Theloderma nebulosum Rowley, Le, Hoang, Dau & Cao, 2011, sp. nov.
Creators: name: Rowley, Jodi J. L.
name: Le, Duong Thi Thuy
name: Hoang, Huy Duc
name: Dau, Vinh Quang
name: Cao, Trung Tien
Publisher: name: Zenodo
Year: 2011
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Rhacophoridae
Theloderma
Theloderma nebulosum

description: Theloderma nebulosum sp. nov.
Holotype: AMS R 173409, adult female, on a leaf of tree,> 10 m from stream, in bamboo/montane evergreen forest in Ngoc Linh Nature Reserve, Dak Glei District, Kon Tum Province, Vietnam (15.06º N, 108. 107.86º E, ~ 2000 m). Collected at ~ 20:00 h on 3 April 2010 by J. J. L. Rowley, Le T. T. D., Dau Q. V. and Hoang D. H.
Other material. UNS 00141/AMS R 173877, eight tadpoles collected inside a water-filled hole (30 cm diameter, 20 cm deep) in a dead tree, approximately 1 m from the ground, on 17 July 2009 in Ngoc Linh Nature Reserve, Dak Glei District, Kon Tum Province, Vietnam (15.08º N, 108. 107.96º E, ~ 1950 m). Collected by Le T. T. D., Dau Q. V., Le V. K. and Nguyen Q. H.
Etymology. specific epithet nebulosus L., meaning misty or cloudy, in reference to the type locality the species, high-elevation forest often shrouded in cloud.
Suggested common name. Misty moss frog (English), Ếch cây sần sương mù (Vietnamese).
Diagnosis. The new species is assigned to the genus Theloderma by the presence of tuberculate skin on the dorsum, a distinct tympanum, terminal phalanx with a Y-shaped distal end, intercalary cartilage between the terminal and penultimate phalanges of digits, tips of digits expanded into large disks bearing circummarginal grooves, absence of co-ossification of the head skin to the skull (Liem 1970), and molecular data. Theloderma nebulosum sp. nov. is distinguished from all other Theloderma by a combination of (1) small body size (29.2 in a single adult female), (2) absence of vomerine teeth, (3) only scattered, minute dorsal asperities, (4) no webbing on hands, (5) brown dorsum with indistinct darker brown longitudinal markings and brownish black lateral surfaces extending from snout to groin, and (6) a bicoloured iris, being pale gold in upper third and dark red in lower two-thirds.
Description of holotype. Body relatively robust; head length equals head width; snout bluntly truncate in dorsal view, rounded in profile; canthus rostralis relatively indistinct, rounded; loreal region slightly concave; interorbital region slightly convex; nostrils oval, only slightly protuberant, without flap of skin laterally, much closer to tip of snout than eye; pupil diamond-shaped, horizontal; tympanum distinct, tympanic rim elevated relative to skin of temporal region, 67% of eye diameter; pineal ocellus absent; skin not co-ossified to forehead; vomerine teeth absent; choanae oval, at margins of roof of mouth; tongue attached anteriorly, wide, notched posteriorly; supratympanic fold present, extending to just beyond level of axilla. Forelimbs slender, 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 width 209% third finger width), disks slightly wider than long (third finger disk width 135% of length), third finger disk width 77% of tympanum diameter; webbing on fingers absent; dermal fringing absent; subarticular tubercles prominent, rounded, formula 1, 1, 2, 2; supernumary tubercles present; two palmar tubercles; accessory palmar tubercles present, small; prepollex with low tubercle. 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 I2 +–3 II2 +–3+ III2 +–3½ IV3 +–2+V; subarticular tubercles rounded, distinct, formula 1, 1, 2, 3, 2; inner metatarsal tubercle distinct, oval; outer metatarsal tubercle indistinct. Dorsal skin very weakly rugose with very sparsely distributed minute, pearly asperities; ventral surface of thighs and posterior surface of belly coarsely granular, chest and throat smooth. Dermal fringes and pointed projection at tibiotarsal articulation absent.
Colour of holotype in life. Dorsum brown with indistinct darker brown longitudinal markings running along midline from snout, including anterior portions of eyelid, to vent; dorsolateral surfaces of body paler brown, slightly mottled; darker spot above cloaca and a darker line slightly anterior to cloaca, running along dorsal midline. Ventral surface of tibiotasus and arms brown with single, indistinct, dark brown band running across ventral surface of tibiotarsus and thigh, and on to body; when leg is adpressed, forms a continuous line. Dark brown inguinal blotches. Ventral surface of hands and feet mostly greyish black with pale bluish grey spots, brown along outside margins; ventral surface of toe-pads brown. Dark brownish black lateral surfaces, extending from snout, through eye and over tympanum to groin; pale bluish white speckling on side of head and on anterior flanks; pale bluish white marbling on posterior surfaces of flanks. Ventral surface of body brownish black with pale bluish white marbling on chest, inguinal region, anterior and posterior surface of thighs; faint white speckling on throat and ventral surfaces of arms. Iris pale gold in upper third, and dark reddish brown in lower two-thirds; division of eye colour matches skin coloration adjacent to eye (Figure 4).
Colour of holotype in preservative. As in life, but paler brown colours on dorsum fade to creamy grey (Figure 5).
Variation. Measurements of the holotype are in Table 3. As was noted for T. licin (McLeod & Norhayati 2007), the new species has the ability to change colour over time, albeit to a lesser degree than was observed in the Langbian Plateau Theloderma specimens (described below). Upon capture (at night), the dorsal pattern was more distinct.
Tadpole. Tadpoles were assigned to the new species based upon the lack of sequence divergence between the tadpole and adults frog at the 16S rRNA gene. Body shape rounded and depressed, eyes dorsal; nares small and nearer to snout than eye; vent tube medial; spiracle sinstral; tail tip broadly rounded. The oral apparatus is anteroventral, almost terminal. The spiracle is ventrolateral and lacks a free tube. The labial tooth row formula (LTRF) is 4(2–4)/3, the marginal papillae have a large dorsal gap. Tadpoles have a uniform dark brown body in preserve, with paler brown tail muscle and transparent tail fin with fine brown speckling (Figure 6). Measurements (in mm) of one tadpole at developmental stage 28 (Gosner 1960): total length 24.6 mm, body length 10.3 mm, tail length 14.3 mm, maximum tail height 4.6 mm, tail muscle height 2.8 mm, interorbital distance 2.1 mm, oral disk width 2.5 mm, oral disk height 0.9 mm. Seven other tadpoles were collected in the same water body, ranging from developmental stage 26–30 and body length 6.0– 13.3 mm.
Theloderma nebulosum sp. nov. Theloderma palliatum sp. nov.
AMS R 173409* AMS R 173130 AMS R 173131* UNS 00140/ AMS R
173508 Distribution and ecology. The new species is known only from montane evergreen forest at ~ 2000 m elevation in Ngoc Linh Nature Reserve (Figure 7). At this elevation, temperatures during the coldest months are 5–6ºC and temperatures during the hottest months are <20ºC (Le et al. 1999). Conditions are also extremely wet, with almost constant mist and approximately 250 rain days per year (Le et al. 1999). Specimens of the new species were collected at 15.8ºC and 90% relative humidity (July 2009) and ~17ºC and 95% relative humidity (March–April 2010).
Comparisons. The small body size; absence of vomerine teeth; minute, very sparsely scattered dorsal asperities; no webbing on hands; brown dorsum with indistinct darker brown longitudinal markings and dark brown lateral surfaces extending from snout to groin; and a distinctly bicoloured iris distinguishes T. nebulosum sp. nov. from all species of Theloderma (Table 1). Theloderma nebulosum sp. nov. appears most morphologically similar to the small T. lateriticum, T. licin, T. rhododiscus, T. truongsonense comb. nov. and the newly collected Langbian Plateau specimens (described below). Theloderma nebulosum sp. nov. differs morphologically from T. lateriticum by having a brown dorsum with indistinct darker brown longitudinal markings (versus a dorsum with a deep brickred wash overtop dark black-green), minute, sparsely scattered dorsal asperities not visible with the naked eye (versus distinctly white dorsal asperities) and a distinctly bicoloured iris (versus uniformly deep brick red iris). From T. licin, T. nebulosum sp. nov. differs by lacking webbing or fringing on the hands (versus lateral dermal fringes and basal webbing on the hands), having a brown dorsum with darker brown longitudinal markings (versus a uniformly coloured dorsal surface), and a distinctly bicoloured iris (versus uniformly red iris). From T. rhododiscus, T. nebulosum sp. nov. differs by having the ventral surfaces of the disks being brown to greyish brown (versus orangered), and a distinctly bicoloured iris (versus uniformly reddish brown). From T. truongsonense comb. nov. (Figure 2), T. nebulosum sp. nov. differs by having a brown dorsum with indistinct darker brown longitudinal markings (versus a pale brown dorsum with small, distinctly darker brown blotches and no longitudinal lines), minute, sparsely scattered dorsal asperities (versus no dorsal asperities), and a black/dark brown belly and ventral surface of thighs, with white/pale brown marbling (versus a grey belly and thighs with white spots). From the Langbian Plateau specimens (described below), T. nebulosum sp.nov. differs by having a brown dorsum with darker brown longitudinal markings (versus a pale to medium brown dorsum with characteristic dark brown blotches), more sparsely scattered dorsal asperities and slightly less webbing on the feet. In addition, Theloderma nebulosum sp. nov. has uncorrected sequence divergences at the 16S rRNA gene of>8.5% for all Theloderma for which homologous 16S rRNA sequences are available.
The tadpole of T. nebulosum is very similar in body shape and proportions to other Theloderma species for which tadpoles have been described: T. asperum (Leong & Lim 2003), T. corticale (Kunz et al. 2010), T. horridum (Boulenger 1903), T. moloch (Annandale 1912), and T. stellatum (Inger et al. 1999; Wassersug et al. 1981). The LTRF of the new species, 4(2–4)/3, is the same as reported in T. stellatum (Inger et al. 1999; Wassersug et al. 1981) and T. moloch (Annandale 1912), but differs from that reported for T. asperum 3(2–3)/3(1) (Leong & Lim 2003).
description: Published as part of Rowley, Jodi J. L., Le, Duong Thi Thuy, Hoang, Huy Duc, Dau, Vinh Quang & Cao, Trung Tien, 2011, Two new species of Theloderma (Anura: Rhacophoridae) from Vietnam, pp. 1-20 in Zootaxa 3098 on pages 8-12, DOI: 10.5281/zenodo.202712