UNIVERSITY OF ILLINOIS LIBRARY AT URBAiMA-CHAMPAIGN BIOLOGY 3 **g o 1 nnnn f s 2 a C/5 m C/3 > 3 co -h z m en o o 00 o O CD S § nnnn TABLE CONT. INDEX FRONT COVER BACK COVER ADS —1 —I (— m 5 CD m XI m O o o > f— i— ■< 3D NAME — University of Illinois/Ur Library Binding Division 1408 W. Gregory Drive Urbana, IL 619 ?1 5 ty z •tj be ro £ H ID O O O O o - 1 si o p 2 I -n 3D m o c: m z o H S 2 Mm I5 d N S TJ T3 ^ O O § o o g m m ^ ■ — ' — o 5 P < =5 5 ™ m — i 35 33 m o & m □ CD o o O z -J o O CO 1 i £ £ i s s ° 2 ^ nn Z -D m m «I •H ' 1 s m j» z CD m cn □ nnnn CO J? 5 1 0 a 01 t- 1-1 1-* tb f CD U) O CTI ID -e — U CO H g 52 y 0 CD CO 1 to Cn 3 U — m O tn 2 H ET O io Q • u> 0 CO CO t* CO 1 Q i to tn lO ID H* cn H c tJ TVJ 0 IC » ■ ca 3T 1 II 3 ct nn en co a n r Zoology i Orlov ogs of Vietnam: Report on New Collections ivricrs.-m FM^AM Zoology NEW SERIES, NO. 92 Frogs of Vietnam: X Report on New Collections Robert F. Inger Zurator Emeritus department of Zoology 7ield Museum of Natural History Xoosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 J.S.A. Nikolai Orlov lya Darevsky department of Herpetology and Ornithology oological Institute ussian Academy of Sciences 19034, St. Petersburg ccepted August 1, 1997 ublished January 29, 1999 ublication 1498 'UBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1999 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Contents Abstract Introduction General Environment Methods 1 1 1 3 Species Accounts 3 Megophryidae 3 Brachytarsophrys carinensis (Boulenger) .... 3 Megophrys feae Boulenger 4 Megophrys lateralis (Anderson) 4 Megophrys palpebralespinosa Bourret 4 Leptobrachium pullum (Smith) 4 Leptolalax tuberosus, new species 5 Leptolalax pelodytoides (Boulenger) 7 Ophryophryne Boulenger 7 Ophryophryne microstoma Boulenger 8 Ophryophryne pachyproctus Kou 8 Ophrynophryne poilani Bourret 9 BUFONIDAE 10 Bufo galeatus Gunther 10 Bufo macrotis Boulenger 10 MlCROHYLIDAE 11 Calluella guttulata (Blyth) 11 Microhyla annamensis Smith 11 Microhyla berdmorei (Blyth) 11 Ranidae 12 Amolops spinapectoralis, new species 12 Amolops ricketti (Boulenger) 13 Huia nasica (Boulenger) 13 Occidozyga martens ii (Peters) 14 Rana attigua, new species 14 I Rana cf. blythii Boulenger 16 Rana johnsi Smith 18 Rana kuhlii Tschudi 19 Rana livida (Blyth) 19 Rana maosonensis (Bourret) 20 Rana milleti Smith 20 Rana montivaga Smith 21 Rana nigrovittata (Blyth) 21 Rana taipehensis Van Denburgh 22 Paa verrucospinosa (Bourret) 22 HACOPHORIDAE 23 Chirixalus doriae Boulenger 23 Chirixalus nongkhorensis (Cochran) 23 Chirixalus palpebralis (Smith) 23 Chirixalus vittatus (Boulenger) 24 Polypedates leucomystax (Gravenhorst) .... 24 Philautus abditus, new species 26 Philautus parvulus (Boulenger) 28 Philautus maosonensis Bourret 28 Rhacophorus annamensis Smith 29 Rhacophorus baliogaster, new species 30 Rhacophorus bimaculatus (Peters) 33 Rhacophorus bipunctatus Ahl 33 Rhacophorus calcaneus Smith 34 Rhacophorus exechopygus, new species ... 35 Rhacophorus reinwardtii (Schlegel) 38 Rhacophorus verrucosus Boulenger 39 Theloderma corticalis (Boulenger) 40 Theloderma gordoni Taylor 40 Theloderma stellatum Taylor 42 Species doubtfully reported from Viet- nam 43 Acknowledgments 44 Literature Cited 44 List of Illustrations 1 . Map of Vietnam 2 2. Leptolalax tuberosus, new species 6 3. Bufo galeatus Gunther 10 4. Amolops spinapectoralis, new species ... 12 5. Rana attigua, new species 15 6. Rana cf. blythii Boulenger 17 7. Rana johnsi Smith 18 8. Rana nigrovittata (Blyth) 22 9. Philautus abditus, new species 27 10. Rhacophorus annamensis Smith 29 1 1 . Rhacophorus baliogaster, new species .... 31 12. Rhacophorus bipunctatus Ahl 33 13. Rhacophorus calcaneus Smith 35 14. Rhacophorus exechopygus, new spe- cies 36 15. Rhacophorus exechopygus, new spe- cies 37 16. Rhacophorus reinwardtii (Schlegel) 38 17. Theloderma corticalis (Boulenger) 41 18. Theloderma stellatum Taylor 42 List of Tables 1. Comparison of coloration in samples of Leptobrachium from Vietnam and Thailand 5 2. Comparison of samples of male Lep- tobrachium from Vietnam and Thai- land 5 3. Comparison of males from popula- tions of Leptolalax pelodytoides 8 4. Sexual dimorphism in body propor- tions of Ophryophryne poilani 9 5. Comparison of Calluella guttulata from Buon Luoi and C. yunnanensis 11 6. Comparison of Rana attigua with sim- ilar species from Southeast Asia 16 7. Comparison of Rana cf. blythii from Vietnam with R. blythii from the Ma- lay Peninsula 17 8. Characters of males from several sam- ples of frogs related to Rana sauteri Boulenger 1 9. Comparison of striped and nonstriped forms of Polypedates leucomystax from Buon Luoi, Vietnam 2 10. Sexual dimorphism of body propor- tions in Rhacophorus annamensis Smith 3 1 1 . Comparison of Rhacophorus verruco- sus and related forms 3 Frogs of Vietnam: A Report on New Collections Robert F. Inger Nikolai Orlov Ilya Darevsky Abstract Bourret's (1942) review of the amphibians of Vietnam was based on collections made in a very few areas of the country. Since that date little has been published on that fauna. We report here on large samples of anurans from central and northern Vietnam obtained by two of us (NO, ID); these samples comprise 60 species, of which six are new and 12 are not previously reported from Vietnam. With this new material the size of the known anuran fauna of Vietnam is increased from 82 to 100 species. Introduction Despite the thorough review of the amphibian :auna of Indochina by Bourret (1942), the content ind distribution of this fauna are still very imper- :ectly known. Bourret's analysis of the anurans of Vietnam, the focal area of the present paper, was >ased primarily on three sets of data: (1) obser- /ations made by himself and associates in the nountainous northern part of the country, (2) col- ections made by Smith (1921, 1922, 1924) in the southern part of the central highlands of Vietnam see map, Fig. 1), and (3) miscellaneous records jkccumulated in the nineteenth century and the first ade of the twentieth century. The last data set vas heavily weighted with species characteristic >f agricultural and other severely disturbed envi- onments (see below). Bourret (1942) reported Vietnamese localities as Tonkin, Annam, and Cochinchine [Cochin :hina]) for 78 species. Dubois (1983, 1987) add- d two new species, and two more are being de- cribed by Lathrop et al. (in press). Papers on the istribution of this fauna have been published in Vietnam (see Literature Cited), but none of these apers has added to the number of species known rom Vietnam or amplified descriptions of poorly nown species. Vietnam consists of two primary physiographic reas: a narrow, mainly coastal zone of lowlands nd a larger core of mountainous terrain with el- vations up to 3143 m (Mt. Fan Si Pan). As re- cently as 1982, Vietnam had approximately 60,000 square kilometers of rain forest and mon- soon forest combined (Collins et al., 1991). An- nual rainfall varies from approximately 1 ,400 mm in the south to 2,700 mm in the north. Given the size of Vietnam (325, 360 km) and its physical attributes, one would expect appreciably more than 82 species of anurans. Recent intensive sampling by two of us (ID, NO) in central and northern Vietnam uncovered 12 species not previously reported from Vietnam and six species new to science, raising the total fauna to 100 species. A number of the other spe- cies found during this sampling have been de- scribed or discussed only once in the literature. The bulk of our sampling was carried out in the central highlands, about 300 km north of the area in which Smith worked (see above). Because so little is known of the amphibian fauna of Vietnam, we will report on all of the species included in these new samples. General Environment The bulk of the specimens on which this report is based come from an area about 40 km long on the Tay-Nguyen Plateau in south-central Vietnam (Fig. 1). The plateau has a north-south axis of about 400 km and mountain ridges running par- allel to that axis, with elevations varying from 500 IELDIANA: ZOOLOGY, N.S., NO. 92, JANUARY 29, 1999, PP. 1^6 23.5°N 1Q0°E CHINA Fig. 1. Map of Vietnam, showing localities of the most important samples. 1: An Khe District; 2: Tarn Dao; 3: Sui Kat; 4: Dalat; 5: Sa Pa; 6: Mao Son. to 2598 m (Mt. Ngoklin). The actual collecting sites were around two villages, Buon Luoi and Tram Lap, and the area of Kon Cha Rang. These sites are 20, 40, and 60 km, respectively, north- west of the town of Kannack (14°20'N, 108°36'E), in the An Khe District (Fig. 1, no. 1), Gia-Lai Province, Vietnam. The dry season in this region lasts from December to April and the rainy season from June to October. Buon Luoi (700-750 m), the locality for most of the specimens, has a well-developed humid ev- ergreen tropical forest, with a closed canopy at about 20-30 m and some emergents (mainly of the families Dipterocarpaceae, Leguminosae, and Moraceae) above 40 m. There are few deciduous trees. The principal stream is the Cha River, which has a moderately steep gradient, a rocky bottom, and several 2- to 4-m waterfalls. The width of the riverbed varies from 4 to 10 m, and the depth during the dry season varied from 30 to 70 cm. Five small perennial tributaries flow through the forest, and each is completely under the canopy (except for the lower end of one). Three of the streams have very steep, rocky banks, and four of them flow through rock outcrops. Their widths vary from 2-3 m to 4-6 m. At the end of the dr season, depths varied from 5 to 50 cm. During th rainy season, depths reached 1 m. Two of th streams have clay beds, with many thick patche of dead leaves; the others have mainly rocky bed? Two of the streams have widened, ponded area; one 10 X 50 m and the other 20 X 150 m. Falle trees block the current in places, and swampy ai eas have developed. Collecting was carried out i Buon Luoi in November 1993 and March-Ma 1995. Tram Lap (ca. 900 m), 20 km north of Buo Luoi, is surrounded by essentially the same kin of forest as Buon Loy, although there are mor deciduous trees. A large river (20-40 m) flow through the area. Portions of the forest have bee cut, and clearings reach the riverbanks in place; Three small forest streams were worked at thi locality; two 2-4 m wide and one 1-2 m wide All three are under canopy. Two flow out of swampy area, with vegetation mainly of Araceat Large tree trunks block the flow and, along on stream, form a swampy area with grassy vegeU tion 8-10 m wide. One stream has a silty bottor and a second has a sandy clay bed. The depth vary from 10 to 100 cm. Collecting was carrie out in November 1993 and April 1995. Kon Cha Rang (1000-1200 m), 20 km north west of Tram Lap, has much the same type c vegetation, but the forest is more intact. Th streams in this area of rough topography hav steep gradients. The major river is the Azun (15 20 m wide), which has a strong current over rocky bed, but wide pools with moderate curren Three small streams (3-10 m wide) on the le bank flow over basalt, have rocky cataracts an waterfalls, and in places flow through rocky car yons. Their beds are rocky. Depths in the dry sei son are 30-50 cm. Two streams on the right ban of the Azun River flow through karst. The large (8-15 m wide) of these two flows below the roc surface in places and has a broken canopy above Collecting was carried out in April 1995. Tarn Dao (21°26'N, 105°39'E) (Fig. 1, no. 2 lies in a mountainous area that includes karst fo mations. Most of the fieldwork was carried out ; 900-1100 m, where the dominant vegetation i humid broad-leaved evergreen forest. Most frog were collected along rocky, steep-gradiei streams 3-5 m wide. There were many cascade! and waterfalls, some as high as 20 m. In som' places the streams run under the surface rock. Th mean temperature in the winter (November: March) is 8°C; it is 20°C in the summer. Rainfa FIELDIANA: ZOOLOG^ from June through September averages 100 mm per month. Collecting was carried out in June 1995 and April-May 1996. Methods Most collecting was done at night between 1800 and 2300 hrs. Specimens were preserved in buffered 10% formalin and transferred to 70% ethanol when they reached museums in the United States. Calls were not recorded. Specimens were measured with electronic calipers. The following abbreviations are used: SVL, snout-vent length; T, tibia length with the hind limb flexed; HW, head width at the tympanum; HL, head length from the rear of the lower jaw to the tip of the snout; TY, horizontal diameter of tympanum; HB, headbody of tadpoles; HBL, headbody length. Measurements are given in millimeters (mm) un- less indicated otherwise. Comparisons of body proportions (e.g., T/SVL) within or between spe- cies were made with the Mann- Whitney [/-test. In synonymies we cite the original description, the source of the present binomial, and a reference to Bourret's (1942) monograph. Specimens were borrowed from The Natural History Museum, London (bmnh), and the Mu- seum of Comparative Zoology (mcz). Other ab- breviations used are fmnh (Field Museum of Nat- oral History), MAS (Malcolm A. Smith), mvz s'Museum of Vertebrate Zoology), and zmh (Zoo- ogisches Museum fur Hamburg). Species Accounts Ten species are characteristic of anthropogeni- :ally modified environments, and all have very vide distributions in Southeast Asia. We collected hem in villages, on roads, and in rice fields and ; )ther agricultural sites. These species (and their orresponding localities) in our samples are Bufo nelanostictus (Buon Luoi, Tram Lap, and Kon ?ha Ran), Kaloula pulchra (Buon Luoi), Micro- tia butleri (Buon Luoi), M. heymonsi (Buon ,uoi. Tram Lap, and Tarn Dao), M. ornata (Buon .uoi), M. pulchra (Tarn Dao), Micryletta inornata Buon Luoi), Hoplobatrachus rugulosa (Buon Aioi), Rana erythraea (Buon Luoi), and R. lim- ocharis (Buon Luoi, Tram Lap, and Kon Cha Lan). These species will not be dealt with further. Megophryidae Brachytarsophrys carinensis (Boulenger) Leptobrachium carinense Boulenger, 1889:748 — Karen Hills, Burma. Brachytarsophrys carinensis Tian and Hu, 1983:42. Megophrys carinensis Bourret, 1942:216. An adult female (SVL 1 18.4) and an adult male (SVL 1 16.8) from Buon Luoi both have a palpe- bral "horn" consisting of a single, broadly based, triangular projection. In this respect they differ from Boulenger's ( 1 908) description of frogs from Burma and from fmnh specimens from northern Thailand and Sichuan, in which the upper eyelid has two to four projecting papillae. In other char- acters, these Vietnamese frogs agree with other specimens. This is the first record of the species from Vietnam, but the range extension is not sur- prising, given the known distribution from Burma through northern Thailand to southern China. Boldly striped megophryid tadpoles with a fun- nel mouth collected at Buon Luoi may belong to this species or M. lateralis, two of which were also collected at Buon Luoi. Coloration of tadpoles in preservative: HB brown dorsally; a ventrolateral black stripe from oral funnel below eye to end of HB above limb buds; a yellow stripe of same width below black one; ventrally HB brown with irregular yellowish spots and a midventral stripe in rear two-thirds; a large yellowish spot below oral funnel; amount of yellowish spotting ventrally varies. Tail muscle mottled brown in upper half, bordered below by narrow dark brown stripe, a wider yellow stripe, and a wider dark stripe; all stripes merge into dark brown coloration of rear two-fifths of tail; dorsal fin mottled brown anteriorly, ventral fin unpig- mented except for several dark marginal spots in proximal half; both fins heavily mottled in distal two-fifths. Mfasurfmrnts — Total length 40.4 (stage 27), 44.0-49.7 (stages 36-37); HBL 12.5 (stage 27), 13.5-15.0 (stages 36-37); tail depth/tail length 0.30 (stage 27), 0.23-0.25 (stages 36-37). This is the first longitudinally striped larval megophryid. Liu and Hu (1961, PI. 14) figured a larval Megophrys that has transverse light and dark banding ventrally and a dark midlateral stripe on the proximal half of the caudal muscle. Al- though Liu and Hu (1961) identified this tadpole VGER ET AL.: FROGS OF VIETNAM as M. carinensis, they did not explain why they assigned it to this species. Megophrys feae Boulenger Megalophrys feae Boulenger, 1887a:512 — east of Bhamo, Khakhien Hills, Burma. Megophrys feae Bourret, 1942:218. A female (SVL 84.0) and two juveniles (SVL 17.2 and 34.0) were collected at Tarn Dao. The tibia of the female (T/SVL 0.40) is slightly longer than head length (HL/SVL 0.38) and shorter than head width (HW/SVL 0.46). The species was re- corded from Tarn Dao by Bourret (1942) and from other areas in northern Vietnam by Tran et al. (1981). Although it was not reported from Yunnan by Yang et al. (1991), the species probably occurs there. Megophrys lateralis (Anderson) Ixalus lateralis Anderson, 1871:29 — Yunnan, China. Megophrys lateralis Liu, 1950:180. Megophrys longipes maosonensis Bourret, 1937:12. Two subadult females (SVL 54.0-54.5) were collected at Buon Luoi and 1 1 adults and one ju- venile at Tarn Dao. These specimens do not differ from descriptions of M. lateralis (Boulenger, 1908; Bourret, 1942; both under the name M. ma- jor Boulenger). Buon Luoi represents an exten- sion of the range of this species southward to cen- tral Vietnam. SVL measurements of the Tarn Dao specimens: females 83.1-91.7 (mean ± standard error [SE] 87.83 ± 1.00, N = 9), males 67.9-75.3 (N = 2); T/SVL 0.48-0.56 (median 0.514, N = 1 1); HW/SVL 0.35-0.39 (median 0.366, N = 11); HL/SVL 0.34-0.40 (median 0.363, N = 11). The ratio HL/SVL in the two males (0.38-0.40) is greater than that in any of the females. There are no obvious differences between the sexes in HW/ SVL or in tibia length. Megophrys palpebralespinosa Bourret Megophrys palpebralespinosa Bourret, 1937:16— Chapa, Tonkin, Vietnam. An adult male (SVL 40.9) collected at Tarn Da agrees with Bourret's (1937) description. Th specimen has brown nuptial pads on the first an second fingers; several short tubercles on the u[ per eyelid and a thicker, triangular one projectin from the edge of the eyelid; an obtusely pointe but strongly projecting snout; large, irregularl shaped dark blotches ventrally separated by na row lighter areas; and distinct, shallow webbin at the base of the toes. T/SVL 0.44, HW/SV 0.33, HL/SVL 0.32. The species is as yet know only from the mountains of northern Vietnai (Tran et al., 1981) and southern Yunnan, Chin (Yang et al., 1991). Leptobrachium pullum (Smith) Megophrys hasseltii var. pullum Smith, 1921 440 — Arbre Broye [Lang Biang], Annan Vietnam; Bourret, 1942:214. Leptobrachium pullus Dubois, 1980:476. Variation among populations of Leptobrachiw in Southeast Asia is known to be complex (Inge 1983), and the taxonomy is unsettled. Two form have been described from Vietnam: L. pullut Smith (type locality 11°54'N, 108°19'E) and I chapaense Bourret (type locality Chapa, Tonki 22°24'N, 103°51'E). Both were treated as varietie of L. hasseltii by the authors. These differ in co oration and in body proportions (Tables 1 and 2 Four adults (three males, one female) from Buo Luoi, approximately 270 km from the type loca ity of L. pullum and 990 km from that of L. chc paense, differ from both forms in coloration an relative head length (Tables 1 and 2). Males fror Buon Luoi are larger than either L. pullum or I chapaense (Table 1). Three newly metamoi phosed juveniles from Buon Luoi, with complete ly resorbed tails but remnants of the operculur still visible, measure 42.2-42.7. These juvenile would almost certainly have achieved sexual ma turity at larger sizes than the males of L. pullut and L. chapaense. Smith (1921) reported the SV] of three females of L. pullum as 44-52; the fou largest females in the type series of L. chapaens were 58-62 (Bourret, 1937); the sole female i the Buon Luoi sample is 64.7 mm. We have also examined specimens of this grou from two localities in northern Thailand — Doi In thanon (18°32'N, 98°32'E) (fmnh 187439-41 and Chiang Dao (19°21'N, 98°59'E) (fmn) 172661, 173973-80). These samples, separated b FIELDIANA: ZOOLOG^ Table 1. Comparison of coloration in samples of Leptobrachium from Vietnam and Thailand. Vietnam Thailand L. pul- lum L. chap- Buon aense Luoi Doi Chi- Inth- ang anon Dao Back* Chest and bellyt Sidest Leg ventrally§ 0 0 4 1 2 1,3 3 3 1 2 1 2 1 4 2 1 1 3 1 1 * Back: 0 = indistinct darker markings; 1 = vermi- form black markings; 2 = black network; 3 = black and yellow speckling; 4 = large round black spots. t Chest and belly: 0 = no spots; 1 = white with large black spots; 2 = dotted light and dark; 3 = black with fine light spots. t Sides: 0 = no spots; 1 = small light spots; 2 = large (half the diameter of the eye) light spots; 3 = large black spots; 4 = small black spots. § Leg ventrally: 0 = white, without spots; 1 = black- ish with light spots; 2 = blackish without spots. about 100 km, differ from one another and from all three of the Vietnamese samples (Tables 1 and 2). The frogs from these five samples represent either one species with great geographic variation or five species. We believe the latter is probably correct, but we adopt a conservative approach and group them all under the name L. pull um pending an analysis involving more samples. The Vietnamese collection also includes one ju- venile from Tarn Dao with SVL 32.6, T/SVL 0.44, and HW/SVL 0.39. It has conspicuous pectoral glands but no opercular scars, thus showing it to )e more advanced than the three juveniles from Buon Luoi. It matches the coloration of L. pullum Table 1), but its body proportions, with tibia ength greater than head width, differ from those )f all of the samples listed in Table 2. Leptolalax tuberosus, new species (Fig. 2) Holotype — fmnh 252844, an adult male, col- lected at Kon Cha Rang Village (1000-1200 m above sea level [ASL]), An Khe District, Gia-Lai Province, Vietnam, 15 April 1995 by Ilya Dar- evsky and Nikolai Orlov. Paratypes — fmnh 252845-59, 14 males and one female from type locality; fmnh 252860, one male from Tram-Lap Village (900 m). An Khe District, Gia-Lai Province, Vietnam. Etymology — Specific name from tuberosus L., meaning full of protuberances. Diagnosis — A small species of Leptolalax, males < 30 mm, female 30 mm; tympanum ob- scured by skin; low, rounded tubercles on snout and eyelids; no ventrolateral glandular ridge; ven- tral surface of thigh with thin black reticulation enclosing round, white spots. Description — Habitus stocky; head width equals length. Snout obtusely pointed, rounded or truncate in profile, not projecting; nostril a bit closer to tip of snout than to eye; canthi rounded, constricted; lores sloping, concave; diameter of eye greater than length of snout; interorbital equal to or narrower than eyelid; tympanum obscured by skin, rim not visible; no vomerine teeth. Tips of fingers blunt; first finger subequal to second; fourth and second fingers equal; subarti- cular tubercles obscured; a large, round inner met- acarpal tubercle, not extending out under first fin- ger; a smaller outer metacarpal tubercle. Tips of toes like fingers; third toe longer than fifth; web- bing confined to bases of toes; toes without lateral fringes; subarticular tubercles replaced by keratin- ous dermal ridges, a relatively short one on each of first two toes, longer ones on third and fifth toes, ridge covering entire underside of fourth toe; Table 2. Comparison of samples of male Leptobrachium from Vietnam and Thailand. Data on L pullum from Jmith (1921); data on L chapaense from Bourret (1937). 1 & : Vietnam Thailand L. pullum (N = 5) L. chapaense (N = 4) Buon Luoi (N = 3) Doi Inthanon Chiang Dao (N = 3) (N = 8) IVL (mm) | VSVL D IW/SVL jIL/SVL 44-49 0.33-0.35 0.41-0.44 0.33-0.36 46-53.5 0.36-0.41 0.45-0.46 0.37-0.40 60.3-64.7 0.36-0.39 0.45-0.47 0.41-0.45 43-47 43-56 0.37-0.41 0.32-0.33 0.43-0.47 0.39-0.43 0.44-0.49 0.38-0.42 1 SVL = snout-vent length; T = tibia length with hind limb flexed; HW = \ ead length from the rear of the lower jaw to the tip of the snout. = head width at the tympanum; HL = il MGER ET AL.: FROGS OF VIETNAM Fig. 2. Leptolalax tuberosus, new species. Holotype, male, 26.1 mm. a small, oval inner metatarsal tubercle, no outer one. Skin above with many scattered small tubercles of varying sizes; eyelid with low rounded tuber- cles; top of snout usually with elevated tubercles; sides and venter smooth; small, weak pectoral and femoral glands; a small, round, white gland just above axilla; no ventrolateral glandular ridge. Color in preservative of dorsal and lateral sur- faces dark gray or black, with obscure light speck- ling, heavier low on sides; usually two dark labial bars; limbs with darker crossbars; venter white, with thin, short black streaks sparse posteriorly, becoming denser anteriorly and on chin and throat forming black network enclosing round white spots; ventral surface of thigh and calf with dark network enclosing round white spots. Males with slit-like vocal sac openings, but no nuptial pads. Measurements — SVL of males 24.4-29.5 (mean ± SE 26.82 ± 0.42, N = 16), of single female 30.2. T/SVL 0.48-0.54 (median 0.500, N = 12); HW/SVL 0.35-0.39 (median 0.363, N = 12); HL/SVL 0.36-0.41 (median 0.381, N = 12). Measurements of holotype: SVL 26.1, T 13.: HW 9.9, HL 10.0, and diameter of eye 4.3. Comparisons — This is the only species of Lei tolalax that has whitish speckling dorsally and tt only one in which the tympanic rim is not clear! visible. In a few individuals of L. tuberosus th tympanum is evident as a smooth, ill-defined are; Leptolalax pelodytoides (Boulenger), from nortl ern Vietnam, Thailand, and Sichuan, China (to a< commodate the expanded definition of the specie by Dubois, 1983) differs from L. tuberosus in i superficial tympanum, low glandular ventrolater; ridge, shorter legs, narrower head, and later; fringes on the toes. Males of L. pelodytoides ha\ T/SVL 0.44-0.51 and HW/SVL 0.30-0.39; di ferences from those values for L. tuberosus ai statistically significant (P < 0.02, Mann-Whitne (/-test). Leptolalax pelodytoides usually has a di; tinct dark pattern on the dorsum and several di; tinct black spots on the sides, but no light spec! ling dorsally and no dark reticulation under th thighs; in each of these features of coloration, 1 pelodytoides differs from L. tuberosus. Bourret (1942: 208-209) listed and illustrate FIELDIANA: ZOOLOG specimens from Chapa, northern Vietnam, as L. pelodytoides. These became types of L. bourreti Dubois (1983); Bourret's drawings show a frog with a superficial tympanum, a distinct dark pat- tern on the back, and distinct black spots on the side, thus differing from L. tuberosus. Dubois (1983) gave the SVL of L. bourreti as 36.3 (male holotype) and 42-45 (female paratypes), i.e., larg- er than L. tuberosus. Two new species from north- ern Vietnam described by Lathrop et al. (in press), L sungi and L. nahangensis, differ from L. tub- erosus in having SVL of males >40 mm, visible tympana, and the undersides of the thighs immac- ulate white. Fei et al. (1990) gave a diagnosis (but no de- scription) for a new species, Leptolalax alpinus, emphasizing those characters that differed from L. pelodytoides; none of the reported differences be- tween L. alpinus and L. pelodytoides involved coloration. Fringes along the toes were explicitly mentioned as a common feature of these two taxa. Consequently, we believe that coloration and toe fringes distinguish L. alpinus from L. tuberosus. Leptolalax tuberosus is smaller than the Bor- nean species L. gracilis (Giinther) (males 31-39), L dringi Dubois (males 30-35), and L. pictus Malkmus (males 30-35). It also differs from L. pictus and L. heteropus (Boulenger) of Peninsular Malaysia in the absence of a clearly defined dorsal pattern, and from L. gracilis, L. dringi, and L. pictus in the absence of black spots laterally. All of the Bornean and Malaysian species mentioned have superficial tympana. Leptolalax pelodytoides (Boulenger) Leptobrachium pelodytoides Boulenger, 1893: 345 — Thao, Karin Bia-Po District, Burma. Leptolalax pelodytoides Dubois, 1983:149. Megophrys pelodytoides Bourret, 1942:208. A large sample of adults was obtained at Tarn Dao. Adult females range from 35.3 to 40.7 mm SVL (mean ± SE 38.26 ± 0.21, N = 22) and males from 27.7 to 30.9 (29.22 ± 0.21, N = 19). Females have slightly shorter legs (T/SVL 0.42- D.49, median 0.460, N = 22) and slightly shorter heads (HL/SVL 0.32-0.39, median 0.363, N = 22) Oian males (T/SVL 0.45-0.51, median 0.48, N = 19; HL/SVL 0.34-0.40, median 0.372, N = 17); differences between the sexes are statistically sig- nificant (in both ratios P < 0.03, Mann-Whitney [/-test). The sexes do not differ in relative head iNGER ET AL.: FROGS OF VIETNAM width (HW/SVL 0.30-0.35; median of males 0.332, of females 0.335). All individuals have black bars on the lip, black markings dorsally, and a broken line of white glands ventrolaterally. The belly is immaculate white or cream-colored (in preservative), and the chin and ventral surface of the calf are dusky with small whitish spots. These specimens are larger than L. pelodytoides from Thailand and Sichuan (including L. mini- mum Taylor and L. oshanensis Liu, to follow the taxonomic arrangement of Dubois, 1983), but are very similar in body proportions (Table 3). Fe- males from Tarn Dao (35.3-40.7) are also larger than those from Thailand and Sichuan (29.2- 35.9). There are slight differences in coloration. Short, dark, vermiform lines occur on the belly in 14 of 15 frogs from Sichuan, but on only four of 19 from Thailand and on none from Tarn Dao. The ventral surface of the thigh in Tarn Dao frogs has fine black network over white, becoming gradually denser toward the rear surface, which is quite dark. In most of those from Thailand ( 1 II 19) and Sichuan (14/15) there is an elongated blackish oval on the ventrolateral surface of the thigh, contrasting sharply with a much lighter area on the anterior half of the ventral surface of the thigh. Leptolalax bourreti Dubois from Chapa (= Sa Pa), northern Vietnam, is clearly larger (females 41-54) than L. pelodytoides from Tarn Dao (see above). The nine females of L. bourreti measured by Bourret (1937, as L. pelodytoides) have longer tibia (T/SVL 0.43-0.53, median 0.490), wider heads (HW/SVL 0.33-0.39, median 0.367), and shorter heads (HL/SVL 0.26-0.34, median 0.326) than females from Tarn Dao; the differences be- tween the two forms in these body proportions are statistically significant (P < 0.01, Mann- Whitney LMest). In L. bourreti the head is wider than long, whereas the reverse is true in L. pelodytoides. Neither Dubois (1983) nor Bourret (1937, 1942) mentions a ventrolateral glandular ridge in L. bourreti. Ophryophryne Boulenger There are three morphotypes in the collection, two from Buon Luoi and one from Tarn Dao. All have the very short gape and short snout charac- teristic of the genus. In all three, the webbing is very reduced, vomerine and maxillary teeth are absent, and the inner metacarpal tubercle extends Table 3. Comparison of males from populations of Leptolalax pelodytoides. SVL Range Mean ± SE N T/SVL Range Median N HL/SVL Range Median N HW/SVL Range Median N Tarn Dao Thailand Sichuan 27.7-30.9 29.22 ± 0.21 19 22.9-28.7 26.91 ± 0.70 10 26.4-29.2 27.66 ± 0.22 14 0.44-0.51 0.477 19 0.44-0.49 0.468 9 0.44-0.49 0.480 14 0.34-0.40 0.372 19 0.37-0.41 0.383 8 0.33-0.38 0.363 14 0.30-0.35 0.332 18 0.33-0.36 0.345 10 0.32-0.36 0.342 14 SE = standard error; N = number of frogs; other abbreviations spelled out in footnote to Table 2. out under the first finger. Each has small orbital "horns." The three forms differ strikingly in ventral col- oration, which is not described for any of the three named taxa to which we assign them. This fact and the slight disparities compared to original de- scriptions leave the assignments to named taxa tentative. However, the recognition of these forms as distinct species is clear. Besides differing in ventral coloration, the three differ in dorsal col- oration, in the length of the orbital horns, and in the pattern of the glandular ridges on the back. There are also minor differences in body propor- tions. Ophryophryne microstoma Boulenger Ophryophryne microstoma Boulenger, 1903: 186 — Man Son Mountains, northern Viet- nam; Bourret, 1942:161. Two mature, but not gravid, females (41.6 and 45.4 mm SVL) and one juvenile (25.8) from Buon Luoi. T/SVL (females only) 0.39-0.42, HW/SVL 0.26-0.28, HL/SVL 0.25-0.26, and eye/HL 0.46- 0.51. Habitus slender; snout projecting, sloping obliquely back to lower jaw; nostril lateral; tym- panum visible, about two-thirds diameter of eye. A distinctly curved supratympanic fold from eye to axilla; back with fine, regular ridges forming a wide, curved V beginning at supratympanic fold and with its apex near center of interscapular n gion and a ) — (-shaped pattern over rear half c trunk; no enlarged tubercles at vent. In preserv; five, gray-brown above with faint occipital V an dark temporal region; ventrally dark grayis brown on throat and chest; abdomen lighter wit scattered small dark spots; ventral surface of hin limb light with denser dark spots. In life, the do sal surface gray-yellow; venter dark gray. This series fits Boulenger's (1903) descriptio of the types from northern Vietnam in dermal o namentation, large, exposed tympanum, and co oration. The one disparity is size; Boulengt (1903) reported that a gravid female measured 5 mm. This species was in a swampy area at Buo Luoi; one specimen was found under dead leave and a second on a broad leaf of an araceous plai during a light night rain. Ophryophryne pachyproctus Kou Ophryophryne pachyproctus Kou, 1985:41 — Zhi shihe, Menghla County, Yunnan, China. Two adult males (32.2-38.6) and two adult ft males (45.3-47.8) from Tarn Dao. T/SVL 0.42 0.45, HW/SVL 0.25-0.27, HL/SVL 0.25-0.2' and eye/HL 0.38-0.45. These are the first specimens recorded since th collection of the type series in Yunnan. One c FIELDIANA: ZOOLOG^ the diagnostic features according to Kou (1985) is the large projecting tubercles on each side of the vent; these are present in the Tam Dao spec- imens, but in varying states. The dorsal coloration and regular pattern of low glandular ridges illus- trated by Kou (1985) are the same as shown in the Tam Dao specimens. Kou reported the SVL of males as 28-30. Habitus moderately stocky; snout truncate or weakly pointed, projecting, sloping obliquely back to lower jaw; nostril lateral; diameter of eye longer than snout; orbital horn twice as long as wide; tympanum visible, about half diameter of eye. Supratympanic fold curved; back with thin glandular ridges forming a V over anterior part of back and an H pattern over rear half of trunk; enlarged tubercles just above vent. General col- oration in preservative grayish brown; an invert- ed, dark, interorbital triangle; entire lumbar region dark; irregular dark spots over rest of back; throat dark brown; chest and abdomen with light net- work surrounding dark blotches; ventral surface of hind limb light with heavy dark mottling. The form of the supra-anal protuberances var- ies, although there was only one on each side: one male had a small tubercle; a second male a thick, rounded tubercle; one female a triangular flap; a second female a large conical tubercle. There is no difference between the sexes in body proportions. Males have nuptial pads on the first two fingers, similar to those in O. poilani. The males lacked vocal sacs. Males were heard calling from low leaves of shrubs along rocky banks of a small stream. Ophryophryne poilani Bourret Ophryophryne poilani Bourret, 1937:8 — Dong Tam Ve, Quang-Tri Province, Vietnam; Bour- ret, 1942:162. The Buon Luoi specimens are the first reported since the collection of the holotype about 1 80 km north of Buon Luoi. Nine males (36.4-43.9 SVL) with nuptial pads and vocal sacs, and four females (53.3-59.9), two With enlarged ova. Habitus moderately stocky; csnout projecting, sloping back to lower jaw; nos- tril lateral; tympanum visible, slightly more than Sialf diameter of eye. Dorsum with low tubercles ind fine, low, short, oblique and transverse ridges t lot forming regular pattern; dorsal tubercles often ( ipped with fine white cones; supratympanic fold Table 4. Sexual dimorphism in body proportions of Ophryophryne poilani Bourret. Males Females (N = 9) (N = 4) T/SVL Range 0.45-0.51 0.41-0.49 Median 0.486 0.435 HW/SVL Range 0.29-0.35 0.28-0.30 Median 0.315 0.291 HL/SVL Range 0.26-0.31 0.25-0.28 Median 0.289 0.272 almost horizontal; supra-anal area with or without large projecting tubercles, one or two per side. In preservative dorsum very dark, almost uniformly black in some individuals, with large obscure black spots in others; ventrally dark grayish brown on throat and chest; abdomen varying from dark brown with fine light speckles to mostly brown with large light areas to mostly light gray with large dark spots; underside of hind limb dark with small light markings. In life males vary from black with sharply demarked yellow spots to gray-brown with obscure lighter spots; females yellowish gray with obscure, small lighter spots. Nuptial pads of males a cluster of very fine black spinules on dorsomedial surface of first fin- ger and a much smaller oval patch on the dorsal surface of the second finger. Usually males with minute brown spinules on the dorsal surface of the head. The sexes differ in body proportions, with the males having relatively longer legs and wider and longer heads (Table 4). Bourret (1937) stated that O. poilani lacked an "orbital horn." This series has a short orbital pro- jection, but it is absent on one side in three indi- viduals. This variation suggests that the holotype represents a deviant individual in this character. The holotype female measured 47 mm, somewhat smaller than the Buon Luoi females. Many individuals of this species were seen in April 1995 calling from rocks and dead branches projecting above the water along streams with steep gradient. Although males were occasionally seen side by side, they were usually spaced 10 m apart. The call is a trill similar to the sound of large orthopterans. NGER ET AL.: FROGS OF VIETNAM Fig. 3. fin/ 72), A. larutensis (Boulen- ger) and A. nepalicus Yang differ in having an INGER ET AL.: FROGS OF VIETNAM outer metatarsal tubercle, and A. chunganensis (Pope) and A. monticola (Anderson) differ in hav- ing dorsolateral folds. Amolops spinapectoralis differs from species in the related genera, Huia Yang and Meristogenys Yang, in having the crossbar of the terminal pha- lanx more than 0.6 times the length of the pha- lanx, as well as in the male's secondary sex char- acters. Amolops ricketti (Boulenger) Rana ricketti Boulenger, 1899:168 — Guandun, Fujian Province, China. Amolops ricketti Yang, 1991:23. Staurois ricketti Bourret, 1942:387. A sample from Tarn Dao, including adult males and females, agrees well with published descrip- tions (Bourret, 1942; Yang, 1991) and specimens from Fujian. The snout is depressed and project- ing, the canthi sharp, lores concave and vertical, the tympanum visible but with obscured rim, and all toes fully webbed to the discs. Males lack gu- lar pouches and vocal sacs, but have a large nup- tial pad consisting of about 100-125 large, white, conical spines. All of the Tarn Dao specimens have the black spotting on the throat that occurs in only one-third of the specimens from Fujian (fmnh 24682-90). The Fujian frogs are larger than those from Tarn Dao. SVL of males from Fujian measure 54.6-58.5 (N = 3), those from Tarn Dao 42.9-46.0 (N = 9); females from Fujian measure 55.0-60.0 (N = 6), those from Tarn Dao 50.0- 56.2 (N = 7). The sexes differ slightly but significantly in cer- tain body proportions. Males have slightly longer legs (T/SVL 0.53-0.57), wider heads (HW/SVL 0.32-0.35), and longer heads (HL/SVL 0.31- 0.36) than females (T/SVL 0.48-0.54, HW/SVL 0.33-0.34, and HL/SVL 0.33-0.34); differences were at the P < 0.05 level (Mann-Whitney U-test) for each comparison. Huia nasica (Boulenger) Rana nasica Boulenger, 1903:187 — Mao Son, Tonkin, Vietnam; Bourret, 1942:352. Huia nasica Yang, 1991:31. Sixteen adult males and one adult female col- lected at Tarn Dao agree with the original descrip- 13 tion in every detail. The strongly depressed and projecting snout and the deeply concave lores are especially distinctive in this species. All of the frogs in this series have a sharply defined black stripe just below the canthus, and the temporal region and lower border of the dorsolateral fold are black; these are color features noted by Bou- lenger (1903). Characters not mentioned by Bou- lenger (1903) or Bourret (1942), who had speci- mens from Tarn Dao, are: males with small, white asperities on the rear of the back; discs of the toes same diameter as those of fingers. SVL measurements of males 44.0-49.8 (mean ± SE 46.33 ± 0.33, N = 16), T/SVL 0.56-0.63 (median 0.590), HW/SVL 0.27-0.30 (median 0.287), and HL/SVL 0.36-0.41 (median 0.390). The single female measured 77.7 mm. Parts of Yang's (1991) description of Huia na- sica were based on two males (fmnh 215970-71) from Kanchanaburi Province, Thailand. These specimens, which are not conspecific with the Tarn Dao series, have much longer legs (T/SVL 0.71), a nonprojecting and nondepressed snout, nostril much closer to the tip of the snout than to the eye (as opposed to equidistant between snout and eye), toe discs larger than those of fingers, and pineal body hidden (visible in Tarn Doa spec- imens). However, in other characters important for generic assignment (Yang, 1991), such as relative lengths of first and second metacarpals, shape of terminal phalanges, and shape of sternum, the Tarn Dao and Thailand frogs are identical. Occidozyga martensii (Peters) Phrynoglossus martensii Peters, 1867:29 — Bang- kok, Thailand. Ooeidozyga laevis (part) Bourret, 1942:401. One adult female from Tram Lap and 11 fe- males, eight males, and three juveniles from Buon Luoi constitute this sample. The males have weak, pale, but distinct nuptial pads on the first finger. Six of the females have enlarged nonpigmented ova. The species is widely distributed in Vietnam (Bourret, 1942). Reasons for placing this species in Occidozyga are given elsewhere (Inger, 1996). Six were collected in rice fields, one along a forest road, and three from the silty bank of a stream where it flowed into a large pond. Rana attigua, new species (Fig. 5) Holotype — fmnh 252775, an adult female, col- lected at Buon Luoi, An Khe District, Gia La Province, Vietnam, 25 April 1995 by Ilya Dar evsky and Nikolai Orlov. Paratypes — fmnh 252776-816, mvz 222932- 34, 222936, 222941-43 from the type locality fmnh 252817-30 from Kon Cha Ran, An Khe District, Gia Lai Province, Vietnam; fmnf 252831-32 from Tram Lap, An Khe District, Gu Lai Province, Vietnam. All were collected by Ily< Darevsky and Nikolai Orlov. Etymology — Specific name from attiguus L. meaning neighboring, referring to its similarity tc R. milleti Smith. Diagnosis — Digit tips expanded, discs of innei fingers without circummarginal groove; first fin- ger longer than second; dorsolateral fold present outer metatarsal tubercle present; males with large humeral gland, but lacking gular pouches; SVL ol males 40-45, of females 55-65; underside of call immaculate white or with faint dark speckling. Description — Habitus moderately stocky; heac longer than broad. Snout obtusely pointed, pro- jecting, rounded in profile; nostril closer to tip ol snout than to eye, lateral; canthi distinct, not con- stricted; lores very slightly oblique, concave; di- ameter of eye equal to snout; interorbital equal tc width of eyelid; tympanum distinct, one-half tc two-thirds diameter of eye, larger in males (see below); vomerine teeth in short oblique rows, well-removed from choanae. Finger tips dilated into distinct, small discs less than twice diameter of penultimate phalanges: discs without circummarginal grooves or with weak grooves on outer fingers; first finger longei than second; subarticular tubercles prominent; su- pernumerary tubercles on three outer fingers. Discs of toes equal to or slightly larger than those of fingers; all with circummarginal grooves; web- bing on first and second toes between subarticulai tubercle and disc, on third and fifth toes narrowl) to disc, on fourth toe between middle and distal tubercles, fourth toe with two to two-and-one- third phalanges free; inner metatarsal tubercle oval, less than half length of first toe, outer meta- tarsal tubercle distinct, round, much smaller thar inner; no flaps of skin on margins of first and fifth toes. Conspicuous, continuous dorsolateral folds: skin of back granular with numerous small tuber- cles; sides with larger tubercles; dorsal surfaces 14 FIELDIANA: ZOOLOGY Fig. 5. Rana attigua, new species. Male, 44.9 mm. of hind limb with tubercles; ventral surfaces smooth. Color in life dorsally medium brown, with or without small darker spots on back; dark brown or black band from snout through eye and tym- panum, sharply defined on snout, less so in tem- poral area; upper lip with white to yellow stripe, extending to axilla; upper half of iris light golden brown; limbs with dark brown crossbars dorsally. In preservative ventral surfaces whitish; brownish spots or mottling on throat and usually on chest and anterior part of abdomen; rear of abdomen without dark pigment; ventral surface of thigh without dark pigment; underside of calf immac- ulate or with very fine dark speckling; rear of thigh dusted with dark pigment, but without clear pattern. Measurements of holotype: SVL 63.7, T 34.3, HW 21.4, HL 24.0, and tympanum 5.5. Males have vocal sac openings at the corners of the floor of the mouth, but lack gular pouches. A velvety nuptial pad covers the medial and dor- sal surfaces of first finger from its base to the proximal end of the penultimate phalanx; the ven- tral margin of the nuptial pad is straight. Males INGER ET AL.: FROGS OF VIETNAM also have a large, conspicuous, flat humeral gland that occupies almost the entire anteromedial as- pect of the upper arm. Variation — The dorsal and temporal areas in about half of the individuals have tubercles set with whitish, spinose tips. This condition is seen in both males and females. SVL of males with nuptial pads 36.2-45.5, only four <39.0 (mean ± SE 41.34 ± 0.28, N = 43), of females with mature oviducts 57.7-63.7 (59.66 ± 0.87, N = 6); T/SVL males 0.51-0.58 (median 0.535, N = 26), females 0.53-0.57 (me- dian 0.554, N = 6); HW/SVL males 0.30-0.34 (median 0.325, N = 27), females 0.31-0.35 (me- dian 0.339, N = 6); HL/SVL males 0.38-0.43 (median 0.396, N = 27), females 0.36-0.38 (me- dian 0.375, N = 6); tympanum/SVL males 0.089- 0.115 (median 0.099, N = 27), females 0.077- 0.089 (median 0.086, N = 6). Comparisons — Rana attigua is very similar to and co-occurs with R. milleti Smith. In preserva- tive, the latter has dark spotting on the ventral surface of the calf and dark speckling or mottling under the thigh, whereas in R. attigua the under- side of the thigh is immaculate whitish and the 15 Table 6. Comparison of Rana attigua with similar species from Southeast Asia. Males only. Data sources: / guentheri, R. leptoglossa, and R. mortenseni from Boulenger (1920); R. cubitalis from Smith (1917); R. spinulos from Smith (1923); other species from present study. Typanum/SVL Sexual dimor- Gular SVL HW/SVL HL/SVL Range phism pouches R. attigua 36-46 0.30-0.34 0.38-0.43 0.089-0.115 Yes Absent R. milleti 34-40 0.30-0.36 0.38-0.44 0.099-0.121 Yes Absent R. guentheri 60-80 0.29-0.33 0.34-0.37 0.067-0.088 No Present R. leptoglossa 51-60 0.34-0.35 0.36-0.37 0.098-0.100 ? Present R. cubitalis 66-68 0.30-0.31 0.31-0.35 0.088-0.091 ? Present R. nigrovittata 44-57 0.36-0.39 0.39-0.45 0.081-0.103 Yes Absent R. spinulosa 37-42 0.33-0.37 0.38-0.41 0.073-0.083 No Absent R. mortenseni 50-71 0.35-0.36 0.35-0.36 0.066-0.085 No Absent R. johnsi 44-49 0.32-0.34 0.38 0.089-0.100 Yes Absent underside of the calf is immaculate or has very faint dark speckling. The rear of the thigh in R. attigua has a light dusting of dark pigment, in contrast to the distinct pattern in R. milleti (see description below). The nuptial pad in male R. milleti has a distinct notch or constriction of the ventral margin, which is straight in R. attigua. The most conspicuous differences between these two involve size. Although the size ranges of males partially overlap, only one of 52 adult males of R. milleti was larger than 40.0 mm, whereas 81% of 43 male R. attigua exceeded 40.0. Size ranges of adult females (determined by condition of the oviduct) of the two species did not overlap; the largest female R. milleti measured 49.6 mm. The difference between mean SVL of males and fe- males in R. attigua (18 mm) is much greater than the corresponding measure in R. milleti (8.25 mm). In a suite of characters — discs of inner fingers without grooves, dorsolateral fold present, first finger much longer than second, outer metatarsal tubercle present — R. attigua and R. milleti resem- ble seven congeners from Southeast Asia: R. cub- italis Smith, R. guentheri Boulenger, R. johnsi Smith, R. leptoglossa (Cope), R. mortenseni Bou- lenger, R. nigrovittata (Blyth), and R. spinulosa Smith. Males of R. cubitalis, R. guentheri, and R. leptoglossa have gular pouches and are larger (Ta- ble 6) than those of R. attigua, and males of R. cubitalis have an extension of the nuptial pad up the arm (Smith, 1930). Rana nigrovittata has sides darker than the back, a much smaller hu- meral gland (only half the length of the upper arm), a much bolder pattern on the rear of the thigh, and full web to the distal tubercle of the fourth toe. Rana mortenseni differs from attigw in having larger males, more extensive webbing head as wide as long, and apparently no sex di morphism in size of tympanum (Table 6). Ram johnsi is sharply different from R. attigua in col oration, having a very dark rhomboidal mark cov ering the temporal region, a dark longitudina streak along the calf and tarsus, but no light strip along the lip; males of R. johnsi lack humera glands. Rana spinulosa lacks sex dimorphism i) size of tympanum and has very warty dorsal sur faces and a wider head than R. attigua. Habitat. — All individuals were found alonj forested streams. About 40% (19) were sitting o) sandy banks, about 30% on dead leaves, and th< rest on rocks. On 7 nights, we collected individ uals of both R. attigua and R. milleti in a singli habitat; in these samples, which were made oi two streams at each of two localities, R. attigm was more numerous in three and R. milleti in foui Most specimens were caught on the banks of tw< streams at Buon Luoi; we collected 33 R. attigm and 21 R. milleti on Stream N5 and 17 R. attigm and 26 R. milleti on Stream N3. Although these data, arranged into a 2 X 2 table, show statisti cally significant departure from randomness (x2 = 4.46, P < 0.05), the segregation by streams i- modest. Most specimens were found on four type of substrate: bare sand or silt, rocks, logs, am dead leaves. The two species show no segregatioi by substrate (x2 = 4.88, df = 3, P > 0.10). Rana cf. blythii Boulenger (Fig. 6) Large samples of this species were collected a Buon Luoi, Kon Cha Rang, and Tram Lap at el 16 FIELDIANA: ZOOLOGY Fig. 6. Rana cf. blythii. Male, 73.8 mm. evations from 700 to 1200 m ASL. Males have enlarged odontoids at the front of the mandible and distinctly enlarged heads, but they lack nup- tial pads and vocal sacs. The tympanum is super- ficial. Although these frogs are clearly part of the R. blythii species group, they differ significantly in a number of ways from typical R. blythii of the Malay Peninsula and Thailand. In the Vietnam form, full webbing reaches only to the distal sub- articular tubercle of the fourth toe (to the base of the swollen tip in R. blythii), SVL and T/SVL are smaller (Table 7), and ripe ova are bicolored with a black animal hemisphere (ova are pale and uni- colored in R. blythii). The canthus is rounded. The three outer fingers have a ridge of skin on the medial border, widest Table 7. Comparison of Rana cf. blythii from Vietnam with R. blythii from the Malay Peninsula. Adults only. Males with enlarged odontoids; females with enlarged, convoluted oviducts. Males Females Vietnam Malay Peninsula Vietnam Malay Peninsula SVL Range N 58.5-93.1 24 66.6-208.0 21 53.7-79.6 28 71.6-156.0 32 T/SVL* Range Median N 0.43-0.56 0.507 20 0.47-0.58 0.541 21 0.47-0.55 0.523 18 0.49-0.59 0.545 32 * Differences between Vietnamese and peninsular samples were statistically significant at P < 0.01 for both sexes (Mann-Whitney IMest). INGER ET AL.: FROGS OF VIETNAM 17 \ •' ' , Fig. 7. Rana johnsi Smith. Female, 46.7 mm. on the second finger. The first and fifth toes have movable flaps of skin externally. The first three toes and the fifth are fully webbed to the base of the swollen tips. There are two wide, black bars from the eye to the lip. The upper half of the tympanum and the supratympanic fold are cov- ered by a black streak. In about one-fourth of the individuals, there is a light vertebral stripe. SVL and T/SVL are given in Table 7. HW/SVL of males 0.39-0.46 (median 0.421, N = 36), of females 0.36-0.41 (median 0.383, N = 21); HL/ SVL of males 0.42-0.50 (median 0.445, N = 35), of females 0.38-0.46 (median 0.410, N = 21). For the largest males (SVL > 80.0, N - 8) HW/SVL 0.40-0.45, HL/SVL 0.42-0.48. Parker (1925) reported Rana macrodon from Bao-Ha (76 m ASL) in the Red River valley of northern Vietnam and, without comment or ref- erence to specimens, gave the range as "Siam and Annam." These and Bourret's (1942) citation of Parker are the only previous mentions of this spe- cies group in Vietnam. Most individuals were seen along the banks of rapid streams, though some were observed along banks of large forest ponds. Rana johnsi Smith (Fig. 7) Rana sauteri johnsi Smith, 1921:434 — Sui Kat Annam, Vietnam. Rana (Hylarana) sauteri johnsi Bourret, 1942 322. The present sample from Buon Luoi, Tram Lap and Kon Cha Ran matches Smith's (1921) de scription and illustration in all details of the dis tinctive color pattern and in having very long leg; (T/SVL of males 0.62-0.65 in Smith's sample 0.63-0.73 in present sample), a distinctly raised continuous dorsolateral fold, full webbing to disc; of third and fifth toes and to distal tubercle o fourth toe, and an inverted V-shaped glandulai fold between the shoulders. They also match ont of Smith's specimens (mcz A8801) that we havt examined. However, there are several conspicuou; differences. Smith (1921) stated that males of R johnsi had internal vocal sacs and brown nuptia pads, whereas the four males from Buon Luo have swollen inner fingers, without the fine spi nules of the typical ranid nuptial pad, and lack 18 FIELDIANA: ZOOLOGY Table 8. Characters of males from several samples of frogs related to Rana sauteri Boulenger. R. sauteri R. johnsi Taiwan Types Buon Luoi SVL 35.3-40.8 40-45 43.5-48.9 T/SVL range 0.55-0.61 0.62-0.64 0.63-0.73 T/SVL median 0.577 0.628 0.652 Vocal sac open- ings Present Present Absent Nuptial pad Testes pigmented Pigmented vo- merine ridges Present Yes No Present 7 No Absent Yes Yes vocal sac openings. Males from Buon Luoi also lack humeral glands. Although these differences may be related to levels of sexual maturity, the Buon Luoi males (SVL 43.5-48.9) are as large or larger than the five males in the type series (40- 45). The four Buon Luoi females having mature, convoluted oviducts measure 49.4-55.5, larger than those in the type series (40-50). All of the Buon Luoi frogs have a heavy concentration of melanophores on the vomerine dental ridges. Smith (1921) did not mention this character, al- though he examined the vomerine teeth of R. johnsi, but the specimen that we examined from his series lacks palatal pigment. Smith (1921) separated the Annamese frogs from typical R. sauteri Boulenger (type locality Taiwan) on the basis of these characters: more pointed snout, longer leg, more prominent dor- solateral folds, and smaller size. Smith (1921) based his comparison on the four females that constituted the type series of R. sauteri. We have examined a more recently collected sample of R. sauteri from Taiwan (fmnh 127569-99), includ- ing nine males. Our specimens from Taiwan have shorter legs (T/SVL 0.55-0.61, median 0.577) and less conspicuously raised dorsolateral folds than R. johnsi, confirming several of Smith's (1921) observations. However, the Taiwanese males mea- sure 35.4-40.8 SVL, and because they are smaller than males in Smith's series, his statement that R. johnsi is smaller than typical R. sauteri is incor- rect. Instead, Taiwanese males are smaller than those from Vietnam (Table 8). Taiwanese males have vocal sac openings and velvety brown or gray nuptial pads, agreeing with Smith's descrip- tion of R. johnsi, but differing from the present Vietnamese sample. None of the Taiwanese frogs has pigment on the vomerine dental ridges. One INGER ET AL.: FROGS OF VIETNAM striking, shared feature of males from Taiwan and Buon Luoi is heavily pigmented testes. Rana kuhlii Tschudi Rana kuhlii Tschudi, 1838:40 — Java; Bourret, 1942:278. Samples were collected at Kon Cha Rang and Tarn Dao. Adult males (with nuptial pads) and females (with enlarged, convoluted oviducts) from Kon Cha Rang measure 40.3-49.2 (N = 2) and 45.4-59.0 (N = 5), respectively. Adult males and females from Tarn Dao measure 41.9-84.8 (N = 8) and 52.0-83.7 (N = 6), respectively. T/SVL 0.41-0.48 (females), 0.42-0.48 (males); HW/SVL 0.36-0.40 (females), 0.39-0.44 (males); HL/SVL 0.35-0.39 (females), 0.35-0.46 (males). The frogs from Tarn Dao have distinct dermal flaps along the lateral margin of the third finger; such flaps are absent in all except two (one male and one female) from Kon Cha Rang. All of the Kon Cha Rang specimens were taken along the banks of streams at about 1 200 m — two on rocks in midstream and the rest on silty banks close to the water's edge. Rana livida (Blyth) Polypedates lividus Blyth, 1856:718 — Tenasser- im, Burma. Rana livida Boulenger, 1 887b:484. Rana (Hylarana) livida Bourret, 1942:371. Two large samples, one from the An Khe Dis- trict (Buon Luoi, Kon Cha Ran, and Tram Lap) and another from Tarn Dao, agree with a sample from Huai Kha Kaeng, Thailand, near the Tenas- serim area of Burma, type locality of the species. All of these frogs have a ridge or flap of skin along the median edge of the third finger and a narrow ridge of skin along the median edge of the first toe. All lack outer metatarsal tubercles. Males have gular pouches. Bourret (1942) suggested that geographic vari- ation in development of a dorsolateral glandular fold divides this species into a "northern" popu- lation, to which Boulenger's (1899) Rana grami- nea applies, and a "southern" one, or Rana livida in the strict sense. That distinction cannot be sup- ported because there is no difference between the northern and southern samples at hand in this 19 character; in fact, there is variation within the Buon Loy sample. However, females from the An Khe District (SVL 74.0-99.9, mean ± SE 86.49 ± 1.81, N = 17) are smaller than those from Tarn Dao (95.5-104.4, 100.70 ± 0.90, N = 11) and have relatively longer tibia length (T/SVL An Khe females 0.58-0.69, median 0.634, N = 18; Tarn Dao females 0.57-0.62, median 0.594, N = 11). The Tarn Dao sample included only a single male, preventing comparisons of this sex. with whitish venter suffused with dark brown; pair of blackish rectangular dark markings at ree of throat. Tran et al. (1981) listed this species from fot provinces in northern Vietnam, and Nguyen et a (1994) reported it from Tarn Dao. Although Yan et al. (1991) did not include it in the fauna c Yunnan, it probably occurs there because the typ locality is on the border. Rutin maosonensis (Bourret) Hylarana maosonensis Bourret, 1937:36 — Mao Son, Tonkin, Vietnam. Rana {Hylarana) maosonensis Bourret, 1942: 351. The present sample consists of 12 males and 16 females from Tarn Dao. Bourret (1937, 1942) did not refer to humeral glands as part of the male suite of secondary sex characters, but weak hu- meral glands are present in the males from Tarn Dao and in two males from the type series (fmnh 123909-10). Males lack gular pouches. The Tarn Dao series matches the original description and the two paratypes in distinctive coloration, spi- nose tuberculation of the skin, habitus, and web- bing. Full webbing reaches just beyond the middle subarticular tubercle of the fourth toe. In the Tarn Dao sample, females are much larger (SVL 5 1 .0- 61.1, mean ± SE 56.29 ± 0.72) than males (37.9- 43.1, 41.49 ± 0.46). Males have slightly narrower heads than females; HW/SVL of males 0.32-0.35, median 0.33; of females 0.33-0.36, median 0.34 (P < 0.05). Males also have slightly longer heads; HL/SVL of males 0.37-0.41, median 0.390; fe- males 0.36-0.39, median 0.378 (P < 0.05). There is no sexual dimorphism in relative length of the tibia (T/SVL of males 0.50-0.57, median 0.527; females 0.50-0.56, median 0.528; P > 0.10) or diameter of the tympanum (TY/SVL of males 0.075-0.099, median 0.086; females 0.063-0.097, median 0.084; P > 0.10). Coloration in life (based on a female) dark red- dish brown dorsally, with irregular, obscure black spots; concavity of lores black; upper lip yellow- ish; upper half of iris dark brown, lower half blackish; tympanum dark brown, with black spot at posterior corner; sides changing from dark brown dorsally through yellow brown to white near belly, with many black spots; limbs with dark brown crossbars. In preservative, Tarn Dao frogs Rana milleti Smith Rana milleti Smith, 1921:432 — Dalat, Annam, Viel nam. Rana {Hylarana) milleti Bourret, 1942:312. These specimens from Boun Luoi, Tram Lap and Kon Cha Ran constitute only the second rec ord of the species, and they agree well wit! Smith's (1921) description, differing in only on important detail. Smith stated that males lacked vocal sac, but a male collected with the type serie (MAS field no. 5110, fmnh 128298) has small round vocal sac openings at the corners of tb mouth; males in the new sample have vocal sa« openings in the same position. The tympanum i larger in males than in females, a character no mentioned by Smith (1921) but clear from th measurements he presented. The nuptial pad ha a noticeable ventral constriction that in a fev males completely divides the pad into proxima and distal halves. Males have a large, flat, whitisl humeral gland. The following notes amplify Smith's (1921) description. Web on first toe to subarticular tubercle or ; little beyond, on second toe between subarticula tubercle and base of disc, on third toe betweei distal tubercle and base of disc, narrowly reachinj base of disc, on fifth toe to base of disc, on fourtl toe to middle tubercle on medial side, betweei middle and distal tubercles on lateral side. Skii between dorsolateral folds granular and witl small tubercles, many of which are capped with ; small spinule; sides with larger tubercles; ventra surfaces smooth. Color in preservative mediun brown dorsally, slightly darker on sides; a sharph defined dark band from tip of snout through eyt to tympanum; upper lip with a wide or thin whitt line extending to axilla; underside of head anc chest usually with dark mottling; entire abdomer often with small dark spots; underside of calf witl heavy dark spotting; rear of thigh dark brown 20 FIELDIANA: ZOOLOGY forming a network around small light spots or blotches. SVL of males with nuptial pads 34.2-40.4 (mean ± SE 37.01 ± 0.18, N = 52), of females with mature oviducts 41.1-49.6 (45.26 ± 0.70, N = 15); T/SVL males 0.51-0.57 (median 0.540, N = 23), females 0.51-0.58 (median 0.531, N = 15); HW/SVL males 0.30-0.36 (median 0.326, N = 23), females 029-0.33 (median 0.313, N = 15); HL/SVL males 0.38-0.44 (median 0.407, N = 23), females 0.35-0.40 (median 0.376, N = 15); tympanum/SVL males 0.099-0.121 (median 0.112, N = 23), females 0.080-0.104 (median 0.090, N = 15). Many of these frogs were collected with R. at- tigua (see above). All were on stream banks, mostly on sand or silt (23) or dead leaves (16). Only five were on rocky substrate. Rami montivaga Smith Rana montivaga Smith, 1921:436 — Dalat, An- nam, Vietnam. Rana (Hylarana) montivaga Bourret, 1942: 346. An adult male (51.9 mm) from Buon Luoi agrees closely with Smith's (1921) description and illustration in general form and coloration. How- ever, it has a gular pouch, whereas Smith reported that males in the type series had "internal vocal sacs." The Buon Luoi male has no humeral gland; because Smith did not mention this structure, we assume topotypic males also lacked them. The Buon Luoi male has the head distinctly longer than wide, whereas the measurements given by Smith show HW > HL. We have examined three immature topotypes (fmnh 135343, mcz 8785-86) collected by Smith at the same time as the types, and these frogs also have HL > HW. The new specimen and the topotypes have a circummargin- al groove around the small disc of the first finger, which is longer than the second; all lack an outer metatarsal tubercle; and all have vomerine teeth in short series widely separated from the choanae. In these characters and in webbing, coloration, and skin surface, these four specimens match Smith's (1921) description. One adult female (64.3 mm) from Tarn Dao agrees closely with Smith's description and illus- tration. The main differences from Smith's de- scription are the presence of a weak outer meta- tarsal tubercle (absent in types), HL slightly great- INGER ET AL.: FROGS OF VIETNAM er than HW, and indistinct light spots on a darker background (the reverse of R. montivaga). Vo- merine teeth in the Tarn Dao specimen are in long series narrowly removed from the choanae, and thus they differ from Smith's description and the four specimens discussed above. Pope (1931) identified frogs from Fujian Prov- ince, China as R. montivaga, although they were later referred to R. swinhoana Boulenger by Liu and Hu (1961). However, Pope's material (ex- amined in fmnh) is not conspecific with R. swin- hoana, differing greatly in size and in secondary sex characters (based on the redescription of R. swinhoana by Wang & Chan, 1977). Pope's (1931) specimens also differ from R. montivaga in having a narrower head (HW/SVL 0.31-0.33 compared to 0.35-0.38 in Smith's types), first fin- ger shorter than second, and a sharply defined, black loreal stripe; the Fujian specimens are also smaller than Smith's types (53-59 mm, vs. 58- 75). Rana nigrovittata (BIyth) (Fig. 8) Limnodytes nigrovittatus Blyth, 1 856:7 18-Pegu. Burma. Rana nigrovittata Sclater. 1892:345. Rana {Hylarana) nigrovittata Bourret, 1942: 317. A large series from Buon Luoi, Tram Lap, and Kon Cha Ran matches Smith's (1922) second morphotype from northeastern Thailand and Laos: SVL often > 55 mm, dark lateral band broken up, and venter often with dark pigment. Males of the present sample have wrinkled skin at the corners of the throat, as noted in this form by Smith, but they do not have gular pouches, such as occur in R. humeralis Boulenger or R. leptoglossa (Cope). Males have gray nuptial pads on the dorsal and medial surfaces of the first finger; a large gland on the proximal fourth or third of the upper arm; and light, cornified spinules on the back and sides. In this sample, males and females do not differ in SVL or T/SVL. SVL of males 43.3-61 .8 (mean ± SE 51.53 ± 0.58, N = 52), of females 45.0- 65.3 (52.92 ± 0.68, N - 53); T/SVL of males 0.50-0.55 (median 0.522, N = 14), of females 0.50-0.56 (median 0.525, N = 16). The sexes do differ in relative size of head and tympanum. HW/ SVL of males 0.35-0.39 (median 0.369, N = 15), of females 0.32-0.36 (median 0.344, N = 16); HL/SVL of males 0.39-0.45 (median 0.414, N = 21 Fig. 8. Rana nigrovittata (Blyth). Male, 52.9 mm. 15), of females 0.37-0.41 (median 0.390, N = 16); tympanum/SVL of males 0.081-0.109 (me- dian 0.093, N = 15), of females 0.074-0.091 (me- dian 0.083, N = 16). Differences between the sex- es in these three body proportions are significant at the P < 0.01 level (Mann- Whitney U-test). Rana taipehensis Van Denburgh Rana taipehensis Van Denburgh, 1909:56-Taipei, Taiwan. Rana (Hylarana) taipehensis Bourret, 1942: 335. Adult males (with nuptial pads) measure 23.6- 30.2 (mean ± SE 27.29 ± 0.79, N = 11); adult females measure 37.2-42.5 (40.07 ± 0.75, N = 7). The whitish, velvety nuptial pad covers the medial and dorsal surfaces of the first finger from its base to the level of the distal edge of the sub- articular tubercle. None of the males has voci sacs. Males have much larger tympana than fe males; tympanum/SVL of males 0.124-0.13 (median 0.130, N = 11), of females 0.086-0.095 median 0.093, N = 7. Specimens in the preser samples were taken at Buon Luoi and Buon Gen Lanh, both in central Vietnam. Smith (1923) re corded this species from southern Annam, al though Bourret (1942) reported it only fron northern Vietnam. Paa verrucospinosa (Bourret) Rana spinosa verrucospinosa Bourret, 1937:26- Chapa, Fan-Si-Pan, and Tarn Dao, Tonkin Vietnam. Rana verrucospinosa Bourret, 1939:8; Bour ret, 1942:295. Eleven specimens from Tarn Dao, four adul 22 FIELDIANA: ZOOLOGY males 95.1-103.6 (mean 97.97), four adult fe- males 72.6-103.8 (mean 92.74), four juveniles 53.3-72.6. The largest juvenile is a male lacking secondary sex characters. The smallest adult fe- male has widened, coiled oviducts but immature ova. Bourret's (1937) description and illustration depict a frog having a very rough back covered by short, thick ridges and round tubercles and sides covered by oval or round tubercles. The Tarn Dao specimens have the same skin rugosity, and all have flattened melanic cones on many of the dorsal ridges. Males have melanic spines covering the entire chest and most of the abdomen, leaving only a patch of smooth skin in the center of the rear third of the abdomen. Similar spines cover the dorsal and medial surfaces of the first two fin- gers. The ventral surfaces of the thighs are smooth. One series of tadpoles was collected at Tarn Dao. They agree very closely to Bourret's (1942) description and have heavily spotted tails, as he illustrated. Denticular formulae vary only slightly: 6(2-6)/3(l) or 6(3-6)/3(l). Total lengths in stages 27-29 were 71.1-75.4. Dubois (1987) considered P. verrucospinosa to be conspecific with P. boulengeri (Giinther). However, in adults of the latter from Sichuan (fmnh specimens), Yunnan (illustrations in Yang et al., 1991), and Guizhou (Wu et al., 1986), Chi- na the center of the back is smooth, and males have the entire abdomen and ventral surface of the thigh covered with black spines. Paa yunna- nensis, which is known from southern Yunnan, close to the border with Vietnam, differs from P. verrucospinosa in having the pectoral spines ar- ranged in two separated groups and in having round rather than elongate ridges on the back. Paa spinosa has scattered rounded tubercles on the sack, and the male lacks melanic spines along the sides of the abdomen. Paa microlineata has wide- ly spaced, narrow ridges on the back and coarsely granular rather that tubercular sides (Bourret, 1942). Rhacophoridae I Chirixalus doriae Boulenger i Chirixalus doriae Boulenger, 1893:341 — Karin Bia-po, Burma; Bourret, 1942:376. il An adult female (33.5 mm) and a subadult (27.0) were collected at Buon Luoi. Although known from Burma, northern Thailand (Taylor, 1962), and southwestern Yunnan (Yang et al., 1991), this is the first record from Vietnam. Chirixalus nongkhorensis (Cochran) Philautus nongkhorensis Cochran, 1927:179 — Nongkhor, southeastern Thailand. Rhacophorus (Chirixalus) nongkhorensis Ahl. 1931:107. Chirixalus nongkhorensis Bourret, 1942:473. An adult male (19.9 mm) collected at Buon Luoi agrees with specimens from eastern Thailand (fmnh 182594-611) in several significant char- acters: the nuptial pad is glandular, lacking fine spinules typical of most rhacophorids; dorsal sur- faces of head, trunk, and limbs with fine, colorless asperities; limbs with dark spots dorsally, but no complete crossbars. It also agrees with the Thai frogs in most other characters. The principal dif- ference is in SVL; the Thai males measure 24-31 mm. The species has also been recorded from Burma (Ahl, 1930, as C. striatus). This specimen was perched in a bush at the edge of a pond. Chirixalus palpebralis (Smith) Philautus palpebralis Smith, 1924:233 — Langbi- an Peaks, Annam, Vietnam. Rhacophorus (Chirixalus) palpebralis Ahl, 1931:104. Chirixalus palpebralis Bourret, 1942:474. Large samples from Buon Luoi and Kon Cha Ran match Smith's (1924) description of the unique type very closely, except that the tympa- num is conspicuous, not "almost hidden." Smith noted that the inner fingers were "partially op- posed" to the two outer ones; this was presum- ably the basis for Ahl's (1931) placement of the species in Chirixalus. Females in the sample from Buon Luoi have bi-pigmented ova, a black hemi- sphere and a brown one; the ova are not greatly enlarged (1.5-1.8 mm). The left ovary of one fe- male held 52 pigmented ova. The pigmentation, size, and number of the ova do not fit the repro- ductive pattern of Philautus, but they confirm Ahl's (1931) generic assignment of the species. Yang et al. (1991) reported C. palpebralis from II NGER ET AL.: FROGS OF VIETNAM 23 Yunnan as a Philautus. We amplify Smith's (1924) description. Full webbing on first toe to distal edge of sub- articular tubercle, on second toe between tubercle and base of disc, on third and fifth toes between distal subarticular tubercle and disc, and on fourth toe to distal edge of middle tubercle or between the two outer tubercles. Males have slit-like vocal sac openings; a whitish, velvety nuptial pad cov- ering dorsal and medial surfaces of the first finger from its base to the level of the subarticular tu- bercle; and many small, whitish pustules dorsally on the trunk. SVL measurements of males 24.4-26.2 (mean ± SE 25.21 ± 0.16, N = 18), of females 29.2- 32.1 (30.90 ± 0.27, N = 10); T/SVL (males only) 0.48-0.53 (median 0.508, N = 15); HW/SVL (males) 0.28-0.32 (median 0.297, N = 15). One tadpole at stage 39 is assigned to this spe- cies. The two inner fingers are opposed to the out- er two, and full webbing of the third and fifth toes almost reaches the discs. The labial tooth formula is 5(2-5)/3(l), and there is a wide median gap in the marginal papillae of the lower lip. The head- body is black dorsally and ventrally. The caudal muscle and fins are black except for a wide ver- tical yellow band immediately behind the body; there is a black spot on the muscle in the center of the light band. Headbody is 13.75 mm; total length is 36.6. The fingers and toes of the tadpole agree with adult C. palpebralis. In larval C. vittatus the distal third of the tail is black and there is a dark lateral stripe on the body (Heyer, 1971b); larval C. nong- khorensis have a blotched tail and only three or four upper rows of labial teeth (Heyer, 1971b); and larval C. doriae have a black stripe on the caudal muscle only, and only three upper rows of labial teeth (Pope, 1931; determined as C. doriae by Heyer, 1971). This species was observed at Buon Luoi in Oc- tober-December 1993 and March-May 1995. Most individuals were perched on broad-leaved Araceae in swampy portions of the banks of a stream, within 1 m of the water's surface. Pairs in amplexus were seen in April-May 1995; most of these were seen after night rains. The tadpole was caught in a ponded portion of a forest stream. Chirixalus vittatus (Boulenger) Ixalus vittatus Boulenger, 1887b:421 — Bhamo, Burma. Chirixalus vittatus Liem, 1970:95. Philautus vittatus Bourret, 1942:462. All specimens were caught at Buon Luoi. SV; of males 19.4-21.4 (mean ± SE 20.46 ± 0.13, 1 = 14), of females 22.5-24.2 (mean 23.42, N « 4). T/SVL (males) 0.47-0.51 (median 0.488, N ■ 11), HW/SVL (males) 0.29-O.32 (median 0.30] N = 11). Males have very fine colorless spinules on do sal surfaces of head and trunk; the back is smoot in females. Males have a white, velvety nupti; pad on the dorsal and medial surface of the fir: finger, from its base to the level of the subarticuh tubercle. This species was seen at Buon Luoi in Octc ber-December 1993 and March-May 1995. Th frogs were perched on branches of shrubs an small trees forming thickets at the edge of a larg forest pond. Perch height was between just abov the water level and 3 m. The species had a ver high density; counts of > 100 were made withi a 10-m strip of bank vegetation. It was never see in the forest away from the pond. Polypedates leucomystax (Gravenhorst) Hyla leucomystax Gravenhorst, 1829:26 — Java. Polypedates leucomystax Tschudi, 1838:75. Rhacophorus leucomystax Bourret, 1942:425. Samples of two morphotypes fitting the gener; definitions of P. leucomystax and P. megacepha us (Matsui et al., 1986) were collected at Buc Luoi. One form has four narrow, dark, dors; stripes; the second only rarely (2/50) has dors, stripes, and those unusual specimens have on] two stripes. The two forms differ in other n spects. In the striped form the rear of the the thig has large light spots circled by a thin black ne work, the throat and chest have small dark spot the sides have dark spots, the disc of the thu finger is almost as wide as the tympanum, ar males lack vocal sacs. In the nonstriped form, tl rear of the thigh has small light spots on a blac background, the throat and chest may be suffuse with dark pigment but lack distinct spots, the sidt have light spots surrounded by a dark networ the disc of the third finger is much narrower th; the tympanum, and the males have vocal sac Differences in body proportions are shown in T ble 9. We also observed differences between the tv, 24 FIELDIANA: ZOOLOG Table 9. Comparison of striped and nonstriped forms of Polypedates leucomystax (sensu lato) from Buon Luoi, Vietnam. Striped Nonstriped SVL males Range Mean ± SE N 49.8-61.6 53.93 ± 0.53 25 SVL females Range Mean ± SE N 71.6-79.3 75.15 ± 0.87 T/SVL males Range Median N 0.50-0.55 0.536 11 T/SVL females Range Median N 0.50-0.57 0.549 9 HW/SVL males Range Median N 0.30-0.33 0.316 14 HW/SVL females Range Median N 0.32-0.35 0.326 9 Pympanum/SVL males Range Median N 0.061-0.071 0.066 13 rympanum/SVL females Range Median N 0.057-0.073 0.068 9 )iscF3t/SVL males Range Median N 0.057-0.064 0.061 13 )iscF3t/SVL females Range 1 Median N 0.054-0.070 0.062 9 49.1-62.4 55.35 ± 0.80 21 71.7-86.4 78.88 ± 1.90 0.46-0.51 0.499 10 0.48-0.54 0.517 9 0.31-0.35 0.326 17 0.31-0.36 0.343 10 0.066-0.093 0.077 17 0.071-0.080 0.075 10 0.045-0.057 0.051 16 0.051-0.059 0.054 10 NS NS <0.01 <0.02 0.03 NS <0.01 <0.01 <0.01 <0.01 * Significance level. For proportions, the Mann- Whitney (/-test was used. NS = not significant. t Width of disc of third finger. tjorrns in habitat distribution at Buon Luoi. The i onstriped form was found mainly in open situa- tions: in rice fields (26 individuals), at the border ;tetween rice fields and forest (8), along a village [ oad (4); only 12 were found in forest. All indi- viduals (31) of the striped form were found in orest at swampy lakes feeding forest streams. i Tie two forms co-occurred at the lake feeding MGER ET AL.: FROGS OF VIETNAM Stream N3 (see General Environment above) — 26 striped, seven nonstriped, and at the lake feeding Stream Nl — four striped and three nonstriped. At Tarn Dao, in northern Vietnam, we found the same two forms. The striped form there has large white spots surrounded by a thin dark net- work on the rear of the thigh, lacks vocal sacs (one adult male), and has body proportions match- 25 ing the striped form at Buon Luoi: T/SVL > 0.54, tympanum/SVL < 0.068, and width of disc of third finger > 0.067 (compare with Table 9). The nonstriped form at Tarn Dao has small light spots on a dark background on the rear of the thigh, has vocal sacs (one adult male), and has body pro- portions matching the same form at Buon Luoi: T/SVL < 0.50, tympanum/SVL >0.071, and disc of third finger <0.056. These two forms behave as distinct species; there are both clear morphological differences and significant ecological isolation. Various authors have tried to clarify the relationship of P. leuco- mystax (Gravenhorst) and P. megacephalus (Hal- lowell), the names commonly applied to these frogs, and several have treated them as subspecies (e.g.. Pope, 1931; Inger, 1966). Matsui et al. (1986) related acoustic and karyological differ- ences between Bornean and Taiwanese popula- tions to morphological differences and concluded that two species were involved; these authors ap- plied the name P. leucomystax (type locality Java) to the striped Bornean frogs, and P. megacephalus (type locality Hong Kong) to the nonstriped Tai- wanese frogs. Matsui et al. (1986) also suggested that there were differences between Taiwanese and mainland Chinese populations and perhaps even differentiation among the mainland popula- tions. In fact, the striped frogs from Buon Loy have the thigh pattern that Matsui et al. (1986) attributed to the nonstriped P. megacephalus, whereas the nonstriped frogs from Buon Loy have the thigh pattern that the same authors attributed to the striped Bornean frogs. Our analysis shares the main weakness of previous taxonomic discus- sions of these tree frogs, namely, failure to in- clude samples from throughout the wide range of P. leucomystax (sensu lato). However, we believe that the demonstration of morphologically and ecologically distinct populations from a single lo- cality is ample proof that at least two species are at issue. The absence of vocal sacs in one of these Buon Luoi populations suggests that acoustic dif- ferences are also likely. Philautus abditus, new species (Fig. 9) Holotype — fmnh 252833, collected at Buon Luoi, An Khe District, Vietnam, 7 May 1995 by Ilya Darevsky and Nikolai Orlov. Paratypes — fmnh 252834 and mvz 222118, 222121, adult males; mvz 222119-20, adult fe- males; fmnh 252835-38 and mvz 222101, juve- niles. All were collected at the type locality o shrubs and grassy vegetation 2-10 m from fo ested streams. Etymology — Specific name from abditus L hidden, concealed, referring to the bold blac spots on the legs that are hidden when the limt are flexed. Diagnosis — A Philautus with extensive wel bing on the foot; tympanum completely hidde under skin; large black spots on anterior and po; terior faces of thigh, not visible when legs ai flexed; no dermal fringes or tubercles on limbs; Description — Habitus stocky; snout rounded, feeble prominence at end; nostril closer to tip c snout than to eye; canthi distinct, rounded, cor stricted behind nares; lores oblique, concave; ey diameter greater than snout; interorbital equal t eyelid; tympanum not visible, but present unde skin; no vomerine teeth. Finger tips with roun discs having circummarginal grooves; disc c third finger wider than tympanum (when skin i lifted over tympanum); three outer fingers wit rudiment of web at base; subarticular tubercle conspicuous. Discs of toes smaller than those c fingers; web of first toe to edge of subarticul; tubercle or slightly beyond that, on second tc between subarticular tubercle and disc, on thir and fifth toes to edge of distal tubercle, to betwee tubercle and disc, or to base of disc; on fourth tc web to distal edge of middle tubercle; inner met; tarsal tubercle low, oval; no outer metatarsal ti bercle. Skin smooth dorsally and laterally; weakly curved supratympanic fold; ventrally fine ly granular; no dermal fringes, flaps, or tubercle on limbs. Color in preservative pinkish brown or gra dorsally and laterally on head and body; tempon region darker than dorsal ground color; wide ve: tical dark bar from center of eye to lip; seconi narrower dark bar from anterior corner of eye t lip; labial bars obscure in some individuals; a obscure dark interobital mark narrowing over oc cipital region; a dark gray mark beginning be tween shoulders, bifurcating in lumbar area, an ending in a large black spot low on side; ventrall head, body, thighs, and tarsi grayish brown; dors; surface of limbs same ground color as body; dar crossbars of thigh and calf ending in large cor spicuous black spots on anterior and posterior sir faces of thigh and ventrally on calf; proxim; black spot on anterior face of thigh, extendin into groin. Males lack nuptial pads; vocal sac opening slightly elongated, but not slit-like; vocal sac me 26 FIELDIANA: ZOOLOG" Fig. 9. Philautus abditus, new species. Paratype, male, 28.1 mm. II3ER ET AL.: FROGS OF VIETNAM 27 dian. The absence of the nuptial pad is probably not a reflection of seasonal reduction because adult males were collected both in the wet and dry seasons. Measurements — SVL of males 26.4-28.1 mm (mean 27.49, N = 4), of females 27.3-29.6 (N = 2); T/SVL 0.46-0.50 (median 0.484, N = 6); HW/ SVL 0.37-0.41 (median 0.391, N = 6); HL/SVL 0.35-0.39 (median 0.364, N = 6). Measurements of holotype: SVL 28.1, T 12.9, HW 10.4, HL 9.8, and eye diameter 4.3. Comparisons — This species lacks the m. cuta- neus pectoris, and its m. geniohyoideus medialis is free of the lateralis portion. These two features, plus the absence of nuptial pads, relate P. abditus to Dring's (1987) P. vermiculatus group. The combination of large black spots on hidden sur- faces of the hind limb and the completely hidden tympanum distinguishes P. abditus from other species in the P. vermiculatus group and from all other Philautus in Southeast Asia. Species with the tympanum completely hidden include P. ban- aensis Bourret, P. gryllus Smith, P. parvulus (Boulenger), P. kempiae (Boulenger), and possi- bly P. annandalii (Boulenger). None of these has the heavy black spots on the flash surfaces of the leg that characterize P. abditus. In addition, P. annandalii, P. parvulus, and P. kempiae differ from P. abditus in having webbing confined to the bases of the toes, and P. banaensis differs in hav- ing dermal fringes along the limbs. The most vari- able of this group in terms of coloration is P. gryl- lus, but P. abditus is invariant in its black spot- ting, a pattern that is not included in the known repertoire of P. gryllus (Smith, 1924; Bourret, 1942). Philautus parvulus (Boulenger) Ixalus parvulus Boulenger, 1893:339 — Karen Bia- Po, Burma. Rhacophorus (Philautus) parvulus Ahl, 1931: 70. Philautus parvulus Bourret, 1942:451. A series from Tarn Dao matches Boulenger's (1893) description point for point except for two characters: (1) the nostril is slightly closer to the tip of the snout than to the eye (midway between eye and end of snout in the types), and (2) the anterior rim of the tympanum is visible (tympa- num hidden in the types). Diagnostic characters are: small size (SVL of adult females 21-25 mm); toes short and web reaching base of distal sub - ticular tubercles of third and fifth toes or not tit far; thigh with a single, wide black bar crossif rear, dorsal, and anterior faces; lumbar region w l white area enclosed by heavy dark brown or bla : spots. We use the Tarn Dao specimens to expal the species description. M. cutaneus pectoris absent. Habitus rath stocky. Snout projecting only slightly, notched t the labial margin of the tip; diameter of eye sho • er than length of snout; tympanum partially c- scured by the skin, posterior rim hidden; no v« merine teeth. Tongue without erect papilla, cl with oval, smooth area near the front of the ce- ter; modified area set off from adjacent tissue a semicircular groove at its anterior margin a I by a narrower groove at its posterior. Disc of tlul finger equal to or wider than tympanum. Oui margins of limbs smooth, no tubercle or spur I heel. Although the webbing reaches the distal su articular tubercles of the third and fifth toes, t| actual extent of webbing is small because the oi- er metatarsals are fused for most of their length SVL measurements of males 19.0-22.1 nj (mean ± SE 20.20 ± 0.38, N = 9), of femali 21.1-25.2 (23.30 ± 0.49, N = 8); T/SVL 0.4 0.52 (median 0.468, N = 17); HW/SVL 0.3- 0.44; HL/SVL 0.38-0.39. Nuptial pad white, y vety, on medial and dorsal surfaces of first finj: from base to beginning of last phalanx. Ferna; with weakly indicated rostral cone. Six ova e cased in gelatinous envelopes on left side of lai est female; diameter of ova 2.4-2.9. Six enlargl ova on left side of smallest female; diameter f ova 2.3-2.75. This is the first record of the species from \\ etnam. The known range through the mountainc; area from Burma through northern Thailand (Tt lor, 1962) to northern Vietnam is a common d tributional pattern in Southeast Asian amphibia; (Inger, 1999). Philautus maosonensis Bourret Philautus maosonensis Bourret, 1937:51 — M Son, Tonkin, Vietnam; Bourret, 1942:467. A specimen from Tarn Dao agrees very close with Bourret's (1937) description. Bourret stat that the female had a smooth dorsum, whereas t male was covered with small tubercles on t head, back, and limbs. The Tarn Dao female i 28 FIELDIANA: ZOOLOC * * ^ % Fig. 10. Rhacophorus annamensis Smith. Female, 84.5 mm. ;mbles the male of the type series in this char- ter. Three characters set this species aside from all incept a few continental Philautus: distinct tym- mum, toes almost completely webbed, fingers ith at most a rudiment of web. In P. maosonen- s the tympanum is about two-thirds the diameter the eye; the disc of the third finger is about l/o-thirds the diameter of the tympanum; the can- us rostralis is rounded or indistinct, the toes (ex- :pt for the fourth) are webbed to the base of the scs; and a large, dark, reddish brown mark oc- ipies most of the back behind the shoulders. " pult females are 29-32 mm SVL. Philautus car- ' ■ensis (Boulenger) (type locality Karen Hills, iirma) has the tympanum half the diameter of s b eye, none of the toes webbed to the discs, and apold ) (-shaped marking on the back; it appears ill be larger (38 mm) than P. maosonensis. Phi- \i\ttus argus (Annandale) (type locality northeast- ern India) has a smaller tympanum (about one- third diameter of eye), a distinct canthus, and a pale reticulation on the back. Measurements of the Tarn Dao female: SVL 29.0, T 15.7, HW 10.3, HL 11.1, eye 4.1, and tympanum 2.4. The female contained large ova that had a black hemisphere. Rhacophorus annamensis Smith (Fig. 10) Rhacophorus annamensis Smith, 1924:229 — Da- ban, Annam, Vietnam. Rhacophorus pardalis annamensis Bourret, 1942:443. The present sample from Buon Luoi, Tram Lap, and Kon Cha Ran is the first record of this species since the collection of the unique male holotype. It agrees closely with Smith's (1924) original de- scription. According to Smith, one distinctive character of R. annamensis is the slope of the (#GER ET AL.: FROGS OF VIETNAM 29 snout in front of the nostrils, but this feature is clearly sexually dimorphic. All of the males in the sample at hand have the snout gently sloping, in profile, to a sharp point projecting beyond the lower jaw. In the females, however, the snout is rounded, not sloping, and not projecting. Most of the features of coloration that were noted by Smith (1924) from the holotype are subject to much variation, as will be shown below; the dark webbing Smith observed appears to be constant. Wolf's (1936) treatment of R. annamensis as a subspecies of R. pardalis was an egregious case of superficial phenetics, especially because he had not seen a specimen and because Smith (1924) had clearly pointed out significant differences be- tween the two species. Besides the differences noted by Smith, the two species are remarkably different in coloration, with red and orange being major components in R. pardalis and brown pre- dominant in R. annamensis. We use this large new series to amplify the original description. Habitus stocky; snout sharply pointed in males, obtusely pointed in females (see also above); nos- tril closer to tip of snout in females and some males, equidistant between eye and tip of snout in most males; canthi distinct, rounded or sharp, weakly constricted; tympanum distinct, less than half diameter of eye. Discs of fingers rounded, those of outer fingers wider than tympanum; web reaching edge of subarticular tubercle of first fin- ger, to disc of three outer fingers; subarticular tu- bercles conspicuous. All toes webbed to base of discs; low inner, but no outer metatarsal tubercle. Heel in half of the sample with a small, bluntly pointed projection; no supra-anal projection; in- fra-anal area usually with two to three long tu- bercles, bases of tubercles rarely fused. Color in life brown dorsally and laterally, with irregular black, red, or green spots on the back and faint dark crossbars on limbs; webbing dark grayish brown dorsally; ventrally ivory or cream; iris silvery, with a pinkish cast and irregular, thin dark lines. In preservative medium to dark gray, with obscure darker markings; lower half of sides white with dark marbling or network; ventrally white, usually with few small dark spots on throat and chest; limbs with obscure dark crossbars, not visible in some individuals; front and rear of thigh brown with irregular lighter markings, these sur- faces rarely pinkish white with black spots; web dark brown or gray. Males with white nuptial pad on dorsal and me- dial surfaces of base of first finger; vocal sac opening round. Table 10. Sexual dimorphism of body proportioi in Rhacophorus annamensis Smith. Range Median N T/SVL Males 0.47-0.54 0.498 9 Females 0.50-0.54 0.521 9 HW/SVL Males 0.30-0.33 0.309 10 Females 0.31-0.33 0.326 9 HL/SVL Males 0.33-0.36 0.340 8 Females 0.32-0.34 0.324 9 SVL of males 57.8-71.7 (mean ± SE 64.04 1.37, N = 10), of females 75.0-86.8 (82.01 1.26, N = 9). Body proportions are subject to se ual dimorphism; the differences between the sex . for each of the ratios (Table 10) are significant : the P < 0.02 level (Mann- Whitney LMest). T sexes do not differ in relative size of the tymp num: tympanum/SVL of males 0.049-0.062, ; females 0.051-0.056. Rhacophorus annamensis was the most coi: monly observed tree frog in the Buon Luoi- Tram Lap — Kon Cha Ran area. Pairs in amplex were seen in both spring and autumn. Nests we placed on rock faces of stream banks above qu: water and on trunks of trees in swampy floo plains. Rhacophorus baliogaster, new species (Fig. 1 Holotype — fmnh 252839, an adult male fro Buon Luoi, An Khe District, Vietnam, collect 27 April 1995 by Ilya Darevsky and Nikolai C lov. Paratypes— mvz 222040-42, 222100, 22210 three adult females and one adult male; FMr 252840, subadult female. All from the type loc; ity. Etymology — Specific name from balios G spotted or dappled, and gaster Gr., belly. Diagnosis — Third finger webbed to edge proximal subarticular tubercle, fourth toe to b tween the subarticular tubercles or to base of di tal tubercle; no dermal flaps or ridges on limlr no dermal appendages around vent; ventral si faces of head and trunk white with black spot females with a strong rostral cone. Description — Snout obtusely pointed, mal with a weak median prominence, females with 30 FIELDIANA: ZOOLOG Fig. 11. Rhacophorus baliogaster, new species. Holotype, male, 33.0 mm. iER ET AL.: FROGS OF VIETNAM B.0L0GY LIBRARY '01 BURRILL HAH 31 strong rostral cone; nostril closer to tip of snout than to eye, in a raised prominence; canthi round- ed, constricted; lores oblique, concave; eye prom- inent, diameter equal to snout in females, longer than snout in males; interorbital about equal to eyelid; tympanum distinct, less than half diameter of eye; vomerine teeth in oblique groups, near to but not touching anterior corner of choanae. Fin- gers with round discs having circummarginal grooves; disc of third finger equal to or smaller than tympanum; fingers webbed at bases; subar- ticular tubercles conspicuous. Discs of toes small- er than those of fingers; webbing on first toe to between subarticular tubercle and base of disc; webbing on second, third, and fifth toes just short of base of discs, on fourth toe to distal subarti- cular tubercle or not quite so far; low inner but no outer metatarsal tubercle. Skin smooth dorsally and laterally, no tubercles on eyelid; supratympanic fold weak, curved; throat smooth, chest and abdomen coarsely gran- ular; limbs without dermal flaps, folds, or con- spicuous tubercles; very weak tubercles on outer edge of tarsus in some individuals; no dermal ap- pendages around vent. Color in life dark brown above, with small ir- regular darker spots; usually blackish spots at shoulder continued as broken streak ending near groin; area below canthus dark brown; a broad dark brown bar below front half of eye; iris gold- en brown in upper third, reddish brown in re- mainder; sides lighter brown than back, black spots low on sides. In preservative gray dorsally with pinkish tinge, darker on snout; an obscure dark interorbital bar; dark markings on back also obscure and variable; most have faint dark streak above shoulder, ending in a large dark blotch on side at groin; venter whitish with conspicuous dark spots, varying in size from smaller than disc of first finger to larger than disc of third finger; limbs dorsally gray or brown, with dark crossbars; underside of leg heavily marbled with brown; rear of thigh gray brown, with or without small irreg- ular light markings. Measurements — SVL of males 33.0-33.3 (N = 2), of females with mature, coiled oviducts 35.8-41.5 (mean 38.57, N = 4); T/SVL 0.44-0.52 (median 0.498, N = 6); HW/SVL 0.34-0.37 (me- dian 0.356, N = 6); HL/SVL 0.34-0.39 (median 0.367, N = 6). Nuptial pad of male white, on dorsal and medial surfaces of first metacarpal. Vo- cal sac openings slightly elongated. Measurements (mm) of holotype: SVL 33.0, T 15.8, HW 11.8, HL 11.4, eye diameter 5.2, tyi panum 2.2. Larvae — Two lots of tadpoles from Buon Lu are assigned tentatively to this species. The; specimens do match any larval species describe by Bourret (1942). The most advanced tadpole, stage 39, has fingers webbed at the bases as i adult R. baliogaster. Headbody oval, slightly flattened above and b low; snout obtusely pointed; eyes dorsolaten; spiracle midway between eye and end of bod, pointed slightly upward, tube not free of boe wall. Oral disc subterminal, one-third of boe width; papillae short, crowded, not interrupted midline; labial teeth 6(2-6)/5(l); jaw sheat; black in marginal third, upper with a weak co vexity. Tail lanceolate; dorsal fin origin at end body, rising slightly to mid-length; tail taperii gradually to rounded tip. Color in preservati gray dorsally; a fine dusting of melanophores ve trally; caudal muscle finely dusted with melan phores, fins unpigmented. HBL 13.3 (stage 35) 15.5 mm (stage 39); total length 34.6-40.3 mi HBL/total 0.36-0.39; tail depth/tail length 0.3 0.36. Remarks — All specimens were collected on tl banks of forest streams perched on grassy veg tation 20-50 cm above the surface and about 3 from the water. The two lots of tadpoles were cc lected in a swampy pond in forest at Buon Loj Comparisons — The reduced webbing betwe< the outer fingers distinguishes R. baliogaster fro R. annamensis Smith, R. pardalis (Gunther), . reinwardtii (Schlegel), R. nigropalmatus Boule ger, R. maximus Boulenger, and R. robinsoni Bo lenger. In addition, all of these fully webbed sp cies differ from R. baliogaster in lacking blat : ventral spotting as the typical pattern and in ha ing conspicuous dermal flaps or ridges on tl limbs. Rhacophorus baliogaster differs from oth Southeast Asian species of Rhacophorus that ha^ less than fully webbed outer fingers — that is, . appendiculatus (Gunther), R. bipunctatus Ahl, . calcaneus Smith, R. notater Smith, R. taronens Smith, R. turpes Smith, and R. verrucosus Bo lenger — in the absence of dermal flaps or fring' on the limbs. Rhacophorus baliogaster also dij fers from all of the previously mentioned speci< (except R. appendiculatus and R. verrucosus) i the presence of a rostral cone in females. The la two species differ from R. baliogaster in lackir black spotting ventrally. 32 FIELDIANA: ZOOLOG Fig. 12. Rhacophorus bipunctatus Ahl. Male, 37.3 mm. Rhacophorus bimaculatus (Peters) Rhacophorus bipunctatus Ahl (Fig. 12) eptomantis bimaculatus Peters, 1867:32 — Agu- son Valley, Mindanao, Philippine Islands. Rhacophorus (Philautus) bimaculatus Ahl, 1931:64. Philautus bimaculatus Bourret, 1942:471. A single adult male (40.2 mm) from Buon Luoi lows comprehensive agreement with R. bimac- 'atus from Borneo and the Philippines. The col- 1 ation of this species is distinctive: medium dark own dorsally; side of head darker with a large ibocular enamel white or bluish spot; flash sur- ces at front and rear of thigh black with small uish white spots. The Buon Luoi frog differs om the described coloration of R. bimaculatus eters, 1867; Inger, 1966) only in having the ven- * heavily suffused with brown (white in Bornean d Philippine frogs). The Buon Luoi frog is larg- than those from Borneo (28-35 mm), but like sm it has a vocal sac but no nuptial pad. This the first report of the species north of peninsular iiailand. tlGER ET AL.: FROGS OF VIETNAM Rhacophorus bimaculatus Boulenger, 1882:90 — Khasi Hills and Assam, India. Rhacophorus bipunctatus Ahl, 1927:46 (substi- tute name). Rhacophorus rhodopus Liu and Hu, 1959: 525 — Meng-yang, Yunnan, China. Rhacophorus reinwardtii bipunctatus Bourret, 1942:446. Liu and Hu (1959) distinguished R. rhodopus from R. bimaculatus Boulenger (/?. bipunctatus Ahl) on the basis of red webbing, pointed snout, and reddish brown dorsal coloration; they as- sumed that Boulenger 's species was green, pos- sibly because Boulenger (1882) reported that it resembled R. reinwardtii. The same comment by Boulenger may have led Liu and Hu to assume that R. bipunctatus had a round snout; in fact, R. bipunctatus has a pointed snout. None of the char- acters that, according to Liu and Hu (1959), sep- arated R. rhodopus from R. bipunctatus do, in 33 fact, distinguish these taxa. All of the characters mentioned in the original description of R. rho- dopus agree with those of R. bipunctatus. Although the present sample (all from Buon Luoi) is the first one reported from Vietnam, it merely fills out the known range: Assam, Burma (Boulenger, 1893), northern and eastern Thailand (Taylor, 1962; Inger & Colwell, 1977), and Yun- nan (Liu & Hu, 1959). In terms of size, webbing, leg length, vomerine teeth, relative size of tym- panum (less than half eye diameter), and other characters, the specimens from Vietnam and Thai- land (fmnh 187509-14, 187516-17) agree very closely with those from Assam (fmnh 72406-07) and Mt. Karen, Burma (zmh Hamburg, Germany [zmh] 736 [3]), a locality from which Boulenger (1893) reported the species. The Vietnamese frogs differ from the Thai frogs in one respect; in the former, the dermal appendage at the tibiotarsal joint is a simple transverse fold, whereas in the Thai specimens the transverse fold is drawn out to a point. Males in the Buon Luoi sample measure (SVL) 33.1-39.8 mm (mean ± SE 36.16 ± 0.24, N = 37), females 46.0-52.2 (mean 47.93, N = 4). TV SVL of males 0.45-0.51 (median 0.484, N = 14), of females 0.48-0.49 (N = 4); HW/SVL of males 0.34-0.38 (median 0.353, N = 14), of females 0.32-0.36 (N = 4). Tadpoles and metamorphs from stages 25-44 have labial teeth 6(2-6)/3(l), short marginal pa- pillae continuous across the lower lip. HBL (stages 31-40) 16.7-18.3 mm, total length (stages 31-40) 45.4-55.4. HBL/total length (stages 31- 40) 0.33-0.38. We found this species at Buon Luoi in forest up to 20 m from a large pond and at Tram Lap along swampy banks of a forest stream. Males called from trees about 1-2.5 m above the surface. Density in both October-December 1993, and March-May 1995 was very high, and it was pos- sible to see >10 males vocalizing in an area of 20-25 m2. Larval samples were taken in ponded portions of forested streams at Buon Luoi. The absence of this abundant species at Kannack (400 m) and Kon Cha Ran (1000-1500 m) suggests that its main zone of distribution in these southern Annam mountains lies in the belt of 600-900 m elevation. Rhacophorus calcaneus Smith (Fig. 13) Rhacophorus calcaneus Smith, 1924:228-Lang- bian Peaks, Annam, Vietnam. These frogs from Buon Luoi (7), Tram Lap ('. and Kon Cha Ran (15) appear to be the first spe imens of R. calcaneus collected since the uniq holotype was described by Smith (1924). Th match Smith's description and figure in diagnosi characters, such as the sharply pointed, slopi); snout, the sharp, light-edged canthi, the loij pointed projection at the heel, and the extent i webbing. None of the new specimens has t black axillary spot and small yellow spots at t rear of the back found on the holotype. Given t variation within the new sample, those differenc appear to fall within the range of individual vaj ation. Smith (1924) thought that R. calcaneus was member of the Rhacophorus bimaculatus Boule . ger (= R. bipunctatus Ahl) species group. Ho\i ever, the sharply pointed, sloping snout and sha \ canthi, webbing, and dermal fringes along tl limbs are more similar to Rhacophorus angu,'. rostris Ahl. The present large sample allows ar plification of the original description. Habitus stocky; snout sharply pointed, slopii downward in front of nostrils, projecting beyoi lower jaw; nostril closer to tip of snout than eye; canthi sharp, weakly constricted; lores slo ing, weakly concave; diameter of eye less th; length of snout; interobital wider than eyelid; tyr panum less than half diameter of eye; vomerii teeth in oblique groups beginning near anteri corners of choanae. Finger tips with large, roui discs, those of outer fingers wider than tymp num; first finger webbed to subarticular tubercl second finger to just below disc, third finger distal edge of distal subarticular tubercle laterall and fourth finger midway between disc and |i bercle; subarticular tubercles conspicuous. Dis< of toes smaller than those of fingers; web to ba: of discs of first three toes laterally and fifth t( medially, narrowly to disc laterally on fourth to a low oval inner but no outer metatarsal tubercl Skin generally smooth above; a straight, stror supratympanic fold extending just beyond level i\ axilla; throat finely granular; belly coarsely grai ular; a smooth-edged ridge of skin along out edge of lower arm and fourth finger, ending elbow in a dull point; a similar, but narrower rid£ along the tarsus and fifth toe; a long, pointed pr< jection at tibiotarsal joint; a low supra-anal ridg- several large, pointed tubercles below vent. Color in life dorsally gray brown to reddis brown, with oval or round dark brown spots, ( with small enamel white spots, or pinkish dorse lateral bands; lower part of sides with thin blac 34 FIELDIANA: ZOOLOG' Fig. 13. Rhacophorus calcaneus Smith. Male, 35.4 mm. letwork enclosing large white spots; iris light tan >r yellowish brown in upper half, darker grayish irown below; front and rear of thigh reddish >rown. In preservative ground color dorsally gray t pinkish; no large black spots laterally in present ample; lateral black network faint in some indi- iduals; supra-anal ridge pinkish white; ventrally /hite usually with fine dark dots or immaculate /hite; limbs with dark crossbars; ventral surfaces f thigh and calf with small black dots; front and ;ar of thigh pale pinkish or flesh-colored, im- laculate (10) or dotted with black (4), or rear of ligh dark brown (11). Male nuptial pad white, on dorsal and medial irfaces of first metacarpal; vocal sac openings ongated. SVL of males 35.2-40.1 mm (mean ± SE 3.20 ± 0.26, N = 10), of females 48.5-55.0 nean 52.93, N = 4). T/SVL 0.49-0.56 (median 516, N = 14), HW/SVL 0.36-0.39 (median 373, N = 14). Five juveniles (females 37.0-40.4 mm) were collected at Tarn Dao. They share all of the di- agnostic features of frogs from the central high- lands: sharp, sloping snout; weak dermal ridge along the lower arm; weak, pinkish supra-anal ridge; discs of outer fingers wider than tympanum; long pointed projection at heel. All of these ju- veniles have black dots on the white venter and underside of the thigh and have the rear of the thigh immaculate flesh-colored. At Buon Luoi and Kon Cha Ran we found this species as single individuals in trees about 2-3 m above ground and 3-15 m from banks of forested streams. Two pairs were caught in amplexus perched on branches 3 m above the ground. Rhacophorus exechopygus, new species (Figs. 14, 15) Holotype — fmnh 252841, adult male collected at Tram Lap, An Khe District, Gia-Lai Province, IJGER ET AL.: FROGS OF VIETNAM 35 Fig. 14. Rhacophorus exechopygus, new species. Holotype, male, 46.5 mm. Vietnam, 21 April 1995 by Ilya Darevsky and Ni- kolai Orlov. Paratypes — fmnh 252842-43, adult males col- lected at the type locality. Etymology — Specific name from exechos Gr., jutting out, and pygos Gr., buttocks, referring to the infra-anal projection. Diagnosis — A Rhacophorus with fully webbed outer fingers; a wide, transverse, white infra- anal projection; forearm and tarsus with wide, weakly crenulated dermal ridge; rear face of thigh black; lower half of side with black spots or band. Description — Habitus stocky; snout pointed, projecting (only males known); nostril about mid- way between eye and tip of snout; canthi round, weakly constricted; lores sloping, concave; eye prominent, diameter less than snout; interorbital wider than eyelid; tympanum distinct, less than half the diameter of the eye; vomerine teeth in transverse groups, touching anterior corner of choanae. Discs of fingers round, with circummar- ginal grooves; disc of third finger larger than tyn panum; three outer fingers webbed to discs, fir finger webbed to subarticular tubercle; subartici lar tubercles conspicuous. Discs of toes small than those of fingers; all toes webbed to disc inner metatarsal tubercle low, oval, no outer met; tarsal tubercle. Skin smooth dorsally, with very fine, colorle: asperities (in males only?); limbs with scattere weak tubercles dorsally; throat smooth or weak] rugose; chest and abdomen coarsely granular; ou er margin of forearm and fourth finger and tarsi and margin of fifth toe with wide, weakly creni late dermal fringe; heel with a narrow transven ridge, usually ending laterally in a weak projei tion; a strong, wide, white-edged, horizontal infr; anal dermal projection. Color in life gray to brown dorsally, unifon or with obscure dark blotch at rear of head; limb with brown or reddish brown crossbars; tubercle on limbs pinkish; iris silvery with fine dark nei 36 FIELDIANA: ZOOLOG^ f Fig. 15. Rhacophorus exechopygus, new species. Paratype, male, 46.4 mm. & vork in upper half, darker in lower half. In pre- tervative uniform purplish gray on all dorsal sur- faces and upper half of side; lower part of side vith black band from axilla to groin, or black at ixilla and groin only, or with a row of large black ipots; one specimen with light spots (blue in life?) fjvithin black area at groin; throat and abdomen . jvhite; rear face of thigh black; front face of thigh ) vhite or black; ventral surface of calf black or i Ivhite; dorsal surface of outer fingers and toes i vith coloration of back, inner digits without dark i figment; web dark on dorsal surface, colorless or j ale reddish ventrally. i Males with white nuptial pad on dorsal and me- !ial surfaces of first finger; vocal sac opening longate. Measurements — Holotype: SVL 46.5, T 22.4, fW 15.6, HL 16.8, snout 7.5, eye 6.7, tympanum .75, and interorbital 5.6. Paratypes: SVL 45.9- vlGER ET AL.: FROGS OF VIETNAM 46.4. T/SVL 0.45-0.48 (N = 2), HW/SVL 0.34 (N = 2), and HL/SVL 0.36-0.38 (N = 2). Remarks — All three specimens were found about 3 m above the ground in trees on swampy banks of a forest stream. Comparisons — The fully webbed outer fingers of Rhacophorus exechopygus match the character in R. annamensis Smith, R. dulitensis Boulenger, R. maximus Boulenger, R. nigropalmatus Boulen- ger, R. partialis Gunther, R. prominanus Smith, R. reinwardtii (Schlegel), and R. robinsoni Boulen- ger. However, R. exechopygus is the only species having a projecting, infra-anal dermal ridge; R. dulitensis, R. prominanus, and R. reinwardtii have a supra-anal dermal ridge, whereas the others list- ed have no dermal appendage in this area. In ad- dition, R. exechopygus is smaller than R. maximus (males 80-86 mm), R. nigropalmatus (75-91), and R. annamensis (55-65). Rhacophorus execho- 37 Fig. 16. Rhacophorus reinwardtii (Schlegel). Female, 85.2 mm. pygus differs from other syntopic species of Rha- cophorus in dermal appendages (R. verrucosus Boulenger, R. bipunctatus Ahl, R. annamensis Smith, R. calcaneus Smith, and R. baliogaster, new species, without a wide infra-anal projection) and coloration (R. baliogaster with boldly spotted abdomen, R. bipunctatus with black spots on the side, R. calcaneus with dark dorsal markings, and R. annamensis with black dotted throat). Other Rhacophorus from Southeast Asia differ from R. exechopygus in (1) the type of dermal appendages (R. appendiculatus [Gunther] with strong crenu- lated fringe or row of conical tubercles along fore- arm and tarsus; R. taronensis Smith and R. turpes Smith without infra-anal projection), and (2) col- oration (none of the listed species has an immac- ulate dorsal pattern). Rhacophorus reinwardtii (Schlegel) (Fig. 16) Hyla reinwardtii Schlegel, 1837:105 — Java (re- stricted by Wolf, 1936). Rhacophorus reinwardtii Tschudi, 1838:73. Although these are the first specimens recorde from Vietnam, their occurrence is not surprisir because the species has been known from Yunns for some time (Yang, 1991). The present samp from Buon Luoi, consisting of eight adult femalt and three adult males, matches specimens froi Java in coloration and dermal appendages. The are green dorsally, have webbing of the three ou er fingers and of the foot black dorsally, and ha\ a large black area in the axilla. The front and re; of the thigh are immaculate cream or white. The have a wide, smooth-edged fringe of skin alon the outer edge of the forearm, a similar, thoug narrower fringe along the tarsus, a squarish fring at the heel, and a conspicuous, transverse projet tion above the vent. The supra- anal flap immed ately distinguishes these frogs from the other larg green species of Rhacophorus in Southeast Asi; R. nigropalmatus Boulenger and R. maximi. Gunther. The Vietnamese specimens are larger than J; van frogs: SVL of males 65.6-69.7 mm (N = 3 of females 81.8-92.7 (mean ± SE 87.66 ± 1.3* N = 8). Two males with vocal sacs from Jav 38 FIELDIANA: ZOOLOG> Table 1 1 . Comparison of Rhacophorus verrucosus and related forms. R. verrucosus Buon Luoi A*, bisacculus Thailand R. appendiculatus Borneo Tam 1 >ao SVL males Range Mean ± SE N 23.8-28.0 26.25 ± 0.40 13 29.2-35.5 6 29.6-37.0 53 29.1-33.5 31.59 ± 0.36 17 T/SVL Range Median N 0.46-0.52 0.480 7 0.51-0.54 0.528* 6 0.50-0.52 0.515 6 0.42-0.51 0.462t 17 HW/SVL Range Median N 0.37-0.41 0.402 8 0.35-0.39 0.361 6 0.31-0.37 0.346 17 rympanum/SVL Range Median N 0.045-0.062 0.056 9 0.063-0.071 0.065* 6 0.060-0.069 0.062* 6 0.051-0.060 0.056 6 * Difference between R. verrucosus and indicated samples significant at P < 0.03; Mann- Whitney (/-test, t Difference between Tam Dao and Buon Loy samples significant at P < 0.05; Mann-Whitney U-test. $ Species not determined. See text discussion under Rhacophorus verrucosus. neasured 46.4-54.6 mm, and six adult females Torn Java were 56.2-65.7 mm (mean 63.15). rogs from Yunnan are the same size as those torn Vietnam: SVL of males 66-72 (N = 10), of emales 74-95 (N = 5) (Yang et al., 1991). The /ietnamese frogs also have relatively shorter legs T/SVL 0.48-0.53, N = 11), narrower heads HW/SVL 0.32-0.35), and shorter heads (HL/ ►VL 0.32-0.34) than Javan frogs (0.50-0.55, 1.36-0.39, and 0.34-0.37 for the three propor- ions, respectively; N = 8). Sexually active frogs planed down to the sur- ace of a large pond from vegetation 3-5 m above round. Pair formation occurred at the water's kirface. Oviposition took place on leaves of [ranches overhanging water. The foamy egg basses were 18-22 cm long. Most nests were pen at the end of April and early May. Vocaliza- : on and breeding activity were observed predom- inantly on rainy nights. 'hacophorus verrucosus Boulenger hacophorus verrucosus Boulenger, 1893:337 — Thao, Karen Hills, Burma. Rhacophorus appendiculatus verrucosus Bourret, 1942:418. Twenty frogs from Buon Luoi and two from 4GER ET AL.: FROGS OF VIETNAM Kon Cha Ran have all of the dermal features of R. appendiculatus (Giinther), except in reduced condition. All of the fringes are less protruding, especially the per-anal one, but they are all pres- ent. Most females of R. verrucosus have a distinct knob on the snout, but it is not as protuberant as in R. appendiculatus. The Thai form R. bisacculus Taylor is clearly closely related to R. appendiculatus. Both of these forms are larger than R. verrucosus and differ from it in certain body proportions (Table 1 1). Fe- males show comparable differences in SVL: R. verrucosus 29.8-35.8 (N = 6), R. appendiculatus (Borneo) 42.4-50.1 (N = 8); no females of R. bisacculus were available. In addition, vomerine teeth, which are always present in R. bisacculus and R. appendiculatus, are usually absent (15 of 20 examined) in Vietnamese R. verrucosus. The infra-anal dermal appendages of these species dif- fer; R. appendiculatus from the Philippines and Borneo has a conspicuous transverse flap with a papillate edge, R. verrucosus has two to six long whitish tubercles that may join basally, and R. bi- sacculus usually has no prominent projections in this area but may have several low, pointed tu- bercles. We amplify the original description on the basis of the Buon Luoi specimens. Finger discs round, that of third finger equal to diameter of tympa- num; first two fingers webbed at base, web of 39 third finger not reaching distal subarticular tuber- cle, web of fourth finger to base of distal tubercle; subarticular tubercles conspicuous. Disc of toes smaller than those of fingers; web to subarticular tubercle of first toe, to between tubercle and disc on second toe, between distal tubercle and disc of third and fifth toes, to just below distal tubercle of fourth toe. Dorsally with small scattered tuber- cles; heel with a low conical tubercle. Nuptial pad white, on dorsal and medial surfaces of first met- acarpal; vocal sac opening a short slit. One female had ovulated, gelatin-encapsulated, yellow ova; an exact count could not be made without dam- age, but there appeared to be fewer than 30 ova on the left side. The ova measured 2.5-3.0 mm. A sample of 21 frogs generally similar to this group of species was collected at Tarn Dao in northern Vietnam. They resemble R. bisacculus in having short tubercles or no dermal appendages below the vent, and they resemble both R. appen- diculatus and R. bisacculus in size (Table 1 1 ; SVL of four adult females 44.7-50.5 mm), in the large rostral cone of the female, and in having vomerine teeth. In each of these characters they differ from the Buon Luoi specimens. However, the Tarn Dao frogs differ from all of the others in this group in having very large dark spots occupying most of the abdomen and in having a shorter leg (Table 11). Specific assignment of the Tarn Dao popu- lation is problematic. They appear to be closest to R. bisacculus, but the intervening geographic po- sition of the Buon Luoi R. verrucosus makes as- signment to R. bisacculus uncertain. Liu and Hu (1959, 1961) and Yang et al. (1991) reported a species of this group from southern Yunnan under the name R. cavirostris (Giinther), which is otherwise known only from Sri Lanka and is considered to be a synonym of Rhacopho- rus microtympanum (Frost, 1985). Zhao and Ad- ler (1993) list this Chinese form as Philautus ca- virostris. SVL measurements of the Yunnan frogs (males 27-33.8, female 43 mm). The frilled tarsi, their lack of long infra-anal tubercles, and their possession of vomerine teeth (Liu & Hu, 1961; Yang et al., 1991) agree with those of the Tarn Dao frogs; unfortunately, there is no information on ventral coloration. Tran et al. (1981) and Nguyen et al. (1994) list- ed R. cavirostris as part of the fauna of northern Vietnam, although they gave no descriptive notes or comments that would confirm the identifica- tion. Their records are probably based on frogs conspecific with the present sample from Tarn Dao. Rhacophorus cavirostris (Giinther) should be removed from the faunal list of Vietnam, b for reasons given above the proper name of the northern Vietnamese frogs is uncertain. Most individuals from Buon Luoi were foui about 1 m above ground on shrubs 20-100 I from any stream; four were caught within 3 m a stream. Theloderma corticalis (Boulenger) (Fig. 17) Rhacophorus corticalis Boulenger, 1903:188- Man-Son Mountains, Tonkin, Vietnam. Theloderma corticalis Liem, 1970:94. Rhacophorus leprosus corticalis Bourn 1942:412. Two adult females (SVL 70.5, 70.6) with d veloping, strongly pigmented ova, and an adi male (SVL 69.2 mm) were collected at Tam Da They agree very closely with Boulenger 's (190 description. They are the same size as the tyj (SVL 70 mm), the finger discs are very wide, tl vomerine teeth are in small groups, and the ma lacks vocal sacs. However, there are slight diffe ences from the original description. Bouleng stated that the fingers lacked webbing and that tl toes were completely webbed; in the Tam D; specimens there is a rudiment of webbing b tween the third and fourth fingers, and full wel bing ends at the distal edge of the distal subart cular tubercle of the fourth toe. The other toes a webbed to the discs. Boulenger also reported th the tympanum was as large as the eye; in the thrt frogs at hand the tympanum is clearly smalL than the eye (see below). In the Tam Dao frogs, the disc of the third fii ger is equal to or wider than the tympanum. Boc proportions are: T/SVL 0.48-0.50, HW/SV 0.41-0.43, tympanum/SVL 0.069-0.078, disc ( third finger divided by SVL 0.069-0.080, tyn panum 0.60-0.69 of diameter of eye. The nupti; pad is a cream-colored mass on the mediodors. surface of the first finger from its base to the lev( of the subarticular tubercle, then medially to ju beyond the tubercle. Theloderma gordoni Taylor Theloderma gordoni Taylor, 1962:51 1— Doi Suthej Chiang Mai Province, Thailand. One female from Buon Luoi, one male froi 40 FIELDIANA: ZOOLOG^ Fig. 17. Theloderma corticalis (Boulenger). Female, 70.0 mm. ram Lap, and one female from Tarn Dao. These e clearly conspecific with the holotype of T. ndoni (fmnh 172248). The skin is extremely ugh, with many pustulose ridges and tubercles l top of the head, temporal region, back, and >rsal and lateral surfaces of the limbs. There is very dense cluster of large spinose tubercles or s behind the tympanum, and the chin and oat have small pustulose tubercles, in sharp ntrast with the smoothly granular chest and bel- The vomerine teeth are in small, widely sep- ted, oblique groups near the anterior corners of ip choanae. The distinct tympanum is slightly spre than half the diameter of the eye. The Tarn iJko female differs from the other two in having ylver of the dorsal warts coalesced into ridges. In life, these frogs were almost black dorsally i laterally, with the crests of some ridges on head and back light brown. In preservative, cluster of warts behind the tympanum is pale >wn. The palmar tubercles and prehallux are nfute (flesh-colored in life?), in sharp contrast to dark color of the remainder of the ventral sur- e of the hand; although the holotype is faded, s still possible to see a contrast between the 4 3ER ET AL.: FROGS OF VIETNAM base of the palm and the rest of the hand. Broad webbing on the first finger reaches between the subarticular tubercle and the disc, on the lateral margins of the second and third fingers and the medial margin of the fifth to between the distal subarticular tubercle and the disc, and on the fourth toe to the middle tubercle or between the middle and distal tubercles. The male measures 47.0 mm (SVL) and the two females 44.2 and 50.9 mm; the larger female has enlarged ova. The male has a nuptial pad of fine spinules cov- ering the medial and dorsal surface of the swollen prehallux and extending in a narrow line along the median edge of the first finger to the base of the disc. There are no vocal sac openings. In all of these characters (except the secondary sex characters of the male), these three specimens match the female holotype (SVL 48 mm) of T. gordoni. However, they also agree with descrip- tions of T. leporosa Tschudi (type locality Padang, Sumatra) except for having slightly less webbing and being somewhat smaller. Theloderma lepo- rosa from the Malay Peninsula reaches 65 mm and has nearly fully webbed toes (Boulenger, 1912). 41 Fig. 18. Theloderma stellatum Taylor. Female, 32.3 mm. One specimen was found on the trunk of a fall- en tree in a small forest clearing. Another was perched on the broad leaf of a shrub about 5 m from a forest stream. Theloderma stellatum Taylor (Fig. 18) Theloderma stellatum Taylor, 1962:514— Sebab, Chantaburi Province, Thailand. Khao Two males and two females from Tram Lap and three males, three females, and four juveniles from Buon Luoi are assigned to this species. The males, all with nuptial pads, measure 30.2-35.1 mm SVL and the females 31.2-35.8 mm. These frogs have the tympanum smaller than the eye, the two outer fingers webbed to the distal edge of the subarticular tubercles, most of the white as- perities scattered and not gathered in rings around a larger one, a conspicuous large, black marking beginning between the shoulders, and males small vocal sac openings. They lack vomu teeth. Taylor (1962) pictured T stellatum with a tinctive dark dorsal marking that is still evide the holotype (fmnh 172249) and in a more re sample (fmnh 183711-15) from Thailand, pattern is exactly as in the Vietnamese frog T. stellatum the two outer fingers are webbed to the edge of the subarticular tubercles. Malt T stellatum from Thailand (holotype and two ers) measure 27.4-32.7 SVL; a single adul male measures 34.8 mm. In these characters in the absence of vomerine teeth, the Vietnai frogs agree with T. stellatum. However, T; (1962) stated that T. stellatum lacked vocal (confirmed in the holotype), and the two ( adult males from Thailand (large nuptial ] present) also lack them. Two other species, T. horridum (Boulen (type locality Pattani, peninsular Thailand) an 42 FIELDIANA: ZOOLC phnnoderma (Ahl) (type locality northern Bur- ma), lack vomerine teeth and have webbing be- tween the fingers. The Vietnamese frogs differ from each of these. Theloderma horridum has more extensive webbing on the hand, with full web reaching the bases of the discs of the two outer fingers (fmnh 186600-02, peninsular Ma- laysia), although Boulenger (1893) merely noted that the fingers were "half- webbed." In addition, the specimens of T. horridum from Malaysia have :learly longer snouts relative to the diameter of lie eye (eye/snout 0.61-0.71) than T. stel latum from Vietnam and Thailand (0.81-0.89). The hree T. horridum from Malaysia measure 38.0- 14.2 and the holotype 40 mm SVL. Theloderma jhrynoderma has no dark dorsal marking, and its ympanum equals the diameter of the eye (Bou- enger, 1893); both of these characters differ from hose of the Vietnamese sample. SVL measure- nents of the types of T. phrynoderma were 45 nm (Boulenger, 1893), about 10 mm longer than hose of the Vietnamese specimens. We assign a sample of tadpoles (mvz 222115) rom Buon Luoi to this species. The headbody is listinctly flattened and oval. The eyes are dorsal nd pointing directly upward. The oral disc is al- most terminal, less than half the width of the ody, and lacks a lateral notch. The labial papillae re long, continuous across the lower lip and at le corners of the upper. The jaw sheaths are com- letely black and finely serrated; the upper jaw heath has a wide median convexity. The labial :eth are 4(2-4)/3. The spiracle is ventrolateral nd lacks a free tube. The caudal muscle is much eeper than the fins. The dorsal fin has its origin t the end of the body. The headbody is black orsally and laterally and a bit lighter below. The 1 dl is completely black. pedes Doubtfully Reported from ietnam kgophrys longipes Boulenger lygalophrys longipes Boulenger, 1885:850 — Per- ak, Malaysia. Megophrys longipes Bourret, 1942:197. Bourret (1941, 1942) recorded this species am Bach Ma. Annam, based on two juveniles | had originally (1939) identified as Ophry- mryne microstoma. The identity of these juve- niles is doubtful. Tran et al. (1981) listed this spe- cies from six provinces in Vietnam, but gave no specimen information or any reason for their iden- tification. Meristogenys jerboa (Gunther) Hylarana jerboa Gunther, 1872:599 — Matang, Sarawak, Borneo. Meristogenys jerboa Yang, 1991:34. Bourret (1937) assigned a juvenile frog from Tarn Dao to this species, but he reported that it had a shorter leg and more reduced webbing than M. jerboa. This species was also listed as part of the fauna of Tarn Dao by Tran et al. (1981) and Nguyen et al. (1994), although it is not clear whether they were merely repeating Bourret's (1937) record or reporting new findings. At any rate, there is no authentic record of this species outside of Borneo (Yang, 1991) and it should be removed from the faunal list of Vietnam. Rhacophorus nigropalmatus Boulenger Rhacophorus nigropalmatus Boulenger, 1895: 170 — Akah River, Sarawak, Borneo. Tran et al. (1981) recorded this species from four provinces. However, it seems evident that they were referring to the form cited by Bourret (1942) as R. nigropalmatus feae Boulenger. Ac- cording to current usage (Frost, 1985), the records of Tran et al. (1981) should be treated as Rhaco- phorus feae Boulenger, and R. nigropalmatus re- moved from the faunal list. Rana miopus Boulenger Rana miopus Boulenger, 1918:11 — Khao Wang Hip and Nakhon Si Thammarat, Thailand. Tran et al. (1981) reported this species from Lang Son Province in extreme northeastern Viet- nam. It was also reported by Ho and Nguyen (1982) from the Tay Nguyen plateau. Neither pa- per gave descriptive notes. The species is other- wise known only from southern Thailand and nearby Peninsular Malaysia. The identity of these Vietnamese specimens must be confirmed before JGER ET AL.: FROGS OF VIETNAM 43 the species can be added to the faunal list of Vi- etnam. Acknowledgments We are grateful to the Institute of Ecology and Biological Resources, Hanoi, for much logistical assistance. Ho Thu Cue, from the Institute, was a member of the field party in the central highlands. In northern Vietnam, we were joined by Dr. Theo- dore Papenfuss, University of California (Berke- ley); we are grateful to him for his contributions to our success in the field. Support for fieldwork came from the National Geographic Society, the Royal Ontario Museum, and the Marshall Field III Fund of the Field Museum of Natural History. Additional support was received from the Robert Bass Fund of the Field Museum. We are grateful to Dr. John Cadle, Museum of Comparative Zo- ology, Harvard University, and Dr. Barry T. Clarke, The Natural History Museum, London, for lending us specimens in their care. Literature Cited Ahl, E. 1927. Zur Systematik der asiatischen Arten der Froschgattung Rhacophorus. Sitzungsberichte der Gesellschaft naturforschender Freude, Berlin, 1927: 35-47. . 1930. Zwei neue Baumfrosche der Familie Polypedatidae. Zoologischer Anzeiger, 87: 228-230. 1931. Amphibia Anura III. Polypedatidae. Das Tierreich. Berlin. 475 pp. Anderson, J. 1871. 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