than its distance from tip of snout, shorter than its distance from nuchal (2.0 mm vs. 3.2 mm), longer than wide (1.8 mm width, 2.6 mm length), touching two anterior supraoculars; frontoparietal single, much larger than interparietal, wider than long (3.0 mm width, 1.9 mm length), touching three supraoculars; parietals in contact posteriorly behind the interparietal; interparietal longer than wide (1.1 mm width, 1.6 mm length), with parietal eye visible posterior to center; a pair of nuchals; seven supralabials on both sides, fifth below center of the eye, first larger than the three following, sixth largest; two loreals, anterior (0.6 mm length, 0.6 mm height) slightly higher, the posterior (0.9 mm length, 0.5 mm height) longer; nasal divided, in contact with the first supralabial, rostral, anterior loreal, and supranasal; nostril in center of nasal; eight supraciliaries, first largest; one enlarged anterior temporal in contact with the sixth and the seventh supralabial; two posterior temporals, lower one smaller and overlapping the upper one, the lower temporal in contact with seventh supralabials; six infralabials, first two in contact with postmental; two pairs of chin shields, first pair medially in contact with each other; ear-opening moderately small, the anterior border with two lobules. Body elongate (AGD/SVL = 0.56); dorsal scales smooth, not larger than lateral and ventral scales, with eight longitudinal rows on the back; 28 midbody scale rows; 71 paravertebral scales; 35 axilla-groin scales; ventral scales smooth, in 55 rows; limbs short (FLL/SVL = 0.18, HLL/SVL = 0.25), pentadactyl, widely separated by ten scales when adpressed; 3, 6, 9, 9, and 7 smooth lamellae beneath fingers I–V, respectively; 4, 7, 13, 13, and 9 smooth lamellae beneath toes I–V, respectively; six enlarged precloacal scales, the two middle ones slightly larger than the others; size of median subcaudal series on the original part of the tail equal to those of its adjacent rows. Coloration. In life, overall dorsal coloration reddish brown with two dorsolateral bright lines on anterior part of dorsum; each dorsal scale of four central rows with a black spot forming four distinct longitudinal lines on dorsum; ear and surrounding area orange; sides dark brown to black intermixed with white, brown and yellow spots; chest and belly yellow (Fig. 3). In preservation, color fades but the four longitudinal dorsal lines remain distinct; orange and yellow faded to cream or white; overall dorsal and lateral coloration black to dark brown with bright spots on lateral sides; venter cream. Variation in paratypes (n = 8). Most morphological characters of paratypes agree with those of the holotype. The following main characters vary: (1) midbody scale rows: vary from 27 to 30; (2) paravertebral and ventral scale rows: range from 55 to 57 and from 55 to 62, respectively; (3) axilla-groin scales: range from 34 to 37; (4) and nuchals: range from one to three scales. Table 3 summarizes variation in size and scalation of the type series. ...Continued on the next page ...Continued on the next page Field notes. All specimens were collected during the daytime, on sandy ground among rotting leaves in Earleaf acacia (Acacia auriculiformis A.Cunn. ex Benth.) plantation, secondary dipterocarp forest, and rubber plantation in lowland coastal areas (Fig. 5). The holotype and paratypes ITBCZ 5843 & 5844 were collected at 15:42, air temperature 30.3 oC, relative humidity 70.2 %; ITCBZ 7038 was collected at 13:30, air temperature 33 oC, relative humidity 75 %; ITBCZ 7046 was collected at 8:54, air temperature 30 oC, relative humidity 80 %; ITBCZ 7056, 7059 & 7070 were collected at 12:10, air temperature 32 oC, relative humidity 65%. To avoid capture, this species can dive into and stay under loose sand. Sexual dimorphism. Adult males mainly differ from adult females in body coloration in life and shape of tail base. Males have yellow color on chest and belly (vs. white, cream or yellowish color in females), orange around the ear (vs. dark brown in females), and distinct bright spots on lateral side (vs. pure dark brown or with small bright dots in females). Ventral side of tail base is slightly swollen in males and flat in females (Figs. 3, 4A–E). Juvenile has bright golden color on the dorsum with four clear dorsal longitudinal stripes (Fig. 4F). Distribution. The new species is currently known only from (1) Ho Tram, Xuyen Moc District, Ba Ria-Vung Tau Province and (2) La Gi Town, Binh Thuan Province, southern Vietnam (Fig. 1). Etymology. The new skink is named after the nation of Vietnam. We recommend “ Vietnam Agile Skink” and “Thằn lằn chân ngắn việt nam” as the common English and Vietnamese names of the new species, respectively. Comparisons. Subdoluseps vietnamensis sp. nov. differs morphologically from other species of the Lygosoma s.l. in Asia as follows: For Subdoluseps, S. vietnamensis sp. nov. differs from S. herberti, S. malayana, S. purthi, and S. samajaya by having a single frontoparietal scale (vs. paired) (Grismer 2019; Karin et al. 2018; Sharma 1977; Smith 1916). In addition, it differs from S. herberti by having a smaller adult size (maximum SVL = 48.7 mm vs. 54.0 mm), presence (vs. absence) of nuchal scales (1–3 vs. 0), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Smith 1916); from S. malayana by having a smaller adult size (maximum SVL = 48.7 mm vs. 65.4 mm), more supralabials (7 vs. 6) with the fifth below the center of the eye (vs. the fourth below the center of the eye), smooth dorsal scales (vs. dorsal with 3–5 strong keels) (Grismer et al. 2019); from S. purthi by having a smaller adult size (maximum SVL = 48.7 mm vs. 67.0 mm), fewer midbody scale rows (27–30 vs. 32–34), and more paravertebral scale rows (55–57 vs. 50) (Sharma 1977); from S. samajaya by having a smaller adult size (maximum SVL = 48.7 mm vs. 70.1 mm), fewer paravertebral scale rows (55–57 vs. 60 or 61), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Karin et al. 2018). Subdoluseps vietnamensis sp. nov. is phenotypically most similar to S. bowringii and S. frontoparietale but can be distinguished from both based on a combination of characters. The new species differs from S. bowringii by having a single frontoparietal scale (vs. paired) (Geissler et al. 2011; Smith 1935; this study), tail length approximately equal to SVL (vs. tail longer than SVL) (mean TL/SVL = 1.04 vs. 1.27 [Smith 1935; this study]), and fewer subcaudal scales (64–74 vs. 76–81) (this study); from S. frontoparietale by having a tail length approximately equal to SVL (mean TL/SVL = 1.04 vs. tail longer than SVL, mean TL/SVL = 1.24 [Taylor 1962; this study, Table 4]), fewer subcaudal scales (64–74 vs. 78–80) [Taylor 1962; this study]), fewer axilla-groin scales (34–37 vs. 39–42) (this study), four distinct longitudinal black stripes on dorsum (vs. six distinct black stripes) (Taylor 1962; this study), and absence (vs. presence) of bright dorsolateral band extending from eye to tail base (Taylor 1962; this study, Fig. 6). From genus Lygosoma s.s., Subdoluseps vietnamensis sp. nov. differs from L. isodactylum, L. siamensis, L. tabonorum, and L. quadrupes (Linnaeus) by having supranasals in contact with each other (vs. not in contact) (Geissler et al. 2011, Heitz et al. 2016, Siler et al. 2018) and from L. corpulentum by having smaller adult size (maximum SVL = 48.7 mm vs. 170.0 mm), frontoparietal scale single (vs. paired), and fewer midbody scale rows (27–30 vs. 36–40) (Smith 1921; Geissler et al. 2011). For the genus Riopa, Subdoluseps vietnamensis sp. nov. differs from R. albopunctata, R. anguinua, R. goaensis, R. guentheri, R. popae Shreve, and R. punctata (Gmelin) by having frontoparietal scales single (vs. paired) (Geissler et al. 2012; Sharma 1976; Smith 1935); from R. lineata and R. lineolata by having fewer paravertebral scale rows (55–57 vs. 104–110 in R. lineata [Smith 1935] and 78–93 in R. lineolata [Siler et al. 2018]); and from R. vosmaeri by having hind-limb with 5 (vs. 4) toes (Gray 1939; Seetharamaraju et al. 2009). For remaining Asian species of the Lygosoma s.l. that were not confirmed by Freitas et al. (2019), Subdoluseps vietnamensis sp. nov. differs from L. boehmei Ziegler, Schmitz, Heidrich, Vu & Nguyen, L. opisthorhodum Werner, L. singha (Taylor), and L. veunsaiensis Geissler, Hartmann & Neang by having frontoparietal scale single (vs. paired) (Geissler et al. 2012; Werner 1910; Ziegler et al. 2007); from L. angeli (Smith) by having a smaller adult size (maximum SVL = 48.7 mm vs. 100.0 mm) and fewer paravertebral scale rows (55–57 vs. 107–115) (Smith 1937; Geissler et al. 2011); from L. haroldyoungi (Taylor) by having a smaller adult size (maximum SVL = 48.7 mm vs. 136.0 mm), fewer paravertebral scale rows (55–57 vs. 141–145), and fewer midbody scale rows (27–30 vs. 38–42) [Taylor 1962; Geissler et al. 2011]; from L. kinabatanganensis Grismer, Quah, Duzulkafly & Yambun by having a smaller adult size (maximum SVL = 48.7 mm vs. 141.0 mm), fewer paravertebral scale rows (55–57 vs. 98), and fewer midbody scale rows (27–30 vs. 42) (Grismer et al. 2018); from L. koratense Smith by having a smaller adult size (maximum SVL = 48.7 mm vs. 105.0 mm) and fewer midbody scale rows (27–30 vs. 32–34) [Smith 1935]; and from L. bampfyldei Bartlett, L. peninsulare Grismer, Quah, Duzulkafly & Yambun, and L. schneideri Werner by having fewer paravertebral scale rows (55–57 vs. 81–85 in L. bampfyldei [Grismer et al. 2018], 87 in L. peninsulare [Grismer et al. 2018], and 95 in L. schneideri [Grismer et al. 2018]).

Le, Manh Van, Nguyen, Vu Dang Hoang, Phan, Hoa Thi, Rujirawan, Attapol, Aowphol, Anchalee, Vo, Thi-Dieu-Hien, Murphy, Robert W., Nguyen, Sang Ngoc (2021): A new skink of the genus Subdoluseps Freitas, Datta-Roy, Karanth, Grismer & Siler, 2019 (Squamata: Scincidae) from southern Vietnam. Zootaxa 4952 (2): 257-274, DOI: https://doi.org/10.11646/zootaxa.4952.2.3

Subdoluseps vietnamensis sp. nov. (Figs. 3 & 4) Holotype. ITBCZ 5842, adult male, collected from Ho Tram Area, Xuyen Moc District, Ba Ria-Vung Tau Province, Vietnam; coordinates 10��29���38.2��� N, 107��27���40.0��� E; elevation 11 m asl by Sang N. Nguyen and Vu D.H. Nguyen, on 21 March 2017 (Fig. 3). Paratypes. Two adult males ITBCZ 5843 and 5844, and one adult female ITBCZ 5856 collected by Sang N. Nguyen and Vu D.H. Nguyen from the same site as the holotype; two adult males, ITBCZ 7038 and 7046, collected by Manh V. Le on 04 June 2020 at Ho Tram Area, Xuyen Moc District, Vietnam, coordinates 10��28���59.8��� N, 107��26���03.6��� E, elevation 10 m asl; two adult females ITCBZ 7056 and 7059, and one juvenile ITBCZ 7070, collected by Manh V. Le on 04 June 2020 at La Gi Town, Binh Thuan Province, Vietnam, coordinates 10��42���38.8������ N, 107��42���10.8��� E, elevation 62 m asl (Fig. 4). Diagnosis. Subdoluseps vietnamensis sp. nov. is distinguished from its congeners by a combination of the following morphological characters: medium size in adults (SVL up to 48.7 mm); tail length approximately equal to SVL; 27���30 smooth midbody scale rows; dorsal scales not enlarged; 55���57 paravertebral scales; 34���37 axilla-groin scales; 55���62 ventral scale rows; 64���74 subcaudal scales; four supraoculars; frontoparietal single; prefrontals not in contact with one another; two loreal scales; seven supralabials, the fifth below the center of the eye; one anterior and two posterior enlarged temporal scales; ear opening with two lobules on the anterior margin; 9 or 10 smooth subdigital lamellae beneath finger III and 12���15 beneath toe IV; six enlarged precloacal scales; and four distinct black stripes on dorsum. Description of holotype. Adult male, SVL 46.5 mm; snout short and obtuse; lower eyelid scaly; body rather robust; tail regenerated. Head scales smooth; rostral convex, distinctly visible from above, broader than long (2.0 mm width, 1.0 mm height); a pair of supranasals forming short median suture, touching nasals and anterior loreals laterally, width equal to length (0.7 mm); prefrontals small, quadrangular, widely separated from each other, each touching both loreals; four supraoculars; frontal truncate anteriorly, longer than its distance from tip of snout, shorter than its distance from nuchal (2.0 mm vs. 3.2 mm), longer than wide (1.8 mm width, 2.6 mm length), touching two anterior supraoculars; frontoparietal single, much larger than interparietal, wider than long (3.0 mm width, 1.9 mm length), touching three supraoculars; parietals in contact posteriorly behind the interparietal; interparietal longer than wide (1.1 mm width, 1.6 mm length), with parietal eye visible posterior to center; a pair of nuchals; seven supralabials on both sides, fifth below center of the eye, first larger than the three following, sixth largest; two loreals, anterior (0.6 mm length, 0.6 mm height) slightly higher, the posterior (0.9 mm length, 0.5 mm height) longer; nasal divided, in contact with the first supralabial, rostral, anterior loreal, and supranasal; nostril in center of nasal; eight supraciliaries, first largest; one enlarged anterior temporal in contact with the sixth and the seventh supralabial; two posterior temporals, lower one smaller and overlapping the upper one, the lower temporal in contact with seventh supralabials; six infralabials, first two in contact with postmental; two pairs of chin shields, first pair medially in contact with each other; ear-opening moderately small, the anterior border with two lobules. Body elongate (AGD/SVL = 0.56); dorsal scales smooth, not larger than lateral and ventral scales, with eight longitudinal rows on the back; 28 midbody scale rows; 71 paravertebral scales; 35 axilla-groin scales; ventral scales smooth, in 55 rows; limbs short (FLL/SVL = 0.18, HLL/SVL = 0.25), pentadactyl, widely separated by ten scales when adpressed; 3, 6, 9, 9, and 7 smooth lamellae beneath fingers I���V, respectively; 4, 7, 13, 13, and 9 smooth lamellae beneath toes I���V, respectively; six enlarged precloacal scales, the two middle ones slightly larger than the others; size of median subcaudal series on the original part of the tail equal to those of its adjacent rows. Coloration. In life, overall dorsal coloration reddish brown with two dorsolateral bright lines on anterior part of dorsum; each dorsal scale of four central rows with a black spot forming four distinct longitudinal lines on dorsum; ear and surrounding area orange; sides dark brown to black intermixed with white, brown and yellow spots; chest and belly yellow (Fig. 3). In preservation, color fades but the four longitudinal dorsal lines remain distinct; orange and yellow faded to cream or white; overall dorsal and lateral coloration black to dark brown with bright spots on lateral sides; venter cream. Variation in paratypes (n = 8). Most morphological characters of paratypes agree with those of the holotype. The following main characters vary: (1) midbody scale rows: vary from 27 to 30; (2) paravertebral and ventral scale rows: range from 55 to 57 and from 55 to 62, respectively; (3) axilla-groin scales: range from 34 to 37; (4) and nuchals: range from one to three scales. Table 3 summarizes variation in size and scalation of the type series. ...Continued on the next page ...Continued on the next page Field notes. All specimens were collected during the daytime, on sandy ground among rotting leaves in Earleaf acacia (Acacia auriculiformis A.Cunn. ex Benth.) plantation, secondary dipterocarp forest, and rubber plantation in lowland coastal areas (Fig. 5). The holotype and paratypes ITBCZ 5843 & 5844 were collected at 15:42, air temperature 30.3 oC, relative humidity 70.2 %; ITCBZ 7038 was collected at 13:30, air temperature 33 oC, relative humidity 75 %; ITBCZ 7046 was collected at 8:54, air temperature 30 oC, relative humidity 80 %; ITBCZ 7056, 7059 & 7070 were collected at 12:10, air temperature 32 oC, relative humidity 65%. To avoid capture, this species can dive into and stay under loose sand. Sexual dimorphism. Adult males mainly differ from adult females in body coloration in life and shape of tail base. Males have yellow color on chest and belly (vs. white, cream or yellowish color in females), orange around the ear (vs. dark brown in females), and distinct bright spots on lateral side (vs. pure dark brown or with small bright dots in females). Ventral side of tail base is slightly swollen in males and flat in females (Figs. 3, 4A���E). Juvenile has bright golden color on the dorsum with four clear dorsal longitudinal stripes (Fig. 4F). Distribution. The new species is currently known only from (1) Ho Tram, Xuyen Moc District, Ba Ria-Vung Tau Province and (2) La Gi Town, Binh Thuan Province, southern Vietnam (Fig. 1). Etymology. The new skink is named after the nation of Vietnam. We recommend ��� Vietnam Agile Skink��� and ���Th���n l���n ch��n ng���n vi���t nam��� as the common English and Vietnamese names of the new species, respectively. Comparisons. Subdoluseps vietnamensis sp. nov. differs morphologically from other species of the Lygosoma s.l. in Asia as follows: For Subdoluseps, S. vietnamensis sp. nov. differs from S. herberti, S. malayana, S. purthi, and S. samajaya by having a single frontoparietal scale (vs. paired) (Grismer 2019; Karin et al. 2018; Sharma 1977; Smith 1916). In addition, it differs from S. herberti by having a smaller adult size (maximum SVL = 48.7 mm vs. 54.0 mm), presence (vs. absence) of nuchal scales (1���3 vs. 0), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Smith 1916); from S. malayana by having a smaller adult size (maximum SVL = 48.7 mm vs. 65.4 mm), more supralabials (7 vs. 6) with the fifth below the center of the eye (vs. the fourth below the center of the eye), smooth dorsal scales (vs. dorsal with 3���5 strong keels) (Grismer et al. 2019); from S. purthi by having a smaller adult size (maximum SVL = 48.7 mm vs. 67.0 mm), fewer midbody scale rows (27���30 vs. 32���34), and more paravertebral scale rows (55���57 vs. 50) (Sharma 1977); from S. samajaya by having a smaller adult size (maximum SVL = 48.7 mm vs. 70.1 mm), fewer paravertebral scale rows (55���57 vs. 60 or 61), and smooth dorsal scales (vs. dorsal with 5 strong keels) (Karin et al. 2018). Subdoluseps vietnamensis sp. nov. is phenotypically most similar to S. bowringii and S. frontoparietale but can be distinguished from both based on a combination of characters. The new species differs from S. bowringii by having a single frontoparietal scale (vs. paired) (Geissler et al. 2011; Smith 1935; this study), tail length approximately equal to SVL (vs. tail longer than SVL) (mean TL/SVL = 1.04 vs. 1.27 [Smith 1935; this study]), and fewer subcaudal scales (64���74 vs. 76���81) (this study); from S. frontoparietale by having a tail length approximately equal to SVL (mean TL/SVL = 1.04 vs. tail longer than SVL, mean TL/SVL = 1.24 [Taylor 1962; this study, Table 4]), fewer subcaudal scales (64���74 vs. 78���80) [Taylor 1962; this study]), fewer axilla-groin scales (34���37 vs. 39���42) (this study), four distinct longitudinal black stripes on dorsum (vs. six distinct black stripes) (Taylor 1962; this study), and absence (vs. presence) of bright dorsolateral band extending from eye to tail base (Taylor 1962; this study, Fig. 6). From genus Lygosoma s.s., Subdoluseps vietnamensis sp. nov. differs from L. isodactylum, L. siamensis, L. tabonorum, and L. quadrupes (Linnaeus) by having supranasals in contact with each other (vs. not in contact) (Geissler et al. 2011, Heitz et al. 2016, Siler et al. 2018) and from L. corpulentum by having smaller adult size (maximum SVL = 48.7 mm vs. 170.0 mm), frontoparietal scale single (vs. paired), and fewer midbody scale rows (27���30 vs. 36���40) (Smith 1921; Geissler et al. 2011). For the genus Riopa, Subdoluseps vietnamensis sp. nov. differs from R. albopunctata, R. anguinua, R. goaensis, R. guentheri, R. popae Shreve, and R. punctata (Gmelin) by having frontoparietal scales single (vs. paired) (Geissler et al. 2012; Sharma 1976; Smith 1935); from R. lineata and R. lineolata by having fewer paravertebral scale rows (55���57 vs. 104���110 in R. lineata [Smith 1935] and 78���93 in R. lineolata [Siler et al. 2018]); and from R. vosmaeri by having hind-limb with 5 (vs. 4) toes (Gray 1939; Seetharamaraju et al. 2009). For remaining Asian species of the Lygosoma s.l. that were not confirmed by Freitas et al. (2019), Subdoluseps vietnamensis sp. nov. differs from L. boehmei Ziegler, Schmitz, Heidrich, Vu & Nguyen, L. opisthorhodum Werner, L. singha (Taylor), and L. veunsaiensis Geissler, Hartmann & Neang by having frontoparietal scale single (vs. paired) (Geissler et al. 2012; Werner 1910; Ziegler et al. 2007); from L. angeli (Smith) by having a smaller adult size (maximum SVL = 48.7 mm vs. 100.0 mm) and fewer paravertebral scale rows (55���57 vs. 107���115) (Smith 1937; Geissler et al. 2011); from L. haroldyoungi (Taylor) by having a smaller adult size (maximum SVL = 48.7 mm vs. 136.0 mm), fewer paravertebral scale rows (55���57 vs. 141���145), and fewer midbody scale rows (27���30 vs. 38���42) [Taylor 1962; Geissler et al. 2011]; from L. kinabatanganensis Grismer, Quah, Duzulkafly & Yambun by having a smaller adult size (maximum SVL = 48.7 mm vs. 141.0 mm), fewer paravertebral scale rows (55���57 vs. 98), and fewer midbody scale rows (27���30 vs. 42) (Grismer et al. 2018); from L. koratense Smith by having a smaller adult size (maximum SVL = 48.7 mm vs. 105.0 mm) and fewer midbody scale rows (27���30 vs. 32���34) [Smith 1935]; and from L. bampfyldei Bartlett, L. peninsulare Grismer, Quah, Duzulkafly & Yambun, and L. schneideri Werner by having fewer paravertebral scale rows (55���57 vs. 81���85 in L. bampfyldei [Grismer et al. 2018], 87 in L. peninsulare [Grismer et al. 2018], and 95 in L. schneideri [Grismer et al. 2018])., Published as part of Le, Manh Van, Nguyen, Vu Dang Hoang, Phan, Hoa Thi, Rujirawan, Attapol, Aowphol, Anchalee, Vo, Thi-Dieu-Hien, Murphy, Robert W. & Nguyen, Sang Ngoc, 2021, A new skink of the genus Subdoluseps Freitas, Datta-Roy, Karanth, Grismer & Siler, 2019 (Squamata: Scincidae) from southern Vietnam, pp. 257-274 in Zootaxa 4952 (2) on pages 263-270, DOI: 10.11646/zootaxa.4952.2.3, http://zenodo.org/record/4674021, {"references":["Taylor, E. H. (1962) New oriental reptiles. The University of Kansas Science Bulletin, 43, 209 - 263. https: // doi. org / 10.5962 / bhl. part. 13346","Grismer, L. L., Dzukafly, Z., Muin, M. A., Quah, E. S., Karin, B. R., Anuar, S. & Freitas, E. S. (2019) A new skink of the genus Subdoluseps (Hardwicke & Gray, 1828) from Peninsular Malaysia. Zootaxa, 4609 (2), 358 - 372. https: // doi. org / 10.11646 / zootaxa. 4609.2.10","Karin, B. R., Freitas, E. S., Shonleben, S., Grismer, L. L., Bauer, A. M. & Das, I. (2018) Unrealized diversity in an urban rainforest: A new species of Lygosoma (Squamata: Scincidae) from western Sarawak, Malaysia (Borneo). Zootaxa, 4370, 345 - 362. https: // doi. org / 10.11646 / zootaxa. 4370.4.2","Sharma, R. C. (1977) A new lizard of the genus Riopa Gray (Scincidae) from Tamil Nadu, India. Records Zoological Survey of India, 73, 41 - 42.","Smith, M. A. (1916) Description of three new lizards and a new snake from Siam. The journal of the Natural History Society of Siam, 2, 44 - 47.","Geissler, P., Nguyen, Q. T., Phung, M. T., Devender, R. W. T., Hartmann, T., Farkas, B., Ziegler, T. & Bohme, W. (2011) A review of Indochinese skinks of the genus Lygosoma Hardwicke & Gray, 1827 (Squamata: Scincidae), with natural history notes and an identification key. Biologia, 66, 1159 - 1176. https: // doi. org / 10.2478 / s 11756 - 011 - 0130 - 2","Smith, M. A. (1935) The fauna of British India, Including Ceylon and Burma. Reptiles and Amphibia, Vol. II. Sauri a. Taylor and Francis, London, 440 pp.","Heitz, B. B., Diesmos, A. C., Freitas, E. S., Ellsworth, E. D., Grismer, L. L., Aowphol, A., Brown, R. M. & Siler, C. D. (2016) New supple skink, genus Lygosoma (Reptilia: Squamata: Scincidae) from the Palawan Faunal Region, Philippines. Herpetologica, 72, 352 - 361. https: // doi. org / 10.1655 / Herpetologica-D- 16 - 00023.1","Siler, C. D., Heitz, B. B., Davis, D. R., Freitas, E. S., Aowphol, A., Termprayoon, K. & Grismer, L. L. (2018) New Supple Skink, Genus Lygosoma (Reptilia: Squamata: Scincidae), from Indochina and Redescription of Lygosoma quadrupes (Linnaeus, 1766). Journal of Herpetology, 52, 332 - 347. https: // doi. org / 10.1670 / 16 - 064","Smith, M. A. (1921) New or little - known Reptiles and Batrachians from Southern Annam (Indochina). Proceedings of the Zoological Society of London, 423 - 440. https: // doi. org / 10.1111 / j. 1096 - 3642.1921. tb 03271. x","Geissler, P., Hartmann, T. & Neang, T. (2012) A new species of the genus Lygosoma Hardwicke & Gray, 1827 (Squamata: Scincidae) from northeastern Cambodia, with an updated identification key to the genus Lygosoma in mainland Southeast Asia. Zootaxa, 3190 (1), 56 - 68. https: // doi. org / 10.11646 / zootaxa. 3190.1.4","Sharma, R. C. (1976) Records of the reptiles of Goa. Records Zoological Survey of India, 71 (1975), 149 - 167.","Seetharamaraju, M., Sreekar, R., Srinivasulu, C., Srinivasulu, B., Kaur, H. & Venkateshwarlu, P. (2009) Rediscovery of Vosmer's Writhing Skink Lygosoma vosmaerii (Gray, 1839) (Reptilia: Scincidae) with a note on its taxonomy. Journal of Threatened Taxa, 1, 624 - 626. https: // doi. org / 10.11609 / JoTT. o 2160.624 - 6","Freitas, E. S., Datta-Roy, A., Karanth, P., Grismer, L. L. & Siler, C. D. (2019) Multilocus phylogeny and a new classification for African, Asian and Indian supple and writhing skinks (Scincidae: Lygosominae). Zoological Journal of the Linnean Society, 186, 1067 - 1096. https: // doi. org / 10.1093 / zoolinnean / zlz 001","Werner, F. (1910) Uber neue oder seltene Reptilien des Naturhistorischen Museums in Hamburg. II. Eidechsen. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 2, 1 - 46.","Ziegler, T., Schmitz, A., Heidrich, A., Vu, N. T. & Nguyen, Q. T. (2007) A new species of Lygosoma (Squamata: Sauria: Scincidae) from the central Truong Son, Vietnam, with notes on its molecular phylogenetic position. Revue Suisse de Zoologie, 114, 397 - 415. https: // doi. org / 10.5962 / bhl. part. 80396","Smith, M. A. (1937) Un nouveau Lezard de Cochinchine. Bulletin du Museum National d'Histoire Naturelle, 9, 366.","Grismer, L. L., Quah, E. S. H., Zaharil, D. & Paul, Y. (2018) On the taxonomy of Lygosoma bampfyldei Bartlett, 1895 (Squamata: Scincidae) with descriptions of new species from Borneo and Peninsular Malaysia and the resurrection of Lygosoma schneideri Werner, 1900. Zootaxa, 4438, 528 - 550. https: // doi. org / 10.11646 / zootaxa. 4438.3.6"]}

Grismer, L. Lee, Rujirawan, Attapol, Termprayoon, Korkhwan, Ampai, Natee, Yodthong, Siriporn, Wood, Perry L., Oaks, Jamie R., Aowphol, Anchalee (2020): A new species of Cyrtodactylus Gray (Squamata; Gekkonidae) from the Thai Highlands with a discussion on the evolution of habitat preference. Zootaxa 4852 (4): 401-427, DOI: https://doi.org/10.11646/zootaxa.4852.4.1

Cyrtodactylus maelanoi sp. nov. Mae La Noi Bent-toed Gecko (Figs. 5���7) Holotype. Adult male ZMKU R 00857 (field tag AA 03726) collected on 22 March 2017 at 2100 hrs by Piyawan Puanprapai, Attapol Rujirawan, Siriporn Yodthong, Natee Ampai, and Elyse S. Freitas from the Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province, Thailand (18.34223��N, 98.02317��E WGS; 991 m in elevation). Paratypes. The paratypes ZMKU R 00852���00856 (field tag AA 03721���25) bear the same data as the holotype. The remaining paratypes ZMKU R 00858���00860 (field tag AA 06195���97) bear the same locality data as the holotype but were collected by Attapol Rujirawan, Siriporn Yodthong, Korkwan Termprayoon, Natee Ampai on 13 June 2018. Diagnosis. Cyrtodactylus maelanoi sp. nov. differs from all species in the C. sinyineensis group by having the combination of 7���9 supralabials; six or seven infralabials; 29���37 paravertebral tubercles; 16���19 longitudinal rows of dorsal tubercles; 27���33 ventral scales ventral scales; 9���12 expanded subdigital lamellae on the fourth toe; 11���14 unmodified subdigital lamellae on the fourth toe; 22���24 total subdigital lamellae on the fourth toe; 24���28 enlarged femoral scales; a total of 8���13 pore-bearing femoral scales in males; 4���8 enlarged precloacal scales; four or five pore-bearing precloacal scales in males; three rows of enlarged post-precloacal scales; approximately five or six jagged dorsal body bands; 11 light-colored caudal bands (n=2); 11 dark-colored caudal bands (n=2); raised and strongly keeled dorsal tubercles that extend beyond base of tail; enlarged femoral and precloacal scales nearly the same size and continuous; pore-bearing femoral and precloacal scales not continuous; medial subcaudals two to three times wider than long and not extending onto lateral side of tail; iris reddish; nuchal loop lacking an anterior azygous notch, and bearing a jagged posterior border; dorsal bands not bearing paravertebral elements, wider than interspaces, bearing lightened centers, edged with white tubercles; dark markings in dorsal interspaces; dark caudal bands wider than light caudal bands; light caudal bands in adults bearing dark-colored markings; light-colored caudal bands not encircling tail; and mature regenerated tail not spotted (Table 5). Description of holotype. Adult male SVL 75.2 mm (Fig. 5); head moderate in length (HL/SVL 0.28), wide (HW/HL 0.73), flat (HD/HL 0.39), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.39), rounded in dorsal profile, broad in lateral profile; eye large (ED/HL 0.27); ear opening oval (EL/HL 0.11); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by supranasals and one internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasals, posteriorly by two postnasals, and ventrally by first supralabials; 8(R,L) rectangular supralabials extending to below midpoint of eye; 6(R,L) infralabials tapering posteriorly to commissure of jaw; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with small tubercles; dorsal superciliaries weakly pointed and directed laterally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals which contact medially for 50% of their length posterior to mental; one row of enlarged chinshields, outermost row bordering first four infralabials; gular and throat scales granular, grading posteriorly into larger, subimbricate pectoral and ventral scales. Body relatively short (AG/SVL 0.39) with well-defined ventrolateral folds; dorsal scales small, raised and interspersed with large, raised, semi-regularly arranged, strongly keeled tubercles; tubercles extend from top of head onto base of tail just beyond the postcloacal swelling; tubercles on nape smaller than those on body; 30 paravertebral tubercles; approximately 17 longitudinal rows of dorsal tubercles; 28 flat, subimbricate, ventral scales larger than dorsal scales; eight enlarged precloacal scales; five pore-bearing precloacal scales not separated on the midline by a poreless scale; three rows of large, post-precloacal scales; and no deep precloacal groove or depression. Forelimbs moderate in stature, relatively short (FL/SVL 0.17); slightly raised, juxtaposed scales of forearm larger than those on body, intermixed with large tubercles; palmar scales slightly raised, juxtaposed; digits welldeveloped, relatively long, inflected at basal, interphalangeal joints; digits narrow distal to inflections; widened proximal subdigital lamellae extend onto palm; slight webbing at base of digit; claws well-developed, sheathed by a dorsal and ventral scale at base; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.21), covered dorsally by small, raised, juxtaposed scales intermixed with large pointed tubercles and bearing flat, slightly larger imbricate scales anteriorly; ventral femoral scales flat, imbricate, much larger than dorsals; one row of 14(R)13(L) enlarged femoral scales and enlarged precloacal scales continuous; enlarged femoral scales nearly equal in size; small, postfemoral scales form an abrupt union with larger, flat ventral scales on posteroventral margin of thigh; 5,6(R,L) pore-bearing femoral scales not continuous with pore-bearing precloacal scales; subtibial scales flat, imbricate; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 11(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that do not extend onto sole, 13(R,L) unmodified subdigital lamellae distal to inflection; and claws well-developed, base of claw sheathed by a dorsal and ventral scale. Tail nearly complete, gracile in proportions, 115.1 mm in length, 8.1 mm in width at base, tapering to a point, TL/SVL (1.42); dorsal scales of tail flat, forming indistinct whorls; median row of transversely expanded subcaudal scales three times as wide as long, not extending onto lateral subcaudal region; three enlarged postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales large, flat. Color pattern (Figs. 5, 6). Dorsal ground color of head, body, limbs, and tail yellowish-brown; top of head and rostrum nearly unicolor, bearing areas of slightly darker, diffuse irregularly shaped markings; nuchal loop smooth posteriorly with two posterior projections, not divided medially; approximately five dark jagged body bands bearing lightened centers, lacking paravertebral elements, edged with whitish tubercles extend from the shoulder to the presacral region; lighter colored interspaces between bands bear darker markings; whitish tubercles scattered on flanks; sacral and postsacral bands continue onto the tail to form five black caudal bands that are wider than the five light-colored caudal bands; light-colored caudal bands bear dark markings and do not encircle tail; limbs bear distinct, dark-colored irregularly shaped markings; posterior one-third of tail regenerated with dark mottled pattern; gular scales bearing only two or three black stipples; black stippling in throat, pectoral region, and anterior portion of belly more dense; subcaudal region darkly mottled, posterior one-third grey with faint mottling. Variation. The paratypes closely resemble the holotype in dorsal banding and nuchal loop pattern (Fig. 6). The nape band are more prominent in ZMKU R 00858���00860 and ZMKU R 00855. Paratypes ZMKU R 00858���00860, 00852, 00855���00857 are darker overall. Paratypes ZMKU R 00853, 00859���00860 have regenerated tails. Paratype ZMKU R 00854 is missing the posterior one-thrid to one-half of the tail. Additional variation in meristic and mensural characters are presented in Table 6. Distribution. Cyrtodactylus maelanoi sp. nov. is known only from the type locality from the Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province, Thailand (18.34223��N, 98.02317��E WGS; 991 m in elevation; Fig. 1). Etymology. The specific epithet ��� maelanoi ��� is a toponym of the type locality Mae La Noi. The recommended vernacular name in English is Mae La Noi Bent-toed Gecko. Natural history. All lizards observed were abroad at night (Fig. 7) in the vicinity of a waterfall (Fig. 8). Lizards were found on the ground, the walls of a building, and on granite rocks and within their cracks. Lizards were also seen on tree trunks as high 70���200 cm above the ground and on a twig approximately 160 cm above the ground. These observations clearly indicate that Cyrtodactylus maelanoi sp. nov. is a habitat generalist. No hatchlings of gravid females were observed indicating that at least the period of March through June is probably outside the reproductive season of this species. characters Clade 2 Clade 1 amphipetraeus aequalis bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. inthanon sinyineensis taungwineensis welpyanensis doisuthep Supralabial scales (SL)mean (��SD) Range9.0 (0.00) 98.1 (0.46) 7���98.0 (0.84) 7���99.4 (0.71) 8���119.0 (0.00) 98.3 (0.58) 8 or 98.3 (0.58) 8 or 98.1 (0.60) 7���911.0 (1.00) 10���129.7 (0.57) 9 or 109.1 (0.86) 7���108.3 (0.6) 8 or 911.0 (1.0) 10���12n9375243339331433Infralabial scales (IL)Mean (��SD) Range7.0 (0.00) 76.8 (0.53) 6���86.2 (0.44) 6 or 77.9 (0.50) 7���97.7 (0.58) 7 or 86.3 (0.57) 6 or 77.0 (0.00) 76.3 (0.50) 6 or 79.3 (0.58) 9 or 108 (0.00) 87.00 (0.39) 6���87.0 (0.00) 79.5 (1.5) 8���11n9375243339331433Paravertebral tubercles (PV)Mean34.732.026.032.531.734. 034.331.1/33.730.331.7/(��SD) Range(0.50) 34 or 35(1.64) 29���36(0.71) 25���27(0.98) 31���36(1.15) 31���33(0.00) 33���35(0.58) 34 or 35(2.37) 29���37/ /(0.00) 33���35(0.84) 29���32(1.5) 30���33/n9375243339/3143/Longitudunal rows of body tubercles (LT)Mean (��SD) Range17.9 (0.78) 17���1920.6 (1.42) 18���2317.4 (01.14) 16���1919.5 (1.47) 17���2214.3 (1.15) 13���1519.0 (1.00) 18���2017.3 (1.54) 16���1815.5 (0.87) 16���1919. 0 (1.00) 18���2015 (0.00) 1518.7 (0.61) 18���2016.0 (0.00) 1619.7 (0.58) 19 or 20n9375243339331433Ventral scales (VS)Mean (��SD) Range28.3 (0.71) 28���3024.5 (1.68) 22���3125.6 (1.52) 24���2825.4 (1.10) 23���2726.7 (1.53) 25���2825.3 (0.58) 25 or 2627.0 (0.00) 2728.4 (1.94) 27���3330.7 (2.89) 29���3428 (0.00) 27���2932.3 (2.02) 30���3629.3 (1.20) 28���3032.0 (3.00) 29���35n9375243339331433nov ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page Comparisons. Cyrtodactylus maelanoi sp. nov. (n=9) differs from all other species of the C. sinyineensis based on various combinations of character states (Table 5). In clade 1, it differs from C. inthanon, C. sinyineensis, and C. taungwineensis in having significantly fewer supralabials and from these species plus C. welpyanensis in having significantly fewer infralabials (Fig. 4; Table 4). It differs further from C. sinyineensis, C. taungwineensis, and C. welpyanensis having significantly fewer precloacal scales; from C. sinyineensis it differs significantly in having fewer enlarged femoral scales; it differs further from C. taungwineensis by having significantly fewer ventral scales (Fig. 4; Table 4). Cyrtodactylus maelanoi sp. nov. may differ further from its sister species C. inthanon in having 24���28 enlarged femoral scales versus 29���32 in C. inthanon and they plot completely separate in the PCA and DAPC analyses (Fig. 3). Increases in sample sizes may indicate that some of these character differences are not statistically significant whereas other differences may be statistically significant. Differences in color pattern among all species of the C. sinyineensis group are listed in Table 5., Published as part of Grismer, L. Lee, Rujirawan, Attapol, Termprayoon, Korkhwan, Ampai, Natee, Yodthong, Siriporn, Wood, Perry L., Oaks, Jamie R. & Aowphol, Anchalee, 2020, A new species of Cyrtodactylus Gray (Squamata; Gekkonidae) from the Thai Highlands with a discussion on the evolution of habitat preference, pp. 401-427 in Zootaxa 4852 (4) on pages 410-424, DOI: 10.11646/zootaxa.4852.4.1, http://zenodo.org/record/4410011

Cyrtodactylus maelanoi sp. nov. Mae La Noi Bent-toed Gecko (Figs. 5–7) Holotype. Adult male ZMKU R 00857 (field tag AA 03726) collected on 22 March 2017 at 2100 hrs by Piyawan Puanprapai, Attapol Rujirawan, Siriporn Yodthong, Natee Ampai, and Elyse S. Freitas from the Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province, Thailand (18.34223°N, 98.02317°E WGS; 991 m in elevation). Paratypes. The paratypes ZMKU R 00852–00856 (field tag AA 03721–25) bear the same data as the holotype. The remaining paratypes ZMKU R 00858–00860 (field tag AA 06195–97) bear the same locality data as the holotype but were collected by Attapol Rujirawan, Siriporn Yodthong, Korkwan Termprayoon, Natee Ampai on 13 June 2018. Diagnosis. Cyrtodactylus maelanoi sp. nov. differs from all species in the C. sinyineensis group by having the combination of 7–9 supralabials; six or seven infralabials; 29–37 paravertebral tubercles; 16–19 longitudinal rows of dorsal tubercles; 27–33 ventral scales ventral scales; 9–12 expanded subdigital lamellae on the fourth toe; 11–14 unmodified subdigital lamellae on the fourth toe; 22–24 total subdigital lamellae on the fourth toe; 24–28 enlarged femoral scales; a total of 8–13 pore-bearing femoral scales in males; 4–8 enlarged precloacal scales; four or five pore-bearing precloacal scales in males; three rows of enlarged post-precloacal scales; approximately five or six jagged dorsal body bands; 11 light-colored caudal bands (n=2); 11 dark-colored caudal bands (n=2); raised and strongly keeled dorsal tubercles that extend beyond base of tail; enlarged femoral and precloacal scales nearly the same size and continuous; pore-bearing femoral and precloacal scales not continuous; medial subcaudals two to three times wider than long and not extending onto lateral side of tail; iris reddish; nuchal loop lacking an anterior azygous notch, and bearing a jagged posterior border; dorsal bands not bearing paravertebral elements, wider than interspaces, bearing lightened centers, edged with white tubercles; dark markings in dorsal interspaces; dark caudal bands wider than light caudal bands; light caudal bands in adults bearing dark-colored markings; light-colored caudal bands not encircling tail; and mature regenerated tail not spotted (Table 5). Description of holotype. Adult male SVL 75.2 mm (Fig. 5); head moderate in length (HL/SVL 0.28), wide (HW/HL 0.73), flat (HD/HL 0.39), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.39), rounded in dorsal profile, broad in lateral profile; eye large (ED/HL 0.27); ear opening oval (EL/HL 0.11); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by supranasals and one internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasals, posteriorly by two postnasals, and ventrally by first supralabials; 8(R,L) rectangular supralabials extending to below midpoint of eye; 6(R,L) infralabials tapering posteriorly to commissure of jaw; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with small tubercles; dorsal superciliaries weakly pointed and directed laterally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals which contact medially for 50% of their length posterior to mental; one row of enlarged chinshields, outermost row bordering first four infralabials; gular and throat scales granular, grading posteriorly into larger, subimbricate pectoral and ventral scales. Body relatively short (AG/SVL 0.39) with well-defined ventrolateral folds; dorsal scales small, raised and interspersed with large, raised, semi-regularly arranged, strongly keeled tubercles; tubercles extend from top of head onto base of tail just beyond the postcloacal swelling; tubercles on nape smaller than those on body; 30 paravertebral tubercles; approximately 17 longitudinal rows of dorsal tubercles; 28 flat, subimbricate, ventral scales larger than dorsal scales; eight enlarged precloacal scales; five pore-bearing precloacal scales not separated on the midline by a poreless scale; three rows of large, post-precloacal scales; and no deep precloacal groove or depression. Forelimbs moderate in stature, relatively short (FL/SVL 0.17); slightly raised, juxtaposed scales of forearm larger than those on body, intermixed with large tubercles; palmar scales slightly raised, juxtaposed; digits welldeveloped, relatively long, inflected at basal, interphalangeal joints; digits narrow distal to inflections; widened proximal subdigital lamellae extend onto palm; slight webbing at base of digit; claws well-developed, sheathed by a dorsal and ventral scale at base; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.21), covered dorsally by small, raised, juxtaposed scales intermixed with large pointed tubercles and bearing flat, slightly larger imbricate scales anteriorly; ventral femoral scales flat, imbricate, much larger than dorsals; one row of 14(R)13(L) enlarged femoral scales and enlarged precloacal scales continuous; enlarged femoral scales nearly equal in size; small, postfemoral scales form an abrupt union with larger, flat ventral scales on posteroventral margin of thigh; 5,6(R,L) pore-bearing femoral scales not continuous with pore-bearing precloacal scales; subtibial scales flat, imbricate; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 11(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that do not extend onto sole, 13(R,L) unmodified subdigital lamellae distal to inflection; and claws well-developed, base of claw sheathed by a dorsal and ventral scale. Tail nearly complete, gracile in proportions, 115.1 mm in length, 8.1 mm in width at base, tapering to a point, TL/SVL (1.42); dorsal scales of tail flat, forming indistinct whorls; median row of transversely expanded subcaudal scales three times as wide as long, not extending onto lateral subcaudal region; three enlarged postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales large, flat. Color pattern (Figs. 5, 6). Dorsal ground color of head, body, limbs, and tail yellowish-brown; top of head and rostrum nearly unicolor, bearing areas of slightly darker, diffuse irregularly shaped markings; nuchal loop smooth posteriorly with two posterior projections, not divided medially; approximately five dark jagged body bands bearing lightened centers, lacking paravertebral elements, edged with whitish tubercles extend from the shoulder to the presacral region; lighter colored interspaces between bands bear darker markings; whitish tubercles scattered on flanks; sacral and postsacral bands continue onto the tail to form five black caudal bands that are wider than the five light-colored caudal bands; light-colored caudal bands bear dark markings and do not encircle tail; limbs bear distinct, dark-colored irregularly shaped markings; posterior one-third of tail regenerated with dark mottled pattern; gular scales bearing only two or three black stipples; black stippling in throat, pectoral region, and anterior portion of belly more dense; subcaudal region darkly mottled, posterior one-third grey with faint mottling. Variation. The paratypes closely resemble the holotype in dorsal banding and nuchal loop pattern (Fig. 6). The nape band are more prominent in ZMKU R 00858–00860 and ZMKU R 00855. Paratypes ZMKU R 00858–00860, 00852, 00855–00857 are darker overall. Paratypes ZMKU R 00853, 00859–00860 have regenerated tails. Paratype ZMKU R 00854 is missing the posterior one-thrid to one-half of the tail. Additional variation in meristic and mensural characters are presented in Table 6. Distribution. Cyrtodactylus maelanoi sp. nov. is known only from the type locality from the Tha Pha Pum Subdistrict, Mae La Noi District, Mae Hong Son Province, Thailand (18.34223°N, 98.02317°E WGS; 991 m in elevation; Fig. 1). Etymology. The specific epithet “ maelanoi ” is a toponym of the type locality Mae La Noi. The recommended vernacular name in English is Mae La Noi Bent-toed Gecko. Natural history. All lizards observed were abroad at night (Fig. 7) in the vicinity of a waterfall (Fig. 8). Lizards were found on the ground, the walls of a building, and on granite rocks and within their cracks. Lizards were also seen on tree trunks as high 70–200 cm above the ground and on a twig approximately 160 cm above the ground. These observations clearly indicate that Cyrtodactylus maelanoi sp. nov. is a habitat generalist. No hatchlings of gravid females were observed indicating that at least the period of March through June is probably outside the reproductive season of this species. characters Clade 2 Clade 1 amphipetraeus aequalis bayinnyiensis chaunghanak- waensis dammathetensis dattkyaikensis naungkayain- gensis maelanoi sp. inthanon sinyineensis taungwineensis welpyanensis doisuthep Supralabial scales (SL)mean (±SD) Range9.0 (0.00) 98.1 (0.46) 7–98.0 (0.84) 7–99.4 (0.71) 8–119.0 (0.00) 98.3 (0.58) 8 or 98.3 (0.58) 8 or 98.1 (0.60) 7–911.0 (1.00) 10–129.7 (0.57) 9 or 109.1 (0.86) 7–108.3 (0.6) 8 or 911.0 (1.0) 10–12n9375243339331433Infralabial scales (IL)Mean (±SD) Range7.0 (0.00) 76.8 (0.53) 6–86.2 (0.44) 6 or 77.9 (0.50) 7–97.7 (0.58) 7 or 86.3 (0.57) 6 or 77.0 (0.00) 76.3 (0.50) 6 or 79.3 (0.58) 9 or 108 (0.00) 87.00 (0.39) 6–87.0 (0.00) 79.5 (1.5) 8–11n9375243339331433Paravertebral tubercles (PV)Mean34.732.026.032.531.734. 034.331.1/33.730.331.7/(±SD) Range(0.50) 34 or 35(1.64) 29–36(0.71) 25–27(0.98) 31–36(1.15) 31–33(0.00) 33–35(0.58) 34 or 35(2.37) 29–37/ /(0.00) 33–35(0.84) 29–32(1.5) 30–33/n9375243339/3143/Longitudunal rows of body tubercles (LT)Mean (±SD) Range17.9 (0.78) 17–1920.6 (1.42) 18–2317.4 (01.14) 16–1919.5 (1.47) 17–2214.3 (1.15) 13–1519.0 (1.00) 18–2017.3 (1.54) 16–1815.5 (0.87) 16–1919. 0 (1.00) 18–2015 (0.00) 1518.7 (0.61) 18–2016.0 (0.00) 1619.7 (0.58) 19 or 20n9375243339331433Ventral scales (VS)Mean (±SD) Range28.3 (0.71) 28–3024.5 (1.68) 22–3125.6 (1.52) 24–2825.4 (1.10) 23–2726.7 (1.53) 25–2825.3 (0.58) 25 or 2627.0 (0.00) 2728.4 (1.94) 27–3330.7 (2.89) 29–3428 (0.00) 27–2932.3 (2.02) 30–3629.3 (1.20) 28–3032.0 (3.00) 29–35n9375243339331433nov ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page ......continued on the next page Comparisons. Cyrtodactylus maelanoi sp. nov. (n=9) differs from all other species of the C. sinyineensis based on various combinations of character states (Table 5). In clade 1, it differs from C. inthanon, C. sinyineensis, and C. taungwineensis in having significantly fewer supralabials and from these species plus C. welpyanensis in having significantly fewer infralabials (Fig. 4; Table 4). It differs further from C. sinyineensis, C. taungwineensis, and C. welpyanensis having significantly fewer precloacal scales; from C. sinyineensis it differs significantly in having fewer enlarged femoral scales; it differs further from C. taungwineensis by having significantly fewer ventral scales (Fig. 4; Table 4). Cyrtodactylus maelanoi sp. nov. may differ further from its sister species C. inthanon in having 24–28 enlarged femoral scales versus 29–32 in C. inthanon and they plot completely separate in the PCA and DAPC analyses (Fig. 3). Increases in sample sizes may indicate that some of these character differences are not statistically significant whereas other differences may be statistically significant. Differences in color pattern among all species of the C. sinyineensis group are listed in Table 5.

Rujirawan, Attapol, Fong, Jonathan J., Ampai, Natee, Yodthong, Siriporn, Termprayoon, Korkhwan, Aowphol, Anchalee (2019): A new karst-dwelling gecko of the Gekko petricolus group (Reptilia: Gekkonidae) from Phitsanulok Provinceı central Thailand. Journal of Natural History 53 (9): 557-576, DOI: 10.1080/00222933.2019.1597937

Odorrana livida (Blyth, 1856) (Figures 2, 3, 4, 5) Polypedates lividus Blyth, 1856: 718 Odorrana livida Fei, Ye and Huang, 1990: 148., Published as part of Rujirawan, Attapol, Stuart, Bryan L. & Aowphol, Anchalee, 2018, Expanded description of Odorrana livida (Blyth, 1856) with notes on its natural history in Thailand, pp. 1581-1600 in Journal of Natural History 52 (21 - 24) on page 1584, DOI: 10.1080/00222933.2018.1481236, http://zenodo.org/record/5174940, {"references":["Blyth E. 1856. Report for October meeting, 1855. J Asiat Soc Bengal. 24: 711 - 723.","Fei L, Ye CY, Huang YZ. 1990. Key to Chinese Amphibia. Chongqing: Chongqing Branch, Publishing House of Science and Technology Materials."]}

Limnonectes coffeatus sp. nov. Holotype: NCSM 77788 (field tag BLS 13957), adult male (Fig. 2), Laos, Champasak Province, Pakxong District, Bolaven Plateau, Dong Hua Sao National Protected Area, Houay Tad Seua Stream, 15.06237°N 106.21075°E, 1,235 m elev., coll. 3 August 2010 by Bryan L. Stuart, Niane Sivongxay, and Sengvilay Seateun. Paratypes: NCSM 77786, adult female, same data as holotype except coll. 30 July 2010. NCSM 77785, NCSM 77787 (Fig. 3–4), two adult females; NUOL 0 0 0 60, one immature female; same data as holotype except coll. 15.06515°N 106.21394°E, 1,245 m elev., 28 July–01 August 2010. FMNH 258440, immature female; FMNH 258441, FMNH 258530, two immature males, same data as holotype except coll. near 15.06528°N 106.21750°E, 1,200 m elev., coll. 22–23 September 1999 by Bryan L. Stuart and Harold F. Heatwole. Referred Specimens: NCSM 77944, larvae (n =13; Fig. 5), same data as holotype except coll. 30 July 2010. Etymology: The specific epithet refers to the coffee plant genus Coffea, in reference to the extensive coffee plantations that are encroaching into the natural habitat of the new species at upper elevations on the Bolaven Plateau of southern Laos. Diagnosis: Assigned to the genus Limnonectes on the basis of its inferred phylogenetic position (Fig. 6), the presence of fang-like odontoid processes on the lower jaw (Emerson et al. 2000; Lambertz et al. 2014), and males with hypertrophied heads (Lambertz et al. 2014). A small-sized Limnonectes having the combination of the single available adult male with SVL 37.9, adult females with SVL 38.1–42.1; male lacking postorbital caruncle; male with hypertrophied head; odontoid processes on anterior margin of lower jaw, larger in male than in females; male with horizontal diameter of tympanum greater than that of eye, females with horizontal diameter of tympanum less than or equal to that of eye; tubercles on dorsum enlarged and rounded, but not arranged in rows; interdigital webbing of foot present; throat uniformly gray in male; tympanum uniformly colored, without distinct dark marking on upper half; flank without distinct spots; and ova with pigmented poles. Description of holotype: Habitus moderately stocky; body tapering to groin. Head broad and depressed; head width greater than head length. Snout obtusely pointed in dorsal view, round in profile, projecting well beyond lower jaw in profile; nostril dorsolateral, much closer to tip of snout than to eye, below canthus; internarial distance greater than interorbital distance; canthus rostralis indistinct, rounded, slightly constricted behind nostrils; lores concave, oblique; eye diameter 55% of snout length, interorbital distance greater than upper eyelid width; pineal ocellus visible; tympanum round, not elevated from side of head, annulus visible, tympanum diameter approximately 117% eye diameter and greater than distance between tympanum and eye; small, slit-like vocal sac openings on floor of mouth near lateral margin of tongue; vomerine teeth on two oblique ridges, closer to choanae than to each other; two large odontoid processes at front of mandible; median triangular protuberance at mandibular symphysis. Forelimbs moderately robust. Fingers relatively slender, without webbing or skin flaps; tip of fingers rounded, weakly expanded into discs; relative finger lengths II = IV Hindlimbs moderately robust. Toes relatively slender; tips of toes rounded, expanded into small discs; relative toe lengths I Skin on dorsum shagreened with large round tubercles, not elongated or arranged in rows; tubercular ridges from posterior margin of eyelid to occiput forming “W”-shape; caruncle absent from interorbital region; distinct supratympanic ridge from posterior corner of eye to axilla; rictal gland absent; dorsolateral fold absent; skin on throat wrinkled, that on remaining ventral surfaces smooth. Color of holotype in life: Dorsum brown with ochre wash; larger tubercles on dorsal surface of head and back orange-red with black margins, those on occiput forming “W”-shape; dark brown triangular marking between eyes with yellow anterior margin; dark brown alternating white-gray bands on upper and lower lips; dorsal surfaces of limbs with indistinct gray-brown bands; dorsal surfaces of thighs with yellowish-wash. Chin and throat gray; ventral surfaces of chest, belly and limbs yellow; ventral surfaces of hands and feet brown; iris bronze. Description of larvae NCSM 77944: Based on Gosner stage 28 individual in series of thirteen larvae (Fig. 5). Body oval in dorsal view, slightly compressed dorsoventrally, maximum body width slightly anterior to level of spiracle. Nares dorsal, without raised rim. Eyes dorsolateral, not visible from below. Spiracular tube single, sinistral on left side, angled slightly dorsally, aperture near midline of side and projecting posteriorly, approximately midway between eye and end of body. Tail slender, tapering gradually in distal one-third to rounded tip, origins of dorsal and ventral fins at end of body, dorsal fin only slightly deeper than ventral fin. Oral disk ventral, subterminal, width about 36% maximum width of body. Anterior labium with single row of papillae on lateral margins; posterior labium with two staggered rows of papillae on lateral and posterior margins; papillae homogenous in length. Labial tooth row formula 2(2)/3(1). All posterior tooth rows subequal in length. Upper and lower jaw sheaths black with serrated margins, upper sheath without median convexity. No distinct cluster of glandules posterior to eye on dorsal surface of body, subequal to interorbital distance. Color in life brown with gold flecking on belly and distinct black spots on dorsal surface of body, caudal muscle, and dorsal and ventral tail fins. In preservative, brown faded to light brown, black faded to dark brown, gold flecking absent. Measurements of larvae are summarized in Table 3. Variation: Females have narrower heads in dorsal view than holotype male (Fig. 4). Females have relatively smaller tympana than holotype male, with the horizontal diameter of the tympanum less than or equal to that of the eye. Females have smaller odontoid processes at front of mandible than holotype male. Female NCSM 77787 has wider black margins on larger tubercles on dorsal surface of head and back than holotype, and with black rather than brown or gray-brown bands on interorbital region, lips, and dorsal surfaces of limbs. Females lack wrinkled skin and have paler coloration on throat than holotype male. Females contain ova with pigmented poles. Measurements of adults are summarized in Table 2. SVL37.938.1–42.1; 40.5 ± 2.141.137.6–46.4; 40.9 ± 2.936.0–41.2; 38.1 ± 2.2HDL17.915.8–17.7; 16.9 ± 1.019.816.0–21.4; 18.9 ± 1.614.4–16.8; 15.7 ± 0.9HDW18.615.2–17.3; 16.6 ± 1.218.017.6–23.2; 20.0 ± 1.813.1–17.7; 15.9 ± 1.7SNT7.65.9–7.3; 6.8 ± 0.86.76.1–9.0; 7.4 ± 0.85.6–6.9; 6.5 ± 0.5EYE4.23.6–4.4; 4.1 ± 0.45.54.3–5.7; 5.0 ± 0.44.2–5.1; 4.7 ± 0.3IOD3.83.2–3.3; 3.3 ± 0.14.44.0–6.2; 5.1 ± 0.72.6–4.6; 3.7 ± 0.7TMP4.93.5–4.3; 3.9 ± 0.45.34.4–7.3; 5.6 ± 0.92.8–3.8; 3.4 ± 0.4IND4.64.3–4.6; 4.5 ± 0.24.84.3–6.2; 5.3 ± 0.63.6–5.2; 4.6 ± 0.6SHK20.219.4–20.0; 19.7 ± 0.320.517.6–22.1; 20.4 ± 1.317.6–20.5; 19.5 ± 1.2TGH20.319.4–21.0; 20.5 ± 0.918.018.9–22.9; 20.9 ± 1.418.5–20.7; 20.0 ± 0.9LAL7.27.9–8.4; 8.2 ± 0.3n/a7.5–8.7; 8.0 ± 0.46.7–8.0; 7.6 ± 0.6HND10.510.0–11.0; 10.4 ± 0.5n/a9.1–12.0; 10.6 ± 0.88.5–10.9; 10.0 ± 0.9FTL19.318.7–20.7; 19.9 ± 1.0n/a17.5–22.4; 20.3 ± 1.516.7–20.1; 19.3 ± 1.4IML3.02.5–2.6; 2.5 ± 0.0n/a1.8–2.7; 2.3 ± 0.31.8–3.1; 2.5 ± 0.4IMW1.20.9–1.2; 1.1 ± 0.1n/a0.8–1.1; 1.0 ± 0.10.7–1.2; 0.9 ± 0.2TMP:EYE1.20.9–1.0; 1.0 ± 0.01.01.0–1.4; 1.1 ± 0.20.5–0.8; 0.7 ± 0.1TMP:SVL0.10.1–0.1; 0.1 ± 0.00.10.1–0.2; 0.1 ± 0.00.1–0.1; 0.1 ± 0.0 Molecules: The aligned dataset contained 1,633 characters. The standard deviation of split frequencies was 0.003805 among the four Bayesian runs, and all parameters had an Estimated Sample Size (ESS) ≥267. The new species was recovered as a member of a clade containing L. doriae, L. plicatellus, L. hascheanus, L. limborgi, and L. macrognathus, and this clade was sister to L. kohchangae (Fig. 6). The exact sister taxon relationship of the new species was unresolved, but the new species was not recovered to be the sister species to L. kohchangae (Fig. 6). The holotype, six paratypes, and referred larvae of L. coffeatus sp. nov. are identical in the 16S gene fragment (differing only by a single insertion-deletion event), but have an uncorrected pairwise divergence of 10.13–11.72% from L. kohchangae (n =11), the species to which it is phenotypically most similar (but not phylogenetically related; Fig. 6). Distribution, natural history, and conservation: Limnonectes coffeatus sp. nov. is only known from wet evergreen forest (Fig. 7) near to the top (≥ 1,200 m elev.) of the Bolaven Plateau in Dong Hua Sao National Protected Area, Pakxong District, Champasak Province, southern Laos (Fig. 1). Most of the known specimens were collected at night on the forest floor within 3 m of the bank, or in shallow water, of small, rocky streams. The immature female (NUOL 00060) was found at night (2215 h) during heavy rain on a forest trail, and one adult female (NCSM 77787) was found during the morning (1030 h) on leaf litter on the forest floor 20 m from a stream. Larvae (NCSM 77944) were collected at night (2130 h) in a 1.5 m wide slow-moving stream, approximately 20 cm deep, with a substrate of sand and algae-covered rocks (Fig. 7). An adult female (NCSM 77786) was collected in shallow water at the same time and place as the larvae. Calls and oviposition sites are unknown. Coffee (Coffea spp.) is the major agricultural export from Laos (for example, it was the largest agricultural export from the country in 2010), and 95% of Lao coffee is grown on the Bolaven Plateau (85% from Pakxong District alone) (Schönweger & Messerli 2015). Rainfall, temperature and physical properties of the soil render the upper elevations of the Bolaven Plateau highly favorable for growing coffee (Trelo-Ges et al. 2010; Schönweger & Messerli 2015), and a system of small landholder coffee production has been in place on the Bolaven Plateau since French colonial times (Delang et al. 2012). Today, approximately 15,000 households (69% of farms) on the Bolaven Plateau depend on coffee production as their primary source of income (Delang et al. 2012; Schönweger & Messerli 2015). Large-scale coffee plantations have also been established in recent years (Delang et al. 2012; Schönweger & Messerli 2015). As such, forest at upper elevations on the Bolaven Plateau has been extensively converted into coffee plantations (Thewlis et al. 1998), with human settlements (and accompanying plantations) encroaching even inside the boundaries of Dong Hua Sao National Protected Area (Delang et al. 2012). Bauxite mining, rubber plantations, and hydroelectric dam construction place additional pressures on forested areas of the Bolaven Plateau (Delang et al. 2012). All known records of the new species were obtained in closed-canopy forest (and not in coffee plantations), and so we expect that the continued conversion of forest into coffee plantations at upper elevations of the Bolaven Plateau likely presents a significant threat to the persistence of the new species. Comparisons: Four other species of Limnonectes in the region (Laos, Vietnam, Cambodia and Thailand) have the combination of males lacking a cephalic caruncle (swollen or cap-like structure; sensu Lambertz et al. 2014); adult SVL L. limborgi (Sclater, 1892), L. hascheanus (Stoliczka, 1870), L. doriae (Boulenger, 1887) and L. kohchangae (Smith, 1922). Limnonectes coffeatus sp. nov. differs from L. limborgi by having webbing on Toe V reaching base of tip (webbing more reduced in L. limborgi); dorsum with rounded tubercles, not arranged in rows (if present, tubercles elongated and arranged in rows in L. limborgi), and having a uniformly gray throat in males (absent in L. limborgi). Limnonectes coffeatus sp. nov. differs from L. hascheanus by having adults with SVL> 26 (adults with SVL L. hascheanus); having males with two large odontoid processes at front of mandible (absent in L. hascheanus); and having ova with pigmented poles (enlarged, non-pigmented ova in L. hascheanus). Limnonectes coffeatus sp. nov. differs from L. doriae by having males with two large odontoid processes at front of mandible (absent in L. doriae); having adults with SVL L. doriae); and males with TMP> EYE (TMP L. doriae). Limnonectes coffeatus sp. nov. is phenotypically most similar (but not phylogenetically related; Fig. 6) to L. kohchangae (Fig. 8). Limnonectes coffeatus sp. nov. differs from L. kohchangae by having dorsum with rounded tubercles, not arranged in rows (distinctly elongated and arranged in rows in L. kohchangae; Fig. 9); by lacking a distinctly bi-colored tympanum, with the upper half heavily pigmented (present in L. kohchangae; Fig. 8); by lacking dark spots on flank (present in L. kohchangae; Fig. 8); and having narrower limb bands, with the maximum shank band width 50% horizontal diameter of eye in L. kohchangae; Fig. 9).

Limnonectes coffeatus sp. nov. Holotype: NCSM 77788 (field tag BLS 13957), adult male (Fig. 2), Laos, Champasak Province, Pakxong District, Bolaven Plateau, Dong Hua Sao National Protected Area, Houay Tad Seua Stream, 15.06237°N 106.21075°E, 1,235 m elev., coll. 3 August 2010 by Bryan L. Stuart, Niane Sivongxay, and Sengvilay Seateun. Paratypes: NCSM 77786, adult female, same data as holotype except coll. 30 July 2010. NCSM 77785, NCSM 77787 (Fig. 3–4), two adult females; NUOL 0 0 0 60, one immature female; same data as holotype except coll. 15.06515°N 106.21394°E, 1,245 m elev., 28 July–01 August 2010. FMNH 258440, immature female; FMNH 258441, FMNH 258530, two immature males, same data as holotype except coll. near 15.06528°N 106.21750°E, 1,200 m elev., coll. 22–23 September 1999 by Bryan L. Stuart and Harold F. Heatwole. Referred Specimens: NCSM 77944, larvae (n =13; Fig. 5), same data as holotype except coll. 30 July 2010. Etymology: The specific epithet refers to the coffee plant genus Coffea, in reference to the extensive coffee plantations that are encroaching into the natural habitat of the new species at upper elevations on the Bolaven Plateau of southern Laos. Diagnosis: Assigned to the genus Limnonectes on the basis of its inferred phylogenetic position (Fig. 6), the presence of fang-like odontoid processes on the lower jaw (Emerson et al. 2000; Lambertz et al. 2014), and males with hypertrophied heads (Lambertz et al. 2014). A small-sized Limnonectes having the combination of the single available adult male with SVL 37.9, adult females with SVL 38.1–42.1; male lacking postorbital caruncle; male with hypertrophied head; odontoid processes on anterior margin of lower jaw, larger in male than in females; male with horizontal diameter of tympanum greater than that of eye, females with horizontal diameter of tympanum less than or equal to that of eye; tubercles on dorsum enlarged and rounded, but not arranged in rows; interdigital webbing of foot present; throat uniformly gray in male; tympanum uniformly colored, without distinct dark marking on upper half; flank without distinct spots; and ova with pigmented poles. Description of holotype: Habitus moderately stocky; body tapering to groin. Head broad and depressed; head width greater than head length. Snout obtusely pointed in dorsal view, round in profile, projecting well beyond lower jaw in profile; nostril dorsolateral, much closer to tip of snout than to eye, below canthus; internarial distance greater than interorbital distance; canthus rostralis indistinct, rounded, slightly constricted behind nostrils; lores concave, oblique; eye diameter 55% of snout length, interorbital distance greater than upper eyelid width; pineal ocellus visible; tympanum round, not elevated from side of head, annulus visible, tympanum diameter approximately 117% eye diameter and greater than distance between tympanum and eye; small, slit-like vocal sac openings on floor of mouth near lateral margin of tongue; vomerine teeth on two oblique ridges, closer to choanae than to each other; two large odontoid processes at front of mandible; median triangular protuberance at mandibular symphysis. Forelimbs moderately robust. Fingers relatively slender, without webbing or skin flaps; tip of fingers rounded, weakly expanded into discs; relative finger lengths II = IV Color of holotype in life: Dorsum brown with ochre wash; larger tubercles on dorsal surface of head and back orange-red with black margins, those on occiput forming “W”-shape; dark brown triangular marking between eyes with yellow anterior margin; dark brown alternating white-gray bands on upper and lower lips; dorsal surfaces of limbs with indistinct gray-brown bands; dorsal surfaces of thighs with yellowish-wash. Chin and throat gray; ventral surfaces of chest, belly and limbs yellow; ventral surfaces of hands and feet brown; iris bronze. Description of larvae NCSM 77944: Based on Gosner stage 28 individual in series of thirteen larvae (Fig. 5). Body oval in dorsal view, slightly compressed dorsoventrally, maximum body width slightly anterior to level of spiracle. Nares dorsal, without raised rim. Eyes dorsolateral, not visible from below. Spiracular tube single, sinistral on left side, angled slightly dorsally, aperture near midline of side and projecting posteriorly, approximately midway between eye and end of body. Tail slender, tapering gradually in distal one-third to rounded tip, origins of dorsal and ventral fins at end of body, dorsal fin only slightly deeper than ventral fin. Oral disk ventral, subterminal, width about 36% maximum width of body. Anterior labium with single row of papillae on lateral margins; posterior labium with two staggered rows of papillae on lateral and posterior margins; papillae homogenous in length. Labial tooth row formula 2(2)/3(1). All posterior tooth rows subequal in length. Upper and lower jaw sheaths black with serrated margins, upper sheath without median convexity. No distinct cluster of glandules posterior to eye on dorsal surface of body, subequal to interorbital distance. Color in life brown with gold flecking on belly and distinct black spots on dorsal surface of body, caudal muscle, and dorsal and ventral tail fins. In preservative, brown faded to light brown, black faded to dark brown, gold flecking absent. Measurements of larvae are summarized in Table 3. Variation: Females have narrower heads in dorsal view than holotype male (Fig. 4). Females have relatively smaller tympana than holotype male, with the horizontal diameter of the tympanum less than or equal to that of the eye. Females have smaller odontoid processes at front of mandible than holotype male. Female NCSM 77787 has wider black margins on larger tubercles on dorsal surface of head and back than holotype, and with black rather than brown or gray-brown bands on interorbital region, lips, and dorsal surfaces of limbs. Females lack wrinkled skin and have paler coloration on throat than holotype male. Females contain ova with pigmented poles. Measurements of adults are summarized in Table 2. SVL37.938.1–42.1; 40.5 ± 2.141.137.6–46.4; 40.9 ± 2.936.0–41.2; 38.1 ± 2.2HDL17.915.8–17.7; 16.9 ± 1.019.816.0–21.4; 18.9 ± 1.614.4–16.8; 15.7 ± 0.9HDW18.615.2–17.3; 16.6 ± 1.218.017.6–23.2; 20.0 ± 1.813.1–17.7; 15.9 ± 1.7SNT7.65.9–7.3; 6.8 ± 0.86.76.1–9.0; 7.4 ± 0.85.6–6.9; 6.5 ± 0.5EYE4.23.6–4.4; 4.1 ± 0.45.54.3–5.7; 5.0 ± 0.44.2–5.1; 4.7 ± 0.3IOD3.83.2–3.3; 3.3 ± 0.14.44.0–6.2; 5.1 ± 0.72.6–4.6; 3.7 ± 0.7TMP4.93.5–4.3; 3.9 ± 0.45.34.4–7.3; 5.6 ± 0.92.8–3.8; 3.4 ± 0.4IND4.64.3–4.6; 4.5 ± 0.24.84.3–6.2; 5.3 ± 0.63.6–5.2; 4.6 ± 0.6SHK20.219.4–20.0; 19.7 ± 0.320.517.6–22.1; 20.4 ± 1.317.6–20.5; 19.5 ± 1.2TGH20.319.4–21.0; 20.5 ± 0.918.018.9–22.9; 20.9 ± 1.418.5–20.7; 20.0 ± 0.9LAL7.27.9–8.4; 8.2 ± 0.3n/a7.5–8.7; 8.0 ± 0.46.7–8.0; 7.6 ± 0.6HND10.510.0–11.0; 10.4 ± 0.5n/a9.1–12.0; 10.6 ± 0.88.5–10.9; 10.0 ± 0.9FTL19.318.7–20.7; 19.9 ± 1.0n/a17.5–22.4; 20.3 ± 1.516.7–20.1; 19.3 ± 1.4IML3.02.5–2.6; 2.5 ± 0.0n/a1.8–2.7; 2.3 ± 0.31.8–3.1; 2.5 ± 0.4IMW1.20.9–1.2; 1.1 ± 0.1n/a0.8–1.1; 1.0 ± 0.10.7–1.2; 0.9 ± 0.2TMP:EYE1.20.9–1.0; 1.0 ± 0.01.01.0–1.4; 1.1 ± 0.20.5–0.8; 0.7 ± 0.1TMP:SVL0.10.1–0.1; 0.1 ± 0.00.10.1–0.2; 0.1 ± 0.00.1–0.1; 0.1 ± 0.0 Molecules: The aligned dataset contained 1,633 characters. The standard deviation of split frequencies was 0.003805 among the four Bayesian runs, and all parameters had an Estimated Sample Size (ESS) ≥267. The new species was recovered as a member of a clade containing L. doriae, L. plicatellus, L. hascheanus, L. limborgi, and L. macrognathus, and this clade was sister to L. kohchangae (Fig. 6). The exact sister taxon relationship of the new species was unresolved, but the new species was not recovered to be the sister species to L. kohchangae (Fig. 6). The holotype, six paratypes, and referred larvae of L. coffeatus sp. nov. are identical in the 16S gene fragment (differing only by a single insertion-deletion event), but have an uncorrected pairwise divergence of 10.13–11.72% from L. kohchangae (n =11), the species to which it is phenotypically most similar (but not phylogenetically related; Fig. 6). Distribution, natural history, and conservation: Limnonectes coffeatus sp. nov. is only known from wet evergreen forest (Fig. 7) near to the top (≥ 1,200 m elev.) of the Bolaven Plateau in Dong Hua Sao National Protected Area, Pakxong District, Champasak Province, southern Laos (Fig. 1). Most of the known specimens were collected at night on the forest floor within 3 m of the bank, or in shallow water, of small, rocky streams. The immature female (NUOL 00060) was found at night (2215 h) during heavy rain on a forest trail, and one adult female (NCSM 77787) was found during the morning (1030 h) on leaf litter on the forest floor 20 m from a stream. Larvae (NCSM 77944) were collected at night (2130 h) in a 1.5 m wide slow-moving stream, approximately 20 cm deep, with a substrate of sand and algae-covered rocks (Fig. 7). An adult female (NCSM 77786) was collected in shallow water at the same time and place as the larvae. Calls and oviposition sites are unknown. Coffee (Coffea spp.) is the major agricultural export from Laos (for example, it was the largest agricultural export from the country in 2010), and 95% of Lao coffee is grown on the Bolaven Plateau (85% from Pakxong District alone) (Schönweger & Messerli 2015). Rainfall, temperature and physical properties of the soil render the upper elevations of the Bolaven Plateau highly favorable for growing coffee (Trelo-Ges et al. 2010; Schönweger & Messerli 2015), and a system of small landholder coffee production has been in place on the Bolaven Plateau since French colonial times (Delang et al. 2012). Today, approximately 15,000 households (69% of farms) on the Bolaven Plateau depend on coffee production as their primary source of income (Delang et al. 2012; Schönweger & Messerli 2015). Large-scale coffee plantations have also been established in recent years (Delang et al. 2012; Schönweger & Messerli 2015). As such, forest at upper elevations on the Bolaven Plateau has been extensively converted into coffee plantations (Thewlis et al. 1998), with human settlements (and accompanying plantations) encroaching even inside the boundaries of Dong Hua Sao National Protected Area (Delang et al. 2012). Bauxite mining, rubber plantations, and hydroelectric dam construction place additional pressures on forested areas of the Bolaven Plateau (Delang et al. 2012). All known records of the new species were obtained in closed-canopy forest (and not in coffee plantations), and so we expect that the continued conversion of forest into coffee plantations at upper elevations of the Bolaven Plateau likely presents a significant threat to the persistence of the new species. Comparisons: Four other species of Limnonectes in the region (Laos, Vietnam, Cambodia and Thailand) have the combination of males lacking a cephalic caruncle (swollen or cap-like structure; sensu Lambertz et al. 2014); adult SVL L. limborgi (Sclater, 1892), L. hascheanus (Stoliczka, 1870), L. doriae (Boulenger, 1887) and L. kohchangae (Smith, 1922). Limnonectes coffeatus sp. nov. differs from L. limborgi by having webbing on Toe V reaching base of tip (webbing more reduced in L. limborgi); dorsum with rounded tubercles, not arranged in rows (if present, tubercles elongated and arranged in rows in L. limborgi), and having a uniformly gray throat in males (absent in L. limborgi). Limnonectes coffeatus sp. nov. differs from L. hascheanus by having adults with SVL> 26 (adults with SVL L. hascheanus); having males with two large odontoid processes at front of mandible (absent in L. hascheanus); and having ova with pigmented poles (enlarged, non-pigmented ova in L. hascheanus). Limnonectes coffeatus sp. nov. differs from L. doriae by having males with two large odontoid processes at front of mandible (absent in L. doriae); having adults with SVL L. doriae); and males with TMP> EYE (TMP L. doriae). Limnonectes coffeatus sp. nov. is phenotypically most similar (but not phylogenetically related; Fig. 6) to L. kohchangae (Fig. 8). Limnonectes coffeatus sp. nov. differs from L. kohchangae by having dorsum with rounded tubercles, not arranged in rows (distinctly elongated and arranged in rows in L. kohchangae; Fig. 9); by lacking a distinctly bi-colored tympanum, with the upper half heavily pigmented (present in L. kohchangae; Fig. 8); by lacking dark spots on flank (present in L. kohchangae; Fig. 8); and having narrower limb bands, with the maximum shank band width 50% horizontal diameter of eye in L. kohchangae; Fig. 9)., Published as part of Phimmachak, Somphouthone, Sivongxay, Niane, Seateun, Sengvilay, Yodthong, Siriporn, Rujirawan, Attapol, Neang, Thy, Aowphol, Anchalee & Stuart, Bryan L., 2018, A new Limnonectes (Anura: Dicroglossidae) from southern Laos, pp. 325-340 in Zootaxa 4375 (3) on pages 329-336, DOI: 10.11646/zootaxa.4375.3.2, http://zenodo.org/record/1158870, {"references":["Emerson, S. B., Inger, R. F. & Iskandar, D. (2000) Molecular systematics and biogeography of the fanged frogs of Southeast Asia. Molecular Phylogenetics and Evolution, 16, 131 - 142. https: // doi. org / 10.1006 / mpev. 2000.0778","Lambertz, M., Hartmann, T., Walsh, S., Geissler, P. & McLeod, D. S. (2014) Anatomy, histology, and systematic implications of the head ornamentation in the males of four species of Limnonectes (Anura: Dicroglossidae). Zoological Journal of the Linnean Society, 172, 117 - 132. https: // doi. org / 10.1111 / zoj. 12171","Schonweger, O. & Messerli, P. (2015) Land acquisition, investment, and development in the Lao coffee sector: successes and failures. Critical Asian Studies, 47, 94 - 122. https: // doi. org / 10.1080 / 14672715.2015.997095","Trelo-Ges, V., Nilavong, K. & Polthanee, A. (2010) Soil quality assessment for coffee production in Pakxong District, Champasak Province of Lao People's Democratic Republic. International Journal of Environmental and Rural Development, 1 - 2, 88 - 93.","Delang, C. O., Toro, M. & Charlet-Phommachanh, M. (2012) Coffee, mines and dams: conflicts over land in the Bolaven Plateau, southern Lao PDR. The Geographical Journal, 2012, 1 - 15.","Thewlis, R. M., Timmins, R. J., Evans, T. D. & Duckworth, J. W. (1998) The conservation status of birds in Laos: a review of key species. Bird Conservation International, 8 (Supplement), 1 - 159. https: // doi. org / 10.1017 / S 0959270900002197","Sclater, W. L. (1892) On some specimens of frogs in the Indian Museum, Calcutta, with descriptions of several new species. Proceedings of the Zoological Society of London, 1892, 341 - 348.","Stoliczka, F. (1870) Observations on some Indian and Malayan Amphibia and Reptilia. Journal of the Asiatic Society of Bengal, 39, 134 - 228. https: // doi. org / 10.1080 / 00222937008696209","Boulenger, G. A. (1887) An account of the reptiles and batrachians obtained in Tenasserim by M. L. Fea of the Genoa Civic Museum. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 25, 474 - 486.","Smith, M. A. (1922) The frogs allied to Rana doriae. The Journal of the Natural History Society of Siam, 4, 215 - 229."]}

Odorrana aureola Stuart, Chuaynkern, Chan-ard, and Inger, 2006 Specimens examined. A single tadpole of O. aureola (ZMKU AM 01138) was collected from Kok Huai Toey Stream, Phu Luang Wildlife Sanctuary, Loei Province (17 �� 19 ' 20.9 "N 101 �� 31 ' 21.6 "E, 1246 m ASL) by Attapol Rujirawan and Natee Ampai on 19 September 2013 (2052 hr). A single adult specimen (ZMKU AM 01137; Fig. 2 A) was collected at the same location (Fig. 2 B) by Anchalee Aowphol, Attapol Rujirawan, Natee Ampai and Somphouthone Phimmachak on 28 July 2013 (2100 hr). Identification. Sequence data from the tadpole (ZMKU AM 01138) and adult (ZMKU AM 01137) were 100 % identical over a fragment of 553 bp. Morphology of the adult specimen of O. aureola matched the original description (Stuart et al. 2006 a). Measurement of tadpole. Morphometric measurements (in mm) at Gosner (1960) developmental stage 30 (Table 1): BH = 3.5; BL = 8.8; BW = 4.7; IND = 1.8; IOD = 2.8; IOS = 1.5; MTH = 4.3; ODW = 2.5; SS = 6.6; TAL = 20.9; TL = 29.7; TMH = 2.7; TMW = 2.4; HUF = 1.5; HLF = 1.2. Species Odorrana aureola Limnonectes isanensis Description of tadpole. Body small and streamlined (Figs. 3, 4). From above, body oval, wider anteriorly than posteriorly, snout acutely rounded; in profile body dorsoventrally depressed, snout rounded, sloping gently from the plane of the eyes, oral disc anteroventral. Nares small with elevated rim, anterodorsally positioned, closer to snout than eye. Eyes positioned at the anterior one-third of the body, dorsolateral, not visible from ventral view. Interorbital distance almost twice internarial distance. Single, sinistral, tubular spiracle opening just below body axis posterior to midpoint of body (Fig. 4 C); SS = 6.6 mm. Vent tube dextral, opens midway between muscle and edge of ventral fin. Tail musculature well developed, round shaped, end rounded. Fin beginning at root of tail, margin of dorsal fin higher than ventral fin and slightly convex, ventral fin parallel to caudal fin, fin tapering to pointed tip, TMH = 2.7 mm. Color pattern. Color in life (Stage 30, Fig. 4): body brown to russet with dense dark brown pigmentation on the dorsum from snout to distal third of tail, clusters of melanophores become less dense towards the venter giving the sides of body and tail a spotted appearance. Dorsum, sides of body and tail flecked with small golden spots arranged almost linearly. Distal third of tail musculature dark brown, tip transparent. Fins flecked with dark brown pigment near musculature, fading to transparent at margins and distal tip of tail. Oral disc pale white. Venter translucent revealing internal organs and centro-sinistral intestines. Coloration in preservative similar to that in life with dark brown pigmentation remaining the same, but body pale grayish, and spots pale golden. Oral disc. Oral disc (Fig. 3 B) anteroventral, emarginated with uniserial array of blunt marginal papillae and 5��� 6 submarginal papillae at intersection of anterior and posterior labia. Upper and lower jaw sheathes medium and serrated with black margins, LTRF: 5 (2���5)/ 4 (1). Ecology. The tadpole was discovered at the bottom of a small, shallow pool in a permanent stream with an accumulation of dead leaves and rocks O. aureola were often found 20���30 cm above the water perched on large rocks. When disturbed, adult frogs escaped to other rocky perches or took cover at the bottom of the stream. Comparisons. The external morphology of some tadpoles in the genus Odorrana has been described previously. Inthara et al. (2005) reported LTRF 5 (2���5)/ 4 (1) for Odorrana livida from Loei Province but did not address other oral features. Songchan (2007) described coloration and body morphology of a Gosner stage 35 tadpole of O. livida from Chiang Dao, Chiang Mai Province, northern Thailand and reported a LTRF consistent with Inthara et al. (2005). Unfortunately, these studies are confounded by the taxonomic confusion over the identity of O. livida complex members and the fact that some species occur in sympatry in Thailand. The lack of genetic data in Inthara et al. (2005) and Songchan (2007) precludes the use of molecular sequence homology to validate the identification. We found the labial tooth row formula of O. aureola to be the same as that of O. livida in Songchan (2007) and Inthara et al. (2005), but we found marginal papillae arranged in a single row not two rows as described by Inthara et al. (2005) and Songchan (2007). A distinctive larval feature of O. aureola appears to the presence of small golden spots arranged almost linearly along the entire brown to russet colored body. This is differs from previous descriptions of O. livida tadpoles, which are described as having scattered small golden or yellow flecks on a gray or dark colored body with some dark spots on the tail muscle. We suggest that the tadpoles described by Inthara et al. (2005) and Songchan (2007) may be those of O. chloronota, but additional studies are required to confirm this., Published as part of Ampai, Natee, Rujirawan, Attapol, Arkajag, Jirachai, Mcleod, David S. & Aowphol, Anchalee, 2015, Description of the tadpoles of two endemic frogs: the Phu Luang cascade frog Odorrana aureola (Anura: Ranidae) and the Isan big-headed frog Limnonectes isanensis (Anura: Dicroglossidae) from northeastern Thailand, pp. 508-520 in Zootaxa 3981 (4) on pages 510-513, DOI: 10.11646/zootaxa.3981.4.3, http://zenodo.org/record/243349, {"references":["Stuart, B. L., Chuaynkern Y., Chan-ard T. & Inger R. F. (2006 a) Three new species of frogs and a new tadpole from eastern Thailand. Fieldiana Zoology, 111, 1 - 19. http: // dx. doi. org / 10.3158 / 0015 - 0754 (2006) 187 [1: tnsofa] 2.0. co; 2","Gosner, K. L. (1960) A simplified table for staging anuran embryos and larvae, with notes on identification. Herpetologica, 16, 183 - 190.","Inthara, C., Lauhachinda, V., Nabhaitabhat, J., Chuaynkern, Y. & Kumtong, P. (2005) Mouth part structures and distribution of some tadpole from Thailand. The Thailand National History Museum Journal, 1 (1), 55 - 78.","Songchan, R. (2007) External morphological characters of some tadpoles as related to different habitat types in Chiang Dao, Chiang Mai Province. M. S. Thesis, Kasetsart University, Bangkok, Thailand, 103 pp."]}

Limnonectes lauhachindai sp. nov. Holotype: NCSM 80222 (field tag AA 01384), adult male (Fig. 1), Thailand, Ubon Ratchathani Province, Sirindhorn District, Kham Khuen Kaew Subdistrict, 15 �� 17 ��� 47.6 ���N 105 �� 28 ���22.0���E, 131 m elev., coll. 29 August 2012 by Anchalee Aowphol, Siriporn Yodthong, Natee Ampai, and Attapol Rujirawan. Paratypes: NCSM 81269, ZMKU AM 01104���09 (seven adult males): same data as holotype. ZMKU AM 0 1111 (one adult male): same data as holotype except coll. 30 August 2012. ZMKU AM 00552��� 53, ZMKU AM 00586��� 92 (nine adult males; Fig. 2): same data as holotype except coll. 19 August 2011 by Attapol Rujirawan, Wut Taksintum, Virayuth Lauhachinda, Anchalee Aowphol, and Siriporn Yodthong. FMNH 266148 / THNHM 0 5185, FMNH 266154 / THNHM 0 5189 (two adult males), FMNH 266147 / THNHM 0 5184, FMNH 266150 / THNHM 0 5025 (two adult females): Thailand, Ubon Ratchathani Province, Na Chaluai District, Phu Jong-Na Yoi National Park, Huay Luang Noi Stream, 14 �� 26 ��� 15.9 ���N 105 �� 16 ��� 48.2 ���E, 360 m elev., coll. 15 September 2004 by Yodchaiy Chuaynkern, Bryan L. Stuart, Chatchay Chuechat, and Sunchai Makchai. FMNH 266151 / THNHM 0 5026 (one adult female): same data as FMNH 266147 / THNHM 0 5184 except 14 �� 25 ��� 43.9 "N 105 �� 16 ���51.0"E, 350 m elev. FMNH 266152 / THNHM 0 5187 (one adult female): same data as FMNH 266147 / THNHM 0 5184 except 14 �� 26 ��� 18.6 "N 105 �� 16 ���04.5"E, 325 m elev. Referred material. ZMKU AM 0 0 593 (one juvenile male), same data as ZMKU AM 0 0 586. Etymology. The specific epithet is a patronym for Associate Professor Dr. Virayuth Lauhachinda, Kasetsart University, and co-collector of the new species, in recognition of his contributions to herpetology in Thailand. Suggested common names. Lauhachinda's Fanged Frog (English), Kob Ngon Arjarn Virayuth (Thai). Diagnosis. Assigned to the genus Limnonectes on the basis of molecular evidence (Fig. 3), the presence of fanglike odontoid processes on the lower jaw (Emerson et al. 2000; Lambertz et al. 2014), and males with hypertrophied heads (Lambertz et al. 2014). A small-sized Limnonectes having males with SVL 30.5 ���42.0, females with SVL 32.9���37.9; males with low-profiled, U-shaped caruncle with free posterior margin that completely occupies, but does not extend beyond, interobital region; males with hypertrophied heads; both sexes with enlarged odontoid processes on anterior margin of lower jaw; and webbing on toes. additions of PWRC = Phu Luang Wildlife Research Centre Museum; RMB = Rafe M. Brown field series; and ZNAC = Anhui Normal University. Description of holotype. Habitus moderately stocky; body tapering to groin. Head broad and depressed; head length and width subequal. Snout obtusely pointed in dorsal view, round in profile, projecting well beyond lower jaw in profile; nostril dorsolateral, much closer to tip of snout than to eye, below canthus, internarial distance subequal to interorbital distance; canthus rostralis indistinct, rounded, slightly constricted behind nostrils; lores concave, oblique; eye diameter 80 % snout length, interorbital distance greater than upper eyelid width; pineal ocellus visible; tympanum round, not elevated from side of head, annulus weakly visible, tympanum diameter about 82 % eye diameter and greater than distance between tympanum and eye; small, slit-like vocal sac openings on floor of mouth near lateral margin of tongue; vomerine teeth on two oblique ridges, equal in distance to each other as to choanae; two large odontoid processes at front of mandible; median triangular protuberance at mandibular symphisis. Forelimb moderately robust. Fingers moderately slender, without webbing; tip of fingers rounded, weakly expanded into discs; relative finger lengths II Color of holotype in preservative. Dorsum dark brown; tips of dorsal warts white, warts encircled with black. Venter cream, chin dark brown, throat with large brown spots, belly and ventral surfaces of limbs with very small dark spots. Color of paratype ZMKU AM 0 0 586 in life. Dorsum brown, with irregular black spots, becoming brassy on dorsal surfaces of limbs and upper flank; continuous black streak under canthus and supratympanic fold, extending from nostril to upper half of tympanum; lips brown with broad black bars; iris bronze; broad cream-yellow vertebral stripe from anterior margin of upper jaw to vent; upper surfaces of hindlimb with broad black bands; lower flank beige and gray (Fig. 2). Variation. Females lack caruncles and have narrower heads in dorsal view than males. The largest male paratype (ZMKU AM 01111; SVL 42.0 mm) is noticeably larger than the second largest (NCSM 81269; SVL 37.7 mm). Seven (NCSM 81269, ZMKU AM 0 1106, ZMKU AM 00552��� 53, ZMKU AM 00586��� 87, ZMKU AM 00593) of 25 (28 %) specimens have a pale vertebral stripe, a polymorphic character seen in other Limnonectes, including L. dabanus and L. kohchangae (Stuart & Emmett 2006; Stuart et al. 2006 b). Measurements are summarized in Table 2. Molecules. The aligned dataset contained 1,624 characters. The new species was recovered as sister to a clade containing L. dabanus and L. gyldenstolpei (Fig. 3). The holotype and six paratypes (NCSM 81269, ZMKU AM 01104���07, ZMKU AM 01109) are identical in the 16 S gene fragment, but have an uncorrected pairwise divergence of 5.98���6.72 % from L. dabanus (n = 2) and 4.25���8.40 % from L. gyldenstolpei (n = 3). Calls. Paratype male ZMKU AM 0 0 589 had two different advertisement call types, referred to here as Type 1 and Type 2 (Fig. 4). Type 1 (n = 30) had 1���10, one-pulsed notes lasting 27.44 to 4342.91 ms; note duration of 10.29���34.50 ms; interval between notes 182.50���972.41 ms; note repetition rate of 0.98 ���4.00 notes per second; and dominant frequency of 1.12���2.33 kHz. Type 2 (n = 3) was a single, multi-pulsed note lasting 1749.06���2148.03 ms with 31���37 pulses; pulse duration of 21.24 ���25.00 ms; interval between pulses 33.74���35.36 ms; pulse rate 16.98��� 17.49 pulses per second; and dominant frequency of 2.15���2.24 kHz (Table 3). The advertisement calls were repeated at a rate of approximately 0.12 calls per second and intercall interval varied from 0.36 to 30.26 s. These calls showed frequency modulation with or without weak harmonics. Distribution and natural history. Limnonectes lauhachindai sp. nov. is only known from Na Chaluai and Sirindhorn Districts, Ubon Ratachathani Province, Thailand (Fig. 5), from 131���360 m elevation. Those from Na Chaluai District were taken on the bank or in the water of a shallow stream flowing over bedrock in semi-evergreen forest on a single day between 1400���2140 h. Those from Sirindhorn District were taken on the ground in deciduous dipterocarp forest with wet grassy understory (Fig. 6). Eggs and larvae are unknown. Comparisons. Limnonectes lauhachindai sp. nov. differs from all other species of Limnonectes except L. dabanus, L. gyldenstolpei, L. macrognathus and L. plicatellus by having mature males with a caruncle on top of the head. Limnonectes lauhachindai sp. nov. further differs from these four species by having males with a lowprofiled, U-shaped caruncle with free posterior margin that fully occupies, but does not extend beyond, the interorbital region (caruncle high-profiled and domed in L. dabanus; caruncle extending beyond interorbital region in L. gyldenstolpei; caruncle lacking U-shape and free posterior margin in L. macrognathus; caruncle high-profiled and horned in L. plicatellus; Fig. 7). Limnonectes lauhachindai sp. nov. further differs from L. dabanus, L. gyldenstolpei, and L. macrognathus by having much smaller males [SVL 52.9���65.5 (mean �� SD 59.8 �� 4.0, n = 15) in L. dabanus; SVL 51.1���68.4 (mean �� SD 62.0 �� 5.7, n = 14) in L. gyldenstolpei; SVL 43.2���48.8 (mean �� SD 45.5 �� 3.0, n = 3, this study), types SVL 47���57 (n = 2; Boulenger 1920) in L. macrognathus], and from L. plicatellus by lacking dorsal rugosities arranged in distinct, longitudinal rows parallel to the body axis (present in L. plicatellus). Limnonectes lauhachindai sp. nov. superficially resembles L. hascheanus (Stoliczka 1870), L. limborgi (Sclater 1892), and L. kohchangae (Smith 1922), but differs from all of them by having cephalic caruncles in males (absent in L. hascheanus, L. limborgi, and L. kohchangae). Limnonectes lauhachindai sp. nov. further differs from L. hascheanus by having larger body size [males with SVL 18.8���25.4, mean �� SD 22.1 �� 1.9, n = 9, females with SVL 20.5 ���25.0, mean �� SD 23.0 �� 1.6, n = 9, in L. hascheanus (Inger & Stuart 2010)], and from L. hascheanus and L. limborgi by having fully webbed toes (greatly reduced toe webbing in L. hascheanus and L. limborgi). Male secondary sexual characters are unknown in L. khammonensis (Smith 1929), which is known only from the female holotype, but females of L. lauhachindai sp. nov. differ by having a distinct tympanum (indistinct in L. khammonensis) and less toe webbing (Toe IV webbed to distal subarticular tubercle, continuing as fringe to base of disc, and all remaining toes webbed to base of disc in L. khammonensis)., Published as part of Aowphol, Anchalee, Rujirawan, Attapol, Taksintum, Wut, Chuaynkern, Yodchaiy & Stuart, Bryan L., 2015, A new caruncle-bearing Limnonectes (Anura: Dicroglossidae) from northeastern Thailand, pp. 258-270 in Zootaxa 3956 (2) on pages 259-268, DOI: 10.11646/zootaxa.3956.2.6, http://zenodo.org/record/233382, {"references":["Emerson, S. B., Inger, R. F. & Iskandar, D. (2000) Molecular systematics and biogeography of the fanged frogs of Southeast Asia. Molecular Phylogenetics and Evolution, 16, 131 - 142. http: // dx. doi. org / 10.1006 / mpev. 2000.0778","Lambertz, M., Hartmann, T., Walsh, S., Geissler, P. & McLeod, D. S. (2014) Anatomy, histology, and systematic implications of the head ornamentation in the males of four species of Limnonectes (Anura: Dicroglossidae). Zoological Journal of the Linnean Society, 172, 117 - 132. http: // dx. doi. org / 10.1111 / zoj. 12171","Stuart, B. L. & Emmett, D. A. (2006) A collection of amphibians and reptiles from the Cardamom Mountains, southwestern Cambodia. Fieldiana, Zoology New Series, 109, 1 - 27. http: // dx. doi. org / 10.3158 / 0015 - 0754 (2006) 187 [1: TNSOFA] 2.0. CO; 2","Stuart, B. L., Sok, K. & Neang, T. (2006 b) A collection of amphibians and reptiles from hilly eastern Cambodia. The Raffles Bulletin of Zoology, 54, 129 - 155.","Boulenger, G. A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum, 20, 1 - 223. http: // dx. doi. org / 10.5962 / bhl. title. 12471","Stoliczka, F. (1870) Observations on some Indian and Malayan Amphibia and Reptilia. Journal of the Asiatic Society of Bengal, 39, 134 - 228. http: // dx. doi. org / 10.1080 / 00222937008696209","Sclater, W. L. (1892) On some specimens of frogs in the Indian Museum, Calcutta, with descriptions of several new species. Proceedings of the Zoological Society of London, 1892, 341 - 348.","Smith, M. A. (1922) The frogs allied to Rana doriae. The Journal of the Natural History Society of Siam, 4, 215 - 229.","Inger, R. F. & Stuart, B. L. (2010) Systematics of Limnonectes (Taylorana) Dubois. Current Herpetology, 29, 51 - 68. http: // dx. doi. org / 10.3105 / 018.029.0201","Smith, M. A. (1929) Descriptions of a new skink from Christmas Island and a new frog from Annam. Annals and Magazine of Natural History, Series 10, 3, 294 - 297. http: // dx. doi. org / 10.1080 / 00222932908672972"]}

Limnonectes isanensis McLeod, Kelly, and Barley, 2012 Specimens examined. Five tadpoles of L. isanensis (ZMKU AM 0 1140 [n= 3], ZMKU AM 0 1141 [n= 1] and ZMKU AM 0 1142 [n= 1]) were collected from Ron Ton Son Stream, Phu Luang Wildlife Sanctuary, Loei Province (17 �� 17 ' 53.2 "N 101 �� 31 ' 22.1 "E, 1,361 m ASL). Specimens were collected by Anchalee Aowphol, Attapol Rujirawan, Natee Ampai, Korkhwan Termprayoon, and Somphouthone Phimmachak on 28 July 2012 (2100 hr) and 11 December 2012 (2045 hr). A single tadpole specimen (ZMKU AM 01142; stage 38) was photographed in life and illustrated (Figs. 6���7). A single adult specimen (ZMKU AM 01139; Fig. 5 A) was collected at Nam San Noi Stream, Phu Luang Wildlife Sanctuary, Loei Province (17 �� 20 '01.0" N 101 �� 30 '35.0"E, 915m ASL) by Anchalee Aowphol, Attapol Rujirawan, Natee Ampai and Somphouthone Phimmachak on 26 September 2012 (2030 hr). Identification. Tadpole (ZMKU AM 01140) has an uncorrected pairwise divergence of 0.18 % from an adult specimen (ZMKU AM 01139). Morphology of the adult specimen of L. isanensis matched the original description (McLeod et al. 2012). Measurement of tadpole. Morphometric measurements (in mm) at Gosner (1960) developmental stage 38: BH = 5.9; BL = 13.4; BW = 7.9; IND = 2.8; IOD = 4.0; IOS = 2.6; MTH = 6.8; ODW = 2.4; SS = 7.7; TAL = 26.9; TL = 40.3; TMH = 3.1; TMW = 3.2; HUF = 2.2; HLF = 1.5. Variation within the series. Mensural and ratio values for external morphology of the tadpole series (Gosner stages 30���38; n = 5) is provided in Table 1. External morphology of the tadpoles remains generally consistent throughout the developmental stages in this series. In all specimens the sinistral spiracle and intestine coils were visible in lateral and ventral views. In early stages 30���35, the tail is weakly mottled whereas in later stages (e.g., Gosner 38), the tail is much more densely mottled. Description of tadpole. Body moderate (Fig. 7 A). From above, body oval, length twice width, wider anteriorly than posteriorly, snout rounded; in profile body depressed, sloping steeply from level of eyes, oral disc anteroventral. Nares are small, rim slightly elevated, anterodorsally positioned, closer to the snout than eye. Eyes dorsolaterally positioned at the end of the anterior quarter of body (closer to nares than spiracle), widely spaced relative to nares (IND /IOD = 0.70), not visible from ventral view. Single, sinistral, tubular spiracle, opening below body axis near midpoint of body SS = 7.7 mm (Fig. 7 A���B). Vent tube dextral, opens midway between muscle and edge of ventral fin; both of fins slightly convex (Fig. 7 B). Tail musculature well developed, tail tapering gradually to pointed tip. Fin beginning at root of tail, margin of dorsal fin slightly higher than ventral fin and convex, ventral fin parallel to dorsal fin, TMH = 3.1 mm. Color Pattern. Color in life (Stage 38, Fig. 7): body pale brown to yellowish with slightly darker brown pigment. Many small, thin, golden-yellow stripes on the sides of body and on dorsum of tail musculature. Dorsum and tail with many dark brown to black spots. Oral disc pale white. Venter of body nearly transparent, lightly flecked with gold; centro-sinistral intestine visible. Ventral surface of tail is semitransparent to white; caudal musculature faintly pale brown in dorsal view, with the upper and lower fins transparent. In preservative, coloration similar to specimen in life, pale yellowish stripes on ventrolateral body, dark brown pigmentation on tail remains distinctly visible. Oral disc. Oral disc (Fig. 6 B), anteroventral, 5���6 marginal papillae at corners of upper labium, papillae on upper labium larger than lower lip, upper and lower jaw sheathes medium, slight medial convexity, margin serrated, black. Separation between individual denticles narrow, LTRF: 2 (1)/ 3 (1); first row without gap, second row sub-equal left and right. The first lower labial tooth row with narrow median gap. Third lower labial tooth row nearly half the length of first and second rows. Ecology. Tadpoles were collected from a shallow pool (12���15 cm deep) at the edge of a permanent stream (4 m wide, 60 cm deep) at 0 845 hr on 11 December 2012 (Fig. 5 B). Ambient air temperature = 22.8 ��C, surface water temperature = 17.9 ��C, water pH = 6.14, and stream current was slow (0.1 m /s). Tadpoles were abundant at the study site during the study period. During the daytime, tadpoles were occasionally seen in the water column, but were predictably found buried in the gravel substrate or hiding beneath clumps of leaves at the bottom of the pools in the stream. When disturbed, tadpoles were observed to swim quickly to new hiding places. At nighttime, tadpoles were active in the water column. Egg masses containing 35���40 ova with transparent external jelly were found in clumps stuck to rocks in stream or to stream bank at the same location as the tadpoles during December (2012) and September (2013). Individual eggs encased in a jelly capsule measured 4���5 mm. Adult males of L. isanensis were found perched either on the ground amidst leaf litter, or on rocks on the bank of the stream at this site. Comparisons. McLeod et al. (2012) described the PLWS endemic, L. isanensis, based on adult morphology, but provided no tadpole information. Our description of the tadpole of L. isanensis closely match the general characteristics of genus Limnonectes tadpoles described by Altig & McDiarmid (1999). McLeod (2008) provided morphometric data from a series of tadpoles and the description of the external morphology of a Gosner stage 40 tadpole of L. megastomias from Sa Kaeo Province, in eastern Thailand. Limnonectes megastomias is the sister taxon of L. isanensis and adults of both species are morphologically very similar (McLeod et al. 2012). Despite this overall adult similarity, the tadpoles of L. megastomias and L. isanensis differ notably in several features. Measurement proportion of two species is shown in Table 2. In overall size, the tadpole of L. isanensis seems to be smaller than L. megastomias, particularly during early stages of development (TL = 27.1���40.3 mm and 31.3���39.8 mm, respectively). Nevertheless, relative to total length, the body of L. isanensis appears to be larger than L. megastomias. When comparing tadpoles of the same developmental stage, Limnonectes isanensis was found to have a significantly greater body height (BH), the ratio of body height (BH) to body length (BL) and consequently in the ratio of body width (BW) to body length (BL) than L. megastomias (P L. isanensis are arranged in single row with about 5���7 papillae per side, two rows of papillae on the lower labium with about 20���22 papillae per side, a very narrow gap in the posterior marginal papillae, and the occurrence of two submarginal papillae ventrally. Tadpoles of L. megastomias have a single row of upper labial papillae, about 11���13 thick papillae of upper labium per side and about 28���30 papillae per side on lower labium arranged in two rows with narrow median interruption (but broader than in L. isanensis). The caudal musculature and fins are more heavily pigmented in L. isanensis (dark brown to black on posterior half of tail) than in L. megastomiasis (faint brown spots scattered over length of tail). Additionally, tadpoles of L. megastomias have a median dark-brown horizontal bar through eye, which L. isanensis lacks., Published as part of Ampai, Natee, Rujirawan, Attapol, Arkajag, Jirachai, Mcleod, David S. & Aowphol, Anchalee, 2015, Description of the tadpoles of two endemic frogs: the Phu Luang cascade frog Odorrana aureola (Anura: Ranidae) and the Isan big-headed frog Limnonectes isanensis (Anura: Dicroglossidae) from northeastern Thailand, pp. 508-520 in Zootaxa 3981 (4) on pages 514-517, DOI: 10.11646/zootaxa.3981.4.3, http://zenodo.org/record/243349, {"references":["McLeod, D. S., Kelly, J. K. & Barley, A. J. (2012) \" Same-same but different \": Another new species of the Limnonectes kuhlii complex from Thailand (Anura: Dicroglossidae). Russian Journal of Herpetology, 19 (3), 261 - 274.","Gosner, K. L. (1960) A simplified table for staging anuran embryos and larvae, with notes on identification. Herpetologica, 16, 183 - 190.","McLeod, D. S. (2008) A new species of big-headed, fanged dicroglossine frog (genus Limnonectes) from Thailand. Zootaxa, 1807, 26 - 46."]}

Ampai, Natee, Rujirawan, Attapol, Arkajag, Jirachai, Mcleod, David S., Aowphol, Anchalee (2015): Description of the tadpoles of two endemic frogs: the Phu Luang cascade frog Odorrana aureola (Anura: Ranidae) and the Isan big-headed frog Limnonectes isanensis (Anura: Dicroglossidae) from northeastern Thailand. Zootaxa 3981 (4): 508-520, DOI: http://dx.doi.org/10.11646/zootaxa.3981.4.3

Limnonectes lauhachindai sp. nov. Holotype: NCSM 80222 (field tag AA 01384), adult male (Fig. 1), Thailand, Ubon Ratchathani Province, Sirindhorn District, Kham Khuen Kaew Subdistrict, 15 ° 17 ’ 47.6 ”N 105 ° 28 ’22.0”E, 131 m elev., coll. 29 August 2012 by Anchalee Aowphol, Siriporn Yodthong, Natee Ampai, and Attapol Rujirawan. Paratypes: NCSM 81269, ZMKU AM 01104–09 (seven adult males): same data as holotype. ZMKU AM 0 1111 (one adult male): same data as holotype except coll. 30 August 2012. ZMKU AM 00552– 53, ZMKU AM 00586– 92 (nine adult males; Fig. 2): same data as holotype except coll. 19 August 2011 by Attapol Rujirawan, Wut Taksintum, Virayuth Lauhachinda, Anchalee Aowphol, and Siriporn Yodthong. FMNH 266148 / THNHM 0 5185, FMNH 266154 / THNHM 0 5189 (two adult males), FMNH 266147 / THNHM 0 5184, FMNH 266150 / THNHM 0 5025 (two adult females): Thailand, Ubon Ratchathani Province, Na Chaluai District, Phu Jong-Na Yoi National Park, Huay Luang Noi Stream, 14 º 26 ’ 15.9 ”N 105 º 16 ’ 48.2 ”E, 360 m elev., coll. 15 September 2004 by Yodchaiy Chuaynkern, Bryan L. Stuart, Chatchay Chuechat, and Sunchai Makchai. FMNH 266151 / THNHM 0 5026 (one adult female): same data as FMNH 266147 / THNHM 0 5184 except 14 ° 25 ’ 43.9 "N 105 ° 16 ’51.0"E, 350 m elev. FMNH 266152 / THNHM 0 5187 (one adult female): same data as FMNH 266147 / THNHM 0 5184 except 14 ° 26 ’ 18.6 "N 105 ° 16 ’04.5"E, 325 m elev. Referred material. ZMKU AM 0 0 593 (one juvenile male), same data as ZMKU AM 0 0 586. Etymology. The specific epithet is a patronym for Associate Professor Dr. Virayuth Lauhachinda, Kasetsart University, and co-collector of the new species, in recognition of his contributions to herpetology in Thailand. Suggested common names. Lauhachinda's Fanged Frog (English), Kob Ngon Arjarn Virayuth (Thai). Diagnosis. Assigned to the genus Limnonectes on the basis of molecular evidence (Fig. 3), the presence of fanglike odontoid processes on the lower jaw (Emerson et al. 2000; Lambertz et al. 2014), and males with hypertrophied heads (Lambertz et al. 2014). A small-sized Limnonectes having males with SVL 30.5 –42.0, females with SVL 32.9–37.9; males with low-profiled, U-shaped caruncle with free posterior margin that completely occupies, but does not extend beyond, interobital region; males with hypertrophied heads; both sexes with enlarged odontoid processes on anterior margin of lower jaw; and webbing on toes. additions of PWRC = Phu Luang Wildlife Research Centre Museum; RMB = Rafe M. Brown field series; and ZNAC = Anhui Normal University. Description of holotype. Habitus moderately stocky; body tapering to groin. Head broad and depressed; head length and width subequal. Snout obtusely pointed in dorsal view, round in profile, projecting well beyond lower jaw in profile; nostril dorsolateral, much closer to tip of snout than to eye, below canthus, internarial distance subequal to interorbital distance; canthus rostralis indistinct, rounded, slightly constricted behind nostrils; lores concave, oblique; eye diameter 80 % snout length, interorbital distance greater than upper eyelid width; pineal ocellus visible; tympanum round, not elevated from side of head, annulus weakly visible, tympanum diameter about 82 % eye diameter and greater than distance between tympanum and eye; small, slit-like vocal sac openings on floor of mouth near lateral margin of tongue; vomerine teeth on two oblique ridges, equal in distance to each other as to choanae; two large odontoid processes at front of mandible; median triangular protuberance at mandibular symphisis. Forelimb moderately robust. Fingers moderately slender, without webbing; tip of fingers rounded, weakly expanded into discs; relative finger lengths II Hindlimb moderately robust. Toes moderately slender; tips of toes rounded, expanded into small discs; relative toe lengths I Skin above shagreened with irregular rows of large oval warts, most concentrated near flank and lower back; low-profiled, U-shaped caruncle with free posterior margin that fully occupies, but does not extend beyond, the interorbital region; distinct supratympanic ridge from posterior corner of eye to axilla; large rictal gland; no dorsolateral fold; skin on venter smooth. Measurements of holotype given in Table 2. Color of holotype in preservative. Dorsum dark brown; tips of dorsal warts white, warts encircled with black. Venter cream, chin dark brown, throat with large brown spots, belly and ventral surfaces of limbs with very small dark spots. Color of paratype ZMKU AM 0 0 586 in life. Dorsum brown, with irregular black spots, becoming brassy on dorsal surfaces of limbs and upper flank; continuous black streak under canthus and supratympanic fold, extending from nostril to upper half of tympanum; lips brown with broad black bars; iris bronze; broad cream-yellow vertebral stripe from anterior margin of upper jaw to vent; upper surfaces of hindlimb with broad black bands; lower flank beige and gray (Fig. 2). Variation. Females lack caruncles and have narrower heads in dorsal view than males. The largest male paratype (ZMKU AM 01111; SVL 42.0 mm) is noticeably larger than the second largest (NCSM 81269; SVL 37.7 mm). Seven (NCSM 81269, ZMKU AM 0 1106, ZMKU AM 00552– 53, ZMKU AM 00586– 87, ZMKU AM 00593) of 25 (28 %) specimens have a pale vertebral stripe, a polymorphic character seen in other Limnonectes, including L. dabanus and L. kohchangae (Stuart & Emmett 2006; Stuart et al. 2006 b). Measurements are summarized in Table 2. Molecules. The aligned dataset contained 1,624 characters. The new species was recovered as sister to a clade containing L. dabanus and L. gyldenstolpei (Fig. 3). The holotype and six paratypes (NCSM 81269, ZMKU AM 01104–07, ZMKU AM 01109) are identical in the 16 S gene fragment, but have an uncorrected pairwise divergence of 5.98–6.72 % from L. dabanus (n = 2) and 4.25–8.40 % from L. gyldenstolpei (n = 3). Calls. Paratype male ZMKU AM 0 0 589 had two different advertisement call types, referred to here as Type 1 and Type 2 (Fig. 4). Type 1 (n = 30) had 1–10, one-pulsed notes lasting 27.44 to 4342.91 ms; note duration of 10.29–34.50 ms; interval between notes 182.50–972.41 ms; note repetition rate of 0.98 –4.00 notes per second; and dominant frequency of 1.12–2.33 kHz. Type 2 (n = 3) was a single, multi-pulsed note lasting 1749.06–2148.03 ms with 31–37 pulses; pulse duration of 21.24 –25.00 ms; interval between pulses 33.74–35.36 ms; pulse rate 16.98– 17.49 pulses per second; and dominant frequency of 2.15–2.24 kHz (Table 3). The advertisement calls were repeated at a rate of approximately 0.12 calls per second and intercall interval varied from 0.36 to 30.26 s. These calls showed frequency modulation with or without weak harmonics. Distribution and natural history. Limnonectes lauhachindai sp. nov. is only known from Na Chaluai and Sirindhorn Districts, Ubon Ratachathani Province, Thailand (Fig. 5), from 131–360 m elevation. Those from Na Chaluai District were taken on the bank or in the water of a shallow stream flowing over bedrock in semi-evergreen forest on a single day between 1400–2140 h. Those from Sirindhorn District were taken on the ground in deciduous dipterocarp forest with wet grassy understory (Fig. 6). Eggs and larvae are unknown. Comparisons. Limnonectes lauhachindai sp. nov. differs from all other species of Limnonectes except L. dabanus, L. gyldenstolpei, L. macrognathus and L. plicatellus by having mature males with a caruncle on top of the head. Limnonectes lauhachindai sp. nov. further differs from these four species by having males with a lowprofiled, U-shaped caruncle with free posterior margin that fully occupies, but does not extend beyond, the interorbital region (caruncle high-profiled and domed in L. dabanus; caruncle extending beyond interorbital region in L. gyldenstolpei; caruncle lacking U-shape and free posterior margin in L. macrognathus; caruncle high-profiled and horned in L. plicatellus; Fig. 7). Limnonectes lauhachindai sp. nov. further differs from L. dabanus, L. gyldenstolpei, and L. macrognathus by having much smaller males [SVL 52.9–65.5 (mean ± SD 59.8 ± 4.0, n = 15) in L. dabanus; SVL 51.1–68.4 (mean ± SD 62.0 ± 5.7, n = 14) in L. gyldenstolpei; SVL 43.2–48.8 (mean ± SD 45.5 ± 3.0, n = 3, this study), types SVL 47–57 (n = 2; Boulenger 1920) in L. macrognathus], and from L. plicatellus by lacking dorsal rugosities arranged in distinct, longitudinal rows parallel to the body axis (present in L. plicatellus). Limnonectes lauhachindai sp. nov. superficially resembles L. hascheanus (Stoliczka 1870), L. limborgi (Sclater 1892), and L. kohchangae (Smith 1922), but differs from all of them by having cephalic caruncles in males (absent in L. hascheanus, L. limborgi, and L. kohchangae). Limnonectes lauhachindai sp. nov. further differs from L. hascheanus by having larger body size [males with SVL 18.8–25.4, mean ± SD 22.1 ± 1.9, n = 9, females with SVL 20.5 –25.0, mean ± SD 23.0 ± 1.6, n = 9, in L. hascheanus (Inger & Stuart 2010)], and from L. hascheanus and L. limborgi by having fully webbed toes (greatly reduced toe webbing in L. hascheanus and L. limborgi). Male secondary sexual characters are unknown in L. khammonensis (Smith 1929), which is known only from the female holotype, but females of L. lauhachindai sp. nov. differ by having a distinct tympanum (indistinct in L. khammonensis) and less toe webbing (Toe IV webbed to distal subarticular tubercle, continuing as fringe to base of disc, and all remaining toes webbed to base of disc in L. khammonensis).

Polypedates discantus sp. nov. Polypedates leucomystax ���Morph B��� Narins, Feng, Yong and Christensen-Dalsgaard, 1998: 129. Polypedates sp. ���Malay Clade��� Kuraishi, Matsui, Hamidy, Belabut, Ahmad, Panha, Sudin, Yong, Jiang, Ota, Thong and Nishikawa, 2013: 1. Holotype. ZMKU AM 0 0 992, adult male (Figs. 4 A and 5), collected at Thung Tam Sao, Hat Yai District, Songkhla Province, Thailand, N06�� 56 ��� 24.9 ������ E 100 �� 15 ��� 15.8 ������, 61 m elevation, on 15 November 2012 with advertisement calls recorded (Table 6; Fig. 3) at 18.35 h at 28.4 ��C by Anchalee Aowphol, Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon, and Natee Ampai. Paratypes. ZMKU AM 00990���00991, ZMKU AM 00997���01000, six adult males, same data as holotype except collected 4 February 2011, 20.20 ���22.00 h, at 24.0 ��C by Anchalee Aowphol, Attapol Rujirawan, Wut Taksintum, Virayuth Lauhachinda, and Somphouthone Phimmachak; ZMKU AM 00993���00996, ZMKU AM 01001���01005, nine adult males, and ZMKU AM 0 1006, one adult female, same data as holotype except collected 15 November 2012, 19.05 ���21.00 h, at 25.0��� 27.5 ��C by Anchalee Aowphol, Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon, and Natee Ampai. Etymology. The specific epithet discantus taken from dis L. for separate and cantus L. for song, in reference to the new species��� differing call from the syntopic P. leucomystax. Suggested common name. Malayan slender tree frog (English). ⊿����k��Dkfln��D: Paad-Reaw-Ma-La-Ewu (Thai), taken from Paad for tree frog, Reaw for slender, Ma-La-Ewu for Malayan. Diagnosis. A medium-sized tree frog (SVL males 48.6���54.7 mm, female 67.5 mm) assigned to the genus Polypedates by having the head longer than broad; vomerine teeth prominent; skin of body and limbs smooth or shagreened; tips of fingers and toes dilated into large, rounded disks having circummarginal grooves; and fingers basally webbed (Liem 1970; Duellman & Trueb 1986; Brown & Alcala 1994), and by its phylogenetic placement based on molecular data (as Polypedates sp. ���Malay Clade��� in Kuraishi et al. 2013). The new species is similar to other members of the P. leucomystax complex, but can be distinguished by the combination of having the skin of head not co-ossified with skull; absent or indistinct white dots on the back of the thigh; presence of vocal sac opening in adult male; smooth skin in orbital region; rounded tubercle on tibiotarsal articulation; and characteristics of the male advertisement call. Description of holotype. Adult male with SVL 49.7 mm; body slender, elongate (SVL:HW = 3.5); head triangular, longer than broad (HL:HW = 1.2); snout obtusely pointed in dorsal view, projecting beyond lower jaw, round at tip; upper eyelid width slightly greater than interorbital distance (ELW:IOD = 1.1); eyes strongly elevated; canthus rostralis distinct; loreal region oblique, slightly concave; pupil horizontal; nostril small and oval, dorsolaterally oriented, much closer to tip of snout than eye; skin of the head not co-ossified with skull; interorbital region slightly convex; tympanum distinct, separated from eye by very narrow distance; tympanum round, diameter smaller than eye (TD:EYE = 0.6); distinct supratympanic fold from eye to shoulder; pineal ocellus absent; vomerine teeth present, in two oblique groups arising on anterior medial margin of choanae, separated by a distance approximate half length of each group; choanae oval, at margins of roof of mouth; lingual papillae absent; tongue attached anteriorly, deeply notched posteriorly; oval vocal sac opening near corner of jaws. Forelimbs slender; relative finger lengths I Color of holotype in life. Above yellowish-brown, below white; dark X-shaped marking weakly visible on interorbital region, neck and shoulders; scattered black blotches on back; weak dark lines running from snout to eyes; narrow dark line extending along canthus rostralis from tip of snout to eye and along supratympanic fold from eye to above shoulder; dorsal surfaces of limbs with narrow dark-cross bands; back of thigh with indistinct white spots (Fig. 6); region around anus black with white-colored tubercles; web of foot dusky; iris grey-brown with dark-brown around the pupil. Color of holotype in preservation. Fading to light grey with dark brown markings. Measurements of holotype. SVL 49.7, HL 17.6, HW 14.5, HD 6.6, ELW 5.6, ED 6.9, IND 3.5, IOD 5.1, SNL 7.7, DNE 5.3, NS 2.3, TD 4.3, FLL 26.2, HLT 16.3, THL 26.5, AGL 24.5, TIL 26.9, FL 20.7, 3 FDW 3.0, 4 TDW 2.1. Variation. Dorsal and ventral patterning is highly variable in this species. Dorsum with dark X-shaped weakly visible in ZMKU AM 00990���00991, 00993���00994, 00996���00997, 0 1002, 0 1004 and 01006; dorsum with four dark longitudinal stripes in ZMKU AM 00999���01001 and 01003; dorsum with two dark longitudinal stripes in ZMKU AM 0 0 995 and 00998; dorsum with scattered black blotches in ZMKU AM 00990���00991, 00993���00994, 00996���01000, 0 1002 and 01005���01006. Chin and anterior chest region with small dark dots in ZMKU AM 00990��� 0 0 991, 00993���00996, 01000���01001, 01003���01004 and 0 1006. ZMKU AM 0 0 995 is missing the terminal phalange in left finger IV and ZMKU AM 0 0 993 has truncated right toe V. Univariate morphological variation in the type series is summarized in Table 7. Calls of holotype. The holotype was calling from a creeping plant 0.7 m above a temporary pond. The advertisement call had three different call types: one-note, two-note and staccato calls (Fig. 3). The one-note call ��� juekk ��� was a single, short, one-pulsed note lasting 0.11 to 0.33 s with dominant frequency of 1.9���2.1 kHz. The two-note call ��� juekk-juekk ��� had two, short, one-pulsed notes lasting 1.63��� 2.02 s; pulse duration of 0.14��� 0.21 s; interval between pulses 1.32��� 1.68 s; pulse repetition rate of 5.23���6.54 pulses per second; and dominant frequency of 2.0��� 2.2 kHz. The staccato call ��� tok-tok-tok ���.......��� was a long call lasting 15.6��� 20.9 s with 11���13, short onepulsed notes; pulse duration 0.05��� 0.08 s; interval between pulses 1.43��� 1.70 s; pulse repetition rate 5.72���6.60 pulses per second; and dominant frequency of 1.4���1.6 kHz (Table 6). Comparisons. Polypedates discantus sp. nov. superficially resembles P. leucomystax (Gravenhorst, 1829), P. megacephalus Hallowell, 1861, P. mutus (Smith, 1940), P. braueri (Vogt, 1911), P. macrotis (Boulenger, 1891), and P. colletti (Boulenger, 1890). Polypedates discantus sp. nov. differs by having indistinct or no white spots on rear of the thigh (distinct spots or marbled pattern present in the other species); skin of head not co-ossified with skull (skin of head co-ossified with skull in P. leucomystax, P. mutus and P. megacephalus); paired vocal sac openings in adult males (absent in P. macrotis and P. mutus); no dark lateral stripe covering tympanum (present in P. macrotis); variable dorsum with X-shaped marking or two or four longitudinal stripes (distinct hour-glass figure beginning from interorbital region to sacrum in P. colletti); smooth skin in orbital region (low tubercles in P. colletti); nuptial pad on first and second fingers (only on first finger in P. colletti); and rounded tubercle on tibiotarsal articulation (conical tubercle in P. colletti; no tubercle in P. leucomystax, P. m a c ro t i s, and P. megacephalus). Size differences among P. discantus sp. nov., P. leucomystax and P. megacephalus are summarized in Table 1. The male advertisement call of P. discantus sp. nov. differs from that of P. leucomystax, P. megacephalus and P. braueri (Table 5 and Kuraishi et al. 2011) by having 1���3 notes per call (one note in P. leucomystax and P. megacephalus); one pulse per note (10���13 pulses in P. leucomystax, 4���7 pulses in P. megacephalus and many fine pulses in P. braueri); pulse duration of 16���37 ms (4���7 ms in P. leucomystax and 6���8 ms in P. megacephalus); interpulse interval of 149���206 ms (5���9 ms in P. leucomystax, 7���9 ms in P. megacephalus); pulse repetition rate of 4���6 s - 1 (79���89 s - 1 in P. leucomystax and 64���75 s - 1 in P. megacephalus); and dominant frequency of 1.7���2.3 kHz (1.1 kHz in P. braueri). Distribution. In Thailand, the new species is known only from the type locality. Narins et al. (1998) reported it as P. leucomystax ���Morph B ��� from Ulu Gombak in Peninsular Malaysia (assigned to P. discantus sp. nov. on the basis of its call characteristics), and Kuraishi et al. (2013) reported it as Polypedates sp. ���Malay Clade��� from Hut. Lip, Kanching Selangor, Kenaboi, Negeri Sembilan, Endau Rompin, Johor, Temerloh, and Pahang, Peninsular Malaysia (assigned to P. discantus sp. nov. on the basis of its 16 S rRNA gene sequence) (Fig. 7). Ecology. The type series was collected around a temporary, road-side pond (Fig. 8). Individuals were found over-hanging the pond on vegetation and creeping plants, 0.3���1.5 m above the pond, co-existing with Microhyla berdmorei, M. heymonsi, M. fissipes, Polypedates leucomystax, P. macrotis, Rhacophorus pardalis and Humerana miopus. No foam-nests or tadpoles were found. Data of P. braueri is from P. leucomystax (Taiwan) in Stejneger (1925) Data of P. braueri is from P. megacephalus (Taiwan) in Matsui et al. (1986), Published as part of Rujirawan, Attapol, Stuart, Bryan L. & Aowphol, Anchalee, 2013, A new tree frog in the genus Polypedates (Anura: Rhacophoridae) from southern Thailand, pp. 545-565 in Zootaxa 3702 (6) on pages 554-559, DOI: 10.11646/zootaxa.3702.6.3, http://zenodo.org/record/284560, {"references":["Narins, P. M., Feng, A. S., Yong, H. S. & Christensen-Dalsgaard, J. (1998) Morphological, behavioral, and genetic divergence of sympatric morphotypes of the treefrog Polypedates leucomystax in peninsular Malaysia. Herpetologica, 54, 129 - 142.","Kuraishi, N., Matsui, M., Hamidy, A., Belabut, D. M., Ahmad, N., Panha, S., Sudin, A., Yong, H. S., Jiang, J. P., Ota, H., Thong, H. T. & Nishikawa, K. (2013) Phylogenetic and taxonomic relationship of the Polypedates leucomystax complex (Amphibia). Zoologica Scripta, 42, 54 - 70. http: // dx. doi. org / 10.1111 / j. 1463 - 6409.2012.00562. x","Liem, S. S. (1970) The morphology, systematics, and evolution of the Old World treefrogs (Rhacophoridae and Hyperoliidae). Fieldiana Zoology, 57, 1 - 145. http: // dx. doi. org / 10.5962 / bhl. title. 2939","Duellman, W. E. & Trueb, L. (1986) Biology of Amphibians. The John Hopkins University Press, Baltimore and London, 670 pp.","Brown, W. C. & Alcala, A. C. (1994) Philippine frogs of the family Rhacophoridae. Proceedings of the Californian Academy of Sciences, 48, 185 - 220.","Smith, M. A. (1940) The amphibians and reptiles obtained by Mr. Ronald Kaulback in upper Burma. Records of the Indian Museum, 42, 465 - 486.","Kuraishi, N., Matsui, M., Ota, H. & Chen, S. L. (2011) Specific separation of Polypedates braueri (Vogt, 1911) from P. megacephalus (Hallowell, 1861) (Amphibia: Anura: Rhacophoridae). Zootaxa, 2744, 53 - 61.","Stejneger, L. (1925) Chinese amphibians and reptiles in the United States National Museum. Proceedings of the United States National Museum, 66, 1 - 115. http: // dx. doi. org / 10.5479 / si. 00963801.66 - 2562.1","Matsui, M., Seto, T. & Utsunomiya, T. (1986) Acoustic and karyotypic evidence for specific separation of Polypedate megacephalus from P. leucomystax. Journal of Herpetology, 20, 483 - 489. http: // dx. doi. org / 10.2307 / 1564245"]}

Rujirawan, Attapol, Stuart, Bryan L., Aowphol, Anchalee (2013): A new tree frog in the genus Polypedates (Anura: Rhacophoridae) from southern Thailand. Zootaxa 3702 (6): 545-565, DOI: http://dx.doi.org/10.11646/zootaxa.3702.6.3