Amy ~athrop,~ Robert W. ~urphy,~ Nikolai L. 0rlov: and Cuc Thu HO~

Transcription

1 Russian Journal of Herpetology Vol. 5, No. 1, 1998, pp TWO NEW SPECIES OF Leptobrachium (ANURA: MEGOPHRYIDAE) FROM THE CENTRAL HIGHLANDS OF VIETNAM WITH A REDESCRIPTION OF Leptobrachium chapaense Amy ~athrop,~ Robert W. ~urphy,~ Nikolai L. 0rlov: and Cuc Thu HO~ Submitted 15 March Two species of Leptobrachium are described from the Central Highlands of Vietnam in Gia Lai Province. These descriptions increase the number of species of Leptobrachium in Indochina from four to six. We also include a redescription of a northern species, Lepfobrachium chapaense, based on recent collections from Tam Dao, Vinh Phu Province, Vietnam. A key to the species of Leptobrachium from Indochina is provided. Key words: Amphibia, Anura, new species, Leptobrachium chapaense, Leptobrachium xanthospilum, Leptobrachium banae, taxonomic key. Our field work in Vietnam for the past five years has led to the discovery of several new species of amphibians (e.g., Lathrop et al., in press), as well as known species not previously recorded from Vietnam. These discoveries include localities collected heavily by pourret. Indeed, at every locality visited thus far, we have found either new species, or new records of species occurrences in Vietnam. Among these discoveries, in the forests of Vietnam we have found two new sympatric species of frogs belonging to the rare megophryid genus Leptobrachium. These terrestrial amphibians may be found in high mountain regions either near streams or on the forest floor. Their broad heads, and often dual-colored iris, easily distinguishes them from other genera of anurans. Until now only four species of Leptobrachium were known from Indochina. One species, L. hasseltii, has a broad distribution (Myanmar, Thailand, Borneo, Java, and the Philippines) and the variation seen throughout its distribution has been carefully detailed by Inger (1954, 1966). Leptobrachium hendrichoni (Taylor, 1962) was described from three specimens from Malaya, Malaysia and Thailand. However, this species was subsequently synonomized Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario, M5S 2C6. AL - AmyL@rom.on.ca; RWM - drbob@rom.on.ca. Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. nbanoms@glas. apc. org. Institute of Ecology and Biological Resources, Nghia Do, Tu Liem, Hanoi, Vietnam. with L. hasseltii because the differences between the two species can be observed across several populations of L. hasseltii alone (Inger, 1966: p. 3 1). A smaller species, L. nigrops, is found on the Malay Peninsula as well as Singapore and Borneo (Berry and Hendrickson, 1963). It is very similar to L. hasseltii, but differs by having a more acurninate snout and broader head. The detailed description of one species, L. pullum, is wanting. All that is known for the species can be traced back to a meager description by Smith (1921: p. 440) in which he describes it as a "variant" of L. hasseltii and comparisons of the former made with L. nigrops (Berry and Hendrickson, 1963). Dubois (1983) elevated two Vietnamese forms, L. hasseltii pullus (Smith, 1921) and L. hasseltii chapaensis, (Bourret, 1937) to full species status without accompanying justification. In light of this taxonomic change, and because we have collected L, chapaense, we believed it desirable to redescribe L. chapaense. Notwithstanding, confirmation of the validity ofl. pullum as a distinct species must wait until type material can be examined. In the interim, the comparisons of our new species with L. pullum are based on the brief description of Smith '(1921). MATERIAL AND METHODS All measurements were taken with digital calipers to the nearest 0.01 mm and rounded to 0.1 rnm. Measurements are summarized in Table 1. Below, snout /98/ O 1998 Folium Publishing Company

2 52 Amy Lathrop et al. vent length has been abbreviated SVL. The formula for toe webbing follows Myers and Duellman (1982). Abbreviations used: ROM) Royal Ontario Museum, Toronto, Canada; MVZ) Museum of Vertebrate Zoology, University of California, Berkeley, USA; KIZ) Kunming Institute of Zoology, Yunnan Province, China; IEBR) Institute of Ecology and Biological Resources, Hanoi, Vietnam. Leptobrachium chapaense (Bourret) Figs. 1,2 Megophlys hasseltii chapaensis Bourret, 1937 Leptobrachium (Leptobrachium ) chapaense Dubois, 1980: p. 476 Diagnosis. A medium-sized Leptobrachium (female mm; male mrn) with short legs; heels do not overlap when legs are at right angles to body; webbing basal on toes. Dorsal color pattern dark brown with small, diffuse orange blotches on the sacral region, flanks, and limbs; ventral coloration of limbs black with white reticulations and banding on the thigh and knee; groin and flanks black with white pustules that change to dark mottling on white near the chest and throat. Axillary gland located in armpit, behind insertion of arm. Dorsal half of iris white. Leptobrachium chapaense has a relatively dark belly with distinct white punctuations; mottling oc- curs on the throat and darker mottling on the legs (Fig. 3) distinguishing it from L. hasseltii, L. hendricksoni, L. pullum, L. nigrops, and the little known L. hainanensis (Ye and Fe) (Ye et al., 1993) that are said to have predominantly light venters (either cream or white) with some dark spotting. The absence of distinct, dark spots on the dorsum distinguishes L. chapaense from most other species of Leptobrachium, including L. pullum, L. nigrops, L. hainanensis, many variants of L. hasseltii, and L. montanum. Description of species (based on 4 males, ROM ; and 3 females, ROM , , IEBR specimen bearing ROM field number 30006). Body short, head 42% in males, 36% females of SVL; head broad, width slightly greater than length, greater than width of body; head dorsoventrally compressed; canthus rostralis rounded, loreal region oblique, slightly concave; interorbital and frontal areas flat and sloping ventrally near snout; nostril to eye distance almost twice that of nostril to tip of snout; snout in dorsal view rounded, in lateral view sloped. Eyes large and protuberant; eye length equal to snout length; tympanum small, half diameter of eye, and separated from eye by a distance equal to or 75% tympanum diameter; supratympanic fold extends from rear comer of eye to articulation of jaw; eyes broadly separated, interorbital distance 50% of TABLE 1. Measurement for Leptobrachiurn from Indochina L. chapaense L. pullum L. hasseltii L, nigrops L. xanthospilum ' L. banae ' Measurements, mm 0" 9 0" 0 0" 0" 0 0" 9 0" 9 n=4 n=3 n=5 n=3 n=6 n=l n=7 n=8 n=2 n=14 n=2 SVL f f 11.3 Head length/svl f f 0.01 Head width/svl f f 0.01 Arm/SVL f f 0.01 Tibia/SVL f f 0.01 Note. Except SVL, where available, each measurement is given as a ratio with the mean k one standard deviation above the range (in parentheses). Numbers in bold are measurements of the holotype. NA) Not available. This study. From Smith (1921: p. 440). From Taylor (1962) for Thailand material. For further data on this species across its distribution see Inger (1954, 1966). From Beny and Hendrickson (1963).

3 Vietnamese Leptobrachium

4 54 Amy Lathrop et al.

5 Vietnamese Leptobrachium

6 56, Amy Lathrop et al. head width; posterior 112 of tongue free, distinctly notched; vomerine teeth absent; choanae separated by a distance slightly less than that between nares. Fingers long (27% SVL), tips not wider than digits; first finger slightly longer than second, latter slightly shorter than fourth; webbing absent, lateral fringes weak; inner metacarpal tubercle, round, not protuberant, same size or slightly larger than outer metacarpal tubercle; midventral ridge incorporating subarticular tubercles of various sizes on each finger, no distinct tubercle at base of digits; webbing on toes basal; inner metatarsal tubercle large, slightly more than half of length of first toe, outer metatarsal tubercle absent; subarticular tubercles on toes 3, 4 and 5; one subarticular tubercle on second toe, none on first; heels do not meet when legs are at right angles to body. Skin on dorsum smooth with microscopic network of ridges that are evident around the sacral region and less evident on head and limbs (in preservation); flanks, belly, chest, and throat, with small white pustules; ventral surface of legs smooth with similar pustules adjacent to vent. Femoral gland distinct, its distance to vent is twice that to knee; chest gland immediately behind insertion of arm; area between jaw articulation and eye with minute tubercles. Coloration in life (based on specimens from Tam Dao). Dorsum and limbs chocolate-brown. Irregularly shaped orange spots on back and limbs; legs with alternating orange and black bands on the inferior surface of the tibiotarsus; feet, ventral side of arms, and posterior thighs with black and white mottling; groin black with small, white spots that are more concentrated on the chest and throat, chest and throat predominantly white with some black mottling. Hands and feet gray, including tubercles; tips of digits white. Dark patch on canthus, just behind nares, and one at posterior corner of eye; lower edge of supratyrnpanic fold bordered by dark line; margin around eyelid black; nictatating membrane dark; dorsal half of iris white. Coloration in preservative. Dorsum dark brown with tan spots on back and limbs; ventral color pattern does not change in preservative. An older specimen (KIZ ) in the collections of the Kunming Institute of Zoology, Yunnan Province, China has retained a similar color pattern, but is slightly faded. The dorsurn is tan with some dark spotting; venter lighter tan, with light-brown mottling. Banding on the legs dark brown. Distribution and ecology. The type material is known from Sa Pa (Chapa), Lao Cai Province (22'22' N, 103'52' E), which is about 30 km south of the Chinese border. Seven specimens, 4 males, ROM ; and 3 females, ROM , , IEBR specimen bearing ROM field number and three specimens, 2 adult females and 1 adult male (MVZ ; 2398 I), were collected in Tam Dao, Vinh Phu Province (21 27'N, 105'38' E), approximately 250 km SE of Sa Pa at an elevation approximately 400 m. Yang (1991) has records for ten individuals collected from five localities in central and southern Yunnan: Tengchong (25'0' N, 98'5' E); Menglian (24'5' N, 98'8' E); Mengla (21'4' E, 101 '5' E); Luchun (23'0' N, 102'4' E); Jingdong (24'4' N, 100'8' E) from elevations between m. Five adult frogs (two males and three' females) collected in Tam Dao on 4 May 1996 were found in reproductive period in the rapid mountain stream, and one couple was observed in amplexus (Fig. 2). This stream full down the steep mountain slope covered by the rain polydominant forest (Fig. 4). Remarks. Bourret (1937, 1942) considered this species to be a subspecies of L. hasseltii. He distinguished it from L. hasseltii hasseltii by relative length of the 3rd finger, greater interorbital space, and the condition and color of the metatarsal tubercle. It was distinguished from L. pullum by the size of the metatarsal tubercle and color pattern. In addition, the white color of the dorsal part of the eye distinguishes it from these two species, both of which have a scarlet-colored dorsal part of the eye. Although there appears to be some variation in the eye color for populations of L. chapaense, the eye color is never scarlet. Yang ( : p. 69) reports that the dorsal portion of the iris is blue. This may be similar to the very dark or black iris described by Bourret (1937,1942). In the live specimens from Tam Dao, the dorsai half of the iris is strikingly white, and very distinguishable from the rest of the eye. This is evident even in our preserved specimens, as they match very well in all other respects to the description and illustration of L. chapaense by Bourret (1942).

7 Vietnamese Leptobrachium 57 Leptobrachium xanthospilum (sp. nov.) Figs. 5,6 Holotype. ROM , an adult female from the forested hills about 2 krn east a farming village (14O20'29" N, 108O28'46" E elevation 850 m), Gia Lia Province, Vietnam, collected on 25 September 1997 by local Ba Na hunters. Paratypes. ROM (female) collected at the same time and in the vicinity of the holotype; ROM (males) from Tram Lap (14O26'24" N, 108O32'58" E elevation 900 m), Gia Lia Province, Vietnam (approximately 35 km NE [by road] of the type locality) collected by N. L. Orlov, R. W. Murphy, and R. Bain on June Diagnosis. A large-sized Leptobrachium (two females mm SVL, eight males rnrn), with long legs, heels overlap when legs are at right angles to body; toes 1/3 webbed. Dorsal color pattern light brown to dark chocolate brown; flanks, posteroventral surface of legs; and sometimes chin with distinct large yellow spots; limbs with or without distinct dark bands, venter uniformly dark with minute white spots on chest, throat and chin. Chest gland located in armpit, behind insertion of arm. Dorsal half of iris white. This large Leptobrachium can be distinguished from all other species of Leptobrachium by the distinct, large yellow (creamy-white in preservation) spots on the flanks, posterior legs, and sometimes on the chin. Furthermore, the uniformly dark venter with scattered white punctuation (Fig. 3) distinguishes it from L. nigrops, L. chapaense, L. hasseltii, and L. hendrichoni (syn. of hasseltii; Inger, 1966) that are not uniformly dark, but predominantly either white marbled or speckled with darker pigmentation. In life (sometimes visible in preserved specimens) the white dorsal part of the iris easily distinguishes it from L. hasseltii and L. pullum that are said to be scarlet (Beny and Hendrickson, 1963: p. 61 8; Smith, : p. 440) and L. nigrops that has a uniformly black iris. The long legs of L. xanthospilum, where the heels broadly overlap when at right angles to the body, distinguishes it from all species ofleptobrachium except Leptobrachium nigrops. Although the latter species has been reported to have relatively short legs (Berry and Hendrickson, 1963: p. 644), it is unclear if this means the heels do not contact when drawn in toward the body. Description of holotype. Body long (SVL 84.8 rnm) head 39% SVL; head large, width equal to length, as broad as body; interorbital and frontal areas flat and sloping ventrally near snout; head dorsoventrally compressed; canthus rostralis rounded, loreal region oblique, slightly concave; nostrils to tip of snout 1/3 distance of nostril to eye; snout in dorsal view rounded, in lateral view sloped. Eyes large and protuberant, diameter greater than snout length; tympanum large, 63% of eye diameter, and separated from eye by a distance 84% of tympanum diameter; supratympanic fold extends from rear comer of eye to articulation of jaw; interorbital distance 55% of head width; posterior 1/2 of tongue free, distinctly notched; vomerine teeth absent; choanae separated by a distance nearly 1.5 times internasal distance. Fingers long (23% SVL), tips not wider than digits; first finger longer than second, latter equal to fourth; webbing absent, lateral fringes weak; inner metacarpal tubercle, oval, protuberant, outer metacarpal tubercle similar shape as inner, but slightly smaller; midventral ridge of subarticular tubercles of various sizes on each finger, no distinct tubercle at base of digits; toes about 1/3 webbed ( IV V); inner metatarsal tubercle large, length equal to first toe, outer metatarsal tubercle absent; subarticular tubercles like those of hand on toes 3, 4, and 5, one on second toe, none on first; heels broadly overlap when legs are at right angles to body. Skin on dorsurn, smooth with microscopic network of ridges that extend onto dorsal surface of limbs and ventral surface of arms; belly, chest, throat, and area adjacent to vent with small pustules. Femoral gland, indistinct, midway between knee and vent; chest gland immediately behind insertion of arm. Color in life. Dorsum and limbs dark brown; legs with darker bands on the posterodorsal surface of thigh and tibiotarsus; flanks with 7-8 large, well-defined yellow spots; venter uniformly dark brown-lavender with evenly dispersed light-brown pustules on belly, chest, and adjacent to vent; more concentrated pustules on throat and chin. Tips of digits and tubercles on hands and feet white. Margin of eyelid partially yellow. Dorsal half of iris white. Color in preservative. Dorsum dark brown with microscopic network of ridges lighter brown; large spots on flanks, chest and chin, creamy-white. Limbs light brown with dark brown bands. Venter brown

8 58 Amy Lathrop dt al. with very small light pustules. Toe tips, metatarsal, and metacarpal tubercles creamy white. Variation. Dorsal color pattern is either dark purple-black (specimens from Krong Pa and the majority of those from Tram Lap) or brown (Tram Lap ROM 32187) with darker diffuse mottling. Venter in all specimens uniformly one color, either very dark (Krong Pa) or lighter brown (Tram Lap). In the lighter colored specimens, the granular nature of the throat and chin can be more easily detected because the granules show up as creamy-white. Two Tram Lap specimens have yellow dots on the chin and chest. Axillary gland more distinct in male Tram Lap specimens. However, all males are from Tram lap and all. females are from Krong Pa and, thus, this may reflect an artifact of collecting. Arms of males are thicker and more stout; females have longer, weaker arms. Banding on arms distinctly obvious in the light brown specimen from Tram Lap, and less distinct in other Tram Lap specimens, and very difficult to see in the dark females from Krong Pa. Interorbital bar on the one light brown specimen from Tram Lap. Some yellow spotting may occur on the head and around the edge of the upper eyelid. Remarks. Two gravid females were collected in late September. As well, two metamorphs (ROM ) (Fig. 7), Gosner (1960) Stage 46, were collected in late June from the same stream where the adults were collected. We are unsure whether they are L. xanthospilum or the other sympatric species, described below. Nonetheless, if these are L. xanthospilum and gravid females are found in September, this suggests that either the L. xanthospilum have an extended larval period lasting nearly nine months, or that the breeding season is long. A similar finding was reported by Pope (1 93 1) for Chinese L. hasseltii in which larvae were found in January, April and July. No conclusions can be drawn until additional collections can be made at different times of the year. Besides the localities noted above, we have observed this species in the vicinity of Buoenloy village (14"20f38" N, 108O36'18" E, elevation 700 m), Gia Lai Province. One adult male was found in October 1993 during the rainy night on the leaf litter in 100 m out from the stream; 26 metamorphs were collected in March- beginning of May 1995 in small mountain stream flowing into Cha River (tributary of Ba River), In the latter locality the tadpoles on the latest develop- mental stages and three couples in amplexus (Fig. 6) were noted on 6-10 May, i.e., in the beginning of the rainy season. Stony streams with Leptobrachium xanthospilum filled by the deciduous leaves, flow in the aerial layered polydominant rain forest; the canopy above the streams is closed (Fig. 8). Etymology. The specific name is derived from the Greekxanthos meaning yellow and the Greek spi- 10s meaning spot. The name refers to the conspicuous yellow spots this frogs bears. Leptobmchium banae (sp. nov.) Fig. 9 Holotype. ROM 32200, an adult female from the forested hills about 2 km east of a farming village, Gia Lia Province, Vietnam (14 20'29" N, 108'28'46" E, elevation 850 m) collected on 2 1 September 1997 by local Ba Na hunters. Paratypes. ROM , (males) and ROM (female) from vicinity of the type locality, collected by N. Orlov, R. Murphy, L. Lowcock, A. Lathrop, and local hunters between June Diagnosis. A large-sized Leptobrachium (two females, and 17 males, mrn) with short legs, heels do not overlap when legs are at right angles to body; basal webbing on toes. Dorsal color pattern black with small red-orange diffuse blotches near groin, flanks and on back; forelimbs and hind limbs with alternating red (sometimes red-orange) and black bands; ventral surface of limbs and flanks black, groin and belly brown with white pustules that extend up to chin. Dorsal half of iris white with a strikingly white nictitating membrane around the eye. Leptobrachium banae has distinct red (gray in preservative) bands on the limbs that distinguish it from L. hendricboni which has no bands on the limbs. The strikingly white porti.on of the iris distinguishes it from all species ofleptobrachium except L. xanthospilum, and some forms of L. chapaense. From these two, it can be identified by its ventral color pattern, as illustrated in Fig. 3. The venter of L. banae is dark with some white mottling and spotting, while L. chapaense is dark with distinct mottling and L. xanthospilum is entirely dark with indistinct spotting. Description of holotype. Body long, head 37% SVL; head large, width slightly greater than length, nearly as broad as body; interorbital and frontal areas

9 Vietnamese Leptobrachium flat and sloping ventrally near the snout; head dorsoventrally compressed; canthus rostralis rounded, loreal region oblique, slightly concave; nostrils half the distance to tip of snout than to eyes; snout in dorsal view rounded, in lateral view sloped. Eyes large and protuberant, eye equal to snout length; tympanum large, 57% of eye diameter, and separated from eye by a distance 81% of tympanum diameter; supratympanic fold extends from rear corner of eye to articulation of jaw; interorbital distance 47% of head width; posterior 3/4 of tongue free, distinctly notched; vomerine teeth absent; choanae separated by a distance slightly more than internasal distance. Fingers long (2 1 % SVL), tips not wider than digits; first finger shorter than second, latter equal to fourth; webbing absent, lateral fringes weak; inner metacarpal tubercle, oval, protuberant, outer metacarpal tubercle similar shape as inner, but slightly smaller; midventral ridge of subarticular tubercles of various sizes on each finger, no distinct tubercle at base of digits; toes 1/3 webbed ( IV4-2V); inner metatarsal tubercle large, length equal to or slightly larger than first toe, outer metatarsal tubercle absent; subarticular tubercles like those of hand on toes 3,4 and 5, two tubercles on second toe, one on first; heels do not overlap when legs at right angles to body. Skin on dorsum smooth with microscopic network of ridges that extend to a lesser extent onto dorsal surface of limbs; flanks, belly, chest and throat with small pustules; ventral surface of legs smooth, sparse pustules on the thighs that extend upward to vent. Femoral gland distinct, more than 50% closer to knee than to vent; chest gland immediately behind insertion of arm; area between jaw articulation and eye with minute tubercles. Color in life. Dorsum and limbs dark brown with red-orange spots on the back and limbs; legs and arms with bright red and black bands; ventral side of arms, legs and belly black, the latter with small pustules that are more or less evenly distributed throughout. Hands and feet gray, including tubercles; tips of digits white. Nictatating membrane white; dorsal half of iris white. Color in preservative: Dorsurn dark brown-black. Orange spots turn cloudy gray in preservative, red stripes on limbs white or gray in preservative; venter dark brown, pustules on belly white. Variation. Among the 19 specimens in the type series, some variation is present in the dorsal color pattern. Almost half the specimens have a uniformly colored dorsum, brown (black in preservative) with only a few spots of red (light gray in preservative) and mottling dorsolaterally. Three specimens are much lighter brown (gray in preservative) with dark mottling, including an interorbital triangle. The remaining eight specimens have a moderate amount of dark mottling on the dorsum with no interorbital triangle. Banding pattern on the limbs is present in all specimens, but more obvious in some, and not consistent with dorsal color patterns. Ventral color pattern is consistent in all specimens. Remarks. Our field work in Krong Pa coincided with the beginning of the rainy season when it rained everyday between September, Following a few days of the heavy rain, choruses of about 5-10 male L. banae could be heard in the distance. Several adults were collected from a forest in which all of the understory had been cut and largely removed by local Vietnamese farmers. Individual frogs were collected near a foot path up to 2 krn from the nearest stream. One male was located by its call, where it was in a hole, buried in mud and leaf litter along with a Bufo galeatus. The call, repeated about every minute, consisted of two loud "crock... crock" followed by 5 to 8 more staccato-like crocks. Etymology. The specific epitaph is a patronym from the word Ba Na which is the name of the ethnic minority people in the area where this species was collected. Their invaluable assistance allowed us to document the herpetofauna of Krong Pa. KEY TO THE SPECIES OF LEPTOBRACHIUM FROM INDOCHINA I. Large conspicuous yellow spots along sides No large yellow spots on flanks 2. Red or orange spots on legs and or flanks Red spots absent on legs and or flanks 3. Ventral surface of tibiotarsus dark with white marbling; iris white, blue or black Ventral surface of tibiotarsus entirely black with no marbling; iris and palpebral membrane white 4. Snout acuminate in dorsal view; iris black Snout rounded in dorsal view, spots on dorsum present or absent; iris scarlet ACKNOWLEDGMENTS L. xan/hospilum L. chapaense L. banae L. nigmps L. pullurn or L. hassel~ii Collecting and export permits were made available through Vietnam's Institute of Ecology and Bio-

10 60 Amy Lathrop et al. logical Resources (IEBR), Hanoi. Import permits for frozen tissues and preserved specimens were issued by Agriculture Canada and all collecting and euthanasia of specimens were performed under approved animal welfare protocols. This study was supported by the Natural Sciences and Research Council (NSERC) of Canada grant A3 148, by the generous assistance of the ROM Sciences Fieldwork Fund, the ROM Future Fund, the ROM Foundation, and the Department of ROM Volunteers. We are extremely grateful to Prof. Cao Van Sung, Director of IEBR, for assist in many ways with on-site arrangements. Critical translations of Chinese descriptions were provided by J. Fu. Dr. Nguyen Van Sang and Pham Duc Tien provided critical assistance in Our heartfelt thanks go to Chief Bou Neu and the Ba Na Minority families from his village. Cathay Pacific Airlines graciously contributed to ow efforts by providing free excess baggage during international travels. Magnalight, Cole man, Johnson Wax, Tilley Endurables, and Benjamin Film contributed significantly to the success of our Vietnam biodiversity project. This is contribution 1 17 from the Centre for Biodiversity and Conservation Biology, ROM. REFERENCES Berry P. Y. and Hendrickson J. R. (1963), "Leptobrachium nigrops, a new pelobatid frog from the Malay Peninsula, with remarks on the genus Leptohrachium in southeastern Asia," Copeia, 1963, Bourret R. (1935), 'Wotes 13erpctologiques sur 1'Indochine Franqaise. X. Les Serpents de la station d'altitude du Tam-dao," Bull. GPn. Instx Publi'ue, 1835, Bourret R. (1937), "Notes HerpCtologiques sur 1'Indochine.Franqaise. XIV. Les Batrachiens de la Collection du Laboratoire des Sciences naturelles de 1'Universit6. Descriptions de quinze especes ou variktb nouvelles. XV. LBzards et Serpents requs au Laboratoire des Sciences naturelles de 1'UniversitC au cours de l'ann6e Descriptions de deux especes et de deux vari6tes nouvelles," Annexe Bull. G6n. Instl: Publique, 1937,l- 80. Bourret R. (1942), "Les Batraciens de l'indochine," Mkm. Inst. Ockanogr: I %rdochine, 6. Dubois A. (1980), "Notes sur la Systematique et la Repartition des Amphibiens Anoures de Chine et des Regions Avoisinantes. IV. Classification Generique et Subgenerique des Pelobatidae Megophryinae," Bull. Mens. Soc. Linn. Lyon, 49, Dubois A. (1983), "Note Preliminaire sur le Genre Leptopalax Dubois, 1980 (Amphibiens, Anoures), avec Diagnose d'une Esp6ce Nouvclle du Vietnam," Alytes, 2, Gosner K. L. (1960), "A simplified table for staging anuran embryos and larvae with notes on identification," Herpetologica, 16, Inger R. F. (1954), "Systematics and zoogeography of Philippine amphibia," Fieldiana Zool., 33, 18 l - 53 l. Inger R. F. (1966), "The systematics and zoogeography of the Amphibia of Borneo," Fieldiana Zool., 52, Lathrop A., Murphy R. W., Orlov N., Ho C. T. (in press), "Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam," Amphibia-Reptilia. Myers C. W. and Duellman W. E. (1982), "A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama," Am. Mus. Novit., 2752, Pope C. H. (1931), "Notes on amphibians from Fukien, Hainan, and other parts of China," Bull. Am. Mus, Nut. Hist.,61, Smith M. A. (1921), "New or little-known reptiles and batrachians from southern Annam (Indo-China)," Proc. Zool. Soc. London, 1921, Taylor E. H. (1962), "The amphibian fauna of Thailand," Univ. Kansas Sci. Bull., 43, Yang D. (1991), Amphibian-Fauna of Yunnan, China Forestry Press, Beijing [in Chinese]. Ye C., Fei L., and Hu S. (1993), Rare and Economic Amphibians of China, Sichuan Science and Technology Publishing House, Chengdu [in Chinese].