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Leptolalax (Lalos) oshanensis (Liu, 1950) Megophrys oshanensis Liu, 1950: 13, 197. Onymophoront, by original designation as “type”: holophoront (holotype), Liu Collection 1000 (now CIB 1000), adult male, SVL 27.0 mm (Liu 1950: 197–198). Onymotope: Emei Shan (29°35’ N, 103°11’ E; alt. 1070 m), Sichuan, China. Collected by Liu Ch’eng-chao, 10 June 1945. Comment. Leptobrachium minimum was considered a synonym of Megophrys oshanensis by Dubois (1981), but in the genetic analysis, onymotopic specimens of both species appear in distinct clades (Fig. 4). Thus we recognize them as a distinct species. The name L. oshanensis should be restricted to Chinese populations. Here we give data from the original description (Liu 1950), from Fei & Ye (1992) and Fei (1999) for morphological diagnosis. Molecular evidence from sequences deposited in GenBank would indicate that the species as currently defined is not monophyletic. Further studies should be conducted to confirm these results as the sequences here included are from GenBank without indication of voucher specimens. Diagnosis. Member of the subgenus Lalos based on the presence of lateroventral glands and molecular phylogenetic relationships (Fig. 4), distinguished from other species included in this subgenus by the following combination of characters: moderate-sized species (adult males 26.6–30.7 mm) (Table 3) without webbing between toes, without lateral fringes on toes, with distinct tympanum; thin irregular glandular ridges on back (Table 4); back reddish brown with pattern including a triangular spot between eyes and lighter glandular spots near vent; a few large dark spots on flanks; throat with dots, belly uniformly white (Table 5). Description of holophoront (rearranged from original description of Liu 1950; numerous characters missing): Size and general aspect. (1) Specimen of small size (SVL 27.0 mm), body moderately stout. Head. (2) Head of medium size, as broad (HW 9.5 mm) as long (HL 9.5 mm). (3) Snout slightly protruding; (eye length EL 4.0 mm). (4) Canthus rostralis well defined, loreal region concave, oblique. (6) Nostrils closer to tip of snout than to eyes. (8) Tympanum visible (TYD 1.9 mm). (11) Tongue large, deeply notched. Forelimbs. (12) Forearm moderately slender (FLL 13.0 mm; HAL 7.0 mm). (14) Relative length of fingers, shortest to longest: I Distribution. China: Guizhou Province, Hubei Province, Sichuan Province. Natural history. As described by Liu (1950: 199): “ L. oshanensis was found under stones in corn fields on hillsides at about 3500 feet. Adults are difficult to find, as I got only two specimens of adult males in three summer’s work. Tadpoles are very abundant in small mountain streams.” The species is known to occur in mountain areas from 700 to 1800 m. Tadpoles were collected from side pools, in pools beneath cascades, on the bottom of pools, in cracks between stones, hiding in vegetation or roots of bamboos and trees at the margin of the water. They stay in shallow running water with the head directed up-stream and the tail curved like an eel (Fei & Ye 1992). Etymology. Adjective as epithet of generic nomen (same grammatical gender), meaning “from O Shan ”, i. e., from Emei Shan, a famous mountain in Sichuan. Placement in molecular phylogeny. Specimens allocated to Leptolalax “ oshanensis ” do not form a monophyletic group. They are members of a highly supported clade that groups L. eos, L. bourreti and Leptolalax sp. from Doi Chiang Dao. L. “ oshanensis ” AY561306, from the onymotope, and thus probably “true” oshanensis, is sister to L. eos whereas L. “ oshanensis ” AY526215 is sister to Leptolalax sp. from Doi Chiang Dao. Further studies are needed to reliably allocate the name oshanensis to probably one of these mitochondrial lineages and to describe the other of these mitochondrial lineages as new species. Conservation status. We confirm the Red List status of this species as Least Concern. In the future, L. oshanensis might be dismantled into various species and then a re-evaluation will be needed., Published as part of Ohler, Annemarie, Wollenberg, Katharina C., Grosjean, Stéphane, Hendrix, Ralf, Vences, Miguel, Ziegler, Thomas & Dubois, Alain, 2011, Sorting out Lalos: description of new species and additional taxonomic data on megophryid frogs from northern Indochina (genus Leptolalax, Megophryidae, Anura) 3147, pp. 1-83 in Zootaxa 3147 (1) on pages 31-32, DOI: 10.11646/zootaxa.3147.1.1, http://zenodo.org/record/5246747, {"references":["Liu, C. - C. (1950) Amphibians of Western China. Fieldiana: Zoological Memoirs, 2, 1 - 400, 10 pl.","Dubois, A. (1981) Notes sur la systematique et la repartition des Amphibiens Anoures de Chine et des regions avoisinantes. V. Megophrys oshanensis Liu, 1950 et Leptobrachium minimum Taylor, 1962. Bulletin de la Societe Linneenne de Lyon, 50, 182 - 192.","Fei, L. (1999) Atlas of amphibians of China. Zhengzhou: Henan Press of Science and Technology. [In Chinese]."]}

Leptolalax (Lalos) ventripunctatus Fei, Ye & Li, 1991 (Fig. 21–24) Leptolalax ventripunctatus Fei, Ye & Li in: Fei et al. 1991: 95, 97, 213, 274. Onymophoront, by original designation: holophoront (holotype), CIB 890063, adult male, SVL 27.0 mm (Fei et al. 1992: 51). Collected by Luo Jiarui, 17 May 1989. Onymotope: Mengla (21°29’ N, 101°15’ E; alt. 850 m), Yunnan Province, China. Material examined. Laos. Phongsaly Province, Phongsaly water supply river (21°40’ N, 102°06’ E; 1000 m): MNHN 2005.0111 – 0116, adult males; MNHN 2006.2576, 1 tadpole (DNA voucher), coll. Annemarie Ohler, 31 January 2005; — Nathen (21°24’N, 101°58’ E; 600 m): MNHN 2005.0137 – 0140, adult males, MNHN 2006.2577, 1 tadpole (DNA voucher), coll. Annemarie Ohler, 9 February 2005; — Long Nai (21°14' N, 101°56' E; 800 m), MNHN 2004.0270 – 0273, MNHN 2005.0117 – 0136, adult males, coll. Annemarie Ohler, 14 and 15 July 2004 and 4 February 2005; — Nam Ou (22°03’ N, 102°16’ E; 520 m), MNHN 2005.0142 – 0144, adult males, MNHN 2006.2578 – 2588, 11 tadpoles, coll. Annemarie Ohler, 13 February 2005; — Houey Phihet (21°60’ N, 102° 17’ E; 500 m): MNHN 2005.0145, adult male, coll. Annemarie Ohler, 16 February 2005; Vietnam. Vinh Yen District, Tam Dao (21°27’ N, 105°38’ E): MVZ 223641–42, coll. Theodore J. Papenfuss, 22 April 1996. Comment. This species was described from extreme southern Yunnan, near the Lao border. We had the opportunity to do fieldwork on the Lao side and discovered this small-sized Leptolalax which is sympatric with the new species L. eos described above. Specimens allocated to the present species resemble the onymophoront in morphology and are therefore assigned to ventripunctatus (no molecular data exist so far from the Yunnan locality). The collection locality in Laos is less than a hundred kilometres from the onymotope. Most of the Lao specimens were included in the molecular analysis and show great overall genetic similarity. They form a clade with specimens from Tam Dao (MVZ 223641 – 42) which are genetically very similar. In consequence, these specimens are included in L. ventripunctatus. Diagnosis. (Based on specimens from Laos, not on onomophoront). Member of the subgenus Lalos based on the presence of lateroventral glands and molecular phylogenetic relationships (Fig. 4), distinguished from other species included in this subgenus by the following combination of characters: rather small-sized species (males 23.7–27.7 mm; females 31.5–35.0 mm) (Table 3) with rudimentary webbing and no fringes on toes; tympanum distinct; dermal ridges under toes distinct; finger tips slightly enlarged; dorsal skin with indistinct glandular warts (Table 4); dorsum grey brown with spots, rather large spots on flanks; ventral side with distinct dark spots on chest and belly; iris copper above, grey brown below (Table 5). Description of the adult male MNHN 2004.0270. Size and general aspect: (1) Specimen of rather small size (SVL 25.2 mm), body moderately slender. Head. (2) Head of medium size, narrower (HW 8.8 mm) than long (HL 10.0 mm; MN 8.4 mm; MFE 6.7 mm; MBE 3.3 mm), rather flat. (3) Snout slightly protruding, its length (SL 4.4 mm) longer than horizontal diameter of eye (EL 3.6 mm). (4) Canthus rostralis distinct, rounded, loreal region concave, vertical. (5) Interorbital space flat, slightly narrower (IUE 2.9 mm) than upper eyelid (UEW 3.1 mm) but larger than internarial distance (IN 2.7 mm); distance between front of eyes (IFE 5.2 mm) about two-thirds of the distance between back of eyes (IBE 7.8 mm). (6) Nostrils rounded, closer to eye (EN 1.9 mm) than to tip of snout (NS 2.1 mm). (8) Tympanum (TYD 1.9 mm) distinct, rounded; about half eye diameter, tympanum-eye distance (TYE 0.9 mm) half the diameter of tympanum. (11) Tongue moderate, cordate. Forelimbs. (12) Forearm rather short, thin (FLL 7.3 mm), longer than hand (HAL 7.0 mm), not enlarged. (13) Fingers I, II and IV rather short and rather strong; finger III long and rather strong (TFL 3.9 mm). (15) Tip of fingers rounded, slightly enlarged. (16) Fingers without dermal fringe. Hindlimbs. (19) Shanks about four times longer (TL 12.0 mm) than wide (TW 3.2 mm), shorter than thigh (FL 12.2 mm) and than distance from base of internal metatarsal tubercle to tip of toe IV (FOL 12.1 mm); heel reaching mid-eye when limbs folded forward. (20) Toe IV (FTL 6.4 mm) about one-third of the distance from base of tarsus to tip of toe IV (TFOL 19.4 mm). (22) Tip of toes rounded, slightly enlarged. (23) Webbing rudimentary, toes without fringes (MTTF 4.9 mm; MTFF 4.9 mm; TFTF 7.3 mm; FFTF 7.7 mm). (24) Dermal fringe along toe V absent. (26) Inner metatarsal tubercle short and very prominent (conical); its length (IMT 1.3 mm) 2.1 times in length of toe I (ITL 2.7 mm). (27) Tarsal fold absent. Skin. (29) Dorsal and lateral parts of head and body: snout, region between eyes and side of head smooth; back with indistinct glandular warts; flanks with glandular warts. (30) Supratympanic fold distinct, from eye to above shoulder. (31) Dorsal parts of limbs: forelimbs with few indistinct glandular warts; thigh, shank and tarsus smooth. (32) Ventral parts of head, body and limbs: throat, chest and belly smooth; thigh with glandular warts. (33) Presence of macroglands: lateroventral glands present as series of glands forming continous glandular ridge; rather large, distinct rounded or oval femoral and axillary glands, and oval suprabrachial glands. Coloration. In alcohol: (34) Dorsal and lateral parts of head and body: dorsal parts of head and dorsum greyishbrown with whitish zones and two black spots between eyes; flanks grey brown with rather large blackish spots; loreal region light brown with distinct dark brown spot; tympanic region including tympanum light brown with blackish line at supratympanic fold; upper lip light brown with brown bands. (35) Dorsal parts of limbs: dorsal part of forelimbs beige with brown crossbands on lower part; dorsal part of thigh, of shank and of foot beige with brown crossbands; posterior part of thigh brown with a few small white spots. (36) Ventral parts of head, body and limbs: throat whitish with traces of brown; margin of throat light brown with whitish spots; chest whitish with few brown spots; belly whitish with dense brown spots; thigh light brown with whitish spots; webbing brown; macroglands white. In life (Fig. 23E–F): Iris copper in upper third, grey brown in lower part. Dorsal parts of head, of back and of upper part of flanks grey brown with darker spots and black spots between eyes; lower part of flanks greyish with black spots; loreal and tympanic region brown with black canthal and supratympanic band; tympanum brown; upper lip brown with blackish bands; upper part of forelimb orange; lower part greyish brown with blackish bands; thigh, shank and foot greyish brown with black crossbands; posterior part of thigh greyish brown with dark brown bands; throat light grey, transparent; border of throat grey with white spots; chest and belly creamy white with brown spots; ventral part of thigh dark grey brown with white dots; webbing grey brown. Macroglands white. Male secondary sexual characters: (38) Vocal sacs present; indistinct on throat; pair, slit-like openings posterior on mouth floor. Variation. Specimens are rather homogenous in colour pattern and show only minor variation in extent of dark pattern on ventral side and presence of spots on dorsum and lateral side. The specimens also show some variation in presence of warts and skin ridges. Female sexual characters: oviduct convoluted, translucent; ovary with small creamy whitish oocytes. The lateroventral glandular ridges are continous in all specimens. There is some variation in size and shape of femoral and axillary glands that are smaller in some specimens and more or less oval or round in shape. Tadpole. The external morphology is based on a DNA voucher specimen in stage 33 (MNHN 2006.2576; BL 16.1 mm). The description of the tail which is missing in the DNA voucher is based on a specimen in stage 36 (MNHN 2006.2578). In dorsal view (Fig. 24A), body elliptical elongate. In lateral view (Fig. 24B), BW 134 % of BH. Eyes moderately sized, ED 9.6 % of BL, bulging. Nares moderately sized, RN 40 % of NP; NN 77 % of PP. Spiracle is a short tube, small, SS 50 % of BL; opening crescent-shaped, situated on an axis just below apex of myotomes of caudal muscle. Tail long; tail musculature, TMH 82 % of BH, TMW 62 % BW, slightly bulging on less than the proximal third, almost reaching tail tip. Upper fin, SU 108 % of BL, almost nonexistent in its proximal third then increasing rather abruptly; lower fin convex, following largely the caudal muscle; point of maximum height of tail located before its proximal third. Anal tube moderately sized. Lateral line system as in L. bourreti but with an additional line running vertically on the lateral sac from the ventral line until the beginning of the ventral part. Glands present but faintly visible. Oral disc (Fig. 24C) moderately sized, ODW 28 % of BL and 52 % of BW. Marginal papillae of lower labium smaller than those of upper labium and more or less round to short, cylindrical, decreasing in size towards centre then increasing again at the extremity of indentation with four larger papillae; five submarginal papillae on left side and two on right side on lower labium at commissure and in front of keratodont rows P1 and P2. No denticulate papillae. KRF 1:4+4/3+3:1, A4> A5> A3> A2> A1; P2 = P1> P3> P4; A1 about half of P4. Details of keratodonts not studied. Jaw sheaths black; upper sheath a large arch with a median convexity formed by the three median serrations, the median one bigger and stronger than the others; lower sheath hidden, arranged deeply under the upper jaw sheath. Coloration in life: Tadpole MNHN 2006.2578 reddish dorsally with indistinct spots; lateral body light grey with distinct grey spots; belly greyish white. Tadpole MNHN 2006.2577 uniformly greyish brown, its tail beige with dark spots; belly bluish white; oral region whitish grey; iris golden. Coloration in preservative: upper part of body sandy orange, the colour fading from before eyes to snout. Flanks (lymphatic sacs) grey ochre. Ventral side of body dirty white. Caudal muscle from sandy for the upper part to ivory for the lower part in gradation; small light brown spots more numerous on the upper part than on the lower part, giving a mottled aspect. Fins translucent with the same small spots as those on the caudal muscle but forming small blotches isolated from each other on the upper fin; lower fin spotless. Rims of nares white. Tadpole variation is described on the basis of 12 other specimens, except when noted (MNHN 2006.2577, 2006.2579–2588, in stages 25–36, TOL 31.0– 55.9 mm and BL 9.0– 16.1 mm). Only the ratio values different from those of the described specimen are given: BW 129–144 % of BH; ED 8.4–10.7 % of BL; RN 35–58 % NP; NN 70–82 % of PP; SS 42–53 % of BL; TAL 212–251 % of BL (n = 11); TMH 63–85 % of BH; TMH 53–70 % of MTH (n = 11); TMW 48–61 % of BW; UF 28–36 % of MTH (n = 11); SU 92–116 % of BL; LF 22–29 % of MTH (n = 11); MTH 115–126 % of BH (n = 11); ODW 25–32 % of BL (n = 11); ODW 43–49 % of BW (n = 11). The more the tadpoles are in advanced stages, the more their upper fin begins close to body end. The upper fin increases at the first quarter and the lower fin can be straight and so increasing in height anteroposteriorly in a few specimens. The KRF varies from 1:2+2/2+2:1 (stage 25) to 1:4+4/3+3:1 (stages 29–36) through the intermediate KRF 1:3+3/2+2:1 (stages 25–34) and 1:3+3/3+3:1 (stages 27–28). The coloration of the tail is more extended on most of the other specimens: proximal half of caudal muscle tan, this coloration grouping in blotches on the distal half with the ground colour beige, similar blotches on the distal third of the lower fin, upper fin smoked with tan pigments. The lateral spots in the tadpole series described herein are less pronounced than in those figured by Fei et al. (2009). We assume that this might be due to the strong contrast between black and white in the drawing of Fei et al. (2009). Distribution. China : Yunnan Province (Mengla County); Laos: Phongsaly Province (Phongsaly District); Vietnam: Vinh Phuc Province (Vinh Yen District) (Fig. 9). Natural history. Specimens were found (July 2004 and January–February 2005) in forests at a distance up to 1 m from small rivulets (Fig. 36A). Most specimens were very close to the water at a distance of 0.2 to 0.5 m. They were sitting either on the ground (stones, sand, earth) or above the ground on leaves or rocks, usually covered by low vegetation. Only a few specimens were sitting in the water. Most of the specimens were in breeding condition. The tadpoles were collected in a puddle of 40 × 30 cm along a stream (but connected to the stream) with a weak current. They were usually motionless on the bottom, hidden within the gravel. Another specimen was collected in a stream of a width of 10 m and a maximum depth of 50 cm, at a depth of 10 cm. The bottom was covered of stones and dead leaves under which the tadpoles were hidden. In these streams, adults of Limnonectes kuhlii and Polypedates “ leucomystax ” were also observed. Placement in molecular phylogeny. Leptolalax ventripunctatus is part of a highly supported clade that groups L. minimus, L. aereus, L. pluvialis and L. nyx. The specimens allocated to L. ventripunctatus form a well supported clade which, with no support, is sister to the clade that goups L. minimus and L. aereus. Conservation status. This work adds more data on distribution and extent of occurence of this species. It is relatively abundant in convenient habitat, within its relatively small range. It might encounter threats by habitat loss through shift and burning agriculture. We propose therefore to change the status of L. ventripunctatus from Data Deficient to Near Threatened.

Ohler, Annemarie, Wollenberg, Katharina C., Grosjean, Stéphane, Hendrix, Ralf, Vences, Miguel, Ziegler, Thomas, Dubois, Alain (2011): Sorting out Lalos: description of new species and additional taxonomic data on megophryid frogs from northern Indochina (genus Leptolalax, Megophryidae, Anura) 3147. Zootaxa 3147 (1): 1-83, DOI: 10.11646/zootaxa.3147.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3147.1.1

Leptolalax (Lalos) pluvialis Ohler, Marquis, Swan & Grosjean, 2000 (Fig. 17–18) Leptolalax pluvialis Ohler, Marquis, Swan & Grosjean, 2000: 74. Onymophoront: holophoront, by original designation as “ holotype ”, MNHN 1999.5675, adult male, SVL 22.0 mm. Other protaxonts: MNHN 1999.5674 and MNHN 1999.5676, 2 adult males (paratypes), SVL 21.3–22.3 mm. Onymotope: Fan Si Pan mountain range (22°19’ N, 103°47’ E; 1900 m), Lao Cai Province, Vietnam. Collected by Annemarie Ohler and Stéphane Grosjean, 28 July 1998. Material examined. Vietnam. Lao Cai Province, Fan Si Pan mountain range (22°19’ N, 103°47’ E; 1900 m): MNHN 1999.5674 – 5676, adult males (hypodigm). Coll. Annemarie Ohler and Stéphane Grosjean, 28 July 1998. Comment. All three specimens of the hypodigm have been included in the molecular analysis. They form the sister group of specimens allocated to a clade that includes L. aereus, L. minimus, L. nyx and L. ventripunctatus. They can be distinguished morphologically from the other representatives of this clade by their smaller size and the dense brown spotting on belly and chest. The holophoront is redescribed to allow easier comparison and in order that all characters used for the new species are described in the same way. Diagnosis. Member of the subgenus Lalos based on the presence of lateroventral glands and molecular phylogenetic relationships (Fig. 4), distinguished from other species included in this subgenus by the following combination of characters: very small-sized species (males 21.0–22.0 mm) (Table 3) with rudimentary webbing and no fringes on toes; tympanum distinct; dermal ridges under toes distinct; finger tips not dilated; dorsal skin smooth (Table 4); dorsum greyish brown with dark pattern; few black spots on flanks; ventral side with dense brown spots; iris dark golden above (Table 5). Description of the holophoront, adult male MNHN 1999.5675. Size and general aspect. (1) Specimen of small size (SVL 22.0 mm), body moderately slender. Head. (2) Head of medium size, narrower (HW 8.1 mm) than long (HL 8.5 mm; MN 6.7 mm; MFE 5.5 mm; MBE 2.2 mm), convex. (3) Snout slightly protruding, its length (SL 4.0 mm) longer than horizontal diameter of eye (EL 3.3 mm). (4) Canthus rostralis angular, loreal region concave, vertical. (5) Interorbital space convex, slightly larger (IUE 2.8 mm) than upper eyelid (UEW 2.7 mm) but narrower than internarial distance (IN 3.0 mm); distance between front of eyes (IFE 4.6 mm) about two-thirds of the distance between back of eyes (IBE 7.5 mm). (6) Nostrils oval, closer to eye (EN 1.4 mm) than to tip of snout (NS 2.0 mm). (8) Tympanum (TYD 1.7 mm) distinct, rounded; about half of eye diameter, tympanum-eye distance (TYE 0.7 mm) two-fifths of its diameter. (11) Tongue moderate, cordate. Forelimbs. (12) Forearm short, thin (FLL 6.2 mm), longer than hand (HAL 6.0 mm), not enlarged. (13) Fingers I–II and IV rather short and rather strong; finger III long and rather strong (TFL 4.0 mm). (15) Tips of fingers not enlarged. (16) Finger II with dermal fringe. Hindlimbs. (19) Shanks about four times longer (TL 11.4 mm) than wide (TW 3.1 mm), longer than thigh (FL 10.9 mm) and than distance from base of internal metatarsal tubercle to tip of toe IV (FOL 10.8 mm); heel reaching nare when limbs folded forward. (20) Toe IV (FTL 6.35) about one-third of distance from base of tarsus to tip of toe IV (TFOL 17.2 mm). (22) Tips of toes not enlarged. (23) Webbing absent (MTTF 3.9 mm; MTFF 3.7 mm; TFTF 6.6 mm; FFTF 6.7 mm). (24) Dermal fringe along toe V absent. (26) Inner metatarsal tubercle rather long and prominent; its length (IMT 1.7 mm) 1.9 times in length of toe I (ITL 2.2 mm). Skin. (29) Dorsal and lateral parts of head and body: snout, region between eyes, side of head and back smooth; flank with flat glandular warts. (30) Supratympanic fold distinct, from eye to shoulder. (31) Dorsal parts of limbs: forelimbs, thigh, shank and tarsus smooth. (32) Ventral parts of head, body and limbs: throat, chest, belly and thigh smooth. (33) Presence of macroglands: lateroventral ridges present as continous ridges of small glands; large, distinct rounded femoral and oval axillary glands; smaller distinct oval suprabrachial glands. Coloration. In alcohol: (34) Dorsal and lateral parts of head and body: dorsal parts of head and dorsum and upper part of flanks greyish-brown with black band between the eyes, a black W-shaped pattern in the shoulder region and a black V in the sacral region; lower part of flanks dark greyish-brown; loreal region blackish-brown; tympanic region including tympanum dark brown with a white band under the tympanum; upper lip dark brown with white and blackish spots. (35) Dorsal parts of limbs: dorsal part of forelimbs, of thigh, of shank and of foot grey-brown with blackish crossbands; posterior part of thigh blackish with small white spots. (36) Ventral parts of head, body and limbs: throat dirty white; margin of throat dark brown with small white spots; chest and belly dirty white with dark brown marbling; thigh brown with some white spots; macroglands white. In life: Iris dark golden above. Upper parts of head and dorsum, flanks and tympanic region grey brown with dark brown pattern, particularly in tympanum and loreal region; upper lip grey-brown with dark brown and whitish-grey bands. Upper part of forelimbs ochre, forearms grey-brown and dark brown; hindlimbs with dark brown and grey-brown crossbands; heels ochre; posterior part of thigh grey-brown with ventrally a dark brown zone. Throat transparent light grey-brown with dark brown margin bearing white spots; chest and belly grey with dark grey marbling; lower part of thigh transparent dark brown with whitish spots. Macroglands whitish grey. Male secondary sexual characters: (38) Vocal sacs inconspicuous, skin of gular region not modified; inner openings of vocal sacs present, located laterally and posteriorly in the floor of the mouth, distinct; rounded. Variation. The two other male protaxonts resemble strongly the holophoront in coloration and morphology. Female sexual characters: Not observed. Distribution. Known only from onymotope (Fig. 9). Natural history. Three adult male specimens of this species were collected calling during heavy rain, while sitting on branches and leaves near a small forest stream. They can be distinguished from sympatric Leptolalax bourreti by their smaller size, ventral coloration and absence of dark spots on flanks. L. bourreti was very abundant in October and November 1997, and was collected in the precise locality of L. pluvialis, but was rather rare in July, when L. pluvialis was breeding. Etymology. From Latin pluvialis: created by the rain, commanding the rain, rainy. Placement in molecular phylogeny. Leptolalax pluvialis is part of a clade that groups with high support L. aereus, L. minimus, L. nyx and L. ventripunctatus. The specimens allocated to L. pluvialis form a well-supported clade which, with high support, is the sister taxon of a poorly supported clade including all the other taxa mentioned. Conservation status. L. pluvialis is still only known from the original description. Its conservation status remains unchanged: Data Deficient., Published as part of Ohler, Annemarie, Wollenberg, Katharina C., Grosjean, Stéphane, Hendrix, Ralf, Vences, Miguel, Ziegler, Thomas & Dubois, Alain, 2011, Sorting out Lalos: description of new species and additional taxonomic data on megophryid frogs from northern Indochina (genus Leptolalax, Megophryidae, Anura) 3147, pp. 1-83 in Zootaxa 3147 (1) on pages 44-46, DOI: 10.11646/zootaxa.3147.1.1, http://zenodo.org/record/5246747, {"references":["Ohler, A., Marquis, O., Swan, S. & Grosjean, S. (2000) Amphibian biodiversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa, 13, 71 - 87."]}

Hylarana guentheri (Boulenger, 1882) Series examined. Tadpoles of Hylarana guentheri (ZFMK 86987-86988: n = 18) were collected in PNKB, Cha Noi region, Vietnam by R. Hendrix from June to August 2006. The larvae were collected throughout day and night time in ponds (bomb craters) in a paddy field and at the forest edge at an altitude of about 300 m above sea level. An adult specimen (ZFMK 86372) was collected at night near a forest stream within Cha Noi region of PNKB, Vietnam, at an altitude of about 300 m above sea level. Identification. The adult specimen (Fig. 1 c) largely agreed with the description provided by Ziegler (2002). Sequence congruence between the adult specimen (ZFMK 86372) and tadpoles (ZFMK 86987, 86988; Fig. 2 c) was 100 %. The following larval description is based on a single tadpole in Gosner stage 27 (Fig. 5; for measurements see Table 6). The tadpole specimen used for description and drawing originates from the same water body as the DNA voucher specimen. In this water body, no other Hylarana species were observed. The DNA voucher specimen was destroyed for DNA extraction but examined morphologically before this procedure, and it fully agreed morphologically with the described and drawn specimen in the diagnostic characters (narrow upper jaw sheath, cone-shaped marginal papillae, one submarginal row of papillae, papillae unpigmented, silver-white colour with dark dorsoventral stripe). The identification of the described specimen is therefore well supported. ��� Ke Bang National Park, Cha Noi region, Vietnam (ZFMK 86987-86988); for abbreviations see material and methods. 27 (1.34 �� 0.13) (1. 4 7 (0. 5 5 (4. 7 3 (9. 53 (15. 6 2 (1. 43 (1.45 �� 0.11) (1. 0 2 �� 0.20) �� 0.09) �� 0.67) �� 0.97) �� 2.03) �� 0.16) �� 0.15) (n= 10) (n= 10) (n= 10) (n= 10) (n= 10) (n= 10) (n= 10) (n= 10) (n= 10) 28 1.4 1.5 0.5 5.5 11.7 19.0 1.7 1.7 1.0 Colour pattern. Colouration of tadpole in preservative: body glassy-white. Pigmentation is strongest from tip of snout to anterior corner of the eye. Behind the eyes, the body is covered with triangular blotches. Furthermore, the body is covered with densely pigmented blotches in the nares region, laterally in the intestinal coils region and ventrally around the heart. Tail slightly pigmented. Colouration in life: body silver to white, with a short dark stripe from the nostril to the anterior corner of the eye and with a dark fleck behind the eye. Tail fin silver to white with small blotches. Ventral side of body is grey, intestinal spiral is slightly visible. Description in dorsal view: Body is oval shaped (maximum body width 0.64 of body length; body length 0.64 of tail length) with a pointed snout. Eyes of remarkable size (maximum diameter of eye 0.15 of body length), dorsolaterally directed and positioned at the beginning of the second third of the body. Interpupilar distance 0.83 of maximum body width. Nares slightly rimmed, anterolaterally positioned and directed. Nares are closer to the tip of snout than to pupil (rostro-narial distance 0.41 of naro-pupilar distance). Internarial distance 0.51 of interpupilar distance. Tail musculature is of moderate size (width of tail musculature at base 0.37 of maximum body width). Description in lateral view: In lateral view body depressed (maximum body height 0.81 of maximum body width), with pointed snout. Spiracle is tubular-shaped, sinistral, laterally positioned and attached to the body wall. Spiracular opening is round, and closer to the end of body than to the snout. Body length 0.63 of tail length. Height of tail musculature at base 0.97 of tail musculature width. Tail musculature is gradually tapering from the proximal to the distal end. Myotomes of tail are V-shaped and from the second third of the tail onwards separated by broad myosepts. Upper fin of tail situated at the base of tail, slightly convex. Maximum height of lower fin 0.78 of maximum height of upper fin. Vent tube is located ventromedially; the cloacal aperture is dextral in dorsal view. Outer wall of vent tube adhered to lower tail fin. Oral disc. Oral disc anteroventrally positioned, trapezoidal, emarginated (oral disc width 0.38 of maximum body width) and nearly surrounded by marginal cone-shaped papillae. Upper labium with large medial gap without papillae. Marginal papillae of the lower labium elongated. Two or three submarginal papillae situated in the corner of the mouth, one row of submarginal papillae situated at the lower labium under the last keratodont row. Keratodont row formula 2 (2)/ 3 (1) or 1 / 3 (1). Upper and lower beaks black, with low and fine serrations. Upper beak is M-shaped, wide with short appendices, lower beak V-shaped. Glandular Zones. Areas with high density of secretory glands are not discernible. Measurements (in mm). BH 3.2 �� 0.2; BL 6.1 �� 1.3; BW 3.9 �� 0.3; ED 0.9 �� 0.1; IND 1.5 �� 0.1; IP 3.2 �� 0.2; LF 0.8 �� 0.1; NP 31 �� 5; NPD 1.3 �� 0.1; ODW 1.5 �� 0.2; RND 0.6 �� 0.1; TAL 9.5 ��1.0; TMH 1.4 �� 0.2; TMW 1.5 �� 0.1; UF 1.0�� 0.2. Variation within the series. Series with 18 tadpoles in stages 25���30. Proportions vary as follows: BW 0.54���1.35 of BL, ED 0.13���0.29 of BL, IP 0.74���0.90 of BW, RND 0.31���0.50 of NPD, IND 0.42���0.60 of IP, TMW 0.30���0.43 of BW, BH 0.71���0.90 of BW, TAL 1.09���2.88 of BL, TMH 0.35���0.53 of BH, TMH 0.42���0.56 of MTH, UF 0.27���0.40 of MTH, LF 0.57���1.33 of UF, ODW 0.29���0.44 of BW. Keratodont row formula 2 (2)/ 3 (1) or 1 / 3 (1). General morphology and colouration in stages 25���30 is homogenous within the series. Previous descriptions. In comparison to the data provided by Pope (1931) for larvae from Nodoa, China (at about stage 39) our series differed only slightly. In our tadpoles, tail fins measured about 1.5 of body length, whereas the tail fins measured at least 2.0 of body length in the Chinese larvae. The first keratodont row P 1 of the lower labium is only slightly interrupted according to Pope���s (1931) description for Chinese larvae, whereas P 1 is distinctly interrupted within the tadpoles from Phong Nha ��� Ke Bang, Vietnam. Natural history. Larvae of H. guentheri were collected in a bomb crater next to a rice field in Phong Nha village and in a bomb crater near the forest edge at Cha Noi region between 10 and 14 August 2006. The larvae were found near the shore area in the shallow water on the sandy ground between leaves, plants and in part also waste. Adults were seen near water bodies in anthropogenic areas such as rice fields, grassland, waysides or forest clearings especially throughout or after rain falls in altitudes up to 300 m a.s.l. Sporadical mating calls of males were heard in the dry season but increased with the begin of the rainy season. Spawn was detected as a layer on the water surface of a pond in early July 2006. The yellow-black eggs measured 1.5���2 mm in diameter., Published as part of Gawor, Anna, Hendrix, Ralf, Vences, Miguel, B��hme, Wolfgang & Ziegler, Thomas, 2009, Larval morphology in four species of Hylarana from Vietnam and Thailand with comments on the taxonomy of H. nigrovittata sensu latu (Anura: Ranidae), pp. 1-25 in Zootaxa 2051 on pages 12-14, DOI: 10.5281/zenodo.186587, {"references":["Ziegler, T. (2002) Die Amphibien und Reptilien eines Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur & Tier Verlag, Munster, 342 pp.","Pope, C. H. (1931) Notes on amphibians of Fukien, Hainan, and other parts of China. Bulletin of the American Museum of Natural History, 61, 397 - 611."]}

Hylarana nigrovittata (Blyth, 1856) from central Vietnam Series examined. Tadpoles of Hylarana nigrovittata (ZFMK 87279: n = 21) were collected in PNKB, Khe Van region, Vietnam by R. Hendrix in July 2006. The larvae were collected throughout day and night time in a primary forest stream at an altitude of about 130 m above sea level. Measurements Males (n = 3) SVL 37.5 ��� 43.0 (39.50 �� 3.04) SL 7.0 ��� 7.5 (7.17 �� 0.29) NL 3.0 ��� 3.5 (3.33 �� 0.29) EN 4.0 ��� 4.5 (4.17 �� 0.29) HW 14.5 ��� 17.0 (15.50 �� 1.32) HL 14.0 ��� 16.0 (15.00 �� 1.00) EE 8.5 ��� 9.0 (8.83 �� 0.29) ED 5.5 TD 4.0 ��� 4.5 (4.30 �� 0.26) HaL 12.0 ��� 12.5 (12.33 �� 0.29) LL 68.0 ��� 73.0 (71.00 �� 2.65) FL 22.0 ��� 23.5 (22.50 �� 0.87) IMT 2.0 ��� 2.5 (2.23 �� 0.25) Identification. Adult H. nigrovittata specimen (ZFMK 71298-71300) from Phong Nha - Ke Bang, Quang Binh Province, Vietnam (Fig. 1 d; for measurements see Table 7) largely agreed with the description provided by Inger et al. (1999) and Ziegler (2002). Sequence congruence between syntopically sampled adult specimens and tadpoles (ZFMK 87279 a���d) was 99.8 %. The following larval description is based on a single tadpole in Gosner stages 35 / 36 (Fig. 6; for measurements see Table 8). The tadpole specimen used for description and drawing originates from the same water body as the DNA voucher specimen (characters of the DNA voucher specimen of series ZFMK 87279: stage 29, BL 8.5 mm, NP 52, NK 35 per 0.5 mm). In this water described specimen is therefore body, also H. guentheri (adults only) was found. The DNA voucher specimens fully agree morphologically with the described and drawn specimen, including the characters that diagnose H. nigrovittata from H. guentheri (narrow upper jaw sheath, finger-shaped marginal papillae, one submarginal row of papillae, papillae pigmented at the basis, reddish to olive brown marbled colour with light blotches on tail). The identification ofwell supported. Colour pattern. Colouration of tadpole in preservative: body marbled, dorsally and laterally covered with brownish to olive-coloured blotches. Pigmentation is strongest around the nares, in the anterior and posterior corners of the eyes, and in the region of the intestinal coils. Single green pigments can be found around the mouth and on the throat region. Chest and belly white to transparent, without pigmentation, intestinal spiral well visible in ventral view. Tail marbled with brown to green pigments. Musculature of tail is white to yellow, upper and lower fin transparent. Hind limbs yellow to white, dorsolaterally with pigments. Colouration in life: body marbled, dorsally and laterally with drab, olive-coloured to brown blotches. Ventral side of body olive-coloured, slightly transparent. Throat and belly silvery. Musculature and lower fin of tail grey to olive with light-coloured blotches. Upper fin of tail is transparent. Description in dorsal view: Body shape oval, elongated (maximum body width 0.60 of body length), with a rounded snout. Eyes (maximum eye diameter 0.11 of body length) directed and positioned dorsolaterally, at the end of first body third, not visible in ventral view (interpupilar distance 0.57 of maximum body width). Nares small and rounded, anterodorsally positioned (rostro-narial distance 0.56 of naro-pupilar distance). Internarial distance 0.63 of interpupilar distance. Tail musculature well developed (width of tail musculature at base 0.49 of maximum body width). Description in lateral view: Body depressed (maximum body height 0.69 of maximum body width). Snout rounded. Spiracle sinister, laterally positioned (distance from snout tip to opening of spiracle 0.67 of body length), conical-shaped, oriented in posterodorsal direction and slightly visible in dorsal view. Tail length 1.78 of body length. Musculature of tail well developed (height of tail musculature at base 0.47 of maximum body height, and 0.45 of maximum tail height). Tail musculature runs parallel, tapering from the base to the middle of tail fin, thereafter continuously tapering towards the tip of the tail. Upper fin situated at base of tail, slightly convex (maximum height of upper fin 0.34 of maximum tail height). Maximum height of lower fin 0.60 of maximum height of upper tail fin. Vent tube medially located at body end, ventrally positioned, cloacal aperture dextral in dorsal view and not adhered to lower tail fin. Oral disc. Oral disc anteroventrally positioned, laterally emarginated (oral disc width 0.38 of maximum body width). Oral disc framed by finger-shaped, pigmented papillae except for a large medial gap on the upper labium. All papillae show a pointed end and are transparent. Marginal papillae of the upper labium elongated. Few submarginal papillae situated on each side between upper and lower labium in the corner of mouth. In addition there is a row of submarginal papillae at the lower labium under the last keratodont row. In total 54 papillae. Keratodont row formula 2 (2)/ 3 (1), with 28 keratodonts per 0.5 mm. Jaw sheath black, and lightly serrated. Upper jaw sheath curved with large appendices, lower jaw sheath V-shaped. continued. (n= 3) - (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) Glandular Zones. Secretory glands are discernible on each side of the body, situated dorsolaterally at midbody, laterally and posteroventrally at the end of body and ventrally under the mouth in the throat region. Measurements (in mm). BH 4.2; BL 10.2; BW 6.1; ED 1.1; IND 2.2; IP 3.5; LF 0.9; MTH 4.4; NK 28; NP 54; NPD 1.8; ODW 2.3; RND 1.0; TAL 18.2; TMH 2.0; TMW 3.0; UF 1.5. Variation within the series. Series with 21 tadpoles in stages 24���41. Proportions vary as follows: BW 0.54���0.70 of BL, ED 0.11���0.15 of BL, IP 0.43���0.63 of BW, RND 0.36���0.62 of NPD, IND 0.53���0.71 of IP, TMW 0.25���0.52 of BW, BH 0.45���0.72 of BW, TAL 1.03���2.10 of BL, TMH 0.45���0.71 of BH, TMH 0.45���0.67 of MTH, UF 0.24���0.42 of MTH, LF 0.40���1.10 times UF, ODW 0.19���0.47 times BW. In early developmental stages, the body skin is more transparent to white with only slight pigmentation, and the keratodont row formula is not definitive (only from stage 25 onwards). The intestinal spiral is always well visible in lateral and ventral view. Previous descriptions. In comparison to the data provided by Smith (1916), Bourret (1942) and Manthey & Grossmann (1997) for larvae from Thailand, and compared with the data provided by Grosjean (2005) for Ben En National Park, southern Vietnam, we found the following differences: According to Smith (1916) the tail fin is about twice as large as the body. This corresponds with larvae in more advanced stages (34���41) from Phong Nha ��� Ke Bang, whereas the tail of younger larvae (stages 25���28) is only slightly longer than the body. The length of tail fins measured by Grosjean (2005) for H. nigrovittata is less than one and a half of body length across all stages of development. In addition, Grosjean (2005) mentions only four pairs of glandular zones which are located dorsally between the eyes, dorsolaterally at midbody, dorsolaterally at the body end in front of the tail insertion, and posteroventrally at the body end. In the larvae from Phong Nha ��� Ke Bang, we could observe an additional pair of glands that is located posteriorly to the mouth in the throat region. Bourret (1942) mentions three uninterrupted keratodont rows [2 (2)/ 3] on the lower labium of H. nigrovitatta larvae, whereas the tadpoles from Phong Nha ��� Ke Bang have an inner interrupted row and two outer continuous keratodont rows [2 (2)/ 3 (1)] on the lower labium which agrees with descriptions by Smith (1916) and Grosjean (2005). Manthey & Grossmann (1997) give keratodont row formulae of 2 (2)/ 3 (1) or 3 (2���3)/ 3 (1). However, the latter formula and an additional row of interrupted keratodonts on the upper labium respectively was neither observed in our specimens nor by Grosjean (2005). Natural history. Larvae of H. nigrovittata were collected during day time in a primary forest stream pool in the Khe Van area of Phong Nha ��� Ke Bang National Park, Vietnam, at 19 July 2006. The stream pools were surrounded by larger rocks and measured up to 80 x 60 cm in diameter with water levels up to 10 cm. The larvae were found near the sandy ground which was completely covered by a thick layer of algae. During the night time (10:00 p.m.) of 19 July 2006, air temperature was 27.2 ��C, relative humidity was 75 %, water temperature was 27 ��C, and pH-value was 6.6. Larval densities were relatively high, with about 40 individuals in Gosner stages 25���41 in a pond of 60 x 40 cm diameter. Throughout day time, the larvae were hidden among the algae. The larvae were found motionless on the layer of algae and escaped into this layer when disturbed. Adult H. nigrovittata were regularly found in and near slowly running forest streams within the vegetation, under rocks and logs and in rock crevices up to 300 m a.s.l. Mating calls of males were regularly heard between 06:00 and 12:00 p.m. except for heavy rain falls. We once counted at least 30 calling males along 100 to 150 m forest stream length. Layers of eggs were seen several times on the water surface in former crab burrows at river banks of slowly running streams between 20 June 2006 and 0 6 August 2006. The whitishblack yolk of eggs measured 1.5 mm in diameter and they always were found on the water surface., Published as part of Gawor, Anna, Hendrix, Ralf, Vences, Miguel, B��hme, Wolfgang & Ziegler, Thomas, 2009, Larval morphology in four species of Hylarana from Vietnam and Thailand with comments on the taxonomy of H. nigrovittata sensu latu (Anura: Ranidae), pp. 1-25 in Zootaxa 2051 on pages 14-18, DOI: 10.5281/zenodo.186587, {"references":["Blyth, E. (1856) Report for October meeting, 1855. Journal of the Asiatic Society of Bengal, Calcutta, 24, 711 - 723.","Inger, R. F., Orlov, N. L. & Darevsky, I. (1999) Frogs of Vietnam: A report on new collections. Fieldiana Zoology, N. S., 92, 1 - 46.","Ziegler, T. (2002) Die Amphibien und Reptilien eines Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur & Tier Verlag, Munster, 342 pp.","Smith, M. A. (1916) Descriptions of five tadpoles from Siam. Journal of the Natural History Society of Siam, London, 2, 37 - 43.","Bourret, R. (1942) Les batraciens de l'Indochine. Institut Oceanographique de l'Indochine, Hanoi, 547 pp.","Manthey, U. & Grossmann, W. (1997) Amphibien und Reptilien Sudostasiens. Natur und Tier Verlag, Munster, 512 pp.","Grosjean, S. (2005) The choice of external morphological characters and developmental stages for tadpole-based anuran taxonomy: a case study in Rana (Sylvirana) nigrovittata (Blyth, 1855) (Amphibia, Anura, Ranidae). Contributions to Zoology, 74, 1 - 22."]}

Hylarana attigua (Inger, Orlov & Darevsky, 1999) Series examined. Tadpoles of Hylarana attigua (ZFMK 87280-87286: n = 7) were collected in PNKB, U Bo region, Vietnam by R. Hendrix in August 2006. The larvae were collected throughout day and night time in a water catchment basin next to a forest road at an altitude of about 700 m above sea level. Adult specimens (ZFMK 86404 -86407, 87469) were collected at night within PNKB in the primary forest of U Bo region at an altitude of about 530 m above sea level. Identification. Adult specimens (Fig. 1 a; for measurements see Table 2) largely agreed with the original description provided by Inger et al. (1999). Sequence congruence between adults (ZFMK 86404 -407, 87469; additional specimens deposited in PNNP) and tadpoles (ZFMK 87280, 87283- 87284; Fig. 2 a) was at least 99.4 %. The following larval description is based on a single tadpole in Gosner stage 36 (Fig. 3; for measurements see Table 3). The tadpole specimen used for description and drawing originates from the same water body as the DNA voucher specimens (characters of three DNA voucher specimens of the series ZFMK 87280 and ZFMK 87283-87284: stages 37-40, BL 14.0-16.0 mm, NP 64-66, NK 27-29 per 0.5 mm). In this water body, also H. guentheri (adults only) and H. maosonensis (larvae) were found. The DNA voucher specimens fully agree morphologically with the described and drawn specimen, including the characters that diagnose H. attigua from H. maosonensis and H. guentheri (narrow upper jaw sheath, finger-shaped marginal papillae, papillae slightly pigmented on tips, one submarginal row of papillae, dark olive brown colour). The identification of the described specimen is therefore well supported. Colour pattern. Colouration of tadpole in preservative: body with dark brown and green pigments. Dorsal and dorsolateral pigmentation of body more dense than ventral pigmentation. Venter semitransparent to white. Oral disc, throat and chest with single green pigments. Hind part of body with distinct ventrolateral pigmentation. Tail marbled with dark drab pigments and with irregular dark round flecks. Musculature of tail yellowish in dorsal view, with the upper and lower fins being transparent to white. Hind limbs dorsolaterally with dark brown pigments. Colouration in life: body marbled, greenish-brown with gold-coloured pigments. Ventral body transparent to white, intestinal spiral slightly visible. Internal gills gleam reddish through the skin. Description in dorsal view: Body shape oval, elongated (maximum body width 0.59 of body length) and with a rounded snout. Anteroventrally positioned mouth. Eyes of remarkable size (maximum eye diameter 0.13 of body length), dorsolaterally positioned at the first third of body, not visible in ventral view (interpupilar distance 0.62 of maximum body width). Nares small, anterodorsally positioned, nearer to the snout than to the pupil (rostro-narial distance 0.37 of naro-pupilar distance). Internarial distance 0.56 of interpupilar distance. Tail musculature well developed, widest at base (width of tail musculature at base 0.52 of maximum body width). 40 (6.60 ± 0.78) (14.67 ± 1.15) (8.70 ± 0.56) (1.87 ± 0.06) (2.97 ± 0.06) (5.33 ± 0.35) 0.8 7.2 (28 ± 1) (64 ± 1) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) 41 6.8 16.0 9.8 1.8 2.8 5.8 1.2 7.2 33 60 Description in lateral view: Body slightly depressed, with maximum height behind the spiracle (maximum body height 0.70 of maximum body width). Snout rounded. Distance from the spiracle opening to centre of eye 5.8 mm. Tubular, tapered spiracle sinistral, laterally positioned at the second third of body, posterodorsally directed and visible in dorsal view. Vent tube medially located at body end, ventrally positioned. Outer wall of vent tube adhered to lower tail fin. Cloacal aperture dextral to caudal fin in dorsal view. Tail large, pointed at the end. Tail length 2.46 of body length. Myotomes of the tail musculature are moderately developed, V-shaped (height of tail musculature at base 0.74 of maximum body height, and 0.67 of maximum tail height). Tail musculature parallel, tapering from the base to the end of the second third of the tail fin, subsequently gradually tapering towards the tip of the tail. Upper fin inserting in the centre of the first third of the tail. Highest point of upper fin at the second third of the tail (maximum height of upper fin 0.28 of maximum tail height). Maximum height of lower fin 0.59 of maximum height of upper tail fin. Oral disc. Oral disc anteroventrally positioned, of trapezoidal shape and laterally emarginated (oral disc width 0.43 of maximum body width). Oral disc framed by finger-shaped, slightly pigmented papillae except for a large medial gap on the upper labium. Few submarginal papillae situated on each side between upper and lower labium in the corner of the mouth. Single submarginal papillae at the lower labium under the last keratodont row. In total 65 papillae. Keratodont row formula 2 (2)/ 3 (1) with about 35 keratodonts per 0.5 mm. Jaw sheath black, and slightly serrated. Upper jaw sheath curved with large appendices, lower jaw sheath Vshaped. Glandular Zones. Areas with high density of secretory glands on each side of body, dorsally between the eyes, in front of the eyes and dorsolaterally at midbody. Further secretory glands were concentrated laterally at the end of the body, ventrolaterally before hind limbs and ventrally below mouth. Measurements (in mm). BH 5.4; BL 13.0; BW 7.7; ED 1.7; IND 2.7; IP 4.8; LF 1.0; MTH 6.0; NK 35; NP 64; NPD 2.3; ODW 3.3; RND 1.1; TAL 32.0; TMH 4.0; TMW 4.0; UF 1.7. Variation within the series. Series with seven tadpoles in stages 35–41. Proportions vary as follows: BW 0.51–0.68 of BL, ED 0.11–0.14 of BL, IP 0.55–0.77 of BW, RND 0.32–0.48 of NPD, IND 0.48–0.58 of IP, TMW 0.43–0.56 of BW, BH 0.69–0.82 of BW, TAL 1.93–2.46 of BL, TMH 0.59–0.74 of BH, TMH 0.56–0.68 of MTH, UF 0.28–0.35 of MTH, LF 0.40–0.59 of UF, ODW 0.34–0.43 of BW. Keratodont row formulae are homogenous within the series. In further developed stages secretory glands extend ventrolaterally at the end of body and ventrally at the throat. Furthermore, there exists a dark-grey dorsolateral stripe on each side of body, from posterior corner of the eye to the end of body. Natural history. Larvae of H. attigua were collected in a water catchment basin next to the highway at km 51 in the U Bo region, Phong Nha – Ke Bang National Park, Vietnam, at 22 August 2006. The tadpoles were discovered between algae on the ground of the water body which had a water depth of only 15 cm. During night time (08: 30 p. m.) of 7 July 2006, water temperature was 26 °C. At day time, this basin was at least in part exposed to the sun. During day time of 22 August 2006 (12:00) the pH value was 5.5. Adult males of H. attigua were found on rocks and under bushes in the U Bo region at night of 7 July 2006. The males were calling next to a forest stream, about 40 cm above the water surface. Air temperature next to the forest stream was 25 °C; relative humidity was 85 % during night time. Another male was discovered in primary forest next to the highway at km 54 in the U Bo region at night of 30 July 2006. The male was in a bush about 60 cm above the forest ground next to a small forest path. In that same night also a female (ZFMK 86406) was found while trying to cross the highway.

Hylarana maosonensis Bourret, 1937 Series examined. Tadpoles of Hylarana maosonensis (ZFMK 87287-87305: n = 19) were collected in PNKB, U Bo region, Vietnam by R. Hendrix from June to August 2006. The larvae were collected throughout day and night time in primary forest streams, in ponds and in a catchment basin next to a forest road at altitudes between 530���700 m above sea level. Adult specimens (ZFMK 86374, 86399- 86400) were collected at night near primary forest streams and ponds within U Bo region of PNKB, Vietnam, at altitudes between 530���700 m above sea level. Identification. Adult specimens (Fig. 1 b; for measurements see Table 4) largely agreed with the descriptions by Bourret (1942), Inger et al. (1999), Ziegler (2002) and Ziegler et al. (2006). Sequence congruence between adults (ZFMK 86373, 86399- 400) and tadpoles (ZFMK 87293 - 295, 87300, 87305; Fig. 2 b) was 100 %. The following larval description is based on a single tadpole in Gosner stage 36 (Fig. 4; for measurements see Table 5). The tadpole specimen used for description and drawing originates from the same water body as the DNA voucher specimens (characters of the five DNA voucher specimens of the series ZFMK 87300, 87305, and 87293-87295: stages 28-40, BL 7.1-15.1 mm, NP 55 in stage 28, 78- 93 in stages 35-40, NK 26-29 per 0.5 mm). In this water body, also H. guentheri (adults only) and H. attigua (larvae) were found. The DNA voucher specimens fully agree morphologically with the described and drawn specimen, including the characters that diagnose H. maosonensis from H. attigua and H. guentheri (broad upper jaw sheath, finger-shaped marginal papillae, two submarginal rows of papillae, papillae strongly pigmented in their distal halfs, dark olive brown colour with broad dark spots and bright blotch on throat). The identification of the described specimen is therefore well supported. Measurements Males (n = 3) SVL 35.5 ��� 37.5 (36.50 �� 1.00) SL 6.0 ��� 7.0 (6.17 �� 0.76) NL 2.5 ��� 3.0 (2.77 �� 0.25) EN 3.0 ��� 3.2 (3.07 �� 0.12) HW 12.5 ��� 13.8 (13.27 �� 0.68) HL 13.0 ��� 14.5 (13.57 �� 0.81) EE 7.0 ��� 8.5 (7.83 �� 0.76) ED 5.5 ��� 6.0 (5.83 �� 0.29) TD 3.5 ��� 4.0 (3.67 �� 0.29) HaL 10.0 ��� 13.0 (11.17 �� 1.61) LL 70.0 ��� 75.5 (72.33 �� 2.84) FL 20.0 ��� 23.0 (21.67 �� 1.53) IMT 1.5 ��� 2.2 (1.90 �� 0.36) Stage BH BL BW ED IND IP LF MTH NK NP 28 2.8 7.1 4.5 0.8 1.9 2.7 - - 29 55 3.7���4.3 8.8���9.2 3.0��� 5.7 1.0��� 1.1 1.0��� 1.1 3.1���3.3 0.9 ���1.0 4.3���4.5 26���28 48���60 30 (4. 0 0 (9. 0 0 (4. 7 3 (1. 0 3 (1. 0 3 (3. 1 7 (0. 9 5 (4. 40 (27 �� 1) (54 �� 8) �� 0.30) �� 0.20) �� 1.50) �� 0.06) �� 0.06) �� 0.08) �� 0.07) �� 0.14) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 3) (n= 2) (n= 2) (n= 3) (n= 2) 32 4.7 10.0 6.3 1.8 2.1 3.7 1.3 5.7 32 87 4.9���5.1 11.1���12.6 6.8 ���7.0 1.5���1.6 2.2���2.6 3.8���4.3 29���30 77���80 35 (5. 0 0 (11. 8 5 (6. 9 0 (1. 5 5 (2. 4 0 (4. 0 5 1.3 5.8 (30 �� 1) (79 �� 2) �� 0.14) �� 1.06) �� 0.14) �� 0.07) �� 0.28) �� 0.35) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) 36 4.7 12.7 7.2 1.6 2.7 4.4 1.2 6.1 34 85 37 6.6 12.8 7.8 1.8 2.8 4.5 1.9 7.2 29 - 38 5.9 12.1 7.6 1.6 2.6 4.6 1.6 6.7 27 90 6.0��� 7.5 13.7���15.1 8.4���10.8 1.8���2.1 2.8���3.1 5.3���6.2 1.0��� 2.2 7.2 ���8.0 27���30 78���98 40 (6. 8 2 (1 4. 5 2 (9. 5 8 (1. 9 0 (2. 9 6 (5. 6 2 (1. 6 4 (7. 58 (28 �� 1) (89 �� 9) �� 0.63) �� 0.54) �� 0.90) �� 0.14) �� 0.11) �� 0.36) �� 0.46) �� 0.30) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) (n= 5) 41 6.5 12.3 8.1 1.9 2.3 5.0 0.8 6.0 29 - 42 6.1 12.3 8.2 2.1 2.3 5.0 1.4 5.2 28 - 5.1���5.2 11.1���11.5 6.40 1.9 ���2.0 1.3���1.4 3.7���3.8 0.5���0.7 4.1���4.2 - 45���62 43 (5. 1 5 (11. 3 0 (6. 3 8 (1. 9 3 (1. 3 5 (3. 7 5 (0. 6 0 (4. 13 - (54 �� 12) �� 0.07) �� 0.28) �� 0.04) �� 0.11) �� 0.07) �� 0.07) �� 0.14) �� 0.04) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) (n= 2) - (n= 2) 2.6���3.2 3.6���4.1 1.0��� 1.5 9.4���10.4 33.0���35.0 47.7���49.8 3.8���4.2 3.8���4.1 2.6 ���3.0 40 (2.80 �� 0.23) (3.86 �� 0.21) (1.18 �� 0.25) (9.80 �� 0.41) (34.00 (48.75 (4.02 �� 0.20) (4.02 �� 0.13) (2.84 �� 0.18) �� 1.41) �� 1.48) 43 (2.23 �� 0.11) (1.86 �� 0.34) (0.58 �� 0.04) - 21.0 32.5 (2.60 �� 0.28) (2.85 �� 0.07) (1.48 �� 0.04) (n= 2) (n= 2) (n= 2) - (n= 2) (n= 2) (n= 2) Colour pattern. Colouration of tadpole in preservative: body semitransparent, dorsolaterally light brown, greyish to yellowish, ventrally whitish. Body marbled with dark brown to dark green irregular spots. Dorsal and dorsolateral pigmentation of body is more dense than on the venter. On the ventral side, the throat and chest region is somewhat stronger pigmented especially around the oral disc. Tail musculature is yellowish, upper and lower tail fin almost transparent. The tail is covered with green to grey pigments and single dark brown irregular spots. Dorsolateral parts of hind limbs with green to grey pigments. Colouration in life: body olive-coloured to brownish with gold-coloured pigments. Furthermore, body and tail covered with remarkable dark flecks. Iris of eye marbled with four black flecks at the outermost margin. Ventral side of body is transparent to white, intestinal spiral slightly visible, with a bright and slightly pigmented fleck at the throat. Internal gills reddish glimmering. Description in dorsal view: Body of tadpole reaches a length of 12.7 mm, oval elongated, with a rounded snout. Maximum body width is between eye and spiracle (maximum body width 0.57 of body length). Nares anterodorsally positioned and directed, nearer to the tip of the snout than to the pupil (rostro-narial distance 0.50 of naro-pupilar distance). Internarial distance 0.61 of interpupilar distance. Eyes dorsolaterally positioned and directed, at the end of the first body third. Eyes are of remarkable size (maximum diameter of eye 0.13 of body length). Maximum interpupilar distance is 0.61 of body width. Muscles of tail are widest at base (width of tail musculature at base 0.44 of maximum body width). Description in lateral view: Body laterally depressed (maximum body height 0.65 of maximum body width). Cone-shaped spiracle sinistral, laterally positioned at the beginning of the last third of body, posterodorsally directed. Vent tube separated from body. Distance of the spiracle opening from centre of eye is 5.4 mm. Vent tube ventromedially located between hind limbs. Outer wall of vent tube adhered to lower fin. Cloacal aperture is situated dextral in dorsal view. Tail length is 1.78 of body length, with a pointed tail tip. Tail musculature distinct (height of the tail musculature at base 0.70 of body height), running parallel from the base to the end of the first third of tail fin, then gradually tapering to the tip of the tail. Upper tail fin situated behind the base of tail, slightly convex, with maximum height at the last third of tail fin. Maximum height of lower fin is 0.55 of maximum height of upper fin. Oral disc. Oral disc anteroventrally positioned (oral disc width 0.46 of maximum body width) and laterally emarginated. Oral disc framed by finger-shaped, marginal papillae except for a large medial gap on the upper labium. Marginal papillae of the lower labium elongated. Single submarginal papillae situated laterally at both ends of the beak between the upper and lower labium. Two additional rows of submarginal papillae situated on the lower labium. Pigmentation is somewhat stronger at the end of papillae. Keratodont row formula 2 (2)/ 3 (1), with about 34 keratodonts per 0.5 mm. Jaw sheaths black, robust, and serrated. Upper jaw sheath remarkably developed and curved with long appendices. Lower jaw sheath V-shaped. Glandular Zones. Areas with a high density of secretory glands are oval-shaped, whitish and with dark brown pigments. Glands are situated on each side of the body, dorsally between the anterior corner of the eyes, dorsolaterally at midbody and laterally at the end of body. Additional secretory glands are situated ventrally on the throat and ventrolaterally before the hind limbs. Measurements (in mm). BH 4.7; BL 12.7; BW 7.2; ED 1.6; IND 2.7; IP 4.4; LF 1.2; MTH 6.1; NK 34; NP 85; NPD 2.2; ODW 3.3; RND 1.0; TAL 22.6; TMH 3.3; TMW 3.2; UF 2.2. Variation within the series. Series with 19 tadpoles in stages 28���43. Proportions vary as follows: BW 0.33���0.73 of BL, ED 0.11���0.18 of BL, IP 0.53���1.08 of BW, RND 0.22���0.69 of NPD, IND 0.34���0.70 of IP, TMW 0.33���0.73 of BW, BH 0.61���1.33 of BW, TAL 1.52���2.36 of BL, TMH 0.47���0.70 of 0.70, TMH 0.48���0.71 of MTH, UF 0.33���0.43 of MTH, LF 0.31���0.76 of UF, ODW 0.25���0.73 of BW. Keratodont row formula is homogenous within specimens. In earlier stages, oral papillae are less developed, the skin of body is more transparent and the internal organs are well visible in lateral and ventral view. Natural history. Larvae of H. maosonensis were collected in a primary forest stream pool below a small cascade during the dry season between 20 June 2006 and 30 July 2006. The tadpoles were hidden on the ground between leaves and rock crevices. Further tadpoles were found in a water catchment basin next to the highway at km 51 and 53 in the U Bo region, Phong Nha ��� Ke Bang National Park, Vietnam, at 20 June, 7 July and 22 August 2006. These larvae were hidden between algae on the ground of the water body during day time (if algae were present, otherwise they were exposed on the sand or gravel of the ground), but were found swimming in the open water during night time. Once disturbed, the larvae tried to escape deep into the mud on the ground. During night time (08: 30 p. m.) of 7 July 2006, water temperature was 26 ��C. During day time, the water catchment basin was at least in part exposed to the sun. At day time of 22 August 2006 (12:00) the pH value was 5.5. Adult H. maosonensis (including calling males) were found during night time on forest stream rocks and cascades as well as around water catchment basins next to the highway at km 51 and 53 in the U Bo region between 20 June and 8 July 2006. Air temperature at night of 8 July 2006 next to a forest stream was 29 ��C. Once disturbed, the adult frogs tried to jump into the water where they remained motionless for some time., Published as part of Gawor, Anna, Hendrix, Ralf, Vences, Miguel, B��hme, Wolfgang & Ziegler, Thomas, 2009, Larval morphology in four species of Hylarana from Vietnam and Thailand with comments on the taxonomy of H. nigrovittata sensu latu (Anura: Ranidae), pp. 1-25 in Zootaxa 2051 on pages 8-11, DOI: 10.5281/zenodo.186587, {"references":["Bourret, R. (1942) Les batraciens de l'Indochine. Institut Oceanographique de l'Indochine, Hanoi, 547 pp.","Inger, R. F., Orlov, N. L. & Darevsky, I. (1999) Frogs of Vietnam: A report on new collections. Fieldiana Zoology, N. S., 92, 1 - 46.","Ziegler, T. (2002) Die Amphibien und Reptilien eines Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur & Tier Verlag, Munster, 342 pp.","Ziegler, T., Ohler, A., Vu, N. T., Le, K. Q., Nguyen, X. T., Dinh, H. T. & Bui, N. T. (2006) Review of the amphibian and reptile diversity of Phong Nha - Ke Bang National Park and adjacent areas, central Truong Son, Vietnam. In: Vences, M., Kohler, J., Ziegler, T. & Bohme, W. (Eds.), Herpetologia Bonnensis II. Proceedings of the 13 th Ordinary General Meeting of the Societas Europaea Herpetologica, Bonn, pp. 247 - 262."]}

Philautus quyeti sp. n. Holotype. ZFMK 82999, adult female from Nuoc Rung area, Dan Hoa Commune, Minh Hoa District, Quang Binh Province, central Vietnam (17 0 47 ’ N; 105 0 41 ’ E, altitude: 1,100 m a.s.l.), collected on 21 April 2004 by Le Khac Quyet (Fig. 1 a, b, c and Fig. 3 e). Paratype. VNUH 160706 (Field No. RH 06095), subadult from Cha Noi area, Phong Nha–Ke Bang National Park, Quang Binh Province, central Vietnam (surroundings of 17 0 38 ’ N; 106 0 05’ E, altitude: 430 m a.s.l), collected on 16 July 2006 by Ralf Hendrix (Fig. 3 a, b, c). Diagnosis. Small rhacophorid (SVL 34 mm), vomerine teeth absent. Head longer than wide. Snout rounded, longer than horizontal diameter of eye. Interorbital distance longer than the width of upper eyelid. Nostrils rounded, closer to tip of snout than to eye. Pupil oval, horizontal. Tympanum distinct, rounded, wider than disc of finger III. Supratympanic fold indistinct. Dermal fringe along outer side of forearm, tibia, tarsus and toe IV absent. Skin not lumpy or spinulous; dorsal surface of head, back and upper portion of flanks covered with small sharp tubercles. Dark brown pattern forming an inverse Y, notably a triangular spot between eyes bifurcating into two bands continuing posteriorly onto the back. Throat and margin of throat and chest yellow-white with brown marbling. Description of holotype. Size. Small rhacophorid (SVL 34 mm), body robust. Head. Head longer than wide (HL 12.5 mm; HW 11.4 mm; MN 11.2 mm; MFE 8.4 mm; MBE 4.5 mm), convex above. Snout rounded, slightly protruding, its length (SL 5.4 mm) longer than horizontal diameter of eye (EL 4.8 mm). Canthus rostralis rounded, loreal region oblique, slightly concave. Interorbital region convex, longer (IUE 4.1 mm) than upper eyelid (UEW 3 mm) and internarial distance (IN 3 mm); distance between front of eyes (IFE 6.9 mm) about 63 % distance between back of eyes (IBE 11 mm). Nostrils rounded without flap of skin laterally, closer to tip of snout (NS 2.1 mm) than to eye (EN 3.5 mm). Pupil oval, horizontal. Tympanum (TYD 2.5 mm) distinct, rounded, 52 % of eye diameter, and wider than disc of finger III (fd 3 1.9 mm); tympanum-eye distance (TYE 1 mm) 40 % of tympanum diameter. Pineal ocellus absent; spinules on upper eyelid absent. Vomerine teeth absent. Tongue moderately, cordate, shallowly emarginate. Tooth-like projections on lower jaw absent. Supratympanic fold indistinct, extending from behind of eye to above shoulder. Parotoid glands absent. Cephalic ridges absent. Co-ossified skin on head absent. Forelimbs (Fig. 2 a). Arm short and gracile; forearm (FLL 7.2 mm) shorter than hand (HAL 10.3 mm), dermal fringe along outer side of forearm absent. Fingers I and II short and thin; fingers III (TFL 8.1 mm) and IV long and thin. Relative length of fingers: I Hindlimbs (Fig. 2 b). Hindlimbs long, heels overlapping when limbs folded at right angles to body. Tibia (TL 20 mm) slightly longer than thigh (FL 18.5 mm) and longer than the distance from base of internal metatarsal tubercle to tip of toe IV (FOL 15.2 mm). Toes moderately long and thin, toe IV (FTL 12.5 mm) 50 % the distance from base of tarsus to tip of toe IV (TFOL 24.8 mm). Relative length of toes: I Skin. Snout and loreals smooth; dorsal skin not lumpy or spinulous, upper head, back and flank rough bearing small sharp tubercles. Dorsolateral folds absent. Dorsal surface of forelimb, thigh, tibia and tarsus smooth. Throat and chest smooth, belly and ventral part of thigh granular. Dermal appendage at vent absent. Coloration in alcohol. Snout and dorsum brownish gray with dark brown pattern forming an inverse Y, notably a triangular spot between eyes bifurcating into two bands continuing posteriorly; lower flank grey; snout, canthus and temporal region; tympanum light grey; upper lip with small white spots on brownish ground. Forelimb, dorsal surface of thigh, tibia and foot yellowish brown with some darker bands, posterior part of thigh below the vent yellowish brown with small white spots. Throat and margin of throat and chest yellow white with brown marblings; belly immaculate yellowish white to white; ventral part of forelimbs and thighs yellowish white; webbing grayish. Coloration in life. Background of dorsum (snout, head and back) brownish to moss-green, a dark, blotching pattern between eyes bifurcating into two bands continuing posteriorly on the back. Forelimb, dorsal parts of thighs, tibia, and foot moss-green with some dark brown bands. Throat and margin of throat and chest white with brown marblings; belly immaculate white; ventral part of forelimbs and thighs yellowish white; webbing grayish; discs of fingers and toes with white spots. Variation. The subadult has a moss-green dorsum; upper side of forelimbs, thighs, tibia, and foot the same as dorsal pattern; dark blotches on back indistinct; throat and margin of throat and chest bluish white with dark marblings; lower part of flank and belly bluish white with distinct granules; outer edge of discs of fingers and toes yellow. The paratype is the same as the holotype in presertative. Comparisons. The generic assignment of small Oriental treefrogs are uncertain and still under debate (Bossuyt & Dubois 2001; Wilkinson et al. 2002; Stuart & Heatwole 2004; Li et al. 2008). Hence, we compare Philautus quyeti with all small-sized rhacophorid treefrogs with a SVL under 45 mm and having reduced finger webbing from Vietnam and adjoining countries, regardless of what genus to which they are currently referred. These include Chiromantis doriae (Boulenger); C. hansenae (Cochran); C. laevis (Smith); C. nongkhorensis (Cochran); C. samkosensis Grismer, Thy, Chav & Holden; C. vittatus (Boulenger); Feihyla palpebralis (Smith); Gracixalus gracilipes (Bourret); G. j i n x i u e n s i s Hu (Fig. 3 d); G. supercornutus (Orlov, Ho & Nguyen); Kurixalus ananjevae (Matsui & Orlov); K. baliogaster (Inger, Orlov & Darevsky); K. bisacculus (Taylor); K. carinensis (Boulenger); K. eiffingeri (Boettger); K. idiootocus (Kuramoto & Wang); K. odontotarsus (Ye & Fei); K. verrucosus (Boulenger); Liuixalus romeri (Smith); Nyctixalus pictus (Peters); Philautus abditus Inger, Orlov & Darevsky; P. andersoni (Ahl); P. banaensis Bourret; P. cardamonus Ohler, Swan & Daltry; P. g r y l l u s Smith; P. hainanus Liu & Wu; P. longchuanensis Yang & Li; P. maosonensis Bourret; P. medogensis Ye & H u; P. menglaensis Kou; P. ocellatus Liu & Hu; P. parvulus (Boulenger); P. p e t e r s i (Boulenger); P. petilus Stuart & Heatwole; P. truongsonensis Orlov & Ho; P. vermiculatus (Boulenger); Rhacophorus appendiculatus (Günther); Theloderma asperum (Boulenger); T. rhododiscus (Liu & Hu), Theloderma ryabovi Orlov, Dutta, Ghate & Kent; and T. stellatum Taylor. The dark-blotched pattern on dorsum of Philautus quyeti sp. n. is similar to Gracixalus jinxiuensis, Kurixalus carinensis, and Philautus cardamonus. However, the new species differs from the latter by having supratympanic fold indistinct (versus distinct in G. jinxiuensis, K. carinensis, and P. cardamonus). Philautus quyeti sp. n. differs from K. carinensis and P. cardamonus by having the length of snout longer than the diameter of eye (snout equal to the diameter of eye in cardamonus and a little shorter than the diameter of eye in carinensis); and having skin on the top of the head and back bearing small sharp tubercles (shagreened in cardamonus and smooth or with small warts in carinensis). It further can be distinguished from cardamonus by having a larger size in adults (SVL 34 mm versus 19.3 mm in cardamonus), brown marbling on throat (absent in cardamonus), and lacking macrogland (present in cardamonus). It differs from G. jinxiuensis from Lai Chau Province by having the head longer than wide (head wider than long in jinxiuensis), tibiotarsal articulation reaching to the snout (behind in original description or to the eye in specimens of jinxiuensis from Lai Chau Province). Philautus quyeti sp. n. differs from Kurixalus ananjevae by having supratympanic fold indistinct (distinct in ananjevae), head longer than wide (head slightly shorter than wide in ananjevae), snout length longer than the diameter of eye (snout length as long as the diameter of eye in ananjevae); by having color pattern on throat and margin of throat and chest yellow white with brown marbling, belly yellowish white to white (ventrally grayish white, without markings in ananjevae). Philautus quyeti sp. n. differs from K. baliogaster by having head longer than wide (head wider than long in baliogaster), snout longer than the diameter of eye (snout equal to or shorter than the diameter of eye in baliogaster); skin on dorsal head and back rough with small sharp tubercles (smooth in baliogaster); lacking vomerine teeth, and venter without conspicious dark spots (present in baliogaster). Philautus quyeti sp. n. differs from K. bisacculus by having snout length longer than the diameter of eye (snout length equal to the diameter of eye in bisacculus); lacking vomerine teeth and dermal fringes or tubercles on outer edges of finger IV, forearm, tarsus, and toe V (present in bisacculus). Philautus quyeti sp. n. differs from Gracixalus gracilipes by having head width 33.5 % of SVL (head width 45 % of SVL in gracilipes), the diameter of tympanum longer than the width of finger III disc (width of finger III disc equal to the diameter of tympanum in gracilipes), and having dark-blotched pattern forming inverse Y (versus forming)(in gracilipes). Philautus quyeti sp. n. differs from Gracixalus supercornutus, Kurixalus odontotarsus, K. verrucosus, Philautus banaensis, P. g r y l l u s, and Rhacophorus appendiculatus by lacking dermal fringes or tubercles on the limbs (present in all latter species). Philautus quyeti differs from G. supercornutus and K. ordontotarsus by lacking sharped tubercles or spines on upper eyelids. It further differs from K. odontotarsus by lacking vomerine teeth. Philautus quyeti sp. n. differs from Chiromantis doriae and C. nongkhorensis by having the diameter of tympanum larger than the width of finger III disc (the width of finger III disc as large as the diameter of tympanum in doriae and nongkhorensis), the length of snout longer than the diameter of eye (the length of snout longer than the diameter in doriae and nongkhorensis). It differs from C. doriae by having dark bands on dorsal thighs (absent in doriae), skin of dorsum with small sharp tubercles (smooth in doriae), and by lacking dark stripes on dorsum (present in doriae). Philautus quyeti sp. n. also differs from nongkhorensis by having supratympanic fold indistinct (distinct in nongkhorensis). Philautus quyeti sp. n. differs from C. laevis, C. samkosensis and C. vittatus by having dark-blotched pattern forming inverse Y on dorsum (absent in laevis, samkosensis and vittatus). It is distinguished from C. hansenae, C. laevis, and C. vittatus in lacking light dorsolateral stripes (present in hansenae, laevis, and vittatus). Philautus quyeti sp. n. has small sharp tubercles on the skin of dorsum (smooth skin in C. laevis, C. samkosensis and C. vittatus). It further differs from C. samkosensis by having dark bands on dorsal aspect of thighs (absent in samkosensis) and not being green and not having geen blood and turquois bones. Philautus quyeti sp. n. differs from C. romeri by having larger size in adult (SVL 34 mm versus 25 mm in romeri) and supratympanic fold indistinct (distinct in romeri). Philautus quyeti sp. n. differs from Feihyla palpebralis by lacking a yellow streak from below the eye to shoulder (present in palpebralis). Philautus quyeti sp. n. differs from Kurixalus eiffingeri and K. idiootocus by having supratympanic fold indistinct (distinct in eiffingeri and idiootocus) and lacking vomerine teeth (present in eiffingeri and idiootocus). It further differs from K. idiootocus in less pointed snout. Philautus quyeti sp. n. differs from Nyctixalus pictus by having snout rounded (pointed in pictus), and having dark-blotched pattern on dorsum (small cream or orange spots on dorsum in pictus). Philautus quyeti sp. n. differs from Philautus abditus by having dorsal skin with small sharp tubercles (smooth in abditus), distinct tympanum (completely hidden in abditus), and lacking large black spots on anterior and posterior faces of thigh (present in abditus). Philautus quyeti sp. n. differs from P. longchuanensis and P. menglaensis by having the diameter of tympanum longer than the width of finger III disc (diameter of tympanum shorter than width of finger III disc in longchuanensis and menglaensis). Philautus quyeti sp. n. differs from P. maosonensis by having the head longer than wide (head wider than long in maosonensis) and lacking a dark spot behind the axilla (present in maosonensis). Philautus quyeti sp. n. differs from P. ocellatus by lacking a round black blotch on the occiput (present in ocellatus). Philautus quyeti differs from P. parvulus by having larger adult size (SVL 34 versus SVL less than 25 mm in parvulus), the snout length longer than the eye length (snout length shorter than the eye length in parvulus), distinct tympanum (hidden in parvulus), and having the nostril close to the tip of snout (nostril midway between eye and tip of snout in parvulus). Philautus quyeti sp. n. differs from P. petilus by lacking vomerine teeth (very small vomerine teeth in petilus), black stripe below edge of canthus extending from tip of snout to flanks and black spot in anterior region of groin (present in petilus), and by having skin of dorsum with small sharp tubercles (smooth in petilus). Philautus quyeti sp. n. differs from P. truongsonensis by having the head longer than wide (head as long as wide in truongsonensis) and lacking black spots on dorsum, flank and lateral part of thigh (present in truongsonensis). Philautus quyeti sp. n. differs from P. andersonii by having the diamter of tympanum 52 % the length of eye (1 / 4 the length of eye in andersonii). Philautus quyeti sp. n. differs from P. hainanus by having larger adult size (SVL 34 mm versus 17.6-18.2 mm in hainanus) and by lacking dark cross-bands on back and lower arm. Philautus quyeti sp. n. differs from P. p e t e r s i and P. vermiculatus by having snout length longer than the eye diameter (snout length equal to or shorter than the eye diameter in petersi and vermiculatus). It differs from P. vermiculatus and P. medogensis by having indistinct supratympanic fold (distinct in vermiculatus and medogensis). Philautus quyeti sp. n. differs from all species of Theloderma by lacking lumpy skin or spinules on dorsal surface. It differs from Theloderma rhododiscus by lacking reddish-orange finger and toe discs (present in rhododiscus). It differs from T. asperum, T. ryabovi and T. stellatum by lacking white blotch on above cloaca (present in asperum, ryabovi and stellatum). Ecological notes. The holotype was found at the beginning of the dry season at 20:00 on leaves (ca. 1.5 m above ground) of a small shrub near a small stream in montane forest in Dan Hoa Commune, Minh Hoa District, Quang Binh Province. The paratype was collected within the dry season at 22: 45 on leaves (ca. 2 m above ground) of a shrub in limestone forest in Cha Noi area, Phong Nha - Ke Bang National Park, Bo Trach District, Quang Binh Province. Due to the altitudes (430-1,100 m a.s.l.) in which the species was found so far, it seems that Philautus quyeti sp. n. is adapted to higher elevations. Distribution. Philautus quyeti sp. n. is only known from the type locality in Quang Binh Province (Fig. 4). Etymology. The name is dedicated to our friend, colleague, and zoologist, Mr. Le Khac Quyet from Fauna and Flora International - Vietnam Conservation Support Programme. Although a primatologist, Le Khac Quyet shows a high interest in the herpetofauna of Vietnam and was involved in the discovery of many different species within this country, such as Amphiesma andreae, Boiga bourreti, Calamaria thanhi, and Shinisaurus crocodilurus. As common names we suggest Quyet’s treefrog (English), Nhai cay quyet (Vietnamese), Rainette de Quyet (French), and Quyets Ruderfrosch (German).

Ziegler, Thomas, Quyet, Le Khac, Thanh, Vu Ngoc, Hendrix, Ralf, Böhme, Wolfgang (2008): A Comparative Study Of Crocodile Lizards (Shinisaurus Crocodilurus Ahl, 1930) From Vietnam And China. Raffles Bulletin of Zoology 56 (1): 181-187, DOI: http://doi.org/10.5281/zenodo.5339533, {"references":["Ahl, E., 1930. Contribution to the amphibian and reptile fauna of Kwangsi. 5. Lizards. Sitzungsberichte der Gesellschaft naturforschender Freunde vom 1. April 1930, Berlin: 329-331. [in German]","Bobrov., V. V., 1997. On the boundary between the Palearctic and Indomalayan faunistic kingdoms in the mainland part of Asia, with special reference to the distribution of lizards (Reptilia, Sauria). Biological Bulletin, 24: 476-487.","Bohme, W., 1988. On the genital morphology of the Sauria: functional and phylogenetic implications. Bonner Zoologische Monographien, 27: 1-176. [In German]","Carranza, S., Arnold, E. N. & J. M. Pleguezuelos, 2006. Phylogeny, biogeography, and evolution of two Mediterranean snakes, Malpolon monspessulanus and Hemorrhois hippocrepis (Squamata, Colubridae), using mtDNA sequences. Molecular Phylogenetics and Evolution, 40: 532-546.","Darevsky, I. S., Orlov, N. L. & T. C. Ho, 2004. Two new lygosomine skinks oft the genus Sphenomorphus Fitzinger, 1843 (Sauria, Scincidae) from northern Vietnam. Russian Journal of Herpetology, 11: 111-120.","David, P., Tong, H., Vogel, G. & M. Tian, in press. On the status of the Chinese pitviper Ceratrimeresurus shenlii Liu & Liang in Liang, 2003 (Serpentes, Viperidae), with the addition of Protobothrops cornutus (Smith, 1930) to the Chinese snake fauna. Asiatic Herpetological Research.","THE RAFFLES BULLETIN OF ZOOLOGY 2008","Fan, T. H., 1931. Preliminary report of reptiles from Yaoshin, Kwangsi, China. Bulletin of the Department of Biology, College of Science, Sun Yatsen University, 11: 1-154.","Good, D. A., Bauer, A. M. & R. Gunther, 1993. An annotated type catalogue of the anguimorph lizards (Squamata: Anguidae, Helodermatidae, Varanidae, Xenosauridae) in the Zoological Museum, Berlin. Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe, 69: 45-56.","Herrmann, H.-W., Ziegler, T., Malhotra, A., Thorpe, R. S. & C. Parkinson, 2004. Redescription and systematics of Trimeresurus cornutus (Serpentes: Viperidae) based on morphology and molecular data. Herpetologica 60: 211-221.","Kumazawa, Y., 2004. Mitochondrial DNA sequences of five squamates: Phylogenetic affiliation of snakes. DNA Research, 11: 137-144.","Le, Q. K. & T. Ziegler, 2003. First record of the Chinese crocodile lizard from outside of China: Report on a population of Shinisaurus crocodilurus AHL, 1930 from North-eastern Vietnam. Hamadryad, 27: 193-199.","Li, Z.-C. & Z. Xiao, 2002. Discovery of Shinisaurus crocodilurus in Guangdong Province. Chinese Journal of Zoology, 37: 76- 77. [In Chinese]","Liu, X., Zhou, F. & G. Pan, 1989. New discovery of localities of Shinisaurus crocodilurus Ahl. Sichuan Journal of Zoology, Chengdu 8: 32-33. [In Chinese]","Macey, J. R., Schulte, J. A. II, Larson, A., Tuniyev, B. S., Orlov, N. & T. Papenfuss, 1999. Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution, 12: 250-272.","Magdefrau, H., 1987. On the situation of the Chinese crocodile lizard Shinisaurus crocodilurus Ahl, 1930. Herpetofauna, Weinstadt 9: 6-11. [In German]","Magdefrau, H., 1997. Biology, keeping and breeding of the crocodile-tailed knobby lizard (Shinisaurus crocodilurus). Zeitschrift des Kolner Zoo 40: 55-60. [In German]","Magdefrau, H. & B. Schildger, 1993. News from Shinisaurus crocodilurus Ahl, 1930. Herpetofauna, Weinstadt 15: 11-16. [In German]","Nguyen, S. V., Nguyen, T. Q. & S. T. Nguyen, 2000. The preliminary results of the survey on herpetofauna in Yen Tu mountain area. Tap Chi Sinh Hoc 22(15): 11-14. [in Vietnamese]","Ohler, A., 2005. Transborder systematics: a critical evaluation of zoogeography and conservation biology at the crossroads between Vietnam, Laos and China. Programme and Abstracts, 13th Ordinary General Meeting, Societas Europaea Herpetologica (SEH), 27 September-2 October, Bonn, Germany: 120-121.","Palumbi, S., Martin, A., Romano, S., McMillan, W. O., Stice, L. & G. Grabowski, 1991. The Simple Fool's Guide to PCR. Version 2, Honolulu, Hawai.","Pesantes, O. S., 1994. A method for preparing the hemipenis of preserved snakes. Journal of Herpetology, 28: 93-95.","Poulakakis, N., Lymberakis, P., Valakos, E., Pafilis, P., Zouros, E. & M. Mylonas, 2005. Phylogeography of Balkan wall lizard (Podarcis taurica) and its relatives inferred from mitochondrial DNA sequences. Molecular Ecology, 14: 2433-2443.","Shen, L. & H. Li., 1982. Notes on the distribution and habits of the lizard Shinisaurus crocodilurus Ahl. Acta Herpetol. Sinica, Chengdu, n. s. 1: 84-85. [In Chinese; translated by D.-T. Yang, 1987, Bulletin of the Chicago Herpetological Society, 22: 4-5]","Vences, M., Wanke, S., Vieites, D. R., Branch, W. R., Glaw, F. & A. Meyer, 2004. Natural colonization or introduction? Phylogeographical relationships and morphological differentiation of house geckos (Hemidactylus) from Madagascar. Biological Journal of the Linnean Society, 83: 115- 130.","Zhang, F., 1986. Studies on morphological characters of hemipenes of the Chinese lizards. Acta Herpetologica Sinica, Chengdu, n. s. 5: 254-259. [In Chinese]","Zhao, E. & K. Adler, 1993. Herpetology of China. Contributions to Herpetology, Number 10. Society for the Study of Amphibians and Reptiles, Oxford, Ohio. 522 pp.","Zhao, E., Zhao, K. & K. Zhou, 1999. Fauna Sinica. Reptilia Vol. 2. Squamata. Lacertilia. Science Press, Beijing. 394 pp.","Zhu, H., Wang, H., Li, B. & P. Sirirugsa, 2003. Biogeography and floristic affinities of the limestone flora in southern Yunnan, China. Annals of the Missouri Botanical Garden, 90: 444-465.","Ziegler, T. & W. Bohme, 1997. Genital structures and mating biology in squamate reptiles, especially the Platynota, with comments on systematics. Mertensiella, 8: 1-207. [In German]","Ziegler, T., Gaulke, M. & W. Bohme, 2005. Genital morphology and systematics of Varanus mabitang Gaulke & Curio, 2001 (Squamata, Varanidae). Current Herpetology, 24: 13-17.","Ziegler, T. & H.-W. Herrmann, 2002. Rediscovery of two horned pit viper species in Vietnam. Mitteilungen Zoologische Gesellschaft fur Arten- und Populationsschutz e.V. 18: 24-26. [In German]"]}

Lycodon sp. Another female Lycodon was collected by us about one month after the discovery of the afore mentioned Lycodon cf. ruhstrati. The discovery took place in the same region at an altitude of about 480 m above sea level. This second Lycodon (Figure 13) was found at night of 30 July 2006, at the end of the dry season, crawling on pebbles near a steep primary forest stream cascade in the U Bo region. The specimen (ZFMK 86453, SVL 505, TaL 147, TL 652 mm) differs from the syntopic Lycodon cf. ruhstrati among others in having smooth dorsal scale rows, bearing enlarged vertebral scales, and having somewhat lower ventral and subcaudal counts (although within the range of L. ruhstrati, see above), as well as a differing banded pattern. The remaining scalation features of the specimen are as follows: eight supralabials, of which third to fifth are in contact with eye; 10 infralabials; one loreal, not in contact with eye; one preocular, not in contact with frontal; prefrontal not entering eye; two postoculars; two anterior temporals, followed by three posterior temporals; 17 dorsal scale rows at midbody; one undivided preventral and 208 ventrals; ventrals distinctly angulate laterally; anal entire; 85 divided subcaudals. Concerning coloration and pattern, the specimen has 20 dark transversal crossbands on its body dorsum, which continue on the venter (except for the neck band). The 19 creamy interspaces of the dark body crossbands are marbled by greyish-brown. Tail dorsum with 10 such light crossbands and 9 dark crossbars (including tail tip), underside of tail dark. On the right maxillary bone seven anterior fang-like teeth increasing in size were discernible, separated by a large toothless interspace from four medium-sized teeth, which were followed by three posterior teeth, the first two ones of which being distinctly enlarged. This Lycodon specimen morphologically resembles the female L. ruhstrati specimen from Ha Tinh province as shown in Ziegler (2002). However, the latter specimen has no enlarged vertebrals and bears minor differences in the banded pattern. The Ha Tinh specimen further shows a distinct light area on the rear head, however, that is still slightly discernible in the larger Lycodon specimen from Phong Nha - Ke Bang as obscured area. As was pointed out above, only future research including currently undertaken molecular methods will prove, whether the mainland " L. ruhstrati " with smooth dorsal scales in fact represent a separate taxon (cf. L. futsingensis) and whether mainland L. cf. ruhstrati (with keeled dorsal scales) represent an unnamed taxon indeed, different from the populations from the terra typica in southern Taiwan.

Pareas macularius Blyth, 1868: Spotted slug snake / Ran ho may dom A single adult female specimen (ZFMK 86446, SVL 371, TaL 74, TL 445 mm) was discovered, that had folded oviducts and oviduct eggs with a maximum diameter of 10.1 mm. The specimen was found during the dry season, at night of 21 June 2006, between the leaves of a small tree in about 1.5 m height in the Cha Noi region at an altitude of about 350���400 m above sea level. Scalation features are as follows: seven supralabials, separated from eye by a longish subocular; seven (right) and eight (left) infralabials; three pairs of chin shields; one large and elongated loreal, on the right side in contact with eye; one preocular; one subocular; two postoculars; prefrontals in contact with the eye; two anterior temporals, followed by three posterior temporals; 15 (keeled) dorsal scale rows at midbody; 158 ventrals; anal plate entire; 45 divided subcaudals. The specimen keys out under Amblycephalus tamdaoensis in Bourret (1936). The coloration and pattern of our specimen (see Figure 15) correspond to the description given in Bourret (1936), however, the only difference we could observe is the presence of only a single postocular (see Bourret 1936). Orlov et al. (2000) treated tamdaoensis as subspecies of Pareas macularius and Nguyen et al. (2005) list Amblycephalus tamdaoensis Bourret as synonym of Pareas macularius. As distribution of the latter, Nguyen et al. (2005) list some northern Vietnamese provinces southwards to Nghe An Province, from where another specimen (ZFMK 82925) of the species was available to us. Thus, our recent finding represents not only the first record for the Quang Binh Province and for Phong Nha - Ke Bang National Park in particular, but also the southernmost record known from Vietnam., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on page 15, DOI: 10.5281/zenodo.176960, {"references":["Bourret, R. (1936) Les serpents de l'Indochine. II. Catalogue systematique descriptif. Henri Basuyau et Cie, Toulouse, 505 pp.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp."]}

Chrysopelea ornata (Shaw, 1802): Golden tree snake / Ran cuom According to Nguyen et al. (2005) the species is distributed all over the country. We photographed this snake (Figure 2) in September 2006 during daytime in secondary shrub vegetation in the National Park's border area. Characteristic were the greenish-yellow head with black markings, the ivory chin and supralabials, the greenish yellow body with some black-edged scales and some entirely black scales that form crossbars, and the yellowish-green ventrals with black spots next to each lateral notch (Cox et al. 1998). Remarkable are the orange dorsal blotches that we already observed in a specimen from Khanh Hoa Province in southern Vietnam, because Manthey & Grossmann (1997) stated such a pattern occurring only in the nominate form from Sri Lanka, India and Bangladesh. However, Pope (1935) mentioned that the coloration in the species is extremely variable (see also Bourret 1936)., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on page 4, DOI: 10.5281/zenodo.176960, {"references":["Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp.","Cox, M. J., van Dijk, P. P., Nabhitabhata, J. & Thirakhupt, K. (1998) A photographic guide to snakes and other reptiles of Peninsular Malaysia, Singapore and Thailand. New Holland Publ. Ltd., London, 144 pp.","Manthey, U. & Grossmann, W. (1997) Amphibien und Reptilien Sudostasiens. Natur und Tier - Verlag, Munster, 512 pp.","Pope, C. H. (1935) The reptiles of China. Turtles, crocodilians, snakes, lizards. Natural History of central Asia. Vol. X. American Museum Natural History, New York, 604 p.","Bourret, R. (1936) Les serpents de l'Indochine. II. Catalogue systematique descriptif. Henri Basuyau et Cie, Toulouse, 505 pp."]}

Pareas hamptoni (Boulenger, 1905): Hampton's slug snake / Ran ho may hamton We found a female specimen (VNUH 30.7.'06- 1, SVL 352, TaL 109, TL 461 mm) with folded oviducts and oviduct eggs with a maximum diameter of 2.7 mm. The snake was seen at night (about 22:00h) of 30 July 2006, during the dry season, in the U Bo region at an altitude of about 525 m above sea level. The specimen climbed in ca. 1.5 m height in a shrub, about 3 m distant from a small forest stream. Scalation features are as follows: seven supralabials, separated from eye by lower preocular (or subocular) and the single postocular; seven (right) and nine (left) infralabials; one loreal; two preoculars (or one preocular and one subocular); one postocular, that is fused on the left side with the lower preocular; prefrontals touch the eye; two anterior temporals, followed by two posterior temporals; 15 dorsal scale rows at midbody (generally smooth, in some scales with indistinct hints of keels); 185 ventrals; anal entire; 82 divided subcaudals. For the characteristic coloration see Figure 14. In Vietnam, the species was previously known from several provinces in the north as well as from southern regions (Ziegler 2002, Nguyen et al. 2005). Our recent finding represents the first record for the Quang Binh Province and for Phong Nha - Ke Bang National Park in particular., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on page 14, DOI: 10.5281/zenodo.176960, {"references":["Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp."]}

Amphiesma sp. On 30 July 2006, at the end of the dry season, we found an adult male specimen (ZFMK 86457, SVL 465, TaL 215, TL 680 mm) in the U Bo region at an altitude of about 510 m above sea level. The snake was captured at night (20:00h– 21:00h) in the leaf litter of a forest stream, only the snout tip was protruding from the water surface. Scalation features are as follows: internasals narrowed anteriorly; dorsolateral nostrils; nine supralabials (4–6 in contact with eye); ten infralabials; one elongated loreal; two preoculars, the uppermost being the larger; two postoculars, the uppermost being the larger; one anterior temporal, followed by one posterior temporal; 19 dorsal scale rows at midbody (keeled, except four outermost rows that bear only keel rudiments, if at all); three preventrals and 171 ventrals; anal divided; 146 divided subcaudals; dorsal tail scales very strongly keeled; TaL / TL ratio 0.32; the coloration and pattern is discernible from Figures 4–5. This Amphiesma obviously represents a species not yet recorded from Vietnam and detailed comparisons as well as a comprehensive discussion concerning the specific assignment will be provided elsewhere (David et al. submitted).

Cryptelytrops albolabris (Gray, 1842): White lipped bamboo snake / Ran luc mep trang During the dry season, on 28 July 2006, we obtained an adult male specimen from local people (ZFMK 86454, SVL 467, TaL 128, TL 595 mm). The species occurs all over the country (Nguyen et al. 2005, Vogel 2006) and the recent finding (Figure 3) was identifiable as C. albolabris among others by its upper lip being sharply demarcated from the upper part of the head, the distinct white stripe down the side of the body, the first supralabial fused to nasal, the internasals being in contact, 21 scales around midbody, and the gular scales arranged in regular pairs (Malhotra & Thorpe 1997, Ziegler 2002). Further characteristic scalation features are as follows: 12 supralabials; 13���14 infralabials; one elongated subocular; two small postoculars; 12 scales in a line between narrow supraoculars; dorsal scale rows keeled; four preventrals and 155 ventrals; anal entire; 75 divided subcaudals., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on page 5, DOI: 10.5281/zenodo.176960, {"references":["Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp.","Malhotra, A. & Thorpe, R. S. (1997) New perspectives on the evolution of south-east Asian pitvipers (genus Trimeresurus) from molecular studies. In: Thorpe, R. S., Wuster, W. & Malhotra, A. (Ed), Venomous snakes. Ecology, evolution and snakebite. Symp. zool. Soc. Lond., Clarendon Press, Oxford, 70, 115 - 128."]}

Liopeltis frenatus (G��nther, 1858): G��nther���s stripe-necked snake / Ran dai ma We found an adult male specimen (VNUH 9.7.'06- 1, SVL 270, TaL 116, TL 386 mm) as roadkill during the dry season at an altitude of ca. 600 m above sea level. The specimen was discovered at noon of 9 July 2006 in the U Bo region. Scalation features are as follows: seven supralabials (only visible at left side), of which third and fourth supralabials are in contact with eye; ca. seven infralabials; one loreal; one preocular; two postoculars (only visible at left side); one anterior temporal, followed by two posterior temporals (only visible at left side); 15 smooth dorsal scale rows at midbody; four preventrals and 126 ventrals; anal divided; 69 divided subcaudals. The ventral number of our specimen (4 + 126) falls distinctly below the minimum value of 140 given in Whitaker & Captain (2004); however, it must be noted that the ventral scales of the in part heavily destroyed roadkill specimen were relatively difficult to count. Despite, the characteristic large eyes of the slender-bodied and long-tailed snake were still well discernible, as well as the two broad black stripes stretching from behind the eyes. The dorsum was olive-brown with in part black-edged scales (see Figure 11). In Vietnam, the species was previously only reported from Lao Cai Province in the North and from the provinces Quang Nam / Da Nang and Gia Lai / Kon Tum in central and southern Vietnam (Nguyen & Ho 1996, Szyndlar & Nguyen 1996, Orlov et al. 2003, Nguyen et al. 2005). In addition, we could record the first L. frenatus from Thua Thien Hue Province, which borders Quang Nam and Da Nang Provinces in the South (Ziegler & Hoang, submitted). Thus, we herein can add with our most recent finding the first record of that species for Quang Binh Province and for the Phong Nha - Ke Bang National Park in particular., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on page 10, DOI: 10.5281/zenodo.176960, {"references":["Whitaker, R. & Captain, A. (2004) Snakes of India, the field guide. Draco books, Chennai, 481 pp.","Nguyen, V. S. & Ho, T. C. (1996) Danh luc bo sat va ech nhai Viet Nam (Amphibians and reptiles of Vietnam). Nha xuat ban khoa hoc va ky thuat, Hanoi, 264 pp.","Szyndlar, Z. & Nguyen, V. S. (1996) Terrestrial snake fauna of Vietnam: distributional records. The Snake, 27, 91 - 98.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp."]}

Lycodon cf. ruhstrati (Fischer, 1886): Ruhstrat's wolf snake / Ran khuyet rutrat A female specimen (ZFMK 86451, SVL 466, TaL 142, TL 608 mm) was discovered at night of 20 June 2006, during the dry season, in the primary forest of U Bo region (Figure 12). The specimen was crawling on the forest ground near a steep cascade at an altitude of about 520 m above sea level. Scalation features are as follows: Eight supralabials, of which third to fifth supralabials are in contact with eye; ten infralabials; one loreal, not in contact with eye; one preocular, not in contact with frontal; prefrontal not entering eye; two postoculars; two anterior temporals, followed by three posterior temporals; 17 dorsal scale rows at midbody, feebly keeled except for outermost rows; vertebral scales not enlarged; two undivided preventrals and 229 ventrals; ventrals distinctly angulate laterally; anal entire; 104 divided subcaudals. The specimen had 19 dark transversal crossbands on its body dorsum. These dark crossbands are largest anteriorly and become shorter towards the cloaca. The 18 interspaces of the dark body crossbands are creamy white in the neck region, but then become distinctly darker. After the first body third the light interspaces consist of a greyish-brown bar that is edged by white and contains in its centre a darker transversal bar. Such a pattern continues on the tail dorsum, which bears ten dark and 11���12 light crossbars, including the tail tip. The body venter is completely yellowish to cream in its anterior fourth, then in part interspersed with a brownish-grey marbling that becomes more intensive in the last body half. The dark dorsal crossbands continue on the underside of the tail. The specimen clearly is a member of the genus Lycodon Boie, due to its pholidosis, a vertically elliptical pupil, and because of its arched maxillary bone, that is bent inwards anteriorly, with 6 anterior fang-like teeth increasing in size and separated by a toothless interspace of about 0.9 mm width at left side from 6 or 7 posterior teeth, the last two ones of which being enlarged (Lanza 1999). In the key provided by Lanza (1999), the specimen from Phong Nha - Ke Bang, with some of its dorsal scale rows keeled along the whole body, clearly keys out as L. ruhstrati at point nine. From the similar L. fasciatus, which is a common snake around the limestone outcrops and caves in the karst forest of Phong Nha - Ke Bang, L. ruhstrati differs e. g. by the loreal not touching the eye, having slightly higher ventral (193���237 vs. 186���220) and subcaudal counts (77���119 vs. 66��� 90), shorter posterior chinshields than the anterior ones (which is vice versa in the species fasciatus), as well as in lacking dark bands that completely encircle the body, as it is the case in L. fasciatus (Bourret 1936, Lanza 1999). Lanza (1999) further distinguishes in his key between the subspecies L. ruhstrati multifasciatus from Japan and the nominate form from southern China and Taiwan, with which our specimen appears to be conspecific at first glance. However, Ota (1988) additionally found distinct meristic differences between the Taiwanese and continental populations of L. r. ruhstrati. These results might support the validity of Dinodon futsingensis Pope, that was described from "Futsing Hsien" in Fujian Province, continental China, and subsequently synonymized by Pope (1935) with L. ruhstrati (Fischer), that was originally described as Ophites ruhstrati from southern Taiwan (Fischer 1886). However, it is noteworthy, that Bourret (1936) states for " Dinodon futsingense ", that the dorsal scales are smooth and not keeled as it is the case in our recent finding from Phong Nha - Ke Bang. Pope's (1928) type specimen of Dinodon futsingensis had smooth scales and also another continental specimen that was studied by us (ZFMK 23363 from Fujian) as well as a specimen from the Vietnamese province Ha Tinh, listed as Lycodon ruhstrati in Ziegler (2002). However, Pope (1929) reported also on specimens from Fujian and Mell (1922) on a specimen from Guangdong, that bear dorsal keels. Taiwanese specimens known so far always had keeled dorsal scales (Fischer 1886: type series O. ruhstrati, Maki 1931, Vogel & Brachtel accepted). Thus, future studies must clarify, whether the coexistence of continental representatives both with keeled and smooth dorsal scales is due to individual variation or rather indicating that different taxa are involved. It is further noteworthy, that female L. ruhstrati from continental China that were studied by Ota (1988) had lower ventral counts (197���227), lower subcaudal counts (77���95), and relatively higher transverse body and tail bands (22���38, 12 ��� 18) compared to our recent finding from Phong Nha - Ke Bang. Nguyen et al. (2005) list both Dinodon futsingensis and Lycodon ruhstrati as occurring in Vietnam: D. futsingensis from Lao Cai Province in the north southwards to Hoa Binh Province, and Lycodon ruhstrati from Cao Bang Province in the north to Da Nang Province in central Vietnam., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on pages 11-13, DOI: 10.5281/zenodo.176960, {"references":["Fischer, J. G. (1886) Herpetologische Notizen. Abh. Naturw. Ver. Hamburg, 9, 1 - 19.","Lanza, B. (1999) A new species of Lycodon from the Philippines, with a key to the genus (Reptilia Serpentes Colubridae). Tropic. Zool., 12, 89 - 104.","Bourret, R. (1936) Les serpents de l'Indochine. II. Catalogue systematique descriptif. Henri Basuyau et Cie, Toulouse, 505 pp.","Ota, H. (1988) Taxonomic notes on Lycodon ruhstrati (Colubridae: Ophidia) from East Asia. J. Taiwan Mus., 41, 85 - 91.","Pope, C. H. (1935) The reptiles of China. Turtles, crocodilians, snakes, lizards. Natural History of central Asia. Vol. X. American Museum Natural History, New York, 604 p.","Pope, C. H. (1929) Notes on reptiles from Fukien and other Chinese provinces. Bull. Amer. Mus. Nat. Hist., 58, 335 - 487, Pl. 17 - 20.","Mell, R. (1922) Beitrage zur Fauna sinica. I Die Vertebraten Suchinas; Feldlisten und Feldnoten der Sauger, Vogel, Reptilien, Batrachier. Arch. Naturgesch. Berlin, (A), 80, 1 - 143.","Maki, M. (1931) A monograph of the snakes of Japan. Dai-Ichi Shobo, Tokyo, 240 pp.","Nguyen, V. S., Ho, T. C. & Nguyen, Q. T. (2005) A checklist of amphibians and reptiles of Vietnam. Nha xuat ban nong nghiep, Hanoi, 180 pp."]}

Calamaria thanhi Ziegler & Le, 2005: Thanh's reed snake / Ran mai gam Thanh Ziegler & Le (2005) described C. thanhi based on a single female specimen from adjacent to Phong Nha - Ke Bang National Park, Dan Hoa commune, Minh Hoa District, Quang Binh Province. Meanwhile a second specimen came to our attention, that was found during April 2005 close to the National Park's border during excavations in a secondary shrub area. Because the species was formerly only known from the heavily destroyed holotype, we herein provide first pictures of a better preserved and at the same time of the first male as well as also largest specimen of the barely known species (Figures 6���9). The new specimen (SVL 415, TaL 46, TL 461 mm), that is deposited in the Science Research Centre of the Phong Nha - Ke Bang National Park (SRC PNKB III '06- 1), largely agrees with the holotype description in pattern and scalation (see Table 2), except for: 1) the rostral portion visible from above being somewhat shorter (1.5 mm) than the prefrontal suture (1.6 mm); 2) the left paraparietal being surrounded by seven instead of six shields and scales, as it is the case in the holotype; 3) the third supralabial being about half the size (instead ��) of second; 4) the new specimen having 184 instead of 198 ventral scales and 28 instead of 21 divided subcaudals; 5) the number of dorsal scale rows is reduced to eight in the male specimen instead of six rows on tail opposite 8 th subcaudal posterior to cloaca; 6) the ratio of tail to total length is 0.099 instead of 0.068 in the female holotype; 7) the male specimen bears five instead of four distinct beige to yellowish dorsal bands in the anterior two body thirds, followed by a sixth, but more vestigial light transversal band, that is only discernible from dorsal view; 8) the light dorsal bands subsequent to the first band follow each with distances of 3���5 cm in the male specimen (distances are extending posteriorly); 9) both yellowish markings on the dorsal tail base and on its end are better developed in the male specimen than it is the case in the female holotype, although both transversal bands do not completely encircle the dorsum; 10) the yellowish tail base bears only few tiny dark flecks in the male; 11) the underside of the head is not brownish-grey as it is the case in the female holotype but resembles the overall yellow venter coloration. We interpret the afore mentioned differences due to individual variation / sexual dimorphism. In addition, we removed the inverted left hemipenis of the male specimen and dissected the thin organ in the inverted condition. Pedicel and truncus of the forked organ were smooth and without detectable spines, and both lobes were covered with a calyces ornamentation., Published as part of Ziegler, Thomas, Hendrix, Ralf, Thanh, Vu Ngoc, Vogt, Martina, Forster, Bernhard & Kien, Dang Ngoc, 2007, The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key, pp. 1-40 in Zootaxa 1493 on pages 7-8, DOI: 10.5281/zenodo.176960, {"references":["Ziegler, T. & Le, K. Q. (2005) A new species of reed snake, Calamaria (Squamata: Colubridae), from the central Truong Son (Annamite mountain range), Vietnam. Zootaxa, 1042, 27 - 38."]}