Rhacophorus vampyrus, Rowley, Jodi J. L., Duong, Le Thi Thuy, Dao, Tran Thi Anh, Stuart, Bryan L. & Huy, Hoang Duc, 2010

Rowley, Jodi J. L., Duong, Le Thi Thuy, Dao, Tran Thi Anh, Stuart, Bryan L. & Huy, Hoang Duc, 2010, A new tree frog of the genus Rhacophorus (Anura: Rhacophoridae) from southern Vietnam, Zootaxa 2727, pp. 45-55 : 46-53

Rhacophorus vampyrus sp. nov.

Holotype. AMS R 173127, adult male, on a tree branch in montane evergreen forest in Bidoup-Nui Ba National Park, Lac Duong District, Lam Dong Province, Vietnam (12.1865 N, 108.7151 E, 1625 m). Collected at 21:00 h on 26 July 2010 by Le T. T. D.

Paratypes. AMS R 173126, female, on a tree branch approximately 2 m above ground in montane evergreen forest (12.1865 N, 108.7150 E, 1625 m), collected at 22:30 h on 26 July 2010 by Le T. T. D. UNS 00103/ AMS R 173128 male, on a leaf approximately 2 m above ground in montane evergreen forest (12.1736 N, 108.6987 E, 1470 m) collected at 22:05 h on 31 March 2009 by Le T. T. D. UNS 00104/ AMS R 173129, male, on a leaf approximately 1.5 m above ground, 0.5 m from rocky stream in montane evergreen forest (12.1865 N, 108.7151 E, 1625 m) collected at 20:30 h on 30 March 2009 by Le T. T. D. AMS R 173507, female, on a leaf approximately 1 m above ground in montane evergreen forest (12.1736 N, 108.6987 E, 1470 m), collected at 21:50 h on 18 August 2009 by Le T. T. D. NCSM 77318, adult male, on a trunk of tree away from water, 1 m above ground in montane evergreen forest (12.1864 N, 108.7149 E, 1627 m), collected at 21:00 h on 6 March 2008 by B. L. Stuart, J. J. L. Rowley, Le T. T. D., Tran T. A. D., and Hoang D. H. UNS 0 0 105, adult female, in montane evergreen forest (12.1736 N, 108.6987 E, 1470 m), collected on 18 March 2010 by Tran T. A. D. ZFMK 91076, adult male, on a tree 1 m above ground near 1 2 m wide, dry stream in montane evergreen forest (12.0096 N, 108.6607 E, 2004 m), collected on 23 March 2010 by Tran T. A. D. All specimens were collected in Bidoup-Nui Ba National Park, Lac Duong District, Lam Dong Province, Vietnam.

Other material. AMS R 173132, six tadpoles, inside small pool of water in an approximately 10 cm diameter tree-hole, approximately 1 m above the ground in montane evergreen forest in Bidoup-Nui Ba National Park, Lac Duong District, Lam Dong Province, Vietnam (12.1736 N, 108.6987 E, 1470 m). Collected at 10:30 h on 28 July 2010 by Le T. T. D.

Etymology. Specific epithet in reference to the unusual tadpole of the new species (see below), applied as a noun in apposition.

Suggested common name. Vampire tree frog (English). Ế ch cây ma cà rông (Vietnamese).

Diagnosis. The new species is assigned to the genus Rhacophorus by the presence of intercalary cartilage between the terminal and penultimate phalanges of digits, Y-shaped distal end of terminal phalanx, tips of digits expanded into large disks bearing circummarginal grooves, webbed fingers, a supracloacal dermal ridge, vomerine teeth, and horizontal pupil ( Brown & Alcala 1994; Duellman & Trueb 1986; Liem 1970). Rhacophorus vampyrus is distinguished from all other Rhacophorus in mainland Southeast Asia by a combination of (1) pale tan to brick red dorsum, (2) white throat, chest and belly, (3) mostly black flanks and anterior and posterior surface of thighs, (4) grey to black webbing between fingers and toes, (5) reduced finger webbing, and (6) pointed projection at tibiotarsal articulation.

Description of holotype. Body dorsoventrally compressed; head length 90% of head width; snout truncate in dorsal view, bluntly truncate in profile, projecting slightly beyond margin of the lower jaw, with slight point on tip of snout visible in ventral view; canthus rostralis distinct, bluntly angular; loreal region sloping, slightly concave; lips slightly flared; interorbital region slightly convex; nostrils oval, slightly protuberant, without flap of skin laterally, much closer to tip of snout than eye; pupil horizontal, tympanum barely visible externally, tympanic rim slightly elevated relative to skin of temporal region, 41% of eye diameter; pineal ocellus absent; skin not co-ossified to forehead; vomerine teeth present in oblique groups, separated by a distance about as long as each group, closer to choanae than wide; choanae oval, at margins of roof of mouth; tongue attached anteriorly, deeply notched posteriorly; tooth-like projections on lower jaw absent; pair of distinct, oval vocal sac openings at base of jaw; external paired subgular vocal sacs; weak supratympanic fold extending to just beyond level of axilla. Forelimbs relatively robust, relative length of fingers I <II <IV <III; tips of all fingers with well-developed disks with distinct circummarginal grooves, disks relatively wide compared to finger width (third finger disk 170% third finger width), third finger disk width greater (163%) than tympanum diameter; webbing formula I 2 – 2 + II 1 + – 2+ III 2 - – 1 IV; subarticular tubercles prominent, rounded, formula 1, 1, 2, 2; palmar tubercle absent; accessory palmar tubercles indistinct; thenar tubercle absent; prepollex enlarged, with low, relatively indistinct tubercle; nuptial pads or nuptial excrescences absent. Relative length of toes I <II <III <V <IV; tips of toes with well-developed disks with distinct circummarginal grooves; disks smaller than those of fingers; webbing formula I 1 – 2 II 1 – 2 III 1 – 2 + IV 2 – 1 V; subarticular tubercles rounded, distal subarticular tubercles distinct, inner less distinct, formula 1, 1, 2, 3, 2; inner metatarsal tubercle low, oval; outer metatarsal tubercle and supernumary tubercles absent. Dorsal skin smooth, ventral surface of thighs and belly coarsely granular, chest and throat smooth. Loose skin on either side of the throat over vocal sacs. Tarsal fold absent. Outer margin of forearm and foot with low dermal ridges; low supracloacal dermal ridge; pointed projection at tibiotarsal articulation present, approximately 1 mm long.

Colour of holotype in life. Dorsal surface pale copper-brown with faint darker brown mottling along back; dorsolateral surfaces with very small, sparse, white and darker brown flecks; dorsal surface of lower arms, thigh and tibiotarsus copper-brown with diffuse darker brown barring; dorsal surface of hands and feet copper-brown proximally, fading distally to pinkish-cream on fingers and toes I II, and grey on fingers III IV and toes III V; dark grey to black webbing dorsally. Black flanks, upper arms, ventral surface of lower arms, anterior and posterior surface of thighs, and ventral surface of crus, with small, irregular white spots within the black on the flanks and upper arms. Ventral surface of throat, chest and belly immaculate except for black mottling extending slightly onto lateral margins of chest at axilla; ventral surfaces of toes and fingers pale grey; ventral surface of webbing grey with dark grey/black margins. Iris pale yellowish gold with a network of fine dark gold reticulations concentrated around the pupil; iris periphery black; posterior periphery of eye blue. Dorsal colouration varied from pale tan (diurnally) to brick red (nocturnally).

Colour of holotype in preservative. As in life, but with dorsum fading to pale tan.

Measurements. Holotype: SVL 43.6, HDL 14.6, HDW 16.3, SNT 5.5, EYE 4.6, IOD 4.8, TMP 1.9, TEY 1.6, IN 3.8, NS 1.2, EN 3.6, TIB 20.7, ML 13.2, PL 19.1, IML 2.4.

Variation. Measurements of the type series are shown in Table 1 . AMS R 173126 has dark brownish-grey marbling on the dorsal surface running from the interorbital region down the back; pinkish coloured webbing, particularly ventrally, between toes III IV; more distinct barring on limbs in preserve; and less dense black colouration on the ventral surfaces of the thighs. AMS R 173507 has a uniformly coloured dorsum, with no black or white spotting or dark patterns; indistinct barring on dorsal surface of tibiotarsus; and grey webbing on hands and fingers. AMS R 173129 has an indistinct, slightly darker brown, irregular pattern running along dorsum; two, 1 mm diameter white spots on the back posteriorly; and completely black posterior and anterior surfaces of thighs, without white spotting. AMS R 173128 is strongly patterned with a darker brown, irregular pattern and scattered darker brown specks on the entire dorsum; very distinct barring on limbs; and less black colouration on ventral surfaces of thighs. NCSM 77318 has pale brown or cream mottling on the dorsal surface of head, back and limbs in life; and a dark brown dorsum with pale dusting only visible in between barring on the dorsal surface of the legs in preserve. ZFMK 91076 is almost uniformly pale tan in life and in preserve; has faint barring on arms and legs; slightly darker blotches in the interorbital region and along back; less extensive black colouration marbled with white on flanks, anterior and posterior surfaces of upper arms, anterior surface of lower arms, and anterior and posterior surface of thighs; and only faint black speckling on ventral surfaces of fingers II IV and finger webbing, ventral surface of crus, and anterior half of pes, including ventral surface of toes I III and toe-webbing. UNS 0 0 105 has an almost uniformly pale tan dorsum in life and in preserve; has faint barring on arms and legs; and the most extensive black colouration, without white spotting on flanks, anterior and posterior surfaces of upper arms, anterior surface of lower arms, ventral surfaces of fingers I IV and finger-webbing, anterior and posterior surface of thighs, ventral surface of crus, and anterior half of pes, including ventral surface of toes I III and toe-webbing. UNS 0 0 105 also has fine black specks concentrated on ventral surface of chest, but also sparsely distributed on the belly and outer margins of the throat. Males vary in the extent and colouration of loose skin in the gular region; UNS 00103/AMS R 173128 has only slightly loose skin in the gular region, only weakly coloured grey; UNS00104/AMS R 173129 has loose skin in the gular region, coloured greyish black; NCSM 77318 and ZFMK 91076 have the most pronounced loose skin in the gular region, with dark grey colouration also extending ventrally along margins of throat. All males lack nuptial pads or asperities.

HDL:SVL 0.33 0.35 0.34 0.35 0.33 0.38 0.37 0.34 TMP:EYE 0.41 0.53 0.45 0.47 0.60 0.61 0.54 0.51 Weight (g) 3.9 - - - - 3.5 4.0 -

* holotype

Tadpole. Tadpoles were assigned to the new species based upon the extremely low sequence divergences between the tadpole and adults frogs (0.0 0.6% at the 16S rRNA gene). The tadpole is elongate, with a tail length about 3 times body length, and the tail is about as tall as the body. The tadpole has a greatly reduced oral disc, only an upper jaw sheath, and most notably, a pair of keratinized hooks on the edge of the lower labium that face away from the mouth. Colour in life dark brownish grey, slightly paler in preservative. A detailed description of the new tadpole will be published separately.

Ecology. The species is a phytotelm breeder, depositing foam nests in small tree-holes, away from streams or ponds. Between May-July 2010, we observed eight foam nests and/or groups of tadpoles consistent with the new species, all in water-filled tree-holes, 0.3 1.2 m above the ground. Adults of the new species were also adjacent to these tree-holes on several occasions. The advertisement call of the species is unknown.

Conservation status. Rhacophorus vampyrus is known only from Bidoup-Nui Ba National Park. The actual distribution of the new species is unknown but probably extends into montane evergreen forest in the Langbian Plateau, including Chu Yang Sin National Park in Dak Lak Province, and Phuoc Binh National Park in Ninh Thuan Province, which are continuous with Bidoup-Nui Ba National Park. Given the available information, we suggest the species should be considered Data Deficient following IUCN’s Red List categories (IUCN 2001).

Comparisons. The pale tan to brick red dorsum, white venter, and mostly black flanks, ventral surface of upper arms and crus, and anterior and posterior surface of thighs distinguishes the new species from all mainland Southeast Asian congeners (Table 2).

TABLE 2. Rhacophorus species known from mainland Southeast Asia. Fields that differ to those of the new species are highlighted in grey.

Pointed projection

Species Dorsum colour Ventral colour Pattern on sides of body Dorsal colour of webbing Finger webbing on heel Source Smith 1924; variable, white and/or yellow; Inger et al. brown with dark scattered black/grey/brown variable; dark greyish 1999; JR pers.

annamensis markings white spots brown/black complete absent obs.

greyish brown with Günther 1858; dark obscure cream coloured; no absent (narrow flap Brown &

appendiculatus mottling spots – – reduced only) Alcala 1994 Ahl, 1927; Bordoloi et al.

bipunctatus green yellow 1–3 black spots red reduced present 2007

chocolate brown with

white markings Andersson anteriorly, whitish 1939; Ohler

burmanus green posteriorly* – – reduced absent 2009

olive/pale green;

small pale yellow yellow, 2 black spots (one side

calcaneus dots pale yellow only) yellow/olive, dark spots reduced present Smith 1924

dark green or

brown; small pale bright yellow with 2 black Orlov et al.

chuyangsinensis spots bright yellow spots bright yellow reduced present 2008

throat and chest white

sand-coloured with with brown speckles; posterior dark brown to

cinnamon coloured belly light yellow to blackish with iridescent blue Manthey &

cyanopunctatus cross bars dark down the flanks spots – reduced absent Seiof 1998 Blanford 1881; dirty white, mottled Bordoloi et al.

dennysi green with dusky none whitish reduced absent 2007

white with variable black

dorsoviridis green white spotting – reduced absent Bourret1937 fleshy with brown lower flanks blackish with Ohler et al.

duboisi dark olive green spots white spots marbled black and white reduced absent 2000 green, with round David 1872 spots of a golden cream yellow, spotted "1871; Liu

dugritei metallic ash with dark grey marbled with cream yellow – reduced absent 1950 mottled grey, Inger et al. brown, and olive white anteriorly; marbled back and white/pale absent (large 1999; JR pers.

exechopygus green yellow posteriorly blue as dorsum complete tubercle only) obs.

The presence of a distinct pointed projection at the tibiotarsal articulation further distinguishes the new species from all mainland Southeast Asian congeners with the exception of Rhacophorus bipunctatus , R. calcaneus , R. chuyangsinensis , R. hoanglienensis , R. kio , R. marmoridorsum , R. rhodopus , and R. spelaeus (Table 2). From these species, R. vampyrus can be distinguished further on the basis of colouration and extent of hand webbing. Rhacophorus vampyrus differs from: R. bipunctatus in having a pale tan to brick red dorsum (versus green), white ventral surface with black margins (versus yellow), black flanks (versus 1 3 black spots), and grey/black webbing on dorsal surface (versus red); from R. calcaneus in having a pale tan to brick red dorsum (versus olive/pale green), white ventral surface with black margins (versus pale yellow), black flanks (versus yellow with black spots), and grey/black webbing on dorsal surface (versus yellow/olive with dark spots); from R. chuyangsinensis in having a pale tan to brick red dorsum (versus dark green or brown with small pale spots), white ventral surface with black margins (versus bright yellow), black flanks (versus bright yellow with black spots), and grey/black webbing on dorsal surface (versus bright yellow); from R. kio in having incomplete finger webbing (versus complete), a pale tan to brick red dorsum (versus green), white ventral surface with black margins (versus lemon yellow), black flanks (versus with one black spot), and grey/black webbing on dorsal surface (versus orange and black); from R. marmoridorsum in having a pale cream to brick red dorsum (versus flesh coloured with marble chocolate pattern), white ventral surface with black margins (versus light cream), and black flanks (versus irregular spots and lines of chocolate colour on cream background); from R. rhodopus in having a white ventral surface with black margins (versus yellow), black flanks (versus 1 black spot), and grey/black webbing on dorsal surface (versus red); and from R. spelaeus in having a pale tan to brick red dorsum (versus greyish brown with dark, irregular spots), white ventral surface with black margins (versus light grey with dark specks), black flanks (versus light grey), and grey/ black webbing on dorsal surface (versus grey-brown with dark speckles).

Tables

TABLE 1. Measurements (mm) of Rhacophorus vampyrus sp. nov. Abbreviations defined in text.

	AMS R 173127*	UNS 00103/ UNS 00104/ NCSM AMS R AMS R 77318 173128 173129	ZFMK 91076	AMS R 173126	AMS R 173507	UNS 0 0 105
Sex	Male	Male Male Male	Male	Female	Female	Female
SVL HDL HDW SNT	43.6 14.6 16.3 5.5	42.5 44.8 44.2 14.9 15.2 15.3 17.0 17.3 17.0 5.8 6.2 6.3	43.3 14.4 17.1 5.8	38.9 15.0 16.8 5.5	41.4 15.3 16.6 6.1	53.4 18.0 20.8 7.3
EYE IOD	4.6 4.8	4.6 4.5 4.7 6.1 5.6 4.7	4.5 5.5	4.2 4.9	4.7 5.2	5.3 5.1
TMP TEY IN	1.9 1.6 3.8	2.4 2.0 2.2 1.2 1.6 1.6 4.1 3.6 4.2	2.7 1.3 4.5	2.6 1.5 3.8	2.5 2.0 3.9	2.7 2.0 4.8
NS EN TIB	1.2 3.6 20.7	2.1 2.4 2.7 3.9 3.9 3.6 20.7 21.3 20.7	2.3 3.5 20.5	1.7 3.6 20.1	2.0 3.7 21.1	3.0 4.4 26.4
ML PL IML HW:HL	13.2 19.1 2.4 1.12	12.5 13.1 14.4 17.4 17.4 19.3 1.9 1.9 2.1 1.14 1.14 1.11	13.8 18.0 1.6 1.19	12.2 17.6 2.0 1.12	13.8 18.2 1.6 1.0 8	16.8 23.1 2.4 1.16
HL:HW TIB:SVL	0.9 0.47	0.88 0.87 0.9 0.49 0.47 0.47	0.84 0.47	0.89 0.52	0.92 0.51	0.87 0.49