ISSN 1345-5834 “ June 2002 SMT AHSON A (OCT 1.8 2uuz LIBRARIES ahi mane nara fas Pe oe | hag =| — CURRENT HERPETOLOGY FORMERLY THE JAPANESE JOURNAL OF HERPETOLOGY Published by THE HERPETOLOGICAL SOCIETY OF JAPAN KYOTO THE HERPETOLOGICAL SOCIETY OF JAPAN Executive Council 2002 President: Masafumi MATSUI, Graduate School of Human and Environmental Stud- ies, Kyoto University, Yoshida Nihonmatsu-cho, Sakyo-ku, Kyoto 606-8501 Japan (fumi@zoo.zool.kyoto-u.ac.jp) Secretary: Tsutomu HIKIDA, Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwake-cho, Sakyo-ku, Kyoto 606-8502 Japan (tom@zoo.zool.kyoto-u.ac.jp) Treasurer: Akira MORI, Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwake-cho, Sakyo-ku, Kyoto 606-8502 Japan (gappa@ ethol.zool.kyoto-u.ac.jp) Managing Editor: Hidetoshi OTA, Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa, 903-0213 Japan (ota@sci.u-ryukyu.ac.jp) Officers: Masami HASEGAWA (PX1M-HSGW @asahi-net.or.jp), Tamotsu KUSANO (tamo@comp.metro-u.ac.jp), Showichi SENGOKU, Michihisa TORIBA (snake-a@ sunfield.ne.jp), Kinji FUKUYAMA (fukuyama@hc.cc.keio.ac.jp) | Main Office: Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa-Oiwake-cho, Sakyo-ku, Kyoto 606-8502 Japan Honorary Member: Richard C. GORIS Current Herpetology.—Current Herpetology publishes original research articles on amphibians and reptiles. It is the official journal of the Herpetological Society of Japan and is a continuation of Acta Herpetologica Japonica (1964-1971) and Japanese Journal of Herpetology (1972-1999). Herpetological Society of Japan.—The Herpetological Society of Japan was estab- lished in 1962. The society now publishes Current Herpetology and Bulletin of the Herpetological Society of Japan, each twice per year. Current Herpetology is printed solely in English and is international in both scope of topics and range of contributors. Bulletin of the Herpetological Society of Japan is printed exclusively in Japanese and publishes more regional topics and announcements about the Herpetological Society of Japan. Membership and Subdscriptions.—Membership (including subscriptions) is open to all interested persons. Current annual fees are 5,000 yen for the regular members who receive both Current Herpetology and Bulletin of the Herpetological Society of Japan. For those who wish to have Current Herpetology only, there is a reduction to 3,000 yen. To subscribe to Current Herpetology and/or Bulletin of the Herpeto- logical Society of Japan, please contact the Treasurer. We accept VISA and Master Card. If you need to use a bank check, please add 2,000 Japanese yen for the handling charge. All other correspondences regarding the society, including the availability and cost of society publications, should be directed to the office of the Secretary. Cover Illustration: A new depressed-bodied Tropidophorus from Vietnam. A photo- graph taken by Nikolai L. Orlov. Current Herpetology 21(1): 1-8, June 2002 © 2002 by The Herpetological Society of Japan Foraging Behavior of Rhabdophis tigrinus (Serpentes: Colubridae) in a Gutter with a Dense Aggregation of Tadpoles Koy] TANAKA Department of Zoology, Graduate School of Science, Kyoto University, Kitashirakawa, Sakyo, Kyoto 606-8502, JAPAN Abstract: Field observations were made on the foraging behavior of Rhabdo- phis tigrinus in a gutter where numerous Hyla japonica larvae aggregated. It seemed that not only chemical cues but also visual cues play important roles in the foraging behavior in this snake. Rhabdophis tigrinus performed predatory behaviors characteristic of generalists, not of aquatic specialists. Examinations of stomach contents revealed that froglets were the dominant prey item although tadpoles were seemingly more abundant in the gutter. It is likely that this biased predation on froglets is attributable to the differential vulnerability to predation among the developmental stages of the frog. The present obser- vations support the idea that R. tigrinus is not well adapted to an aquatic life. Possible significance of success ratio of predatory attempt as an index is discussed. Key words: prey; Ratio of successful strike INTRODUCTION Snakes are all carnivorous predators that eat various types and sizes of prey, ranging from small invertebrates to large mammals (Mushinsky, 1987; Greene, 1997). From the viewpoint of feeding habits, each species is placed at a certain position on a con- tinuum between two extremes, “general- ist” and “specialist”. The specialist often exhibits physiological, morphological, and/ or behavioral adaptations to feed on a specific prey type (Gans, 1983; Mushinsky, 1987; Greene, 1997). To clarify the extent Tel: + 81-75-753-4099; Fax: + 81-75-753-4113; E-mail address: koji@ethol.zool.kyoto-u.ac.jp Rhabdophis tigrinus; Foraging behavior; Anuran larvae; Aquatic of the specialization in feeding habits in a particular species, field observation is desir- able because natural surroundings influence animal behavior and place constraints on it that are often difficult to control in the laboratory. Rhabdophis tigrinus is a medium-sized, diurnal colubrid snake commonly seen on the main islands (exclusive of Hokkaido) and adjacent islets of Japan (Stejneger, 1907). For this snake, there are a number of records in the literature that indicate its natural diet as consisting mostly of frogs, toads, and fish (Mori and Moriguchi, 1988). In regard to feeding behavior, Mori et al. (1992) reported a peculiar foraging behavior of R. tigrinus in the field, that is, a nocturnal ambush predation on the forest green tree frog, Rhacophorus arboreus. Under experi- mental conditions, Mori (1997) reported prey-handling behavior of the snake using frogs and fish as prey animals. However, although Rhabdophis tigrinus is generally considered semi-aquatic, little is known about how the snake forages for aquatic prey (e.g., fish, tadpoles) in the field and to what extent the snake is specialized for an aquatic life. In the present paper, I report foraging by R. tigrinus on aquatic and semi-aquatic prey observed in the field. MATERIALS AND METHODS Field observations were made sporadi- cally during the ecological survey of snakes on Yakushima Island (30°20’N, 130°32’E), Japan. Foraging behavior of R. ftigrinus was observed in the gutter along the road named Seibu-rindo (Western Woodland Road). The road runs close to the western coast of the island and is surrounded by secondary forests. The gutter was 40 cm wide and contained up to 10 cm depth of still water, which I refer to as “the pool” (Fig. 1). Water of the pool was moderately clear (the bottom was clearly visible). The pool extended approximately 20 m along Ee litter and mud concrete Current Herpetol. 21(1) 2002 the gutter, and had the paved road on one side and a steep mountain slope on the other side. One end of the pool was over- grown with weeds, which I call “the weed area” (Fig. 1), and the opposite end was bare wet space covered with leaf litter and mud. At the center of the pool, leaf litter and mud formed another wet area, which I call “the land” (Fig. 1). Numerous Hyla Japonica larvae of various developmental stages were observed in the pool. | In the initial three observations, I captured the snakes as soon as I saw them without observing their behavior. For these snakes, therefore, only the results of examination of the stomach contents are presented. In the other observations, behavior of the snakes was directly observed at distances of more than 2 m. These observations did not seem to disturb the subjects because no abrupt changes of behavior were evident. Each observation was terminated if the snake began to crawl out of the gutter or to enter the weed area. After most observations, the snakes were captured and their body temperatures were measured immediately to 0.1 C by inserting a thermistor bulb into the cloaca in the shade. Air and water tempera- tures were also measured. Stomach contents were forcibly ejected. Unless otherwise ——- HIGy ol: Schematic diagrams of cross (A) and longitudinal sections (B) of the gutter. In the latter, a large part of the right pool is omitted (indicated by slanted lines) so as to depict all microenvironments in the gutter. Bold lines indicate the water surface level. Snake and anuran larvae are out of scale. TANAKA—FORAGING BEHAVIOR OF RHABDOPHIS TIGRINUS 3 mentioned, recovered prey items were fixed in 10% formalin and preserved in 70% ethanol. At initial capture, each individual was brought to the field station, where it was measured for snout-vent length (SVL, measured to 1 mm by tape scale) and body mass (BM, up to 100 g, weighed to 0.1 g on an electric toploading balance; above 100 g, weighed to 1 g with a spring scale). The snake was then marked with paint on the head and by ventral scale clipping for individual recognition, and was released on the following day at the site of capture. Recaptured individuals were released imme- diately after body temperature measurement and examination of stomach contents. RESULTS A total of seven observations were made for three individuals (Table 1). Observation 1: A snake, marked as no. 62, contained one tadpole of H. japonica and part of the forefoot of a Bufo japonicus Japonicus in its stomach. Observation 2: A snake, marked as no. 63, contained one tadpole and two froglets (defined here as Gosner’s [1960] stages 43- 45) of H. japonica in its stomach. Observation 3: A snake, marked as no. 65, had no stomach contents. Observation 4: No. 65 was found swim- ming in the pool. The snake reached the land, where it anchored its tail to dead twigs and started to peer into the pool. Suddenly, the snake lunged with the anterior part of the body (for convenience, I refer to this as a normal strike) into the water and swept its head from side to side several times with its mouth open. The snake seemingly responded to movements of a few tadpoles that were swimming near its head. Then the snake resumed peering, and after a few minutes it struck again, but this time it raised its head and neck immediately. After several minutes of wandering on the land, the snake exhib- ited another normal strike and succeeded in biting a tadpole at midbody (near developing hindlimbs), which it then swallowed tail first. Until capture, the snake continued prowling on the land or swimming in the pool with vigorous tongue flicking. The snake con- tained two tadpoles and nine froglets of H. japonica in its stomach. All strikes were made from the land, and success ratio of predatory attempts (SRPA: the number of successful strikes/the total number of strikes) was 0.33 (1/3). Observation 5: No. 65 was found swim- ming in the pool. The snake crawled onto the dead leaves floating on the water surface and struck at a tadpole, but missed. Then the snake continued to swim with occasional tongue flicking. After several minutes, the snake crawled onto the land, and three nor- mal strikes (two missed and one succeeded) at tadpoles followed the peering posture. Until disappearing into the weed area, the snake was swimming around in the pool or prowling on the land. SRPA was 0.25 (1/4). Observation 6: No. 65 was found lying on the land with its body loosely coiled and peering into the pool. Suddenly, the snake moved its head slightly forward, probably in response to the movements of tadpoles. Then, the snake wandered about on the land or swam in the pool, and made two normal strikes at tadpoles from the land but neither of them succeeded. After a brief swim following these failures, the snake suddenly posed with its throat region con- tacting the wall of the gutter and with the remaining portion of the body floating in the pool. The anterior and posterior parts of the body formed an angle of 90°. From this posture, the snake protruded its head (I refer to this as floating strike), and captured a froglet clinging on the wall. After this success, three additional strikes were made at tadpoles that were either swimming in the pool or trapped in the shallow part. Of these strikes, two were made from the land and one from the pool with serpentine diving (mid-water diving with medium to fast loco- motion and sinuous movements of head and body: Drummond, 1983). 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At this time, no. 62 was seen approaching no. 65. When the two snakes reached a point about 10 cm from each other on the land, no. 65 started to crawl rapidly, passed by the side of no. 62 and went away. I captured no. 62 at this time. No. 65 continued to swim in the pool and wander on the land, and the seventh normal strike (failure) was made from the land. No. 62 contained no prey in its stom- ach, whereas no. 65 contained one tadpole and two froglets of H. japonica in its stom- ach. These prey items were re-fed to the snake. SRPA for no. 65 was 0.29 (2/7). Observation 7: No. 65 was found swim- ming in the pool. The snake made a float- ing strike at a froglet clinging on the wall, but the prey escaped. Then, it briefly repeated the pose with its snout touching the wall of the gutter several times while swim- ming until it made a successful normal strike at a froglet in the shoal. After this success, the snake repeated the snout touching behavior described above, and made a float- ing strike (failure) at a froglet clinging on the wall. The snake contained 15 froglets of H. japonica in its stomach. I judged that the prey items were all newly ingested (i.e., not including the two froglets that were re- fed on the previous day: see above) by the degree of digestion. All prey were re-fed to the snake. SRPA was 0.33 (1/3). Total SRPA of no. 65 throughout the observation period was 0.29 (5/17). DISCUSSION The importance of chemical cues in the snake feeding behavior has been demonstrated by numerous experimental studies (see Burghardt [1990]; Halpern [1992] for review). Snakes transfer chemicals to chemoreceptive organs that are situated on the roof of the oral cavity by either tongue flicking or snout touching (Halpern and Kubie, 1980). It is, therefore, likely that the tongue flicking and snout touching behavior observed in the present study indicates that the snakes were searching for prey using chemical cues. On the other hand, both laboratory experiments and field observations indicate that visual cues also play an important role in the feeding behavior of snakes (e.g., Czaplicki and Porter, 1974; Herzog and Burghardt 1974; Drummond, 1985; Ota, 1986; Heinen, 1995). The present observations suggest that in the predatory behavior of R. tigrinus, visual cues, especially those from prey move- ments, are also important, because on several occasions normal strikes occurred immedi- ately after movements of the tadpoles. Hyla japonica larvae of various develop- mental stages formed dense schools in the pool. Froglets were the dominant prey recov- ered from the snakes’ stomachs, although tadpoles were seemingly more abundant (about 10 times or more) than froglets in the pool. Both laboratory and field studies have indicated that anurans are particularly vulnerable to snake predation during the meta- morphic transition (Wassersug and Sperry, 1977; Arnold and Wassersug, 1978). This vulnerability during the metamorphic transi- tion is attributed to the locomotor inepti- tude of transforming anurans: froglets can neither swim nor hop as effectively as tad- poles or frogs, respectively (Wassersug and Sperry, 1977; Arnold and Wassersug, 1978). In this study, predatory events actually observed were only against three tadpoles and two froglets. Therefore, it may be possi- ble that the prey recovered from the snakes’ stomachs were taken at some other place where froglets were more abundant than in the gutter. Such a situation, however, is actually unlikely because (1) the area sur- rounding the gutter was relatively dry and lacked possible habitats for tadpoles and metamorphosing froglets, such as swamps and streams; (2) in froglets recovered from the snakes’ stomachs, the tail was not yet fully absorbed, making it unlikely that they had already dispersed far away from the tadpoles’ habitat when eaten by the snakes; and (3) the degree of digestion was almost identical in all prey exclusive of B. japonicus, indicating that all predatory events involving H. japonica tadpoles and froglets had occurred in a fairly short period. Thus, I believe that all Hl. japonica tadpoles and froglets recovered from the snakes’ stomachs were taken in the gutter, and that the biased predation toward froglets (five tadpoles vs. 28 froglets) was the consequence of a greater vulnerability to predation in froglets than in tadpoles in H. japonica. Drummond (1983) described aquatic for- aging in garter snakes and a water snake. In his study, “generalists” exhibited charac- teristic behaviors such as open-mouth search- ing and serpentine diving in mid-water, whereas “aquatic specialists” exhibited other characteristic behaviors such as substrate- crawling and mid-water pursuit (rapid loco- motion toward prey in mid-water). Rhab- dophis tigrinus in the present observations performed open-mouth searching and serpen- tine diving, but substrate-crawling and mid- water pursuit were never observed. Under experimental conditions, R. tigrinus was inef- ficient in capturing fish, and fish once grasped by the snake frequently escaped during handling (Mori, 1997). Furthermore, the snake always handled and swallowed fish on land (Mori, 1997). Mori (1997) assumed that these behavioral propensities reflect incomplete adaptation of this snake to an aquatic life. The present observations seem to support this postulation, although further field observations on more individuals are definitely needed to draw firm conclu- sions on this problem. Several studies reported foraging behavior and its efficiency in actively foraging snakes including both dietary generalists and special- ists in the field. Wendelken (1978) observed that Thamnophis proximus (a putative spe- cialist on amphibian prey: Rossman et al., 1996) chased 10 frogs but none were cap- tured during 45 min. Patterson and Davies (1982) reported that in spite of a high abun- dance of fish in a river, only one out of many individuals of Natrix maura (a putative Current Herpetol. 21(1) 2002 specialist on fish prey: Hailey and Davies, 1986; Santos and Llorente, 1998) there caught a fish in several hours’ observation. Hailey and Davies (1986) reported that 48 individu- als of N. maura made 124 strikes at fish throughout a total of 921 min observation, and only two made contact. Ota (1986) reported that SRPA against lizards by Elaphe quadrivirgata (a putative dietary generalist: Mori and Moriguchi, 1988) was 0.125 (2/16). Balent and Andreadis (1998) reported that Nerodia sipedon (a putative aquatic and dietary specialist on fish prey: Drummond, 1983) did not succeed in capturing aquatic prey in any of 17 attempts throughout a total of 44.75 min observation. In the present observations, total SRPA of R. tigrinus (no. 65) was slightly higher (0.29) than in these previous studies. However, three out of the five successful strikes involved either a tadpole trapped in a shoal or froglets, and these prey would have been captured more easily than tadpoles in free water because of their locomotory impediment. Therefore, it seems that SRPA of R. tigrinus against aquatic prey is not so high as expressed by the above figures. Can we, however, infer the extent of specialization of the subject species only on the basis of SRPA as an indicator? In the studies cited above, foraging success is not higher in the specialists than in the generalists. This suggests that low SRPA does not always indicate less adaptation to a particular type of microhabitat (e.g., aquatic, terres- trial, arboreal) or of prey, because various external (e.g., temperature, light condition, prey density) and internal factors (e.g., age and body condition of either prey or predator) would affect foraging efficiency. For better understanding of the biological and evolu- tionary significance of SRPA, further field observations on both active foragers and ambushers are desirable. ACKNOWLEDGMENTS I am grateful to A. Mori for valuable TANAKA—FORAGING BEHAVIOR OF RHABDOPHIS TIGRINUS 7 comments on the manuscript and T. Hirai for helping with identifications of items regurgitated by snakes. I also thank two anonymous reviewers for helpful comments on an early version of the manuscript, and S. Horiuchi, H. Kudo, T. Teramura, and R. Tsujino for their hospitality during my stay at the Kyoto University Yakushima Field Station during the fieldwork. LITERATURE CITED ARNOLD, S. J. AND R. J. WASSERSUG. 1978. Differential predation on metamorphic anurans by garter snakes (Thamnophis): social behavior as a possible defense. Ecology 59(5): 1014- 1022. BALENT, K. L. AND P. T. ANDREADIS. 1998. The mixed foraging strategy of juvenile northern water snakes. J. Herpetol. 32(4): 575-579. BURGHARDT, G. M. 1990. Chemically mediated predation in vertebrates: diversity, ontogeny, and information. p. 475-499. In: D. W. McDonald, D. Muller-Schwarze, and S. E. Natynczuk (eds.), Chemical Signals in Vertebrates, Vol. 5. Oxford Univ. Press, New York. CZAPLICKI, J. A. AND R. H. PORTER. 1974. Visual cues mediating the selection of goldfish (Carassius auratus) by two species of Natrix. J. Herpetol. 8(2): 129-134. DRUMMOND, H. 1983. Aquatic foraging in garter snakes: a comparison of specialists and gener- alists. Behaviour 86(1/2): 1-30. DRUMMOND, H. 1985. The role of vision in the predatory behavior of natricine snakes. Anim. Behav. 33(1): 206-215. GANS, C. 1983. Snake feeding strategies and adaptations: conclusion and prognosis. Amer. Zool. 23(2): 455-460. GOSNER, K. L. 1960. A simplified table for stag- ing anuran embryos and larvae with notes on identification. Herpetologica 16(3): 183-190. GREENE, H. W. 1997. Snakes: the Evolution of Mystery in Nature. Univ. California Press, Barkeley California. 351 p. HAILEY, A. AND P. M. C. DAVIES. 1986. Diet and foraging behavior of Natrix maura. Herpetol. J. 1(2): 53-61. HALPERN, M. 1992. Nasal chemical senses in reptiles: structure and function. p. 423-523. In: C. Gans and D. Crews (eds.), Biology of the Reptilia, Vol. 18. Physiology E: Hormones, Brain, and Behavior. Univ. Chicago Press, Chicago. HALPERN, M. AND J. K. KUBIE. 1980. Chemical access to the vomeronasal organs of garter snakes. Physiol. Behav. 24(2): 367-371. HEINEN, J. T. 1995. Predator cues and prey responses: a test using eastern garter snakes (Thamnophis s. sirtalis) and American toads (Bufo a. americanus). Copeia 1995(3): 738-741. HERZOG, H. A., JR. AND G. M. BURGHARDT. 1974. Prey movement and predatory behav- ior of juvenile western yellow-bellied racers, Coluber constrictor mormon. Herpetologica 30(3): 285-289. Mor], A. 1997. A comparison of predatory behav- ior of newly hatched Rhabdophis tigrinus (Ser- pentes: Colubridae) on frogs and fish. Jpn. J. Herpetol. 17(2): 39-45. Mor!I, A. AND H. MORIGUCHI. 1988. Food habits of the snakes in Japan: a critical review. Snake 20(2): 98-113. Mor!, A., M. ToDA, S. KADOWAKI, AND H. MORIGUCHI. 1992. Lying in ambush for noc- turnal frogs: field observations on the feeding behavior of three colubrid snakes, Elaphe quadrivirgata, E. climacophora, and Rhabdo- pDhis tigrinus. Jpn. J. Herpetol. 14(3): 107-115. MUSHINSKY, H. R. 1987. Foraging ecology. p. 302-334. In: R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes: Ecology and Evo- lutionary Biology. Macmillan Publ. Co., New York. OTA, H. 1986. Snake really an able hunter?: predatory behavior of Japanese striped snake, Elaphe quadrivirgata, in the field. J. Ethol. 4(1): 69-71. PATTERSON, J. W. AND P. M. C. DAVIES. 1982. Predatory behavior and temperature relations in the snakes Natrix maura. Copeia 1982(2): 472-474. ROSSMAN, D. A., N. B. FORD, AND R. A. SEIGEL. 1996. The Garter Snakes: Evolution and Ecology. Univ. Oklahoma Press, Norman and London. i-xx + 332 p.+15 pls. SANTOS, X. AND G. A. LLORENTE. 1998. Sexual and size-related differences in the diet of the snake Natrix maura from the Ebro Delta, Spain. Herpetol. J. 8(3): 161-165. STEJNEGER, L. 1907. Herpetology of Japan and adjacent territory. Bull. U.S. Nat. Mus. (58): 1-577. WASSERSUG, R. J. AND D. G. SPERRY. 1977. The relationship of locomotion to differential pre- Current Herpetol. 21(1) 2002 dation on Pseudacris triseriata (Anura: Hylidae). Ecology 58(4): 830-839. WENDELKEN, P. W. 1978. On prey-specific hunting behavior in the western ribbon snake, Tham- nophis proximus (Reptilia, Serpentes, Colubridae). J. Herpetol. 12(4): 577-578. Accepted: 11 April 2002 Current Herpetology 21(1): 9-23, June 2002 © 2002 by The Herpetological Society of Japan Three New Depressed-bodied Water Skinks of the Genus Tropidophorus (Lacertilia: Scincidae) from Thailand and Vietnam TSUTOMU HIKIDA"™, NIKOLAI L. ORLOV’, JARUJIN NABHITABHATA;3, AND HIDETOSHI OTA‘ ! Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto 606-8502, JAPAN * Zoological Institute, Russian Academy of Sciences, Universitetskaya Nab. 1, St. Petersburg 199034, RUSSIA 3 Thailand Natural History Museum, National Science Museum, Technopolis, Khlong 5, Khlong Luang, Pathumthani 12120, THAILAND 4 Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903-0213, JAPAN Abstract: Three new species of the genus Tropidophorus, characterized by distinct depression of body, strongly keeled lateral body scales, and saxicolous habits, are described from Indochina. Of these, two moderately depressed species, one with undivided frontonasal and widened paravertebral scales, and the other with divided frontonasal and unwidened paravertebral scales, were collected from small areas in northeastern and eastern Thailand, respectively. The remaining species with extremely depressed head and body was found from one limited area in northern Vietnam. The three species most resemble T. baviensis Bourret, 1939 from northern Vietnam among the known conge- neric species in body size, body shape, and scutellation. However, body depression in 7. baviensis is not so prominent as in the present three species. Considering that most specimens of these species were collected from rock crevices, their characteristic body shapes may represent certain stages of adap- tation to life in crevices. Key words: Tropidophorus, Scincidae, Thailand, Vietnam, New species, Taxonomy, Body depression, Rock crevices INTRODUCTION Tropidophorus is a group of approxi- * Corresponding author. Tel: +81-75-753- 4091; Fax: +81-75-753-4114; E-mail address: tom@zoo.zool.kyoto-u.ac.jp mately 20 species of small to moderate sized lygosomine skinks distributed in Bangladesh, southern China, and Southeast Asia including the Philippines (Welch et al., 1990; Wen, 1992). This genus is characterized by super- ficial location of the tympanum, and most of its species are known to prefer semiaquatic 10 habitats along forest streams (e.g., Taylor, 1963; Brown and Alcala, 1980). During recent herpetological surveys in Indochina, specimens of skinks, characterized by distinctly depressed bodies with strongly keeled lateral scales, were collected from three localities, two in Thailand and one in Vietnam (Fig. 1). These skinks possessed superficial tympanums and thus were identi- fied as members of the genus 7ropidophorus. They most resembled 7. baviensis Bourret, 1939 from northern Vietnam in body size, body shape, and scutellation, but differed from the latter or any other congeneric species so far described by distinct depression of body. Also, specimens from the three localities showed substantial differences from each other in a number of morphological charac- ters. We thus describe three new species of Tropidophorus on the basis of these speci- mens. 100°E pe i. e ere Thanh caine Cuc Phuon O “Phu Wua_ > Lf? Fic. 5. (C) views of head scales of Tropidophorus matsuii sp. nov. (holotype, TNHM-R-60006). Lateral (A), dorsal (B), and ventral Paratypes and other specimens None. Diagnosis A Tropidophorus with moderately depressed head, body, and tail; scales on dorsal surface of head smooth as a whole, but those in temporal region keeled; frontonasal divided; eight superciliaries; paravertebral scales smooth or feebly keeled, subequal to neighboring scales in size; 65 paravertebral scales; dorso- lateral and lateral scales distinctly keeled; 34 midbody scale rows. See Discussion for comparisons with other congeneric species. Description of holotype Snout rounded, rostral partly visible from above; no supranasals; frontonasal divided with right element smaller than left one, overlapped by rostral, nasals, and upper anterior loreals, and overlapping prefrontals; prefrontals overlapped by frontonasals, upper anterior, and posterior loreals, and overlapping frontal, first supraoculars, and first superciliaries; right prefrontal narrowly overlapped by left one; frontal large, nar- rowing posteriorly, overlapped by prefrontals, and overlapping first and second supraocu- lars and frontoparietals; supraoculars four, overlapped by superciliaries; eight supercili- aries; interparietal smaller than frontal, nar- rowing and slightly concave posteriorly, and overlapping parietals; small transparent spot on interparietal, showing location of parietal foramen; parietals separated by interparietal; nuchals lacking; nostril piercing nasal; nasal overlapped by rostral and first supralabial, and overlapping frontonasal and two anterior loreals; both anterior loreals overlapped by nasal, and overlapping posterior loreal; lower anterior loreal overlapping upper anterior loreal and very slightly second supralabial, and contacting first supralabial at point; upper anterior loreal overlapping frontonasal and prefrontal; supralabials six, three anterior to, one just beneath, and two posterior to orbit; shallow groove running on loreal-labial border, posteriorly crossing subocular in obliquely downward direction; two presuboculars, anterior one larger than posterior one, over- lapped by posterior loreal and supralabials; lower eyelid with six scales, posterior three extremely enlarged, separated from labials by three rows of granular scales; three small postoculars, overlapped by fourth supraocular and palpebrals, and overlapping postsuboc- ulars; postsupraocular overlapped by fourth supraocular and postoculars, and overlapping parietal, primary temporal, and postsubocu- lars; postsuboculars five, first one smooth, overlapped by fourth supralabial and over- lapping fifth supralabial, remaining four postsuboculars keeled; temporals in seven 16 rows, those in secondary and tertiary rows more or less enlarged, uppermost rows largest, smooth, overlapped by parietal, the others smaller, keeled; temporals in the other rows keeled, subequal to body scales in size, upper ones directed straight backward, lower ones directed obliquely downward; tympanum superficial; mental overlapping first infrala- bials and postmental; postmental undivided, overlapped by mental and first infralabials, and overlapping first chinshields; chinshields in three pairs, first right one overlapping first left one, second pair separated by single scale, third pair separated by three scales; six infralabials; one postgenial following each of third chinshields; 34 midbody scale rows; 15 scale rows at position of tenth subcaudal on tail; paravertebrals 65, subequal in size to neighboring scales, with two very weak keels on neck, smooth on body and base of tail, and with one moderate keel on the remaining portion of tail; scales in row adjacent to paravertebral row on each side with two weak keels on neck, and with single weak keel on body and tail; dorsolateral and lateral scales distinctly keeled; eight rows of mid-ventral scales smooth, scales in outer row on each side feebly keeled; preanals two, enlarged, right one overlapped by left one; subcaudals smooth anteriorly, weakly keeled posteriorly, first one four times as broad as neighboring scales, remaining ones only two times as broad as neighboring keeled scales; scales on ventral surfaces of hindlimbs only feebly keeled, those on the other portions of hindlimbs and on forelimbs distinctly keeled; 22-23 subdigitals on fourth toe. Presacral vertebrae 26. Testes regressed, 6.6 mm in longer axis; epididymides regressed. Measurements of holotype (mm) SVL, 94.1; tail length, 113.0; snout to forelimb length, 34.2; head length, 15.9; head width, 14.7; head depth, 8.9; snout length, 6.1; eye length, 5.5; eye to tympanum length, 7.6; snout to tympanum length, 18.0; tym- panum height, 3.6; tympanum width, 2.2; Current Herpetol. 21(1) 2002 axila to groin length, 50.2; midbody width, 20.1; midbody depth, 9.6; forelimb length, 26.1; hindlimb length, 35.7; fourth toe length, 11.0. Color in preservative Dark brown on dorsal and lateral surfaces of head, body, and tail; three, nine, and 23 transverse bands, pale brown in color and somewhat irregular in shape, on dorsal sur- faces of neck, body, and tail, respectively; several irregular pale brown spots on lateral sides of body (Fig. 2); several pale brown spots on supralabials and infralabials; yel- lowish ivory on gular and venter; ventral surface of tail yellowish ivory with indistinct dark flecks. Color in life. Dorsal and lateral surfaces slightly darker than after preservation; portions light brown in preservative (see above) with somewhat orange tint. Etymology The name is dedicated to Prof. Masafumi Matsui of Kyoto University, the project leader of the herpetological survey in Thailand, during which the present species was discov- ered. Natural history The single type specimen was found in a crevice of the lower portion of a sandstone outcrop, surrounded by relatively humid ever- green forest. Two gekkonid lizards, Cyrto- dactylus papollionoides and Gekko petricolus, were also found in crevices of sandstone rocks near the collecting site. Tropidophorus murphyi sp. nov. (Figs. 6-8) Holotype Adult male, ROM 41227 (ROM field No. 27044), from Quang Thanh Village (22°37'43 N, 105°54'46 E, altitude 700- 750 m), Nguyen Binh District, Cao Bang HIKIDA ET AL.—NEW DEPRESSED-BODIED TROPIDOPHORUS Mg) Province, northern Vietnam, collected by R. W. Murphy, N. L. Orlov, A. Lathrop, T. Mason, S. Riabov, and T. C. Ho in May 1998. Paratypes Four males, ROM 41220 (ROM Field No. 26739), 41222 (Field No. 26960), 41223 (Field No. 26961) and 41226 (Field No. 27002), and seven females, ROM 41221 (ROM Field No. 26959), 41224 (Field No. es Yi Wy, 27000), 41225 (Field No. 27001), 41228 (Field No. 27045), 41229 (Field No. 27058), 41230 (Field No. 27059) and KUZ R58270 (Field No. 27003), with same sampling data as the holotype. Diagnosis A Tropidophorus with extremely depressed head, body, and tail; head scales smooth on dorsal surface, but rugose on lateral surfaces; frontonasal undivided; 6-8 superciliaries; oe FIG. 6. Dorsal (A) and ventral (B) views of Tropidophorus murphyi sp. nov. (holotype, ROM 41227). 18 => FIG. 7. (C) views of head scales of Tropidophorus murphyi sp. nov. (holotype, ROM 41227). Lateral (A), dorsal (B), and ventral paravertebral scales smooth or feebly keeled, subequal to neighboring scales in size; 55-67 paravertebral scales; dorsolateral and lateral scales distinctly keeled; 30-32 midbody scale rows. See Discussion for comparisons with other congeneric species. Description of holotype Snout rounded, rostral partly visible from above; no supranasals; frontonasal undivided, overlapped by rostral, nasals, and upper anterior loreals overlapping prefrontals; pre- frontals contacting each other at point, overlapped by frontonasals, upper anterior and posterior loreals, and overlapping fron- tal, first supraoculars, and first supercili- aries; frontal large, narrowing postriorly, overlapped by prefrontals, and overlapping first and second supraoculars and frontopa- Current Herpetol. 21(1) 2002 rietals; supraoculars four, overlapped by superciliaries; superciliaries six in left, seven in right; interparietal smaller than frontal, narrowing and slightly concave posteriorly, overlapping parietals; small transparent spot on interparietal, showing location of parietal foramen; parietals separated by interparietal; three pairs of nuchals; nostril piercing nasal; nasal overlapped by rostral and first suprala- bial, and overlapping frontonasal and two anterior loreals; both anterior loreals over- lapped by nasal, and overlapping posterior loreal; lower anterior loreal very slightly overlapped by second supralabial, and over- lapping upper anterior loreal, but not contacting first supralabial; upper anterior loreal overlap- ping frontonasal and prefrontal; supralabi- als six, three anterior to, one just beneath, and two posterior to orbit; shallow groove running on loreal-labial border, posteriorly crossing subocular in obliquely downward direction; two presuboculars, anterior one larger than posterior one, overlapped by posterior loreal and supralabials; lower eyelid with seven scales, separated from labials by two or three rows of granular scales; two postoculars, overlapped by fourth supraocular and palpebrals, and overlapping postsuboc- ulars; postsupraocular overlapped by fourth supraocular and postoculars, and overlapping parietal and primary temporal; postsubocu- lars four, smooth, first one overlapped by fourth supralabial, and overlapping fifth supralabial; temporals in seven rows, those in secondary and tertiary rows more or less enlarged, uppermost secondary temporal divided, overlapped by parietal together with uppermost tertiary temporal; temporals in the other rows subequal to body scales in size; tympanum superficial; mental overlap- ping first infralabials and postmental; post- mental undivided, overlapped by mental and first infralabials, and overlapping first chin- shields; chinshields in three pairs, first right one overlapped by first left one, second pair separated by single scale, third pair separated by three scales; five infralabials; one scale broadly overlapping third left chinshield, and HIKIDA ET AL.—NEW DEPRESSED-BODIED TROPIDOPHORUS 19 FIG. 8. Male (A) and female (B) Tropidophorus murphyi sp. nov. in life. Note relatively broad body in the latter. overlapped by fourth and fifth infralabials; tebrals 62, subequal in size to neighboring 30 midbody scale rows; 13 scale rows at scales, smooth or feebly keeled on neck, body, position of tenth subcaudal on tail; paraver- and base of tail, and moderately keeled on 20 the remaining portion of tail; scales in row adjacent to paravertebral row on each side weakly keeled on neck, body and tail; dor- solateral and lateral scales distinctly keeled; six rows of mid-ventral scales smooth, scales in outer row on each side feebly keeled; preanals two, enlarged, right one overlapped by left one; subcaudals smooth, four times as broad as neighboring scales, remaining ones only two times as broad as neighboring keeled scales; scales on ventral surfaces of hindlimbs only feebly keeled, those on the other portions of hindlimbs and on forelimbs distinctly keeled; 24 subdigitals on fourth toe. Presacral vertebrae 26. Left testis 4.7 mm in longer axis. Measurements of holotype (mm) SVL, 85.1; tail length, 101.0 (tail tip lost); snout to forelimb length, 30.5; head length, 14.3; head width, 12.3; head depth, 6.4; snout length, 5.7; eye length, 5.0; eye to tympanum length, 6.5; snout to tympanum length, 16.7; tympanum height, 3.1; tympanum width, 2.3; axila to groin length, 42.8; midbody width, 14.7; midbody depth, 5.5; forelimb length, 22.9; hindlimb length, 33.0; fourth toe length Fe Color in preservative Dark brown or dorsal and lateral surfaces of head, body, and tail; three, seven, and 17 transverse bands, pale brown in color and rather irregular in shape, on dorsal surfaces of neck, body, and tail, respectively; several pale brown spots on supralabials and infralabials; yellowish ivory on gular and venter; ventral surface of tail yellowish ivory with indistinct dark flecks. Variation Of paratypes, three adult males and one young male measured 62.4-85.1 mm and 55.2 mm in SVL, respectively, while SVL in five adult and two immature females ranged from 92.2 to 96.3 and from 56.1 to 76.8, respectively. Relative breadth of body was distinctly greater in adult females than in Current Herpetol. 21(1) 2002 adult males (midbody width/SVL*100: 18.6- 22.0, vs. 17.7-18.2) (Fig. 8). The paravete- bral number was also greater in females than in males (mean and range: 59.6 and 60-67, vs 62.6 and 55-62). No intersexual differ- ences were evident in other scale characters. The number of midbody scale rows ranged from 30 to 32. Upper and lower anterior loreals were fused on one side in two speci- mens and on both sides in one specimen. The arrangement of nuchals was highly vari- able, two on both sides in five specimens, none on both sides in two specimens, two on left and three on right in two specimens, two on left and none on right in one speci- men, two on left and four on right in one specimen, and three on both sides in one specimen. Supralabials were invariably six. Infralabials were usually five, but rarely six. Superciliaries were usually six or seven, but rarely five or eight. Frontal contacted usually two, but rarely three supraoculars. The number of subdigitals ranged from 20 to 25. Scale rows on tail at position of tenth sub- caudal were invariably 13. Etymology The name is dedicated to Dr. Robert W. Murphy of the Royal Ontario Museum, the project leader of the herpetological survey in Vietnam, during which the present species was discovered. Natural history All specimens were collected in humid rocky areas along a stream flanked by steep rocky slopes with bush. There were many moist crevices in the rocks, in which the skinks hid themselves during the daytime. Active individuals were observed only after sunset, and this strongly suggests that this species is nocturnal. Because individuals, prevented from direct contact with moist substrates, showed rapid dehydration even under high atmospheric humidity, it is likely that 7. murphyi is highly vulnerable to drought. Another congeneric species, 7. sinicus occurred sympatrically, but was found under HIKIDA ET AL.—NEW DEPRESSED-BODIED TROPIDOPHORUS ZA rocks closer to the stream. Female paratypes had 3-S eggs in oviducts, which had no recognizable embryos. A non-type female from the type locality, kept in captivity at Tula Exotarium, Russia, gave birth to a juvenile, 28 mm in SVL, on 30 March 2002. It is thus obvious that this species is viviparous like several other con- generic species (Smith, 1923; Taylor, 1963). DISCUSSION Smith (1923), in a revision of the genus Tropidophorus from the continental region of Southeast Asia, recognized nine species. Later, T. baviensis and T. guanxiensis were described from northern Vietnam, and Guanxi, China, respectively (Bourret, 1939; Wen, 1992), mak- ing the total number of recognized continental species of Zropidophorus 11. Of these, T. baviensis most resembles 7. latiscutatus, T. matsuil, and 7. murphyi in having keeled scales on the temporal and smooth scales on the remaining dorsal and lateral surfaces of the head, and smooth or feebly keeled dorsal scales on the body. These three species, however, differ from 7. baviensis or any other congeneric species so far described in having a distinctly depressed body. Moreover, the numbers of paravertebrals in the present species are distinctly greater than that in 7. baviensis (Table 1). Of the three depressed-bodied species described above, the degree of depression, particularly of the head, is much greater in T, murphyi than in the two Thailand species (Table 1, Fig. 9). Tropidophorus matsuii differs from T. /atiscutatus in having greater numbers of midbody scale rows and paraver- tebrals. Also, 7. matsuii is distinct from T. latiscutatus, as well as from T. murphyi and T. baviensis, in having a divided frontonasal. Tropidophorus latiscutatus is distinct from the other species in having distinctly widened paravertebrals (Table 1). Species characterized by distinctly depressed bodies have been reported for a few other lizard families, such as Gekkonidae, Iguanidae, and Xenosauridae. Because many of these species are strongly associated with saxi- > 9 a) mae ) ak’ A a Oa ¢ aq 7 oo, 8 6 @ o¢ . 4 5 e ys 923 ee eee eee 50 60 70 80 90 100 110 SVL Midbody Depth ano N Oo oO 50 60 70 80 90 100 110 SVL FIG. 9. Two dimensional plots of head depth (A) and depth at midbody (B) against snout-vent length (SVL), showing interspecific variations in degrees of head and body depressions, respectively. See Fig. 1 for explanations of symbols. TABLE 1. Comparisons in external characters of the three depressed-bodied 7ropidophorus and T. baviensis. Character T. baviensis T. latiscutatus T. matsuii T. murphyi Body depression slight moderate moderate extreme Midbody scale rows 28-30 28-30 34 30-32 Frontonasal undivided undivided divided undivided Paravertebral widened no yes no no Paravertebral number 49-53 58-63 65 59-67 22 Current Herpetol. 21(1) 2002 colous habitats, such body depression is often regarded as a kind of adaptation to the use of narrow rock crevices as shelters for pred- atory avoidance (Vitt, 1981; Doughty and Shine, 1995; Ballinger et al., 2000). Known members of the genus 7ropidophorus, includ- ing 7. baviensis, are usually found beneath rocks and leaf litter on the forest floor (occasionally close to a stream: Taylor, 1963; Brown and Alcala, 1980), or in burrows on banks (7. baviensis: Ngo et al., 2000). Dis- covery of the three exceptionally depressed- bodied Tropidophorus in rock crevices, a type of habitat also exceptional to the genus, offers a first substantial support from the Scincidae for the assumption regarding the enhancement of body depression by this type of habitat. Several authors have assumed that in lizards the physical constraint from the crevice- dwelling habits provides an evolutionary force to some reproductive traits, such as relative clutch mass and frequency of clutch produc- tion (Vitt, 1981, 1993; Doughty and Shine, 1995). However, relevant hypotheses have not yet been sufficiently assessed by appropriate comparative approaches (e.g., see Ballinger et al. [2000]). The present species and T. baviensis, seemingly representing differential stages of an adaptation to life in rock crevices, would offer a good opportunity to examine the evolutionary consequences of crevice-dwelling habits in reproductive and other ecological and physiological traits. ACKNOWLEDGMENTS T. Hikida and H. Ota thank M. Matsui for providing opportunities to visit Thailand, and S. Panha, M. Matsui, K. Araya, M. Toda, T. Chen, and M. Honda for helping with field work there. Likewise, N. L. Orlov is very grateful to R. W. Murphy for enabling him to make field surveys in Vietnam, and to A. Lathrop, T. Mason, Ho Thu Cuc, Nguyen Van Sang, S. Ryabov, and E. Rybal- tovsky for their assistance during the field work. We are also much indebted to I. Ineich and A. Dubois (MNHN) for allowing us to examine the type specimen of 7. baviensis under their care, to M. Hori for facilities for autoradiography, to S. Sakata for helping with preparations of line drawings and radiographs, and to N. B. Ananjeva for various help in preparing the draft of this manuscript. Sampling of lizards in Thailand was carried out with the permission of the National Research Council of Thailand (NRCT), Bangkok, and with financial support from a Monbusho International Scientific Program (Field Research No. 08041144; project leader: M. Matsui). For the Vietnamese specimens, permission for collecting and exporting was obtained from the Vietnam Institute of Ecology and Biological Resource (IEBR), Hanoi. LITERATURE CITED BALLINGER, R. E., J. A. LEMOS-ESPINAL, AND G. R. SMITH. 2000. Reproduction in females of three species of crevice-dwelling lizards (genus Xenosaurus) from Mexico. Stud. Neotrop. Fauna Env. 35(3): 179-183. BOURRET, R. 1939. Notes herpétologiques sur l’Indochine Francaise. XVII. Reptiles et Batra- ciens recus au Laboratoire des Sciences Naturelles de l’?Université au cours de l’année 1938. Description de trois espéces nouvelles. Ann. Bull. Gén. Inst. Publ. 6: 13-34. BROWN, W. C. AND A. C. ALCALA. 1980. Phil- ippine Lizards of the Family Scincidae. Silli- man University Natural Science Monograph Series No. 2. Silliman University Press, Dum- aguete. 264 p. DOUGHTY, P. AND R. SHINE. 1995. Life in two dimensions: natural history of the southern leaf-toed gecko, Phyllurus platurus. Herpeto- logica 51(2): 193-201. GREER, A. E. 1982. A new species of Leiolo- pisma (Lacertilia: Scincidae) from Western Australia, with notes on the biology and rela- tionships of other Australian species. Rec. Aust. Mus. 34(12): 549-573. Noo, A., R. W. MURPHY, N. ORLOV, I. DAREVSKY, AND V. S. NGUYEN. 2000. A HIKIDA ET AL.—NEW DEPRESSED-BODIED TROPIDOPHORUS 3, redescription of the Ba Vi water skink Tropido- phorus baviensis Bourret, 1939. Russ. J. Her- petol. 7(2): 155-158. PETERS, J. A. 1964. Dictionary of Herpetology. A Brief and Meaningful Description of Words and Terms Used in Herpetology. Hafner Publ., New York and London. 393 p. SMITH, M. A. 1923. A review of the lizards of the genus Jropidophorus on the Asiatic main- land. Proc. Zool. Soc. London, 1923: 775- 781. SMITH, M. A. 1935. The Fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia. II. Sauria. Taylor and Francis, London. 440 p. TAYLOR, E. H. 1936. A taxonomic study of the cosmopolitan scincoid lizards of the genus Eumeces, with an account of the distribution and relationships of its species. Univ. Kansas Sci. Bull. 24: 1-643. TAYLOR, E. H. 1963. Lizards of Thailand. Univ. Kansas Sci. Bull., 44(14): 687-1077. VITT, L. J. 1981. Lizard reproduction: habitat specificity and constraints on relative clutch mass. Am. Nat. 117(4): 506-514. VITT, L. J. 1993. Ecology of isolated open-for- mation Tropidurus (Reptilia: Tropiduridae) in Amazonian lowland rain forest. Can. J. Zool. 71: 2370-2390. WELCH, K. R. G., P. S. COOKE, AND A. S. WRIGHT. 1990. Lizards of the Orient: A Checklist. R. E. Krieger Publ. Co., Malabar, Florida. 162 p. WEN, Y.. 1992. A new species of the genus Tropidophorus (Reptilia: Lacertilia) from Guangxi Zhunag Autonomous Region, China. Asiatic Herpetol. Res. 4: 18-22. APPENDIX Specimens examined for comparisons See text for institutional acronyms. Tropidophorus baviensis—-MNHN 1948.63 (holotype), Mt. Ba Vi, Ha Tay Province, northern Vietnam; ZISP 22251 (N-37), Ba Vi National Park, Ha Tay Province, northern Vietnam; ZISP 19805, Cuc Phuong National Park, Hoa Binh Province, northern Vietnam; ZISP 21009-1, 21009-2, Myiongte, Lai Chau Province, northern Vietnam. Accepted: 28 May 2002 Current Herpetology 21(1): 25-34, June 2002 © 2002 by The Herpetological Society of Japan Taxonomic Relationships of an Endangered Japanese Salamander Hynobius hidamontanus Matsui, 1987 with H. tenuis Nambu, 1991 (Amphibia: Caudata) MASAFUMI MATSUI", KANTO NISHIKAWA!, YASUCHIKA MISAWA2, MASAICHI KAKEGAWA:?, AND TAKAHIRO SUGAHARA‘ ! Graduate School of Human and Environmental Studies, Kyoto University, Yoshida- Nihonmatsu-cho, Sakyo-ku, Kyoto 606-8501, JAPAN * Civil Engineering and Eco-Technology Consultants, Higashi-ikebukuro 2-23-2, Toshima-ku, Tokyo 170-0013, JAPAN 3 Matsugaya 3-2-8, Taito-ku, Tokyo 111-0036, JAPAN 4 Hazama-cho 1740-1, Hachioji, Tokyo 193-0941, JAPAN Abstract: We assessed the taxonomic relationships of an endangered Japanese small salamander, Hynobius hidamontanus Matsui, 1987, and its close relative H. tenuis Nambu, 1991 electrophoretically and found that they were not clearly distinguished from each other. This result, together with available morphological and ecological information, strongly indicates that H. tenuis Nambu, 1991 is a subjective junior synonym of H. hidamontanus Matsui, 1987. By this conclusion, the total distribution range of H. hidamontanus is greatly expanded, but its endangered status and the necessity of its conservation is not be changed since the habitats of this species are fragmented and not continuous. The distribution pattern of this species is interesting from the viewpoint of biogeography. Key words: Hynobiidae; Allozyme; Specific status; Conservation; Biogeography; Japan INTRODUCTION Reliable taxonomic identification of animals forms a fundamental basis for estimating spe- cies richness, which forms the most important basis in conserving animal biodiversity (May, 1995). Through the use of biochemical tech- niques, the presence of many new species has been elucidated in various animals, and this has rapidly increased the known species diver- sity of animals including amphibians (e.g., * Corresponding author. Tel/Fax: + 81-75-753- 6846; E-mail address: fumi@zoo.zool.kyoto-u.ac.jp Glaw et al., 1998). Pertinent taxonomic identification of pop- ulations of a species also affects appropriate measures of conservational and protective sta- tus of each species. Currently, species formerly considered to be widespread in their distri- bution and omitted from “Red List” tend to be split into several distinct species, and each of these needs new conservation measures because of its more restricted range of dis- tribution (e.g., Nishikawa et al., 2001). However, though the number of cases is fewer, more than one species hitherto con- sidered to be distinct from others has been 26 proven to be identical to another species through extensive biochemical studies. We herewith report such a case found in two species of Japanese hynobiid salamanders. Matsui (1987) described a salamander, H. hidamontanus, from a montane region of Nagano Prefecture, central Honshu, mainland of Japan, on the basis of morphological and biochemical evidence. The species was later listed as an endangered species in IUCN (IUCN, 1996) and Japanese (Matsui, 2000a) Red Data Books. On the other hand, Nambu (1983) reported the occurrence of hynobiid salamanders from montane regions of Toyama and Niigata Prefectures, surrounding Nagano. Although Matsui (1987: 62) pointed out that these salamanders share diagnostic charac- teristics with H. hidamontanus, Nambu (1991) described them as a separate species, H. tenuis, on the basis of a specimen from Arimine, Ooyama-machi, Toyama Prefecture. Nambu (1991), while also considering this species to be closely related to H. hidamon- tanus Matsui, 1987 in sharing several mor- phological characteristics, such as the lack of the fifth toe, the small number of vomerine teeth, and similar body proportions, argued for its difference from the latter in skull morphology (Nambu, 1991: 995). Nambu (1991), however, did not make any quantitative analyses such as statistical com- parisons of morphological characteristics using a large number of specimens or electro- phoretic assessment of genetic differentiation, both of which are routine in the taxonomic study of hynobiid salamanders (e.g., Matsui and Miyazaki, 1984; Matsui, 1987; Matsui et al., 1992). The taxonomic validity of H. tenuis, there- fore, remains dubious (Matsui, 1996b, 2000a), but no study has been conducted to reassess the taxonomic relationship of this species and H. hidamontanus. Because, as noted above, H. hidamontanus is now considered to be endangered (IUCN, 1996; Matsui, 2000a), elucidation of the taxonomic relationships of H. tenuis and H. hidamontanus has become Current Herpetol. 21(1) 2002 urgent from the viewpoint of species conser- vation. Small salamanders of the genus Hynobius exhibit high inter- and intraspecific variations in morphology, and, therefore, species iden- tification is usually difficult without locality information (Matsui and Miyazaki, 1984). For studies of intra- and interspecific variations and taxonomy among morphologically similar urodelan species, electrophoretic studies con- tribute the best data (e.g., Matsui et al., 1992, 2000; Jackman and Wake, 1994; Highton, 1999). Indeed, recent genetic analyses by electrophoresis have elucidated many taxo- nomic problems in Hynobius (e.g., Matsui 1987; Nishikawa et al., 2001). Herein, we examined the taxonomic validity of H. tenuis chiefly by means of this technique. MATERIALS AND METHODS The small Japanese salamanders generally occur allopatrically (Matsui, 1996a), and only One species has been recorded from each of the type localities of H. hidamontanus and H. tenuis (Matsui, 1996b). Therefore, there is little possibility that samples collected from each of these localities will include more than One species. In addition to this, H. hida- montanus closely resembles H. lichenatus in external morphology (Matsui and Matsui, 1980), but is reported to form a genetic group not with the latter but with H. nebu- losus (Matsui, 1987). We therefore used both H. lichenatus and H. nebulosus for compar- isons. Further, because populations of many small Japanese salamanders are in decline (Matsui, 2000a), we need to refrain from collecting a large number of specimens. Bearing this in mind, we used a total of 46 specimens to evaluate genetic relationships among four species: H. tenuis from Toyama (sample 1, topotypic population, N=13), and Gifu (sample 2, N=7) Prefectures, H. hida- montanus from Nagano Prefecture (sample 3, topotypic population, N=16), and one population each of H. nebulosus from Shiga Prefecture (sample 4, N=4) and H. lichenatus MATSUI ET AL.—TAXONOMY OF ENDANGERED SALAMANDER 27 Fic. 1. 500km A map of northern Central Honshu, Japan, showing sampled localities of hynobiid species used in this study. For sample numbers, refer to text. Coarsely dotted area >1000 m asl. Finely dotted area >2000 m asl. from Yamagata Prefecture (sample 5, N=6) (Fig. 1). In the laboratory, liver samples were removed from fully anesthetized salamanders and stored at -84 C. Voucher specimens were fixed in 10% formalin, preserved in 70% ethanol, and deposited in the Graduate School of Human and Environmental Studies, Kyoto University (KUHE) (see Appendix). Supernate fraction and homogenates were examined by standard horizontal starch gel electrophoresis (Shaw and Prasad, 1970; Ayala et al., 1972), containing Starch Art (Starch Art Corp., Smithville, USA) and Connaught starch (Connaught Lab., Ontario, Canada) mixed in a 4:1 ratio and then suspended in buffer at a concentration of 11.5%. Enzymes exam- ined, locus designations, and buffer systems employed are listed in Table 1. Staining methods, genetic interpretations of allozyme data, enzyme nomenclature, E. C. numbers, abbreviations, and isozyme designations follow Nishikawa et al. (2001). We studied 19 enzyme systems coded by 24 presumptive loci (Table 1). Genetic vari- ability for each sample was assessed by the standard analyses: percentage loci polymor- phic (P, a locus is considered as polymor- phic unless the frequency of the most common allele exceeds 0.95), the mean heterozygosity 28 Current Herpetol. 21(1) 2002 TABLE 1. among Hynobius species. Enzymes Enzymes, presumptive loci, and buffer systems used in the analyses of allozyme variations Aconitate hydratase Aspartate aminotransferase Aspartate aminotransferase Alcohol dehydrogenase Fumarate hydratase Glucose-6-phosphate isomerase Glycerol-3-phosphate dehydrogenase Glutamate dehydrogenase Guanine deamidase 3-Hydroxybuthyrate dehydrogenase Isocitrate dehydrogenase L-Lactate dehydrogenase L-Lactate dehydrogenase Malate dehydrogenase Malate dehydrogenase Malic enzyme** Malic enzyme** Peptidase (leucyl-glycine) Phosphoglucomutase Phosphoglucomutase Phosphogluconate dehydrogenase Sorbitol dehydrogenase Superoxide dismutase Xanthine dehydrogenase E.C. numbers Locus Buffer system* 4.2.1.3 mAcoh-A TC8 2.6.01 mAat-A CAPM6 226 hel sAat-A CAPM6, TC7 1S Adh-A TBE8.7 4.2.1.1 Fumh-A TBE8.7 3321.9 Gpi-A CAPM6 LAS G3pdh-A TC8 1.4.1.3 Gtdh-A TC8 3.5.4.3 Gda-A TBE8.7 Pele .30 Hbdh-A CAPM6 Pele 42 mIdh-A TCT Pel 1.27 Ldh-A CAPM6, TC7 1 hee) Ldh-B CAPM6, TC7 Ja eS es ig mMdh-A CAPM6, TC8 Pt.1e37 sMdh-A CAPM6, TC8 1.1.1.40 mMdhp-A TC7 1.1.1.40 sMdhp-A TC7 3.4.11.- Pep-A TBE8.7 5.4.2.2 Pgm-A TC7 5.4.2.2 Pgm-C TC? 1.1.1.44 Pgdh-A TC7 1.1.1.14 Sdh-A CAPM6 ja) ses lad | Sod-A TBE8.7 1.1.1.204 Xdh-A TC8 * Buffer systems—CAPM6: Citrate-aminopropylmorpholine, pH=6.0 (Clayton and Tretiak, 1972), TC7: Tris- citrate, pH=7.0 (Shaw and Prasad, 1970), TC8: Tris-citrate, pH=8.0 (Clayton and Tretiak, 1972), TBE8.7: Tris-borate-EDTA, pH=8.7 (Boyer et al., 1963). ** NADP-dependent malate dehydrogenase. by direct count (H), and the mean number of electromorphs per locus (A). In order to estimate overall genetic diver- gence among samples, we calculated two genetic distances: Nei’s (1978) unbiased genetic distance and modified Rogers’ distance (Wright, 1978). We inferred patterns of phenetic simi- larities among samples from Nei’s (1978) distance clustered by the UPGMA algorithm (Sneath and Sokal, 1973), and modified Rogers’ distance clustered by the Neighbor- joining (NJ) procedure (Saitou and Néi, 1987). Hynobius lichenatus (sample 5) was designated as an outgroup in the NJ method. We ran these analyses by employing BIO- SYS-1 (Swofford and Selander, 1981) and PHYLIP vers. 3.5 C computer programs (Felsenstein, 1993). RESULTS Fifty-two alleles were detected at 24 puta- tive loci of which 16 (other than mAat-A, mAcoh-A, Fumh-A, Gtdh-A, sMdhp-A, Pgm- A, Sdh-A, and Sod-A) were variable (Table 2). The most variable loci were Gda-A, MATSUI ET AL.—TAXONOMY OF ENDANGERED SALAMANDER AS) Gpi-A, and Pgdh-A, each with four alleles, followed by Adh-A, G3pdh-A, mIdh-A, Ldh- B, sMdh-A, and Xdh-A, each with three alleles. Adh-A in H. nebulosus, and sAat-A, Adh-A, Hbdh-A, and sMdh-A in H. lichenatus were fixed by unique alleles that were not shared with other species. We found no unique alleles in either H. tenuis or H. hidamontanus. However, we found some differentiations of allelic frequencies in Gda-A, Gpi-A, and Ldh- A, among H. tenuis and H. hidamontanus (samples 1-3). The mean number of electromorphs per locus (A) varied from 1.1 to 1.3, the per- centage of polymorphic loci (P) from 4.2 to 25.0, and the mean heterozygosity (H) from TABLE 2. Allele frequencies at 16 polymorphic loci of Hynobius samples examined. For sample numbers, refer to Fig. 1 and text. A=mean number of alleles per locus; P=percentage of loci; H=mean heterozygosity by direct count. Species and sample number (N) tenuis tenuis hidamontanus nebulosus lichenatus Locus 1 (13) 2 (7) 3 (16) 4 (4) 5 (6) sAat-A al .000 al .000 al .000 al .000 b1.000 Adh-A al .000 al .000 al .000 c1.000 b1.000 Gda-A a0.923 a0.143 al .000 b1.000 b1.000 d0.077 c0.429 d0.429 G3pdh-A b1.000 b1.000 a0.063 c1.000 c1.000 b0.938 Gpi-A b0.462 b1.000 b1.000 a0.375 b0.833 c0.192 c0.500 d0.167 d0.346 d0.125 Hbdh-A al.000 al.000 al .000 al .000 b1.000 mIdh-A a0.962 al .000 al .000 a0.500 a0.833 b0.038 c0.500 b0.167 Ldh-A a0.077 al.000 b1.000 al.000 b1.000 b0.923 Ldh-B c1.000 c1.000 c1.000 a0.500 c1.000 b0.500 mMdh-A b1.000 b1.000 b1.000 b1.000 a0.167 b0.833 sMdh-A c1.000 c1.000 c1.000 a0.250 b1.000 c0.750 mMdhp-A b1.000 b1.000 b1.000 b1.000 a0.167 b0.833 Pep-A b1.000 b1.000 b1.000 al .000 al .000 Pgdh-A c1.000 c1.000 c1.000 al.000 a0.083 b0.083 c0.083 d0.750 Pgm-C al.000 al .000 a0.938 a0.125 al .000 b0.063 b0.875 Xdh-A a0.923 al .000 a0.969 a0.750 al .000 c0.077 b0.031 c0.250 A 1.3 1.1 Vt 1.3 8) P 16.4 4.2 8.3 25.0 20.8 H 0.010 0.000 0.003 0.031 0.021 30 Current Herpetol. 21(1) 2002 0.00 to 0.03 (Table 2). The highest A, P, and H values were found in AH. nebulosus (sample 4), while the lowest A value was seen in H. tenuis from Gifu Prefecture (sample 2) and H. hidamontanus (sample 3), and the lowest P and H values were shown by H. tenuis from Gifu Prefecture (sample 2). As shown in Table 3, the highest Nei’s (1978) and modified Rogers’ distances were obtained between H. tenuis (sample 2) and H. lichenatus (sample 5) (0.46 and 0.60, respectively). By contrast, the lowest distances were found between H. tenuis from Toyama of H. tenuis were not clustered as one group. In the UPGMA tree, relationships among the three groups, the tenuis-hidamontanus group, H. nebulosus (sample 4), and H. lichenatus (sample 5) were not solved clearly, but dis- tances among them were substantially large. TABLE 3. Matrix of Nei’s (1978) unbiased genetic distance (above diagonal) and modified Rogers’ distance (Wright, 1978: below diagonal). Diagonal numbers show the numbers of loci fixed by unique allele(s). Samples 1 y) 3 4 5 Prefecture (sample 1) and H. hidamontanus (sample 3) (0.01 and 0.10, respectively). 1 0 0.067 0.009 0.429. 0.419 Results of UPGMA and NJ analyses are shown in Fig. 2. In both phenograms, H. 2 veo eae OOO ae tenuis from Toyama Prefecture (sample 1) 3 0.102 0.255 0 0.457 0.406 joined with H. hidamontanus (sample 3) 4 0.573 0.551 0.592 1 0.433 first, and then with AH. tenuis from Gifu Prefecture (sample 2): the two populations 2 we ee USS) veils 4 A 3 hidamontanus 1 3 tenuis 2 4 nebulosus 5 lichenatus 0.4 0.3 0.2 0.1 0 Nei's (1978) D B 0.1 Modified Rogers' D (Wright, 1978) 3 hidamontanus 1 tenuis 2 4 nebulosus 5 lichenatus FIG. 2. UPGMA tree based on Nei’s (1978) unbiased genetic distances (A) and an NJ tree based on modified Rogers distances (Wright, 1978) rooted by the outgroup (sample 5) (B), among five samples studied. For sample numbers, refer to text. MATSUI ET AL.—TAXONOMY OF ENDANGERED SALAMANDER 31 DISCUSSION Recent biochemical reassessments of the Japanese small salamanders have resulted in elucidating the presence within a named spe- cies of many genetically distinct populations that require taxonomic splitting (cf. Matsui, 2000b). However, the present biochemical study of H. hidamontanus and H. tenuis has provided an opposite conclusion, 1.e., the two species are genetically conspecific. As shown by the values of A, P, and H, each sample of H. hidamontanus and H. tenuis did not exhibit particularly great genetic differentiation within a locality, and so there was little possibility of there being a mixture of more than one species. The topotypic populations of H. tenuis from Toyama and H. hidamontanus from Nagano Prefectures first formed a group with the smallest genetic distance (Nei’s D=0.01) among all pairs compared, and the other population of H. tenuis from Gifu Prefecture had a sister relationship to this group. This very small genetic differentiation between the topotypic populations offers little support for taxonomic separation of these two species. Matsui (1987) reported the Nei’s (1978) D between H. hidamontanus and H. nebulosus to be 0.30, which value is smaller than what we found in this study (0.46), notwithstanding the use of the same populations in the two studies. This discordance might have been induced by differences in specimens used, number of loci examined, and the electro- phoretic conditions, especially the larger num- ber of buffer systems employed in this study which might have separated more electro- morphs. In this way, the absolute genetic distances derived from allozymic data them- selves are variable, and were not directly used for outlining species boundaries. Relative distances from allozymic data, however, show a good taxonomic standard (Matsui, 2000b). Hynobius tenuis and H. hidamontanus are genetically not considered to be different species because their genetic distances are much smaller than are found among already named species or even within one species of Hynobius (D>0.22: e.g., Matsui 1987; Matsui et al. 2000; Nishikawa et al. 2001). Allelic compositions of Gda-A, Gpi-A, and Ldh-A show some genetic fragmentation among H. tenuis and H. hidamontanus (samples 1-3). Matsui (1987) already reported genetic varia- tions in H. hidamontanus. Namely, he found fixed differences in two loci and Nei’s (1978) D of 0.20 between two populations of this species. However, he treated them as One species because of their morphological similarities. Nambu (1991) argued that AH. tenuis is specifically distinct from H. hidamontanus on the basis of different conditions in five skull characters (angle of articulation of maxilla and premaxilla, length and shape of maxilla, shape of premaxilla, shape of vomer, and proportion of vomerine teeth series). However, the skull morphology sometimes greatly varies even within a single species in Hynobius salamanders (Ebitani, 1952). Our preliminary examinations of morpho- logical variations in H. tenuis and H. hida- montanus specimens including those from their type localities revealed the absence of tangible differences in these two species (Matsui, 1996b; Matsui et al., unpublished data). In addition to these morphological similarities, descriptions in the literature (Matsui and Matsui, 1980; Nambu, 1983) and our extensive ecological observations in the field strongly indicate that they live and spawn in very similar habitats (slowly flowing water in marshes and swamps near montane forests), and share a similar ecological niche. From these lines of evidence, we have no other choice but to conclude that AH. tenuis Nambu, 1991, is a subjective junior synonym of H. hidamontanus Matsui, 1987, and that separation of the two species even at the subspecific level is not necessary. Although no further studies have been made, the pop- ulation of a salamander from O-umi, Niigata Prefecture (Matsui, 1987) should also be included in this species from its diagnostic characteristics (see Matsui, 1987: 62). 32 A case similar to that reported in the present paper has been known in some plethodontid salamanders; Feder et al. (1978) found little allozymic variation between Plethodon dunni and P. gordoni and also found that none of the diagnostic characters mentioned in the original description of P. gordoni were reli- able. Consequently, they (Feder et al., 1978) synonymized P. gordoni as a color variant of P. dunni. The discovery of H. hidamontanus (Matsui and Matsui, 1980; Matsui, 1987) and H. tenuis (Nambu, 1983, 1991) filled the distributional gaps of lentic breeding salamanders known previously (Sato, 1943; Nakamura and Uéno, 1963; Matsui and Miyazaki, 1984). The geohistory of the Japanese islands including the formation of the Fossa Magna (=Itoigawa- Shizuoka tectonic line) was supposed to have strongly affected the formation of patterns in distribution and divergence of these animals (Matsui, 1987; Matsui et al., 2000; Nishikawa et al., 2001). Until now, the high montane region of the Hida Mountains could be considered a geographic barrier that had promoted the divergence of H. hidamontanus and H. tenuis, occurring on its eastern and western sides, respectively. The close genetic similarities demonstrated in the present study, however, indicate the invasion either (1) from west of the population ancestral to the present eastern population (H. hidamontanus sensu stricto), or (2) from east of the ancestral population of the present western population (H. tenuis), possibly through a route north of the north- ernmost edge of the Hida Mountains. The relationships among H. hidamontanus (sensu lato), H. nebulosus, and H. lichenatus were not clearly resolved in the present study, but through the use of more extensive samples, Matsui (1987) considered that H. hidamon- tanus belongs to one of the two large lin- eages of Japanese lentic breeding Hynobius and is phylogenetically closer to H. nebulosus from western Japan than to the eastern lineage including H. lichenatus. If this is the case, the first hypothesis advanced above seems Current Herpetol. 21(1) 2002 more plausible. Hynobius hidamontanus (now including H. tenuis) occurs on the western side of the Fossa Magna and H. lichenatus occupies its eastern side (Matsui, 1987). If these two species actually belong to different lineages, as stated above, the possible dichot- omy in the lentic breeding Hynobius in older times might have prevented further eastwards invasion of H. hidamontanus. The present results greatly expand the range of distribution of H. hidamontanus, which was formerly limited to around the type locality in Hakuba Village, Nagano. Prefecture (Matsui, 1987, 2000a). The species (sensu stricto) has been listed in Red Data Books for the reason that its habitats are limited and being destroyed (IUCN, 1996; Matsui, 2000a). We need to maintain this conserva- tional status of H. hidamontanus because, even after synonymizing H. tenuis with it, the actual habitats, though ranging over a seemingly large area, are largely isolated from each other and the size of each local population is obviously very small (Matsui, 2001: 198). ACKNOWLEDGMENTS We would like to thank T. Abe and S. Tanabe for help in collecting the specimens examined. 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APPENDIX Materials examined Voucher specimens used for electrophoresis are stored at the Graduate School of Human and Environmental Studies, Kyoto University (KUHE). Sample 1: Hynobius tenuis from Ooyama- machi, Toyama Prefecture (KUHE 13236- 13248). Sample 2: H. tenuis from Shirakawa-mura, Gifu Prefecture (KUHE 13262-13263, 13265- 13269). Sample 3: H. hidamontanus from Hakuba- mura, Nagano Prefecture (KUHE 9481, 9484- 9485, 9498-9510). Sample 4: H. nebulosus from Hino-cho, Shiga Prefecture (KUHE 9599-9600, 16937- 16938). Sample 5: HA. lichenatus from Oguni-cho, Yamagata Prefecture (KUHE 18421, 18426- 18427, 18432, 18436-18437). Accepted: 29 May 2002 Current Herpetology 21(1): 35-41, June 2002 © 2002 by The Herpetological Society of Japan Karyotypes of Four Agamid Lizards from Southeast Asia HIDETOSHI OTA!", CHEONG-HOONG DIONG?’, ENE-CHOO TAN?, AND HOI-SEN YONG‘ ! Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa, 903-0213, JAPAN 2 Natural Sciences, National Institute of Education, Nanyang Technological University, I Nanyang Walk, SINGAPORE 637616 3 Defence Medical Research Institute, Clinical Research Centre, 10 Medical Drive, SINGAPORE 117597 4 Institute of Biological Sciences, University of Malaya, 50603, Kuala Lumpur, MALA YSIA Abstract: We karyotyped four lizards, Acanthosaura armata, Bronchocela cristatella, Calotes emma, and C. versicolor, all belonging to the tropical Asian clade of the family Agamidae. The karyotype of A. armata consisted of 12 metacentric macrochromosomes and 20 microchromosomes, whereas B. cris- tatella had 14 metacentric macrochromosomes and 20 microchromosomes. Except for the presence of 22 microchromosomes, the karyotypes of the two Calotes species were similar to that of A. armata. The 20 microchromosome state in the A. armata karyotype may have emerged in the ancestral lineage common to Gonocephalus robinsonii, whose karyotype also exhibits a 12M + 20m format. Comparison of the present results with previously published information suggests the presence of cryptic taxonomic diversity in B. cristatella and C. ver- sicolor. Key words: Karyotype; Chromosomal variation; Agamidae; Acanthosaura armata; Bronchocela cristatella; Calotes emma; C. versicolor; Southeast Asia. Macey et al., 2000). With respect to the INTRODUCTION The family Agamidae is an Old-World representative of the iguanian lizards, and is much diverged in Australia, Africa, tropical and temperate Asia, and the Indo-Australian Archipelago. Recent molecular studies yielded a few tenable hypotheses for early stages of divergence in this family (Honda et al., 2000; * Corresponding author. Tel: + 81-98-895-8937; Fax: + 81-98-895-8966; E-mail address: ota@sci.u- ryukyu.ac.jp phylogeny and systematics at generic and infrageneric levels, however, much is yet to be done for the Agamidae. Chromosomal approaches to phylogeny have been receiving persistent, formidable criticism chiefly on the ground of adequacy in analytical procedures adopted and in presumptions regard- ing the mode and direction of character trans- formation (e.g., Kluge, 1994). In some cases, however, chromosomal data have been provid- ing very convincing evidence for monophyly in some agamid groups (e.g., Diong et al., 36 Current Herpetol. 21(1) 2002 2000; Honda et al., 2002). Also, chromosomal investigation may clarify biological species that are cryptic to morphological approaches (King, 1993: see Ota [1988] for example of an agamid group). In consideration of these merits, we have been conducting a chromosomal survey of Southeast Asian agamid lizards (Diong et al., 2000). In this paper, we describe karyotypes of four species obtained in this survey. MATERIALS AND METHODS A total of 16 specimens representing four species, Acanthosaura armata, Bronchocella cristatela, Calotes emma, and C. versicolor, were karyotyped (Table 1). All of these species belong to group V of the Agamidae sensu Moody (1980) (see Honda et al. [2000, 2002] and Macey et al. [2000: as the South Asian clade] for the monophyly of this group). Mitotic cell preparations were made by the bone-marrow air-dry method following Diong et al. (2000). Then they were stained in 6% Gurr Giemsa (BDH) solution, and were pho- tographed with a Nikon Optiphot 2 Photo- micrography camera using Kodak TMAX ASA 100 film. Karyotypes were determined for each indi- vidual lizard on the basis of 10-16 well-spread metaphase cells. Chromosomes in each pho- tograph were paired according to the similarity in size and shape, and resultant pairs were arranged in order of decreasing size. For the calculation of arm ratio for each chro- mosome pair, the lengths of chromosome arms were measured with a CALCOM digitizer. Terminology for chromosomal description fol- lows Levan et al. (1964) as modified by Green and Sessions (1991), and the karyotype formula follows Peccinini-Seale (1981). Voucher speci- mens were deposited in the Zoological Ref- erence Collection, Department of Biological Sciences, National University of Singapore (ZRC). RESULTS The karyotype of male Acanthosaura armata consisted of 16 pairs. Of these, the six largest pairs were all metacentric macrochromosomes, and the remaining 10 were microchromosomes (Fig. 1A). Fundamental numbers (N. F.) thus equaled 44. In the macrochromosome group, there was a marked size gap between pairs 5 and 6 (also see Appendix). The karyotype of male Bronchocela cristatella was similar to that of A. armata in the clear division of component chromosomes into metacentric macrochromosomes and micro- chromosomes. However, the macrochromo- some group of the B. cristatella karyotype had an additional pair, which made its diploid and fundamental numbers 34 and 48, respec- tively (Fig. 1B). Also, the karyotype of B. cristatella differed from that of A. armata in the presence of a distinct size gap between pairs 1 and 2, besides between pairs 5 and 6 (Fig. 1B, Appendix). TABLE 1. Localities, sizes, and sexual composition of samples of four agamid species examined in this study. N Species Males Females Total Locality Acanthosaura armata 3 0 3 Pulau Tioman, near Peninsular Malaysia Bronchocela cristatella 4 0 4 Singapore Calotes emma 1 0 1 Banding, Peninsular Malaysia Calotes versicolor 5 3 8 Singapore OTA ET AL.—KARYOTYPES OF AGAMID LIZARDS by) “4 *o ‘#64 LL 8 Fic. |. The karyotypes of Calotes emma and C. versicolor were also similar to that of A. armata in the number (six pairs), shape (meta- centric), and size arrangement (prominent gap only between pairs 5 and 6) of macrochromo- somes (Fig. 2, Appendix). However, they dif- fered from the A. armata karyotype by having an additional pair of microchromosomes, which made the diploid and fundamental numbers 34 and 46, respectively. In all karyotypes examined, no sex chro- mosome heteromorphism, or intraspecific chro- mosomal variations were evident, although female karyotypes were not available for A. armata, B. cristatella, and C. emma. “@ os ae o*% . | / ‘eRe nba a SSDI lett tips o A LL SE athe A et Delp RINLn io et LtR Resta NaN tEteR atte a Male katyotypes of (A) Acanthosaura armata, and (B) Bronchocela cristatella. Bar equals 5 um. DISCUSSION This is the first chromosomal description for the genus Acanthosaura. The karyotype of A. armata differs from most other known karyotypes of the group V species (Moody, 1980; Honda et al., 2000) in having only 20 microchromosomes, because most other mem- bers of this tropical Asian clade have another pair of microchromosomes (e.g., Moody, 1980; Witten, 1983; Solleder and Schmid, 1988; Ota and Hikida, 1989). Within this group, the 20 microchromosome state is shared only with Bronchocela cristatella (see Solleder and Schmid [1988] and below), Calotes versi- 38 Current Herpetol. 21(1) 2002 FIG. 2. Male katyotypes of (A) Calotes emma, and (B) C. versicolor. Bar equals 5 um. color in one report (see below), and Gono- cephalus robinsonii (see Diong et al., 2000). In contrast, this state is common in the clade consisting of groups II-IV of Moody (1980) (see Honda et al. [2000, 2002] and Macey et al. [2000: as the Australian-New Guinean clade]) as pointed out by Witten (1983). Diong et al. (2000), recognizing the common occurrence of 20 microchromosomes in G. robinsonii and some Australian-New Guinean species, sur- mised that G. robnsonii might have been derived from endemic Australian radiation, followed by long dispersals like Physignathus cocincinus (Honda et al., 2000). This view, however, was negated by a molecular phylo- genetic investigation by Honda et al. (2002), which clearly indicated the allocation of G. robinsonii in group V. Likewise, recent molec- ular phylogenetic studies (Honda et al., 2000, 2002; Macey et al. 2000) negate a close affinity of Acanthosaura with the Australian-New Guinean agamids, although none of these studies examined A. armata. Relatively close allocation of Acanthosaura (as represented by A. crucigera) with G. robinsonii on the molecular phylogenetic tree of Honda et al. (2002) suggests that the deletion of a pair of microchromosomes may have occurred in their common ancestral lineage. This assumtion needs substantial verification on the basis of additional chromosomal data for relevant taxa, particularly A. crucigera and other Acan- thosaura and Phoxophrys species (Honda et al., 2002). The karyotypes of B. cristatella, C. emma, and C. versicolor were already reported in OTA ET AL.—KARYOTYPES OF AGAMID LIZARDS | 39 TABLE 2. Karyotypes of the four agamid species examined in this study. M=macrochromosomes; m=microchromosomes. Sources are as follows: 1, this study; 2, Solleder and Schmid (1988); 3, Moody (1980); 4, De Smet (1981); 5, Singh and Bhatnagar (1987); 6, papers cited in Das and Ota (1998) exclusive of 4 and 5. Species 2n Arm nos. in macrochromosomes Chromosomal formula Source Acanthosaura armata 32 24 12M + 20m 1 Bronchocela cristatella 34 28 14M + 20m le? 48 28 28M + 20m 3 Calotes emma 34 24 12M + 22m 10K 97 Calotes versicolor 34 24 12M + 22m 1,6 32 24 12M + 20m 4 34-62 24 . 12M + (22-50)m 5 some previous studies (Table 2). However, absence of locality data for materials used in most of those studies makes our data deserving of publication, because our data may con- tribute to the detection of cryptic taxonomic diversity in those species (Ota et al., 2001). For example, Moody (1980), on the basis of unpublished information from W. P. Hall, listed the karyotype of B. cristatella as con- sisting of 28 acrocentric macrochromosomes and 20 microcromosomes, an arrangement dramatically different from the “conspecific” karyotype subsequently reported by Solleder and Schmid (1988), and in the present study (Table 2). It is thus likely that B. cristatella in the current definition actually contains more than one species. In C. versicolor, the microchromosome number seems to vary, because, although most authors reported the number to be 22, De Smet (1981) described one male of unknown locality as having 20 microchromosomes only. Furthermore, Singh and Bhatnagar (1987) reported a remarkable variation in the micro- chromosome number in some Indian materials (Table 2). Although one or both of these records may simply have resulted from the miscounting of those tiny elements, it is not surprizing if C. versicolor in the current def- inition is actually a composite of more than one cryptic species with different microchro- mosome numbers, considering the extensive morphological variation in this broadly dis- tributed species (Auffenberg and Rehman, 1993). In both cases, absence of reliable locality information in most crucial works (Hall in Moody [1980], and Solleder and Schmid [1988] for the B. cristatella karyotype; and De Smet [1981], and a few other works [Table 2] for the C. versicolor karyotype) imposes serious difficulties in utilizing their chromosomal data for unequivocal solutions of taxonomic prob- lems implied therein. Accumulation of chro- mosomal data with reliable locality information and voucher specimens is definitely needed for the appropriate estimation of taxonomic diversity in these and other agamid species from Southeast Asia. ACKNOWLEDGMENTS We thank the Institute of Biological Sci- ences, University of Malaya, Kuala Lumpur, for provision of facilities for experiments. This study was partially supported by an Academic Research Grant from the Nanyang Technological University of Singapore (No. RP21/97, to CHD). LITERATURE CITED AUFFENBERG, W. AND H. REHMAN. 1993. Studies on Pakistan reptiles. Pt. 3. Calotes versicolor. Asiatic Herpetol. Res. 5: 14-30. Das, I. AND H. OTA. 1998. A checklist of chro- 40 mosome numbers of South Asian reptiles. Hamadryad 23(2): 179-193. DE SMET, W. H. O. 1981. Description of the orcein stained karyotypes of 27 lizard species (Lacertilia Reptilia) belonging to the families Iguanidae, Agamidae, Chameleontidae and Gekkonidae (Ascalabota). Acta Zool. Antverp. 76: 35-72. DIONG, C.-H., M.-H. Low, E.-C. TAN, H.-S. YONG, T. HIKIDA, AND H. OTA. 2000. On the monophyly of the agamid genus Gono- cephalus Kaup, 1825 (Reptilia: Squamata): a chromosomal perspective. Curr. Herpetol. 19(2): 71-79. GREEN, D. M. AND S. K. SESSIONS. 1991. Nomen- clature for chromosomes. P. 431-432. In: D. M. Green and S. K. Sessions (eds.), Amphibian Cytogenetics and Evolution. Academic Press, San Diego, California. HONDA, M., H. OTA, M. KOBAYASHI, J. NABHITABHATA, H.-S. YONG, S. SENGOKU, AND T. HIKIDA. 2000. Phylogenetic relation- ships of the Family Agamidae (Reptilia: Igua- nia) inferred from mitochondrial DNA sequences. Zool. Sci. 17(4): 527-537. HONDA, M., H. OTA, S. SENGOKU, H.-S. YONG, AND T. HIKIDA. 2002. Molecular evaluation of phylogenetic significances in the highly divergent karyotypes of the genus Gonocephalus (Reptilia: Agamidae) from tropical Asia. Zool. Sci. 19(1): 129-133. KING, M. 1993. Species Evolution: The Role of Chromosome Change. Cambridge University Press, Cambridge. 336 p. KLUGE, A. G. 1994. Principles of phylogenetic systematics and the informativeness of the karyotype in documenting gekkotan lizard relationships. Herpetologica 50(2): 210-221. LEVAN, A., K. FREDGA, AND A. A. SANDBERG. 1964. Nomenclature for centromeric position . | Current Herpetol. 21(1) 2002 on chromosomes. Hereditas 52: 201-220. MACEY, J. R., J. A. SCHULTE, II, A. LARSON, N. B. ANANJEVA, Y. WANG, R. PETHIYAGODA, N. RASTEGAR-POUYANI, AND T. J. PAPENFUSS. 2000. Evaluating trans-Tethys migration: an example using acrodont lizard phylogenetics. Syst. Biol. 49(2): 233-256. Moopy, S. M. 1980. Phylogenetic and Historical Biogeographical Relationships of the Genera in Family Agamidae (Reptilia: Lacertilia). Unpub- lished doctoral dissertation, University of Michi- gan, Ann Arbor, Michigan. 373 p. OTA, H. 1988. Karyotypic differentiation in an agamid lizard, Japalura swinhonis swinhonis. Experientia 44: 66-68. OTA, H. AND T. HIKIDA. 1989. Karyotypes of three species of the Genus Draco (Agamidae: Lacertilia) from Sabah, Malaysia. Jpn. J. Her- petol. 13(1): 1-6. OTA, H., T. HIKIDA, J. NABHITABHATA, AND S. PANHA. 2001. Cryptic taxonomic diversity in two broadly distributed lizards of Thailand (Mabuya macularia and Dixonius siamensis) as revealed by chromosomal investigations (Rep- tilia: Sauria). Nat. Hist. J. Chulalongkorn Univ. 1(1): 1-7. PECCININI-SEALE, D. 1981. New developments in vertebrate cytotaxonomy. IV. Cytogenetic studies in reptiles. Genetica 56(2): 123-148. SINGH, S. AND V. S. BHATNAGAR. 1987. Karyo- typic anomaly in the common Indian garden lizard, Calotes versicolor Daudin. Nucleus 30(1/ 2): 28-30. SOLLEDER, E. AND M. SCHMID. 1988. Cytoge- netic studies on Sauria (Reptilia). I. Mitotic chromosomes of the Agamidae. Amphibia- Reptilia 9(3): 301-310. WITTEN, G. J. 1983. Some karyotypes of Aus- tralian agamids (Reptilia: Lacertilia). Aust. J. Zool. 31(4): 533-540. OTA ET AL.—KARYOTYPES OF AGAMID LIZARDS 41 APPENDIX Quantitative morphological data for macrochromosomes of the four agamid species examined in this study (x+SD). Abbreviations are: AA, Acanthosaura armata; BC, Bronchocela cristatella; CE, Calotes emma; CV, Calotes versicolor; RR, relative length (%); AR, arm ratio (or centromeric ratio). Macrochromosome pairs 1 2 3 4 5 6 q AA RR 23.08 20.22 1792 15.79 13:37 9.65 +1.14 1323 +1.14 tA) 2 +1.50 +129 AR 118 1.26 1.19 1.18 ht 1215 +0.15 a). 22 +0.14 +0.08 +0.11 +0.17 BC RR 22.38 16.52 15.40 14.33 12.55 10.07 8.60 1,52 +1.48 +0.80 +0.85 +0.77 +099 +1.15 AR 1.51 1.37 1.29 1.28 je 1.29 1.20 +0.26 +0.38 +0.31 a7 +0.16 +0.28 20216 CE RR 22,311 21.48 16.82 16.03 13.75 9.61 +1.07 +0.38 nua 0 5) +0.90 +1.15 +0.04 AR 1.08 1.33 108 1.12 1.20 Ne 1 +0.08 a(),22 +0.05 +0.08 +0.10 +0.07 CV RR 23.31 20.86 16.86 16:12 13.36 9.49 +0.50 +1.74 +0.48 A202 +0.67 1.13 AR 1-31 1.38 1.13 1.17 1.17 1:12 £-().32 +0.29 +0.21 +0.11 +0.21 +0.10 Accepted: 31 May 2002 Current Herpetology 21(1): 43-50, June 2002 © 2002 by The Herpetological Society of Japan Early Growth of Elaphe quadrivirgata from an Insular Gigantic Population AKIRA MORI!” AND MASAMI HASEGAWA2 ! Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto 606-8502, JAPAN * Department of Biology, Faculty of Science, Toho University, Funabashi, Chiba 274- 8510, JAPAN Abstract: Elaphe quadrivirgata on Tadanae-jima Island shows a clear insular gigantism. Based on data from captive animals, we compared growth rates over the first two years after hatching between this population and a conspe- cific population from the Japan mainland. The purpose of this experiment was to test the hypothesis that early growth rates in snakes in the insular gigan- tism population are higher than those in conspecific non-gigantic populations under the same feeding schedule. Growth rates in snout-vent length and body mass of Tadanae-jima snakes were not higher than those of main island snakes, and thus, the hypothesis was rejected. This result suggests that the gigantism in FE. quadrivirgata on Tadanae-jima Island is not caused by a genetically based modification for rapid growth before maturation. Key words: Snake INTRODUCTION Body size is a fundamental character that affects almost all life history traits of animals (Schmidt-Nielsen, 1984). Intraspecific geo- graphic variations in body size would, there- fore, offer important opportunities to study microevolution of organisms. To understand the process of microevolution, it is essential to elucidate the proximate factors that cause the phenotypic variation in body size. Insular gigantism is a notable example of geographic body size variations and has been * Corresponding author. Tel/Fax: + 81-75-753- 4075; E-mail address: gappa@ethol.zool.kyoto- u.ac.jp Elaphe quadrivirgata; Growth; Izu Islands; Insular gigantism; reported in many animal taxa including snakes (Case, 1978; Schwaner, 1985; Hasegawa and Moriguchi, 1989; King, 1989; Kohno and Ota, 1991; Mori, 1994; Mori et al., 1999). Prac- tically, gigantism is determined by comparing standard sizes of animals among populations, i.e€., average or maximum sizes of mature individuals. In snakes, which show asymptotic growth after maturity, gigantism can be caused by several mechanisms and proximate factors that are not mutually exclusive: differences in age structure, growth rates, size at maturity, or asymptotic size (Stamps, 1993). Each of these differences may stem from either genet- ically based local adaptations or phenotypi- cally plastic variations largely due to local environmental conditions such as high abun- +4 Current Herpetol. 21(1) 2002 dance of food resources. Elaphe quadrivirgata is a common colubrid snake widely distributed in Japan including small islands adjacent to the main islands. Insular body size variation in this snake has been documented for populations of the Izu Islands, located off the south of the central part of Honshu Island, the largest island of Japan (Hasegawa and Moriguchi, 1989). Among these, snakes on Tadanae-jima Island, an uninhabited islet with an area of 0.01 km’, show a remarkable gigantism in adult body size, being more than three times larger in body mass than snakes of other adjacent islands (Hasegawa and Moriguchi, 1989) and of Honshu Island (Fukada, 1992; Kadowaki, 1996; Mori, unpublished data). As a part of a long term ecological and behavioral study of EF. quadrivirgata of the Izu Islands, we investigated possible mech- anisms of the gigantism of EF. quadrivirgata on Tadanae-jima Island. Higher growth rate is one of the possible factors that causes the gigantism and could result from differences in food supply (even only during early life, see Madsen and Shine, 2000), food type, and/ or physiological efficiencies. Here, we tested the hypothesis that the juvenile E. quadriv- irgata of Tadanae-jima island show higher growth rates than those of Honshu Island. The underlying hypothesis for differential growth rates is a genetically based modifica- tion in physiological traits, such as higher energy assimilation and lower energy expen- diture (Angilletta, 200la, b) of the former. We reared hatchlings from Tadanae-jima and Honshu under the same feeding schedule for approximately two years and compared their growth patterns. MATERIALS AND METHODS The subjects were neonate EF. guadrivirgata hatched in the laboratory from eggs laid by wild-caught gravid snakes. A male and a female hatchling were randomly selected from each of eight clutches oviposited by females (snout-vent length: SVL, range, 1007-1292 mm, x=1147 mm, body mass: BM, range, 361-972 g, x=631 g) collected from Tadanae-jima Island (34°13'N, 139°12’E) in June 1994. Six hatch- lings (two males and four females) from a single clutch of a female (SVL=680 mm, BM=147 g) collected from Tokai-mura, Ibaraki Prefecture (36°30’N, 140°30’E), Honshu, were used for comparisons. Although quantitative body size data of E. quadrivirgata in this population is not available, we assumed, based on the body size data of EF. quadriv- irgata in various areas of Honshu and the Izu Islands (Hasegawa and Moriguchi, 1989; Fukada, 1992; Kadowaki, 1996; Mori, unpub- lished data), that average body size of the snakes in Tokai-mura is well within the size range of FE. quadrivirgata in Honshu and considerably smaller than that of the Tadanae-jima population. Newly hatched snakes were weighed (BM), measured (SVL), and then housed individu- ally in white polypropylene (190x140x70 mm) cages, each containing a water dish and paper floor covering. As the snakes grew larger (ca. one year after hatching), they were moved to larger plastic cages (320x180x260 mm). The temperature varied between 25 and 30 C except winter, when the snakes were induced to hiber- nate. Illumination was provided by sunlight. During the first month after hatching in 1994, small frogs (Hyla japonica and Rana limnocharis) or lizards (Eumeces okadae) were offered to the snakes. Thereafter, because of unavailability of small live food, the snakes were raised by force feeding (Frye, 1991) until 11 May 1995. In this period, beef liver mixed with a multivitamin supple- ment and calcium powder and lubricated with raw egg was provided to each individual. From 16 May 1995, live suckling mice (Mus musculus) were provided. If the snakes did not spontaneously eat the mice a day after their introduction, the mice were fed to the snakes by force feeding. Food items were provided basically twice a week except for the hibernation period (from mid November to mid April). Although the weight of food varied during the study period (0.4-6.4 g), MORI & HASEGAWA—GROWTH OF ELAPHE QUADRIVIRGATA 45 approximately the same amount of food was offered to each of the snakes on each feeding day. In 1994, SVL and BM were measured immediately before hibernation. In 1995 and 1996, these measurements were taken approx- imately every month except during the hiber- nation period. This study was focused on the growth pattern before maturation, which can occur as early as the age two years (see Fukada, 1992), and therefore the study was terminated before the beginning of the third hibernation. Growth rates in SVL and BM were calculated for each measurement day —e— Tadanae-jima --%-- Honshu 700 Male 600 € = 900 25 > 400 Y) 300 200 700 Female 600 = = 500 = => 400 YW) 300 200 Hatch Dec May Nov Apr Nov 1994 1995 1996 FIG. 1. Growth trajectories of captive Elaphe quadrivirgata from populations on a Japan main island (Honshu) and an islet of the Izu Islands (Tadanae-jima). Each point shows the mean snout-vent length (SVL). Vertical bars show range. 46 Current Herpetol. 21(1) 2002 —e— Tadanae-jima --%-- Honshu Male Female Hatch Dec May Nov Apr Nov 1994 1995 1996 Fic. 2. Growth trajectories of captive Elaphe quadrivirgata from populations on a Japan main island (Honshu) and an islet of the Izu Islands (Tadanae-jima). Each point shows the mean body mass (BM). Vertical bars show range. as: increment in SVL (BM)/SVL (BM) at ences in SVL and BM were detected either the last measurement/elapsed days. in Tadanae-jima or Honshu snakes (all P>0.20), and so the sexes were pooled for the following analysis. Hatchlings of Tadanae- jima were significantly larger in SVL and At hatching, no significant sexual differ- BM than those of Honshu (SVL: Tadanae- RESULTS 47 MORI & HASEGAWA—GROWTH OF ELAPHE QUADRIVIRGATA vere —e— Tadanae-jima --*-- Honshu 5 x (A) zi eo See i Cp) 1p = 3 e ) es ed ! \ © fe ee ie2 ei, cle aii x S ‘g * = { Mixcy ; . : 9 v a l ie 0 ~\ wa ».4 x 10° = aa) = ob) oa | } a ap = O } ‘e) Hatch Dec May Nov Apr Nov 1994 1995 1996 Fic. 3. Growth rates in snout-vent length (SVL: A) and body mass (BM: B) for captive Elaphe quadrivirgata from populations on a Japan main island (Honshu) and an islet of the Izu Islands (Tadanae-jima). Each point shows mean growth rate. Growth rates were calculated as: increment in SVL (BM)/SVL (BM) at the last measurement/elapsed days. 48 jima, x=290 mm, Honshu, x=249 mm, Mann- Whitney U-test, U=94, P<0.001; BM: Tadanae- jima, x=11.2 g, Honshu, x=6.5 g, Mann-Whit- ney U-test, U=96, P<0.0005). Growth patterns were very similar between the two populations both in SVL and BM (Figs. 1 and 2). No higher growth rates were evident in the Tadanae-jima snakes (Fig. 3). In contrast, growth rates seemed to be higher in Honshu snakes. Accord- ingly, the between-population differences in SVL and BM, apparent at the hatching stage, gradually decreased toward the end of the study (Figs. 1 and 2). Both SVL and BM at the final measurement (November 1996) were not significantly different between the two populations (SVL: Tadanae-jima, x=627 mm, Honshu, x=582 mm, Mann-Whitney U-test, U=34, P=0.051; BM: Tadanae-jima, x=89.3 g, Honshu, x=81.8 g, Mann-Whitney U-test, U=28, P>0.25: sexes were pooled for these statistical tests because there were no signifi- cant sexual differences at this age). Two and three snakes of Tadanae-jima and Honshu, respectively, died during the course of the study. A close inspection of Fig. 3 revealed a subtle difference in growth pattern between Tadanae-jima and Honshu snakes. In both SVL and BM, average growth rates of Tadanae- jima snakes were greater than those of Honshu snakes only before hibernation (i.e., October to November). This trend was observed in all three years. DISCUSSION Differences in body sizes among popula- tions of E. quadrivirgata could arise from several factors. They could, for example, result from genetic differences among popu- lations due to selective pressures, genetic drift, or both. Alternatively, phenotypic differences could directly result from more proximate differences in environmental conditions such as food availability, duration of activity sea- son, and temperature. Even if snakes of different populations have the same growth Current Herpetol. 21(1) 2002 trajectories, differences in average body size could arise as a mere result of difference in age structure: 1.e., average body sizes of snakes from populations consisting of older indi- viduals could be larger than those of younger ones. Our experiment was aimed to test the hypothesis that gigantism of E. quadrivir- gata on Tadanae-jima island is caused by genetically based high growth rates before maturation. The present result does not support this hypothesis, as the growth rates of the two samples compared were basically similar. Average growth rates were even higher in the Honshu snakes than in the Tadanae-jima snakes. Non-facilitated growth rates of Tadanae-jima snakes were also con- firmed when their growth rates were com- pared with those of EF. quadrivirgata under natural conditions in Kyoto Prefecture, Honshu Island (Fukada, 1992). Therefore, it is likely that the gigantism of E. quadriv- irgata on Tadanae-jima is attributable to greater and/or continuous growth of adult snakes with abundant food resources, such as eggs and nestlings of sea birds (Hasegawa and Moriguchi, 1989). Nonetheless, differ- ential body size among populations of the Izu Islands may not be sufficiently explained on the grounds of pure phenotypic plasticity, and the possibility that other growth param- eters, such as maturity and asymptotic sizes, are genetically determined cannot be excluded at present (see Wikelski et al., 1997). Several experimental studies have been conducted to clarify which of the above mentioned factors are responsible for geo- graphic size differences in snakes. Barnett and Schwaner (1985) reported that neonates from an insular gigantic population of Notechis ater reared under laboratory conditions with ad libitum feeding grew faster than those of a mainland conspecific population under natural conditions. In addition, Schwaner (1985) briefly reported that N. ater from another gigantic island population grew faster than that from a dwarfed island population when reared with the same amount of food, MORI & HASEGAWA—GROWTH OF FLAPHE QUADRIVIRGATA 49 concluding that geographic size variation is genetically controlled. Similarly, Bronikowski (2000) showed, based on the results of a common-garden growth experiment, that field variation in growth in Thamnophis elegans has a genetic basis. Forsman (1991) attrib- uted the difference in body size between the island populations of Vipera berus to differ- ence in growth rate rather than difference in age structure, based on the growth data of wild snakes. On the other hand, Madsen and Shine (1993) compared growth rates of mainland Natrix natrix with those of an island dwarf population under laboratory conditions over six years and concluded that the differences in adult body size between these populations result from direct influ- ence of prey availability without any genetic modification. These results, coupled with the present one, indicate the presence of diverse mechanisms responsible for geographic variations of body size in snakes. Such diversity may be partially attributable to the differences in local envi- ronmental conditions and historical back- grounds. Studies on geographic variations of body size in snakes would offer splendid Opportunities to unravel the adaptive micro- evolution and evolutionary significance of phenotypic plasticity. Two observations should be noted here. First, the subtle but consistent difference in growth patterns between Tadanae-jima and Honshu snakes, that is, higher growth rates in the former only before hibernation, may imply that some genetic-based differences in growth pattern are present between them. Light cycle, time of the year, or temperature (although room temperature was kept between 25 and 30 C during the feeding period, average temperature varied seasonally within this range) may have interacted with genetically deter- mined physiological traits of the source populations to influence growth pattern (Bronikowski, 2000). Second, we emphasize that explicit discrim- ination between gigantism of adult snakes and large size of hatchlings should be made when we examine the causal mechanisms of geographic size variations in snakes. Although it is likely that neonate size is affected by size and nutritional conditions of maternal snakes (e. g., Ford and Seigel, 1989), larger hatchling size of Tadanae-jima population than populations of adjacent islands and Honshu seems to be, at least partially, genetically determined, because allometric relationships between maternal snake sizes and their offspring sizes are different between Tadanae-jima and the other populations. (Hasegawa, unpublished data). Because of gape limitation in snakes, larger hatchlings having larger gape size would have the advan- tage of being able to exploit a wider size range of prey. This may be especially true on Tadanae-jima Island, where potential prey animals available to small snakes are limited (Hasegawa and Moriguchi, 1989). On the other hand, larger body size of adults on Tadanae-jima might reflect a phenotypic response to the high food availability, as well as an adaptation for delayed maturation that facilitates growth and enables them to exploit food resources available only for large indi- viduals (eggs and nestlings of sea birds). This scenario considers larger hatchling size as a local adaptation, and gigantism in adult snakes as representing a complexity of phe- notypic plasticity and local adaptation. In any event, hatchling size and adult size (standard size) should be considered sepa- rately when mechanisms of geographic body size variations are studied, especially in snakes, where ingestible prey are quite different between hatchlings and adults because of gape limi- tation. ACKNOWLEDGMENTS We wish to thank many colleagues in Kyoto University for their assistance in keeping ani- mals. M. Honda provided a snake collected in his home town. This study was partially supported by grants from the Fujiwara Nat- ural History Foundation, the Pro-Natulra Foundation, and the research project of the 50 Current Herpetol. 21(1) 2002 National Institute of Environment on con- servation methods of subtropical island eco- systems awarded to M. Hasegawa. LITERATURE CITED ANGILLETTA, M. J., Jr. 2001la. Variation in metabolic rate between populations of a geo- graphically widespread lizard. Physiol. Biochem. Zool. 74(1): 11-21. ANGILLETTA, M. J., Jr. 2001b. Thermal and physiological constraints on energy assimilation in a widespread lizard (Sceloporus undulatus). Ecology 82(11): 3044-3056. BARNETT, B. AND T. D. SCHWANER. 1985. Growth in captive born tiger snakes (Notechis ater serventyi) from Chappell Island: implica- tions for field and laboratory studies. Trans. R. Soc. S. Aust. 109(2): 31-36. BRONIKOWSKI, A. M. 2000. Experimental evidence for the adaptive evolution of growth rate in the garter snake Thamnophis elegans. Evolution 54(5): 1760-1767. CASE, T. J. 1978. A general explanation for insular body size trends in terrestrial vertebrates. Ecol- ogy 59(1): 1-18. FORD, N. B. AND R. SEIGEL. 1989. Phenotypic plasticity in reproductive traits: evidence from a viviparous snake. Ecology 70(6): 1768-1774. FORSMAN, A. 1991. Variation in sexual size dimorphism and maximum body size among adder populations: effects of prey size. J. Anim. Ecol. 60: 253-267. FRYE, F. L. 1991. A Practical Guide for Feeding Captive Reptiles. Krieger Publ. Co., Malabar, Florida. 171 p. FUKADA, H. 1992. Snake Life History in Kyoto. Impact Shuppankai Co., Tokyo. 172 p. HASEGAWA, M. AND H. MORIGUCHI. 1989. Geographic variation in food habits, body size and life history traits of the snakes on the Izu Islands. p. 414-432. In: M. Matsui, T. Hikida, and R. C. Goris (eds.), Current Herpetology in East Asia. Herpetol. Soc. Japan, Kyoto. KADOWAKI, S. 1996. Ecology of a Japanese snake community-Resource use patterns of the three sympatric snakes, Rhabdophis tigrinus, Elaphe quadrivirgata and Agkistrodon b. blomhoffii. Bull. Tsukuba Univ. Forests (12): 77-148. (in Japanese with English summary) KING, R. B. 1989. Body size variation among island and mainland snake populations. Her- petologica 45(1): 84-88. KOHNO, H. AND H. OTA. 1991. Reptiles in a seabird colony: herpetofauna of Nakanoka- mishima Island of the Yaeyama Group, Ryukyu Archipelago. Island Studies in Okinawa, 9: 73- 89. MADSEN, T. AND R. SHINE. 1993. Phenotypic plasticity in body sizes and sexual size dimor- phism in European grass snakes. Evolution 47(1): 321-325. MADSEN, T. AND R. SHINE. 2000. Silver spoons and snake body sizes: prey availability early in life influences long-term growth rates of free- ranging pythons. J. Anim. Ecol. 69(6): 952-958. MorI], A. 1994. Ecological and morphological characteristics of the Japanese rat snake, Elaphe climacophora, on Kammuri-jima Island: a pos- sible case of insular gigantism. Snake 26: 11-18. Mor!I, A., H. OTA, AND N. KAMEZAKI. 1999. Foraging on sea turtle nesting beaches: flexible foraging tactics by Dinodon semicarinatum (Serpentes: Colubridae). p. 99-128. In: H. Ota (ed.), Tropical Island Herpetofauna, Origin, Current Diversity, and Conservation. Elsevier, Amsterdam. SCHMIDT-NIELSEN, K. 1984. Scaling. Why Is Animal Size So Important? Cambridge Univ. Press, Cambridge. 241 p. SCHWANER, T. D. 1985. Population structure of black tiger snakes, Notechis ater niger, on offshore islands of South Australia. p. 35-46. In: G. Grigg, R. Shine, and H. Ehmann (eds.), Biol- ogy of Australasian Frogs and Reptiles. Royal Zool. Soc. New South Wales, New South Wales. STAMPS, J. A. 1993. Sexual size dimorphism in species with asymptotic growth after maturity. Biol. J. Linn. Soc. 50: 123-145. WIKELSKI, M., V. CARRILLO, AND F. TRILLMICH. 1997. Energy limits to body size in a grazing reptile, the Galapagos marine iguana. Ecology 78(7): 2204-2217. Accepted: 13 June 2002 Current Herpetology 21(1): 51-53, June 2002 © 2002 by The Herpetological Society of Japan On the Authorship of Babina (Ampbibia: Ranidae) HIDETOSHI OTA!” AND MASAFUMI MATSULP ! Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903-0213, JAPAN 2 Graduate School of Human and Environmental Studies, Kyoto University, Yoshida-Nihonmatsu-cho, Sakyo-ku, Kyoto 606-8501, JAPAN Abstract: The genus-group name Babina, origi- nally proposed as a full genus for Rana holsti (type species) and R. subaspera, is usually attrib- uted to “Van Denburgh, 1912”. However, it is obvious from the chronological order of publica- tion of relevant papers that the authorship of Babina should be “Thompson, 1912”, not “Van Denburgh, 1912”. Key words: Babina; Ranidae, Anura; Nomencla- ture; Authorship; Priority The genus-group name Babina was first published in 1912 by two different authors, Thompson (1912a) and Van Denburgh (1912a). In both of these papers, the name was given to a new full genus described to accommodate two endemic frogs of the central Ryukyus, Rana holsti Boulenger, 1892, and R. subaspera Barbour, 1908, with the former being the type species. Also, both Thompson (1912a) and Van Denburgh (1912a) highlighted a sharp, spine-like metacarpal on the inner side of the first finger in these two species as the prominent character distinguishing Babina from other ranid genera. Van Denburgh published a more detailed description of the genus, without referring to Thompson’s (1912a) description, in his famous work on the East Asian herpeto- * Corresponding author. Tel: + 81-98-895-8937; Fax: +81-98-895-8966; E-mail address; ota@sci.u- ryukyu.ac.jp fauna published later in the same year (Van Denburgh, 1912b). Probably because both of the preceding descriptions were privately published by the respective authors and thus suffered limited availability, Van Denburgh (1912b) seems to have been regarded as the only source of information on the original description of Babina by most subsequent authors. Some of them (e.g., Okada [1930], p. 154) even erroneously referred to Van Denburgh (1912b) as the original description of the genus, although it was unequivocally stated on Van Denburgh’s (1912b) second page that a number of his new taxa (includ- ing Babina) were originally described in Van Denburgh (1912a). Of the subsequent authors, some consid- ered Babina as invalid (Inger, 1947; Dubois, 1981), others continued to use the name as a valid full genus (Okada, 1930, 1966), whereas most recent authors regard Babina as a subgenus of Rana (Nakamura and Uéno, 1963; Kuramoto, 1972; Matsui and Utsunomiya, 1983; Frost, 1985; Maeda and Matsui, 1989; Dubois, 1992; Duellman, 1993). In any event, authorship of this genus-group name has invariably been given as “Van Denburgh, 1912” (e.g., Okada, 1930, 1966; Inger, 1947; Nakamura and Ueno, 1963; Dubois, 1992; Duellman, 1993). In 1912, Thompson and Van Denburgh separately described a number of East Asian amphibians and reptiles on the basis of the same series of specimens (and sometimes even on the basis of exactly identical holo- type specimens) in rivalry with each other (Zhao and Adler, 1993: p. 32), and this resulted in a “most regrettable tangle of names” (Barbour, 1917; Nakamura and Uéno, 1963; Zhao and Adler, 1993). Nakamura and Uéno (1963) referred to this confusing situation with an example of the authorship of Hyla hallowellii, a species described by both Thompson (1912b) and Van Denburgh (1912a). While severely criticizing Thomp- son’s actions, Nakamura and Uéno (1963) argued that the name, usually given as “Hyla hallowellii Van Denburgh, 1912” to 52 Current Herpetol. 21(1) 2002 that date (e.g., Okada, 1930; Inger, 1947), should be attributed to “Thompson, 1912”, recognizing that Thompson (1912b) had preceded Van Denburgh (1912a) by approxi- mately one month. Nakamura and Uéno (1963), nevertheless, continued to regard “Van Denburgh, 1912” as the author of Babina, although Thompson (1912a: Herpe- tological notices 1) should have preceded Thompson (1912b: Herpetological notices 2) in publication date. Based on the date printed in each of the relevant papers, Zhao and Adler (1993) confirmed the chronological order of their publications as Thompson (1912a: on 15 June), Thompson (1912b: 28 June), Van Denburgh (1912a: on 29 July), and Van Denburgh (1912b: on 16 December). It is thus obvious from the principle of priority of the International Code of Zoological Nomenclature (2000) that Thompson (1912a) should be regarded as the author of the original description of Babina. Thus, the authorship of this genus-group name should be “Thompson, 1912”, not “Van Denburgh, 1912”. ACKNOWLEDGMENTS We thank K. Adler and R. I. Crombie for the provision of pertinent literature. LITERATURE CITED BARBOUR, T. 1917. A most regretable [sic] tangle of names. Occ. Pap. Mus. Zool. Univ. Michigan (44): 1-9. DUBOIS, A. 1981. Liste des genres et sous-genres nominaux de Ranoidea (Amphibiens Anoures) du monde, avec identification de leurs espéces- types: conséquences nomenclaturales. Monit. Zool. Ital., New Ser. 15 (suppl.): 225-284. DUBOIS, A. 1992. Notes sur la classification des Ranidae (Amphibiens Anoures). Bull. Mens. Soc. Linn. Lyon 61(10): 305-352. DUELLMAN, W. E. 1993. Amphibian species of the world: additions and corrections. Univ. Kansas Mus. Nat. Hist. Spec. Publ. (21): 1- 372. FROST, D. (ed.) 1985. Amphibian Species of the World: A Taxonomic and Geographical Refer- ence. Allen Press, Lawrence. 732 p. INGER, R. F. 1947. Preliminary survey of the amphibians of the Riukiu Islands. Fieldiana: Zool. 32(5): 297-352. INTERNATIONAL CODE OF ZOOLOGICAL NOMEN- CLATURE. Fourth Edition. 1999. International Trust for Zoological Nomenclature, The Natural History Museum, London. I-xxix + 306 p. KURAMOTO, M. 1972. Karyotypes of the six species of frogs (genus Rana) endemic to the Ryukyu Islands. Caryologia 25(4): 547-559. MAEDA, N. AND M. MATSUI. 1989. Frogs and Toads of Japan. Bun-ichi Sogo Shuppan, Tokyo. 207 p. MATSUI, M. AND T. UTSUNOMIYA. 1983. Mating call characteristics of the frogs of the subgenus Babina with reference to their relationship with Rana adenopleura. J. Herpetol. 17(1): 32-37. NAKAMURA, K. AND S.-I. UENO. 1963. Japanese Reptiles and Amphibians in Colour. Hoikusha, Osaka. 214 p. (in Japanese) OKADA, Y. 1930. A Monograph of Japanese Tail-less Batrachians. Iwanami-shoten, Tokyo. 234 p.+ 27 pls. (in Japanese) OKADA, Y. 1966. Fauna Japonica: Anura (Amphibia). Tokyo Electronic Engineering Col- lege Press, Tokyo. 12+ 234 p. THOMPSON, J. C. 1912a. Herpetological Notices No. 1. Prodrome of a Description of a New Genus of Ranidae from the Loo Choo Islands. Privately published by the author, San Fran- cisco. 3 p. THOMPSON, J. C. 1912b. Herpetological Notices No. 2. Prodrome of Descriptions of New Species of Reptilia and Batrachia from the Far East. Privately published by the author, San Francisco. 3 p. VAN DENBURGH, J. 1912a. Advance Diagnoses of New Reptiles and Amphibians from the Loo Choo Islands and Formosa. Privately published by the author, San Francisco. 8 p. VAN DENBURGH, J. 1912b. Concerning certain species of reptiles and amphibians from China, Japan, the Loo Choo Islands, and OTA AND MATSUI—AUTHORSHIP OF BABINA 53 Formosa. Proc. California Acad. Sci. Ser. 4, China. Society for the Study of Amphibians 3: 187-258. and Reptiles, Oxford, Ohio. 522 p. ZHAO, E. AND K. ADLER. 1993. Herpetology of Accepted: 12 May 2002 > in Pe ees Pe - : 5 INSTRUCTION TO CONTRIBUTORS Scope and Manuscript Categories Current Herpetology is the official English journal of the Herpetological Society of Japan, and publishes Reviews, Original Articles, and Short Notes dealing largely or exclusively with the biology and diversity of amphibians and reptiles. Reviews are usually invited by the Managing Editor. Those who wish to submit Reviews should consult with the Managing Editor in advance. Original Articles are full-length reports usually greater than nine manuscript pages. Short Notes are more limited in scope and usually fewer than 10 manuscript pages. Manuscripts may be moved from one category to another at the discretion of the Managing Editor. Manuscript Submission Submissions of papers from both members and non-members of the Society are welcome. 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(in Japanese) HONDA, M, Y. YASUKAWA, AND H. OTA. In press. Phylogeny of the Eurasian fresh- water turtles of the genus Mauremys Gray, 1869 (Testudines). J. Zool. Syst. Evol. Res. KAMEZAKI, N. 1989. The nesting sites of sea turtles in the Ryukyu Archipelago and Taiwan. p. 342-348. In: M. Matsui, T. Hikida, and R. C. Goris (eds.), Current Herpetology in East Asia. Herpetological Society of Japan, Kyoto. LEVITON, A. E. AND R. H. GIBBS, Jr. 1988. Standards in herpetology and ichthyology. Standard symbolic codes for institution resource collections in herpetology and ichthyology. Supplement no. 1: additions and corrections. Copeia 1988(1): 280-282. LEVITON, A. E., R. H. GIBBS, Jr., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985(3): 802-832. MATSUI, M. 1987. Isozyme variation in salamanders of the nebulosus-lichenatus complex of the genus Hynobius from eastern Honshu, Japan, with a description of a new species. Jpn. J. Herpetol. 12(2): 50-64. MATSUI, M., H. IWASAWA, H. TAKAHASHI, T. HAYASHI, AND M. KUMAKURA. 1992a. Invalid specific status of Hynobius sadoensis Sato: electrophoretic evidence (Amphibia: Caudata). J. Herpetol. 26(4): 308-315. MATSUI, M. AND K. MIYAZAKI. 1984. Hynobius takedai (Amphibia, Urodela), a new species of salamander from Japan. Zool. Sci. 1(6): 665-671. MATSUI, M., T. SATO, S. TANABE, AND T. HAYASHI. 1992b. Electrophoretic analyses of systematic relationships and status of two hynobiid salamanders from Hokkaido (Amphibia: Caudata). Herpetologica 48(4): 408-416. MOODY, S. M. 1980. Phylogenetic and Historical Biogeographical Relationships of the Genera in the Family Agamidae (Reptilia: Lacertilia). Unpublished doctoral dissertation. University of Michigan, Ann Arbor. 373 p. 58 TAKENAKA, T. 2000. Extinction of the naturalized freshwater turtle in Chichijima-Island of Ogasawara (Bonin) Islands, South Japan. Bull. Herpetol. Soc. Japan 2000(1): 4-7. (in Japanese with English abstract) ZHAO, E. AND K. ADLER. 1993. Herpetology of China. Contribution to Herpetology, 10. Society for the Study of Amphibians and Reptiles, Oxford, Ohio. 522 p. In the text, references to papers by one or two authors should give their surnames; papers with more than two authors are referenced by the first author’s surname followed by “et al.”. Strings of references should be placed in chronological order. When there are two or more references published in a same year, they should be arranged in alphabetical order. Two or more references by the same senior author (for papers by one or more than two authors) or by the same senior and junior authors (for papers by two authors) with the same year of publication should be designated by lowercase letters: e.g., (Matsui et al., 1992a, b). 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(number), yr (year[s]), mo (month[s]), wk (week[s]), h (hour[s]), min (minute[s]), s (second[s]), P (probability), df (degrees of freedom), SD (standard deviation), SE (standard error), NS (not signifi- cant), | (liter), kg (kilogram), g (gram), m (meter), cm (centimeter), mm (millimeter), um (micron), C (degrees Celsius, not °C), asl (above sea level; given as, e.g., 100 m asl), °,', and" (degrees, minutes, and seconds in geography, respectively), N, S, E, and W (north and south latitudes, and east and west longitudes, respectively, but only when preceded by values with appropriate geographical units; e.g., 15°25'N, 121°43'E). 60 In figures and tables, names of months can be abbreviated as: Jan (January), Feb (February), Mar (March), Apr (April), May (May), Jun (June), Jul (July), Aug (August), Sep (September), Oct (October), Nov (November), Dec (December). In the text, however, they should be fully spelled. 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The following changes can be made on the proof: correc- tions of typographical errors, addition of publication information for a reference previously in an “In press” state, and addition of the present address of the author due to the change of his/her address after acceptance of a manuscript. Other changes may be charged to the author. CURRENT HERPETOLOGY MANAGING EDITOR Hidetoshi OTA Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 903-0213, JAPAN (ota@sci.u-ryukyu.ac.jp) ASSOCIATE EDITORS Kraig ADLER, Department of Neurobiology and Behavior, Cornell University, Seeley G. Mudd Hall, Ithaca, New York 14853-2702, USA (kka4@cornell.edu) Aaron M. BAUER, Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA (aaron.bauer@villanova.edu) Ilya S. DAREVSKY, Zoological Institute, Russian Academy of Sciences, St.Petersburg 199034, RUSSIA (Darevsky@herpet.zin.ras.spb.ru) Indraneil DAS, Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, MALAYSIA (idas@mailhost.unimas.my) Richard C. GORIS, Hatsuyama 1-7-13, Miyamae-ku, Kawasaki 216-0026, JAPAN (goris@ twics.com) Ivan INEICH, Laboratoire des Reptiles et Amphibiens, Museum National d’Histoire Naturelle, 25 rue Cuvier 75005 Paris, FRANCE (ineich@cimrs1.mnhn.fr) Ulrich JOGER, Hessisches Landesmuseum Darmstadt, Zoologische Abteilung, Friedensplatz 1, D-64283 Darmstadt, GERMANY (u.joger@hlmd.tu-darmstadt.de) Tamotsu KUSANO, Department of Biological Science, Graduate School of Science, Tokyo Metropolitan University, Minami-ohsawa, Hachioji, Tokyo 192-0397, JAPAN (tamo@ comp.metro-u.ac.jp) Masafumi MATSUI Graduate School of Human and Environmental Studies, Kyoto University, Yoshida Nihonmatsu-cho, Sakyo-ku, Kyoto 606-8501, JAPAN (fumi@zoo. zool.kyoto-u.ac.jp) Colin McCARTHY, Department of Zoology, The Natural History Museum, Cromwell road, London SW7 5BD, UK (cjm@nhm.ac.uk) Michihisa TORIBA, Japan snake Institute, Yabuzuka-honmachi, Nitta-gun, Gunma 379-2300, JAPAN (snake-a@sunfield.ne.jp) Home Page of THE HERPETOLOGICAL SOCIETY OF JAPAN http://zoo.zool.kyoto-u.ac.jp/~herp/ DATE OF PUBLICATION Current Herpetology, Vol. 20, No. 2, was mailed 28 December 2001. IWAN 39088 01298 1403 CONTENTS Original articles Foraging behavior of Rhabdophis tigrinus (Serpentes: Colubridae) in a gutter with a dense aggregation of tadpoles er aliohfella| io! nile mlerelnteticldyatay cilecatatsleneie: «etedetadatstelte ts isis ietelel tel dier ar eiasieteta Koji Tanaka eaeiaioae i Three new depressed-bodied water skinks of the genus 7Zropidophorus (Lacertilia: Scincidae) from Thailand and Vietnam gee 928 Tsutomu Hikida, Nikolai L. Orlov, Jarujin Nabhitabhata, and Hidetoshi Ota ------ 9 Taxonomic relationships of an endangered Japanese salamander Hynobius hidamon- tanus Matsui, 1987 with H. tenuis Nambu, 1991 (Amphibia: Caudata) oes Gaye aoe og a- Saas 2 Masafumi Matsui, Kanto Nishikawa, Yasuchika Misawa, Masaichi Kakegawa, and Takahiro Sugahara ------ 2a Karyotypes of four agamid lizards from Southeast Asia ee Hidetoshi Ota, Cheong-Hoong Diong, Ene-Choo Tan, and Hoi-Sen Yong ------35 Early growth of Elaphe quadrivirgata from an insular gigantic population SSO OC eG i ihn ris Om Ciseic sb ce ona Seosd ooo ee Aes Akira Mori and Masami Hasegawa Soy ie | Short note On the authorship of Babina (Amphibia: Ranidae) w sie miio\'nle\s(avein a) allclleyaite 'e lelleyelepelelini siete alonsheiatsic cue ataietotaieta ete ielaiel ollste te tet a aie Hidetoshi Ota and Masafumi Matsui eB | FUTURE MEETING Hokkaido Tokai University, Sapporo, Hokkaido, Japan, 5-6 October 2002 (Sen Takenaka, Chair)