Philautus maosonensis Bourret, 1937

BAIN, RAOUL H. & TRUONG, NGUYEN QUANG, 2004, Herpetofaunal Diversity of Ha Giang Province in Northeastern Vietnam, with Descriptions of Two New Species, American Museum Novitates 3453, pp. 1-43 : 24-32

Philautus maosonensis Bourret, 1937

This species was collected while calling on vegetation between 0.5 and 2 m off the ground in the forest near (but not directly adjacent to) any water source. This frog called alone or in syncopated choruses. Choruses can be as dense as one individual male per 4 m 2. Males were found calling on tops of leaves, though never on the topmost leaf of the plant. The color of these specimens is uniformly brown during the day (orangeypeach at night), thighs brown, except flash colors, which are yellow; tips of some fingers yellow; venter mottled white on black. This species has been reported in Mau Son, Lang Son, and Cao Bang Provinces on the northeast border with China, and from Tam Dao in Vinh Phu Province ( Bourret, 1937; Inger et al., 1999; Orlov et al., 2002).

MEASUREMENTS OF MALES (in mm, N = 6, AMNH 163904–163909; mean, ± standard deviation, range in parentheses): SVL 30.60 ± 2.42 (25.97–32.88), HDL 12.15 ± 0.83

(11.18–13.44), HDW 10.92 ± 0.78 (9.43– 11.54), SNT 4.73 ± 0.57 (3.69–5.16), TIB 15.42 ± 0.99 (13.53–16.46), FTL 22.02 ± 1.80 (18.79 –23.98), HND 10.09 ±0.85 (9.11–11.58), EYE 4.65 ± 0.35 (4.15–5.23), TMP 1.79 ± 0.21 (1.57–2.06), IOD 3.71 ± 0.34 (3.30–4.14); median HDL:HDW 1.12 (1.03–1.23), median TIB:SVL 0.50 (0.49– 0.52), median TMP:EYE 0.39 (0.37–0.42), median SNT:HDL 0.39 (0.29–0.46).

Philautus odontotarsus Ye and Fei, 1993

Figure 16

This species was found at the forest edge, beside large wet rice paddies at 600 m. It was found atop vegetation and branches less than 2 m above the ground.

Philautus odontotarsus is part of a taxonomically problematic group including Rhacophorus appendiculatus , R. bissaculus , R. cavirostris , and R. verrucosus (see discussions in Inger et al., 1999: 39–40; Orlov et al., 2001: 39–40; Frost, 2002; Orlov et al., 2002: 97–98). Although Philautus odontotarsus has been reported from Lao Cai Province (Ohler et al., 2000), this is the first stated record east of the Red River. In addition to the Lao Cai specimens (Ohler et al., 2000), these specimens are possibly conspecific with unidentified specimens from Tam Dao ( Inger et al., 1999: 40) and specimens reported as Rhacophorus verrucosus from Lao Cai (Orlov et al, 2001).

We base our identification on the specimens’ similarity with the original description of P. odontotarsus and the proximity of Ha Giang Province to its type locality (Ye and Fei, 1993). The type locality of P. odontotarsus is in Yunnan Province, China, but the type locality of other frogs of the group is much farther away: R. appendiculatus , the Greater Sunda Islands; R. bissaculus, Phu Kading , Loei Province, Thailand; R. cavirostris , Sri Lanka; and R. verrucosus , Myanmar. We include a translation of the original description of P. odontotarsus from Chinese in appendix 2.

Our specimens are identified as P. odontotarsus based on the following characters: male SVL between 28 and 35 mm; vomerine teeth present in short oblique rows; relative finger lengths I ‹ II ‹ IV ‹ III; finger tips with expanded disks; disks with circummarginal grooves; disk on I small; disk on III smaller than tympanum diameter; fingers with lateral fringes and basal webbing; five to six sawteeth tubercles on outer edge of forearm; no outer metatarsal tubercle; saw­teeth tubercles on tarsus along outside of toe V to tibiotarsal joint; skin rough; dorsum with irregularly shaped tubercles, more on upper eyelids; dorsum olive and gray; flanks grayish­green; back of forelimbs with black­brown transverse bars; flash colors orange­red; chest and belly with scattered spots; males with creamy white nuptial pad at the base of finger I.

Our specimens differ from the original description in female size (SVL 43 mm for the Chinese population, 34.2 and 35.64 for Ha Giang specimens), male TMP:EYE (0.50– 0.66 for Chinese populations, 0.34–0.47 for Ha Giang specimens), and foot webbing (in Chinese population web full to toe tips, except toes I and V; for Ha Giang specimens web full to between distal subarticular tubercle and disk for toes I, V, and postaxial sides of toes II and III, web basal [below subarticular tubercle] for preaxial side of II, web full to between medial and distal subarticular tubercles for toe IV, and the postaxial side of III, web reaching all disks as a fringe, except for preaxial side of II). We agree with Inger et al. (1999: 40) and Orlov et al. (2000: 39– 40) that our identification is uncertain and that a stable identification (both generic and specific) requires a detailed analysis of this group of species from India, Myanmar, Thailand, Indochina, and China.

Philautus odontotarsus is known from China: Tibet Autonomous Region, and Yunnan, Guizhou, Hainan, Guangxi Provinces; and Vietnam: Lao Cai, Ha Giang Provinces ( Fei, 1999; Ohler et al., 2000; this paper) .

MEASUREMENTS OF FEMALES (in mm, N = 2, A­163977, 163978): SVL 34.2, 35.64; HDL 11.94, 13.35; HDW 11.49, 12.77; SNT 5.31, 6.09; TIB 17.44, 17.48; FTL 22.69, 24.36; HND 10.0, 9.83; EYE 4.67, 4.97; TMP 1.59, 2.79; IOD 4.18, 3.96); HDL: HDW 1.04. 1.05, TIB:SVL 0.49, 0.51, TMP: EYE 0.34, 0.56, SNT:HDL 0.44, 0.46.

MEASUREMENTS OF MALES (in mm, N = 4, A­163975, 163976, 163979, 163980; mean, ± standard deviation, range in parentheses): SVL 31.59 ±1.01 (30.58–32.84), HDL 11.68 ± 0.50 (11.13–12.19), HDW 10.82 ± 0.41 (10.45–11.39), SNT 5.25 ± 0.68 (4.44– 5.99), TIB 16.61 ± 0.76 (15.48–17.05), FTL 22.12 ± 0.66 (21.14–22.59), HND 9.71 ± 0.96 (8.36–10.57), EYE 4.45 ± 0.48 (4.01– 4.96), TMP 1.74 ± 0.13 (1.59–1.90), IOD 3.57 ± 0.08 (3.48–3.65). median HDL:HDW 1.08 (1.05–1.13), median TIB:SVL 0.53 (0.51–0.55), median TMP:EYE 0.39 (0.34– 0.47), median SNT:HDL 0.45 (0.39–0.54).

Philautus rhododiscus Liu and Hu 1962

Figure 17–19

A single specimen was found at 0545 hours on a tent at the 1400 m camp on Mt. Tay Con Linh II (site 2). This represents the first record of this species outside of China (Guangxi, Fujian Provinces) ( Fei, 1999). The specimen from Ha Giang matches the original description precisely (Liu and Hu, 1962); diagnosis: toes with disks orange­red on the ventral side, tea­brown dorsally; dorsum with white tubercles interweaved as a network. A translation of the original description of P. rhododiscus is in appendix 3 .

We enhance the description, based on AMNH 163892/IEBR 59, an adult male. Palmar tubercle appears to have a free edge. Relative toe length I ‹ II ‹ V = III ‹ IV, which agrees with the illustration in the original description (Liu and Hu, 1962: fig. 11a, b). Webbing reaches the base of every toe disk as a fringe, except for toe V, where it reaches subarticular tubercle (reaches second articulation on toe IV of holotype). Preaxial fringe on toe I runs from the base of the disk to the top of the subarticular tubercle; toe V without a postaxial fringe. The dorsal longitudinal, pointy ridges are as in the description of the holotype. The Ha Giang specimen exhibits spinules on every dorsal and lateral surface (torso, head, appendages, and digits), except the tympanum. The dorsum is olive brown with long black spots on the appendages, middorsal, and sacral areas. Venter is black with a white marbled network. Chin is dark black with white spots. White nuptial pad on the thumb.

MEASUREMENTS OF MALE (in mm, N = 1, AMNH 163892/IEBR 59): SVL 26.60, HDL 11.70, HDW 8.30, SNT 4.00, EYE 1.90, IOD 2.85, TMP 1.85, HND 8.4, FPW 1.72, TIB 14.9, FTL 20.25, TPW 1.19.

Polypedates dugritei David, 1872 ‘‘1871’’

Figures 6E, F , 20–22

Polypedates dugritei David, 1872 ‘‘1871’’.

Rhacophorus hui Liu, 1945 .

Rhacophorus pleurostictus batangensis Vogt, 1924 .

Hyla bambusicola (Barbour, 1912) .

Polypedates davidi Sauvage, 1877 .

Four male specimens referable to P. dugritei were found at 1700 m (fig. 6E, F). All were calling atop broadleaves or bamboo 1– 3 m above temporary forest pools (in the same microhabitat as Chirixalus gracilipes ). No females or foam nests were found. Polypedates dugritei is known from southern Sichuan, Guizhou, Hubei, and Yunnan Provinces, China, and the Hoang Lien Mountains of northwestern Vietnam (Wu and Zheng, 1994; Fei, 1999; Ohler et al., 2000; Orlov et al., 2001). The specimens from Ha Giang are the first recorded east of the Red River.

The series from Ha Giang closely matches the brief original description ( David, 1872 ‘‘1871’’) and a subsequent amplification of the description based on Hoang Lien Mountain specimens (Orlov et al., 2001). The color in life (and in preservative) of Ha Giang P. dugritei is as follows: dorsum light green (livid blue), dorsum, flanks, and limbs heavily spotted red­brown to digit tips, each spot with thick black border (brown with black border); red­brown band on supratympanic fold present or absent; axillary region, flanks, posterior and anterior surface of thighs thickly marbled black and white; inner thighs tinged red; white line along lateral edge of toe V continuous with the lateral margin of the tarsus and across anus, thin black line immediately below transmedial white line (fig. 6F); a similar, but less distinct line on the lateral edge of finger IV along the forearm to the flanks; thick black on white marbling across the entire venter (black on white), chin with more brown mottling (gray), chest yellow­white (dull white); webbing and tips of feet red; dorsal one­third of iris gold, orange­red ventral two­thirds.

Specimens of P. dugritei from Ha Giang were compared with specimens from the Hoang Lien Mountains and syntypes from Mouping, China (appendix 1) (figs. 20–22). All three series closely match each other in shape, although the Hoang Lien Mountain specimens are the smallest of all published accounts of P. dugritei , its synonyms, and similar species (table 4). Tubercles on the hands and feet of the syntypes are very pronounced (fig. 21). The inner metatarsal tubercle is high and sharply projecting in the syntypes, low and ovoid in the Vietnamese specimens (fig. 22). Orlov et al. (2001: 23) commented on the possible conspecificity of P. dugritei from the Hoang Lien Mountains with P. puerensis ( He, 1999) . The description of P. puerensis closely matches that of the syntypes and Vietnamese series of P. dugritei , except that P. puerensis has a glandular wart at end of the lip that forms a triangle with supratympanic fold and the supratympanic fold continues from the eye to connect to a weak, obscured fold on the flank (not found in P. dugritei ). We did not examine the type series of P. puerensis , but in light of the marked similarity between the two species, we include a translation of the original description of P. puerensis in appendix 4.

Polypedates dugritei shows a high level of sympatric variation (Pope and Boring, 1940; Liu, 1950; Wu and Zheng, 1994; Fei, 1999; Ohler et al., 2000; Orlov et al., 2001), including: (1) vomerine teeth vary from being absent to distinctly present; (2) red­brown dorsal spots vary from lightly scattered to a dense mosaic; (3) red­brown coloration of supratympanic fold present or absent; (4) venter varies from immaculate to marbled to deeply mottled brown; (5) white transmedial line along lateral edge of toe V continuous with the lateral margin of the tarsus, across anus, and continuous with lateral edge of finger IV along the forearm (created when individual sits still [fig. 6F]) present or absent; (6) inner three toes orange­red or not; (7) groups of adults with distinct size differences. Some differences are also noted across allopatric populations: the size range of P. dugritei is 32–47 mm for males and 46–66 mm for females (table 4), and specimens are described as being rough, with tubercular skin (Liu, 1950; Fei, 1999), whereas the syntypes and Vietnamese specimens are smooth, with or without tubercles.

Wu and Zheng (1994) reported that four Chinese populations of P. dugritei within 100 km of each other exhibit two distinct karyotypic patterns. They described a new species, P. zhaojuensis from Sichuan and Hubei Provinces, based on differences in the karyotype ( P. zhaojuensis with secondary constriction in the middle of long arm of chromosome 10; P. dugritei with satellite on the terminal end of arm 10), and snout coloration (yellowish­brown in P. zhaojuensis ). This suggests that P. dugritei is a complex of species and would account, at least in part, for its widespread range and highly variable nature (Pope and Boring, 1940; Liu, 1950; Ohler et al., 2000; Orlov et al., 2001). The confusion that this variation presents is also evidenced by the five junior synonyms of P. dugritei . A thorough investigation of all known specimens (including types of junior synonyms) is necessary to help resolve this confusion.

MEASUREMENTS OF MALE POLYPEDATES DU­ GRITEI (in mm, N = 4 males, AMNH 163910, 163911, 163912/IEBR 62, 163913/IEBR 63; mean, ± standard deviation, range in parentheses): SVL 42.59 ± 2.87 (40.11–46.63), HDL 15.50 ± 0.77 (14.43–16.10), HDW 15.92 ± 0.70 (15.13–16.75), SNT 7.11 ± 0.40 (6.67–7.64), EYE 5.02 ± 0.63 (4.26– 5.79), IOD 5.28 ± 0.33 (4.93–5.65), TMP 2.39 ± 0.19 (2.20–2.57), HND 13.85 ± 0.92 (13.18–15.20), TIB 18.03 ± 0.58 (17.43– 18.80), FTL 28.39 ± 1.35 (26.61–29.68);

TABLE 4 Published Snout­Vent Lengths (in mm) of Polypedates dugritei, Its Junior Synonyms ( Rhacophorus bambusicola and Rhacophorus pleurostictus batangensis ), and Polypedates puerensis Measurement means with ranges are shown for series.

median HDL:HDW 0.97 (0.93–1.02), medi­ an TMP:EYE 0.48 (0.44–0.53), median SNT: HDL 0.46 (0.44–0.48), median TIB:SVL 0.42 (0.40–0.45).

Rhacophorus hoanglienensis Orlov et al., 2001

This single specimen was collected on a broad leaf 2 m above the ground in the forest. This is only the second specimen and the first female recorded for this species and is the first record east of the Red River (Orlov et al. 2001). Some additional color information for this specimen: dorsum orange­red with irregular black and brown spotting; white line runs from the supra­tympanic fold, over the eyelid, across the canthus rostralis to tip of snout; brown line running between the eyes; inner thigh black with white speckling, white increases along sides; webbing gray; venter evenly dark brown to tibia.

MEASUREMENTS (in mm, N = 1, AMNH 163767/IEBR 57): SVL 57.89, HDL 21.45, HDW 22.06, SNT 8.85, EYE 7.17, IOD 7.84, TMP 3.45, HND 17.38, TIB 31.8, FTL 40.20.

Figures

Fig. 16. Philautus odontotarsus.

Fig. 17. Philautus rhododiscus (AMNH 163892/IEBR 59), male: dorsal (left) and ventral (right) views. Scales equal 10 mm.

Fig. 18. Head of Philautus rhododiscus (AMNH 163892/IEBR 59), male. Scale equals 5 mm.

Fig. 19. Philautus rhododiscus (AMNH 163892/IEBR 59), male: hand (left) and foot (right). Scales equal 5 mm.

Fig. 6. Three frog species from Ha Giang Province, Vietnam: (A) paratype of Rana iriodes, new species (AMNH 163928), adult male, SVL 42.80 mm; (B) paratype of Rana iriodes, new species; (C) paratype of Rana tabaca, new species (AMNH 163915), adult male, SVL 59.18 mm; (D) paratype of Rana tabaca, new species (AMNH 163920), adult female, SVL 103.40 mm; (E) Polypedates dugritei (AMNH 163910), adult male, in profile; and (F) posterior view showing transmedial white line, SVL 40.50 mm.

Fig. 20. Syntypes of Polypedates dugritei: clockwise from top left: MNHNP 1994.2641, male, SVL 42.40 mm; MNHNP 1994.2361, female, SVL 55.92 mm; MNHNP 1994.2362, female, SVL 59.24 mm; and MNHNP 5563, male, SVL 39.80 mm.

Fig. 21. Hands and feet of Polypedates dugritei: hand (A) and foot (B) of MNHNP 1994.2361, syntype; hand (C) and foot (D) of AMNH 163910. Scales equal 5 mm.

Fig. 22. Feet of Polypedates dugritei: (left) foot of MNHNP 5663, male syntype, with taller tubercles (especially the metatarsal tubercle) than those from Vietnam, as shown in the foot of AMNH 163910 (right).