HARVARD UNIVERSITY Ernst Mayr Library of the Museum of Comparative Zoology *«0*ae ISSN 1051-3825 Hiaz, R. E., M. T. Leong, L. L. Grismer, and N. S. Yaakob. A New Species of Dibamus (Squamata: Dibamidae) from West Malaysia Grismer, L. L., J. L. Grismer, and T. M. Youmans. A New Species of Leptolalax( Anura: Megophryidae) FROM PULAU TlOMAN, WEST MALAYSIA * * Leong, T. M. and L. L. Grismer. A New Species of Kukri Snake, Oligodon (Colubridae), from Pulau Tioman, West Malaysia 12-16 Stuart, B. L. and H. Heatwole. A New Philautus (Amphibia: Rhacophoridae) from Northern Laos 17-21 Diesmos, A. C., G. V. A. Gee, M. L. Diesmos, R. M. Brown, P. J. Widmann, and J. C. Dimalibot. Rediscovery of the Philippine Forest Turtle, Heosemys leytensis (Chelonia; Bataguridae), from Palawan Island, Philippines 22-27 Feldman, C. R. and J. F. Parham. Molecular Systematics of Old World Stripe-Necked Turtles (Testudines: Mauremys ) 28-37 Hutchison, J. H„ P. A. Holroyd, and R. L. Ciochon. A Preliminary Report on Southeast Asia’s Oldest Cenozoic Turtle Fauna from the Late Middle Eocene Pondaung Formation, Myanmar 38-52 Joyce, W. G. and C. J. Bell. A Review of the Comparative Morphology of Extant Testudinoid Turtles (Reptilies: Testudines) 53-109 Le, Minh, T. Hoang, and D. Le. Trade Data and Some Comments On the Distribution of Mauremys ANNAMENSIS (SlEBENROCK, 1 903) 110-113 Perala, J. and R. Bour. Neotype of Testudo terrestris ForsskAl, 1775 (Testudines, Testudinidae) 114-119 Schilde, M., D. Barth, and U. Fritz. An Ocadia sinensis x Cyclemys shanensis hybrid (Testudines: Geomydidae) 120-125 Shi, H„ Z. Fan, F. Yin, and Z. Yuan. New Data on the Trade and Captive Breeding of Turtles in Guangxi Province, South China 126-128 Stuart, B. L. and S. G. Platt. Recent Records of Turtles and Tortoises from Laos, Cambodia, and Vietnam 129-150 Auffenberg, K., K. L. Krysko, and W. Auffenberg. Studies on Pakistan Lizards: Cyrtopodion stoliczkai (Steindachner, 1867) (Gekkonidae: Gekkoninae) 151-160 Dulger, B., j. H. Ugurta§, and M. Sevinq:. Antimicrobial Activity in the Skin Secretion of Bufo viridis (Laurenti, 1768) 161-163 Du§en, S., M. Oz, and M. R. Tunq:. Analysis of the Stomach Contents of the Lycian Salamander Mertensiella luschani (Steindachner, 1891) (Urodela: Salamandridae), Collected from Southwest Turkey 164-167 Ebrahimi, M., H. G. Kami, and M. Stock. First Description of Egg Sacs and Early Larval Development in Hynobiid Salamanders (Urodela, Hynobiidae, Batrachuperus ) from North-Eastern Iran 168-175 Jarrar, B. M., and N. T. Taib. Histochemical Characterization of the Lingual Salivary Glands of the House Gecko, Ptyodactylus hasselquistii (Squamata: Gekkonidae) 176-181 Kami, H. G. The Biology of the Persian Mountain Salamander, Batrachuperus persicus (Amphibia, Caudata Hynobiidae) in Golestan Province, Iran 182-190 Khan, M. S. Annotated Checklist of Amphibians and Reptiles of Pakistan 191-201 (Continued on inside back cover) 1 IU Asiatic Herpetological Research Volume 11 • 2008 Chengdu Institute of Biology of the Chinese Academy of Sciences Asiatic Herpetological Research Society at the Museum of Vertebrate Zoology, University of California Honorary Editor-in-Chief Er-mi Zhao Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, Sichuan, China Editor-in-Chief Yue-zhao Wang Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, Sichuan, China Associate and Managing Editor Associate Editor Raul E. Diaz Theodore J. Papenfuss University of Kansas Medical Center; University of Museum of Vertebrate Zoology, University of Kansas Museum of Natural History, Kansas, USA California Berkeley, California, USA Consulting Editor James F. Parham California Academy of Sciences Editorial Board Kraig Adler Cornell University, Ithaca, New York, USA Natalia B. Ananjeva Zoological Institute, St. Petersburg, Russia Steven C. Anderson University of the Pacific, Stockton, California, USA Aaron Bauer Villanova University, Villanova Pennsylvania, USA Christopher Bell University of Texas, Austin, Texas, USA Leo Borkin Zoological Institute, St. Petersburg, Russia I-Jiunn Cheng Institute of Marine Biology, National Taiwan Ocean University, Keelung, Taiwan, China Wen-hao Chou National Museum of Natural Science, Taichung, Taiwan, China Ilya Darevsky Zoological Institute, St. Petersburg, Russia Indraneil Das Madras Crocodile Bank, Vadanemmeli Perur, Madras, India William E. Duellman University of Kansas, Lawrence, Kansas, USA Jinzhong Fu Department of Integrative Biology, University of Guelph, Guelph, Ontario, Canada Robert F. Inger Department of Zoology, Field Museum of Natural History, Chicago, Illinois, USA Xiang Ji Hangzhou Normal College, Hangzhou, Zhejiang, China; Nanjing Normal University, Nanjing, Jiangsu, China Pi-peng Li Shenyang Normal University, Shenyang, Liaoning, China Robert W. Murphy Royal Ontario Museum, Toronto, Ontario, Canada Goren Nilson University of Goteborg, Goteborg, Sweden Nikolai Orlov Zoological Institute, St. Petersburg, Russia Hidetoshi Ota Department of Biology, University of the Ryukyus, Nishihara, Okinawa, Japan Soheila Shafii University of Shahid Bahonar, Kerman, Iran Hai-tao Shi Hainan Normal University, Haikou, Hainan, China Xiao-ming Wang Department of Zoology, East China Nonnal University, Shanghai, China Yehudah Werner Hebrew University, Jerusalem, Israel Xiao-mao Zeng Chengdu Institute of Biology. Chinese Academy of Sciences, Chengdu, Sichuan, China Asiatic Herpetological Research (AHR) Volume 1 1 is published by the Chengdu Institute of Biology of the Chinese Academy of Sciences (CIB) and the Asiatic Herpetological Research Society (AHRS) at the Museum of Vertebrate Zoology, University of California. The editors encourage authors from all countries to submit articles concerning, but not limited to, Asian herpetology. All correspondence should be sent via email to the editorial office at ahr@cib.ac.cn. Authors should consult Guidelines for Manuscript Preparation and Submission at the end of this issue and on the web. Website For more information with regards to subscription, manuscript submission, contacts, back issues and general questions visit AHR’s website at http://www.Asiatic-Herpetological.org. Asiatic Herpetological Research Previous volumes were published by the Asiatic Herpetological Research Society (AHRS) and the Chinese Society for the Study of Amphibians and Reptiles (CSSAR) as follows: Vol. 10 (2004), Vol. 9 (2001), Vol. 8 (published in 1999), Vol. 7 (1997). Vol. 6 (1995), Vol. 5 (1993), Vol. 4 (1992), Vol. 3 (1990), and Chinese Herpetological Research Vol. 2, which was published at the Museum of Vertebrate Zoology, 1988-1989, as the journal for the Chinese Society for the Study of Amphibians and Reptiles. Volume 2 succceeded Chinese Herpetological Research 1987, published for the Chengdu Institute of Biology by the Chongqing Branch Scientific and Technological Literature Press, Chongqing, Sichuan, China. Acta Herpetologica Sinica ceased publication in June, 1988. 2008 Asiatic Herpetological Research, Vol. 1 1 pp. 1-9 A Preliminary Study of the Lizard Fauna and Their Habitats in Northwestern Iran F. Ahmadzadeh1’*, B. H. Kiabi2, H. G. Kami3 and V. Hojjati4 MCZ LIBRARY 1 Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti University, Evin, Tehran, Iran, 2 Department of Biology, Faculty of Sciences, Shahid Beheshti University, Tehran, Iran, 3 Department of Biology, Faculty of Sciences, Agricultural Sciences and Natural Resources University, Gorgan, Iran, 4Damghan Islamic Azad University, Damghan, Iran. * Corresponding author E-mail: f_ahmadzade@sbu.ac.ir MAY 07 2008 HARVARC UNIVERSIT Abstract.- Northwestern Iran has unique geographical and climatic conditions that support a rich flora and fauna. In view of the lack of in-depth studies on the lizards of the region, an investigation was started in the northern part of Ardabil Province for an inventory of this component of the fauna and their habitats. Collections were made from October 2003 to June 2005 and 165 specimens were collected and identified. Five families, 12 genera and 15 species are represented, including Agamidae: Laudakia caucasia, Phrynocephalus persicus, Trapelus ruderatus; Lacertidae: Lacerta media media, Lacerta strigata, Lacerta brandtii, Darevskia raddei raddei, Eremias strauchi strauchi, Eremias arguta, Ophisops elegans; Scincidae: Mabuya aurata transcaucasica, Eumeces schneiderii princeps, Abelepharus bivittatus\ Anguidae: Pseudopus apodus and Gekkonidae: Cyrtopodion caspium caspium. Comparing this list to the data provided by Anderson (1999), it seems that most of the lizards are being reported for the Province for the first time. The families Gekkonidae and Anguidae are newly recorded, and the gecko Cyrtopodion caspium is first recorded from the west and northwest of Iran. With seven species represented in the area, lacertids have the high- est species diversity among the lizard families and need further study. Habitat features also have been given for all species. Keywords.- Iran, Ardabil, fauna, lizard, Lacerta. Introduction General information about the herpetofauna of Iran has been provided by Mertens (1957), Anderson (1966), Tuck (1971, 1974), Latifi (1984, 1991), Balouch and Kami (1995) and Kami and Vakilipoure (1996a, 1996b). Furthermore, a handbook of amphibians and reptiles of the Middle East has been published by Leviton et al. (1992), a book on the Lizards of Iran was recently pub- lished by Anderson (1999) and an updated checklist to the lizards of Iran was provided by Firouz (2000). Despite these publications, the lizards of Iran are still poorly-known and infrequently collected, with many new species still being discovered (Rastegar-Pouyani, 1996; Rastegar-Pouyani and Nilson, 1998). Studies on the lizards of Ardabil Province are also very limited (Ahmadzadeh, 2004). The aim of this study is to determine in detail the lizard fauna and their habitat features in the northern part of Ardabil Province, which is of particular signifi- cance considering the unique geography and vegetation of the region. Moreover, this study will collect baseline population data for future management. Materials and Methods The area of study is in the Northwest part of Iran, specif- ically, the northern part of Ardabil Province (38° 15' E, to 39° 40' E, 47° 30' N to 48° 00' N). The region is sur- rounded by the Alborz Mountains and the Caspian Sea to the east, Aras village is to the north. Arasbaran pro- tected area and Gare-Dagh Mountain to the west and the Sabalan Mountain chains to the south (Fig. 1). Altitude ranges between 20 m in the Moghan steppe to 4,888 m on the Sabalan Mountain. The study was carried out between October 2003 and June 2005. All of the samples were caught by hand and some lizards which are active and difficult to catch, such as green lizards (e.g., Lacerta m. media and L. strigata ), were captured by dust shot. Locality data and their habitat features were recorded for all species encountered during the study. However, all have been preserved in accordance to standard methods (Formalin 1 0%) and voucher specimens are stored in the Biodiversity and Ecosystem Management Department Collection (BEMD) at Shahid Beheshti University of Iran. Specimens were identified with Leviton et al. (1992) and Anderson (1999) using morphometric meas- © 2008 by Asiatic Herpetological Research 2 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 1. The study area, the northern part of Ardabil Province of Iran. urements, coloration and pholidosis features (including the number, structure and range of plates). Results A total of 165 samples were collected in the study area, comprising 15 species in 12 genera and 5 families. The species composition is given in Table 1. Distribution of species are presented in Figure 2. Family: Agamidae Laudakia caucasia caucasia (Eichwald, 1831) Laudakia c. caucasia is widely distributed in the study area, preferring montainous habitats and eroded sand canyons in flooded plains adjacent to mountains, rock cliffs, old houses and stony walls near roads. At 6 km2, 210 specimens were recorded on 10 June 2004 in Meshkinshar. This species was also collected on 24 November and 3 March in the Meshkinshahr and Arshagh areas at elevations between 500-2,800 m. Specimens were light olive to dark gray in ground color with adult snout- vent length of 152 mm in males and 155 mm in females. Phrynocephalus persicus persicus De Filippi, 1863 This species was rarely encountered in the study area. Four specimens were captured in Arshagh, Alma village - a semi-arid area with ephemeral plants in spring and loamy soil. Xerophyte vegetation, both plants and bush- es, grow in these areas. Dorsal coloration was light brown with three dark transverse marks, within which on the hind limbs were enlarged tubercular scales. The largest female was 40 and 48 mm snout-vent and tail length, respectively. During the study period, no males were found. Trapelus ruderatus ruderatus (Olivier, 1 804) The small agamid lizard Trapelus r. ruderatus was found on open stony ground and in cultivated fields with sparse weed vegetation in autumn. On sunny summer days, it hides under weeds such as Euphorbia spp., Chenopodium spp. and Chrozophora tinctoria. Its activ- ity appears to begin in early June and extends to late September. In total, 10 specimens were collected on the harvested wheat and barley fields in Gooshe area at approximately at 10:30 AM. Ground color was typically grayish-brown with five dark transverse bars on the trunk which were interrupted by a series of light ovoid vertebral spots (Fig. 3). The largest male examined with distinct callous preanal scales, had a 65 mm snout-vent and 74 mm tail length. The largest female had measure- ments of 63 mm and 75 mm, respectively. Family: Anguidae Pseudopus apodus (Pallas, 1775) Pseudopus apodus occurs throughout the Hyrcanian for- est of northern Iran. It has recently been collected from the Arasbaran protected area, but there are no records for Ardabil Province. This species was found in grassland and shrubby vegetation near streams. On a sunny day, four P. apodus were observed in a pond. The ground color of the dorsum was gray with zig-zagging blackish- brown stripes in the juvenile. The head was light yellow- ish-brown in adults with the remainder of the body dark brown. The longest adult had a 520 mm snout-vent length and a 670 mm tail. Family: Gekkonidae Cyrtopodion caspium caspium (Eichwald, 1831) An isolated population of Cyrtopodion caspium was found in the Moghan Steppe for the first time, represent- ing a new family record for northwestern Iran. One spec- 2008 Asiatic Herpetological Research, Vol. 11 3 Table 1. Lizard species collected from the sfudy area. Family Species Common Name Agamidae Laudakia caucasia Caucasian agama Phrynocephalus persicus Persian toad agama Trapelus ruderatus Olivier’s agama Anguidae Pseudopus apodus Glass-snake, sheltopusik Gekkonidae Cyrtopodion caspium Caspian bent-toed gecko Lacertidae Darevskia r. raddei Azarbaijan lizard Eremias arguta Steppe-runner Eremias strauchi Strauch’s racerunner Lacerta brandtii Persian lacerta Lacerta media Three-lined lizard Lacerta strigata Caspian green lizard Ophisops elegans Snake-eyed lizard Scincidae Ablepharus bivittatus Two-streaked snake-eyed skink Eumeces schneiderii Schneider’s skink Mabuya aurata Transcaucasian grass skink imen was collected at night on walls of an old house at 20 m elevation. After sunset they fed on various noctur- nal insects around lights. Dorsal scales are strongly keeled. Dorsal body coloration was light gray with five dark transverse bars on the body and 11-12 on the tail (Fig. 4). In Pars-Abad, one male specimen with a snout- vent length of 75 mm and a tail length of 70 mm was measured. Family: Lacertidae Darevskia raddei raddei (Boettger, 1892) Darevskia r. raddei is common in rocky areas where Laudakia caucasia is also frequently found. In March, this species was seen on vertical surfaces of rocks in the Kapas Mountains near Meshkin-Shahr. Darevskia r. rad- dei is various shades of light brown dorsally and more common in the rocky habitats than other lacertids. Specimens were found below altitudes of 900 m in Meshkinshar, but in the Salavat and Arshag Mountains, it was collected at altitude up to 2,400 m. The relation- ship of this subspecies to D. r. vanensis in northwestern Iran requires further study. The largest male had a snout- vent of 71 mm and tail length of 131 mm. One adult female had a 70 mm snout-vent length and 130 mm tail length (Fig. 5). Lacerta brandtii De Filippi, 1863 Lacerta brandtii was collected under stones, on foothills and in the burrows of other animals in open arid bushy and stony habitats in the Razeye area. Large numbers of this lizard were also found in Samian District, 100 km from the study area on a foothill surrounded by cultivat- ed land. The relationship between the two Iranian popu- lations of this species in Esfahan Province and east Azarbijan Province remains problematic. This species is less active in comparison to other lacertid lizards such as Darevskia raddei. The dorsal surface was olive-gray with small black spots (Fig. 6) and the ventral surface had 8 longitudinal rows of plate. The longest male spec- imen had total length of 192 mm. The tail of this species displays autotomy (approximately 60% of total speci- mens). Eremias arguta (Pallas, 1773) Eremias arguta has a limited distribution in the study area: one adult specimen was collected in a harvested barley field on a sunny day near to the Ardabil-Meshkin road and two juvenile specimens were captured in the Ardabil Airport area in August 2004. This lizard had a white belly and a dorsum with white spots edged with black on a grayish background (Fig. 7) that sometimes formed transverse bands in the adults. Our adult speci- mens had a 95 mm snout-vent length and a 1 10 mm tail. Eremias strauchi strauchi Kessler, 1 878 There are two subspecies of this lizard in Iran - Eremias s. strauchi and E. s. kopetdaghica , of which only the first was found in the study area. The specimen was col- lected in the eastern part of the study area in the Arshag plain under wheat straw in a dry, stony harvested field. Eremias s. strauchi is active and hides in shrubby vege- tation. Five eggs of this lizard were found under an Artemisia sp. shrub on 14 June 2004 in Amir-Abad vil- lage. In the study area one male specimen was measured 4 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 2. Distribution of species in the study area: (a) L. caucasia, (b) Ph. persicus, (c) T. ruderatus , (d) P. apodus , (e) C. caspium, (f) D. raddei. 2008 Asiatic Herpetological Research, Vol. 11 5 Figure 2. (continued) (g) E. arguta, (h) E. strauchi, (i) L. brandtii, 0) L. media, (k) L. strigata, (I) O. elegans. 6 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 2. (continued) (m) A. bivitattus, (n) E. schneiderii, (o) M. aurata. with a 71 mm snout-vent length and had a 128 mm tail. It’s olive-gray color pattern does not vary greatly among populations (Fig. 8). Lacerta media media (Lantz and Cyren, 1 902) Lacerta m. media was very common in the grassy and shrubby areas along the Khyave Chaye and Garesoo river banks. One male specimen was captured under a stone near a bean field. Specimens were observed at an altitude of 2,100 m and males were seen on stony walls near the roads at the end of the winter. The dorsal sur- face of the adult male, unlike juveniles, was green with- out any light lines or spots. Females were dark olive- brown with large lateral spots that disappeared with age. The largest lacertid collected during the study was one male specimen with a snout-vent of 117 mm and a tail length of 272 mm. This species exists in two differently- spotted morphs, with specimens from cultivated fields being larger than those from other habitats. Lacerta strigata Eichwald, 1831 Lacerta strigata was most frequently found in the Hyrcanian Forest in northern Iran and in some bushy and wooded streams banks associated with this forest, such as the Arax River in the northern part of the study area. Large numbers of this species were seen in Pars- Abad, Bilasovar and Germi near streams with dense Tamarix and Rubus vegetation. One specimen was cap- tured far from the Hyrcanian Forest in an open harvested wheat field on 25 August 2004. Most collection sites represent new locality records. The general color of the dorsum was light green in males and dark-olive to brown in females; it was more strongly spotted than Lacerta m. media. Females were also smaller with more numerous dark spots. The largest male had a snout-vent length of 160 mm and a tail length of 100 mm, while the 2008 Asiatic Herpetological Research, Vol. 11 7 Figure 5. Darevskia raddei raddei. Figure 6. Lacerta brandtii. Figure 7. Eremias arguta. Figure 9. Eumeces schneiderii princeps. Figure 8. Eremias strauchi. Figure 10. Mabuya aruata transcaucasica. 8 Asiatic Herpetological Research, Vol. 1 1 2008 largest temale had measurements of 97 mm and 187 mm. Ophisops elegans Menetries, 1832 Ophisops elegans is widely distributed, but is most com- mon on the Moghan Steppe. A large isolated population was found in Amir-Abad village. Specimens were active and encountered almost everywhere, particularly in dry stony habitats. Males and females both showed different color patterns during the reproductive period. Dorsal coloration was generally olive-green to brownish with two light dorsolateral stripes that disappear in the adult. An adult female from the Moghan Steppe had a snout- vent length of 64 mm and a tail length of 1 10 mm, and a male from Amir-Abad village had a snout-vent of 70 mm and a tail length of 120 mm. Family: Scincidae Ablepharus bivittatus (Menetries, 1832) The Two-streaked Snake-eyed Skink, Ablepharus bivit- tatus, has only been found in Amir-Abad village on Ardabil-Germi Road on a slope with large spiny cushion vegetation where it was sympatric with Ophisops ele- gans and Eremias strauchi. This active lizard has a high population density in the Neur Lake area in southern part of Ardabil Province reaches. Body coloration on the dorsum and tail is bronze-brown. The largest adult female specimen reached 60 mm in snout-vent length. Eumeces schneiderii princeps (Eichwald, 1839) This species lives on sand dunes, stony hills and dry river beds. We captured one male on a foothill in Meshkinshahr on 16 June 2004 at 08:00 AM, where Mabuya aruata transcaucasica was also found. This lizard is very active, hides in burrows and can jump approximately 2 m. In comparison to other scincid lizards, Eumeces schneiderii occurs in relatively few localities - overgrazing and destruction of habitat is threatening extirpation of this species in the study area. The dorsum was brownish with a narrow creamy-white lateral line from the posterior labial through the ear along the sides to the groin (Fig. 9). Total length (snout- vent + tail length) of the captured male was 240 mm. Mabuya aurata transcaucasica (Chernov, 1 926) Mabuya aurata transcaucasica lives in sandy areas and small hills that are covered with Astragalus and Acantolimon vegetation. This lizard often jumps from stone to stone for hunting insects especially grasshop- pers. The sympatric occurrence of Mabuya aurata tran- scaucasica, Darevskia raddei raddei and Laudakia cau- casia caucasia has been documented on Salvat Mountain in a rocky habitat. On the Arshagh Mountains, juveniles with blue tails were found in cliffs, but we could not find adult specimens at this locality. Dorsal coloration is olive-brown with dark spots in longitudinal rows. These spots disappear on the tail and head (Fig. 10). In the study area a specimen with a 1 15 mm snout- vent length and a 125 mm tail was collected. Literature Cited Ahmadzaheh, F. 2004. Preliminary studies of the lizard's fauna and their habitats in Meshkinshahr district. Enviromental Sciences 1(2): 39-44. Anderson, S. C. 1999. The lizards of Iran. Society for the study of Amphibians and Reptiles, 442 pp. Anderson, S. C. 1996. The turtles, lizards, and amphib- ians of Iran. Ph.D. Thesis. Stanford University. 660 pp. Baloutch, M. and H. G. kami. 1995. Amphibians of Iran. Tehran University Publication, Tehran. 177 pp. Firouz, E. 2000. A Guide to the Fauna of Iran (In Per- sian). Iran University Press, Tehran. 491 pp. Kami, H. G. and A. Vakilipoure. 1996a. Geographic dis- tribution: Bufo bufo. Herpetological Review 27(3): 148. Kami H. G. and A. Vakilipoure. 1996b. Geographic dis- tribution: Rana camerani. Herpetological Review 27(3): 150. Latifi, M. 1984. The snakes of Iran. Iran Department of the Environment, Tehran. 221pp. Latifi, M. 1991. The snakes of Iran. Society for the Study of Amphibians and Reptiles. Contributions to Herpetology 7.viii + 159 pp. Leviton, A. E., S. C. Anderson, K. A. Adler and S. A. Minton 1992. Hand book to Middle East Amphibians and Reptiles. Oxford, Ohio. Vii + 252 pp. Mertens, R. F. W. 1957. Weitere Unterlagen zur Herpetofauna von Iran 1956. Jahreshefte des Vereins fur vaterlandische Naturkunde in Wurtemberg 112(1): 118-128. Rastegar-Pouyani, N. 1996. A new species of Asaccus (Sauria: Gekkonide) from the Zagros Mountain, Kermanshahan Province, western Iran. Russian Journal of Herpetology 3(1): 11-17. 2008 Asiatic Herpetological Research, Vol. 11 9 Rastegar-Pouyani, N. and G. Nilson 1998. A new species of Lacerta (Sauria: Lacertidae) from the Zagros Mountain, Esfahan Province, west-central Iran. Proceeding of the California Academy of Science, ser. 4 50(10): 267-277. Tuck, R. G. 1971. Amphibians and reptiles from Iran in the United State National Museum Collection. Bulletin of the Maryland Herpetological Society 7(3): 48-36. Tuck, R. G. 1974. Some amphibians and reptiles from Iran. Bulletin of the Maryland Herpetological Society 10: 59-65. Submitted: 07 August 2006 Accepted: 24 August 2007 pp. 10-12 Asiatic Herpetological Research, Vol. 11 2008 A Second Record of Ptyctolaemus gularis (Peters, 1864) from Bangladesh M. Farid Ahsan*, Ghazi S. M. Asm at and S. Chakma Department of Zoology, University of Chittagong, Chittagong 4331, Bangladesh. * Corresponding author E-mail: mfaridahsan@yahoo.com Abstract.- Ptyctolaemus gularis (Peters, 1864), the blue-throated lizard, was collected from a hilly stream in Rangamat District in Bangladesh in July 2003 and November 2004, representing the second recorded occurrence of this species in Bangladesh. Keywords.- Ptyctolaemus gularis , blue-throated lizard, occurrence, habitat, ecology, morphology, Bangladesh. Introduction The blue-throated lizard, Ptyctolaemus gularis (Peter, 1864), has been previously described from Meghalaya, Assam, the Chittagong Hill Tracts, Tibet and China (Boulenger, 1890; Hora, 1926; Smith, 1935; Zhao and Adler, 1993). Boulenger (1890), who developed the lizard taxonomy of the Indian Subcontinent, examined two specimens of P. gularis, the type specimen from Calcutta, preserved in the Berlin Museum, and a speci- men from Sadiya, Assam, in the British Museum. Following Boulenger (op. cit.), Hora (1926) reported ten specimens from Assam and Nainimukh (correctly spelled Mainimukh), Chittagong Hill Tracts, which are presently deposited in the Zoological Survey of India. The single specimen from Nainimukh represented the first record of this species in Bangladesh. This record has been subsequently overlooked by other authors, including Ahsan, 1998; Khan, 1982; Sarker and Sarker, 1988. Observations and Discussion During a herpetological survey of Bangladesh, one spec- imen of Ptyctolaemus gularis was collected from Rangamati District (part of the Chittagong Hill Tracts) on 18 July 2003 (Fig. 1). Two other specimens were later collected from Rampahar about 50 km east of Chittagong City in Kaptai National Park (22.30.425' N, 092.10.446' E), Rangamati District, on 25 November 2004 (Fig. 1). The first specimen was collected from Rupkari Chara (23.12.126' N, 092.10.628' E), a hilly stream of the Rupkari Union Parishad under Baghaichari Upazila. The collection site is approximately 7 km northwest from the Baghaichari Upazila headquarter. At the time of collection, approximately 1300 h, the speci- men was observed on a large stone hunting insects. This specimen has been deposited in the departmental muse- um of Zoology, University of Chittagong, Chittagong, Bangladesh (Fig. 2A, B). The other two specimens, cur- rently in the collection of S. Chakma, were collected approximately 150 m apart between 1300 and 1400 h. These animals were also collected while they were hunt- ing for insects. One of us (MFA) also observed this species in Chittagong at the Chunati Wildlife Sanctuary in 1990. With these new records, it is likely that Ptyctolaemus gularis also occurs in the hills of Sherpur, Jamalpur, Hobiganj, Moulvibazar, Sylhet (i.e., British- Indian Assam), Khagrachari, Bandarbans (part of Chittagong Hill Tracts) and Cox’s Bazar Districts in Bangladesh, which share similar habitats. Habitat and Ecology Ptyctolaemus gularis is a terrestrial, diurnal species that is frequently found south of the Brahmaputra River in India (Smith, 1935; Daniel, 2002) and uncommonly encountered in the southeastern hilly forests of Bangladesh. It is most often observed in search of food on land, stones and logs near streams and water-logs. The first collection locality visited in 2003 was a narrow stony stream, with the hills on both sides covered with bamboo brakes (muli [Melocanna bambusoides ]), gameri ( Gmelina arborea) and teak ( Tectona grandis) trees. Ferns and some natural herbs grew between the stones. The second and third specimens collected in 2004 were found on the slopes of a hilly, stony stream close to a waterfall. The upper canopy was dominated by garjan (Diptero carpus spp.) and gutgutia (P rotium ser- ratum) trees, and the lower canopy and forest floor were densely covered by shrubs and herbs. © 2008 by Asiatic Herpetological Research 2008 Asiatic Herpetological Research, Vol. 1 1 11 84' 28 "7~ t. bb V ~. ,-V V- -O v. •'t 26° 24 22 NEPAL LEGEND 1= Sherpur 2= Jamalpur 3= Sylhet 4= Hobiganj 5= Moulovibazar 6= Khagrachari 7= Rangamati 8= Kaptai \ 9= Chittagong (Chunoti) 10= Cox's Bazar ?= Not Confirmed #= Confirmed 84l 86 88 Figure 1. Map showing collection localities. SCALE 40 80 Mi lt r1 0 40 80 120 Km Identification The collected specimens can be most readily separated from congeners by having three parallel longitudinal folds on each side of the throat that converge posteriorly (cf. Boulenger, 1890; Smith, 1935) (Fig. 2C). Other use- ful characters include an olive-brown dorsum with dark transverse bars and/or spots, two curved dark brown cross-bars between the eyes separated by a central light bar, a dark stripe below the eye to the angle of the mouth, dark blue throat folds, and limbs and a tail with dark cross-bars above and yellowish-white cross-bars below (cf. Boulenger, 1890; Smith, 1935). The head is also rather long and narrow with unequally-sized upper scales that are strongly keeled. The dorsal body scales are also unequally-sized, with large, strongly-keeled scales and smaller feeble ones. Several mid-dorsal rows also point backwards and upwards and the ventral scales are strongly keeled and mucronate. The limbs are moderate in size; the third and fourth fingers are equal while the fourth toe is much longer than the third. The tail is rounded, slender and covered with sub-equal keeled scales (cf. Boulenger, 1890; Smith, 1935). Table 1 compares the lengths of the present specimens with those collected previously. Acknowledgments We wish to thank Dr. M. A. G. Khan and an anonymous reviewer(s), who kindly reviewed an earlier version of this manuscript. Mr. M. S. Islam and Mr. M. A. W. Chowdhury kindly helped in drawing the map. Table 1. Comparison of recently collected specimens with those from earlier collections. Snout-vent length (mm) Tail length (mm) Source 76.3* (77,69, 83) 167.7* (157, 166, 180) Present report 80 170 Smith 1935 45.7 162.5 Hora 1926 69.85 (2.75”) 158.75 (6.25”) Boulenger 1890 * Mean and raw data within brackets 12 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 2. Dorsal (A) and ventral (B) aspects of Ptyctolaemus gularis collected. Inset (C) shows the gular region folds converging posteriorly, diagnostic of this species. Literature Cited Ahsan, M. F. 1998. Country report for Bangladesh- Herpetofauna of Bangladesh: present status, distri- bution and conservation. Pp. 9-17. In: A. de Silva (ed.), Biology and Conservation of Amphibians, Reptiles and Their Habitats in South Asia (Proceedings of the International Conference in Biology and Conservation of the Amphibians and Reptiles of South Asia, held at the Institute of Fundamental Studies, Kandy and University of Peradniya, Sri Lanka, August 1-5, 1996). Amphibia and Reptile Research Organization of Sri Lanka (ARROS). Boulenger, G. A. 1890. The fauna of British India including Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis, London. 541 pp. Daniel, J. C. 2002. The book of Indian reptiles and amphibians. Oxford University Press, Oxford. 238 pp. Hora, S. L. 1926. Notes on lizards in the Indian Museum: II. On the unnamed collection of lizards of the Family Agamidae. Records of the Indian Museum 28: 415-420 + 1 plate. Khan, M. A. R. 1982. Wildlife of Bangladesh: a check- list. The University of Dhaka, Dhaka. 173 pp. Sarker, M. S. U. and N. J. Sarker. 1988. Wildlife of Bangladesh (a systematic list with status, distribu- tion and habitat). The Rico Printers, Dhaka. 59 pp. Smith, M. S. 1935. Fauna of British India including Ceylon and Burma: Reptilia and Amphibia, Vol. II- Sauria. Taylor and Francis Ltd., London. 441 pp. + 1 map + 1 plate. Zhao, E-M. and K. Adler. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Oxford, Ohio. 522 pp. Submitted: 11 November 2006 Accepted: 22 September 2007 2008 Asiatic Herpetological Research, Vol. 11 pp. 13-16 Observations on the Ovipositional Behavior of the Crest-less Lizard Calotes liocephalus (Reptilia: Agamidae) in the Knuckles Forest Region of Sri Lanka A. A. Thasun Amarasinghe1’2 and D. M. S. Suranjan Karunarathna1’3 [The Young Zoologists' Association of Sri Lanka, National Zoological Gardens, Dehiwala, Sri Lanka. Corresponding authors E-mails: 2 aathasun@gmail.com; 3 dmsameera@gmail.com Abstract.- A mature female Calotes liocephalus lying on the ground in Pitawala in the Knuckles Forest Region of Sri Lanka. This is the first described observation of the ovipositing of Calotes liocephalus. The ovipositional behavior consisted of digging a hole to lay eggs, laying the eggs, scraping soil to bury the eggs, filling of the spaces between the eggs, the tight compression of the soil and camouflaging the nest. Keywords.- Agamidae, Calotes, egg-laying behavior, Knuckles, Sri Lanka, conservation. Introduction There are eighteen species of agamid lizards in Sri Lanka, fifteen of them are endemic to the island (Bahir and Surasinghe, 2005; Manamendra-Arachchi et al., 2006; Samarawickrama et al., 2006). Seven species belong to the genus Calotes. Five of them (C. ceylonen- sis Muller, 1887; C. liocephalus Gunther, 1872; C. liolepis Boulenger, 1885; C. nigrilabris Peters, 1860; C. desilvai Bahir and Maduwage, 2005) are endemic. The remaining two Calotes (C. calotes [Linnaeus, 1758); C. versicolor (Daudin, 1802]) are probably widespread species throughout South East Asia. According to the published literature, Calotes liocephalus is a largely arboreal species found only in parts of the Knuckles Forest Region in Sri Lanka (Manamendra-Arachchi and Liyanage, 1994). Its conservation status is Rare and Endangered (Bahir and Surasinghe, 2005). It can be dis- tinguished from its congeners by the presence of an oblique fold in front of the shoulder, a lower jaw that is rather short, a head without spines (or rarely a rudimen- tary spine above the ear), enlarged supraocular scales and poorly-developed dorsinuchal crests on the head and lower neck (Manamendra-Arachchi, 1 990). Adults have a snout to vent length of 9 1 mm, a head length of 37 mm, a tail length of 261 mm and an axilla to groin length of 43 mm (Deraniyagala, 1953). Location of observation.- Observations were made approximately 1 km from Matale-Pallegama Road in Pitawala in the Knuckles Forest Region (altitude: 783 m) in Matale District, Central Province. The habitat con- sisted mainly of disturbed home gardens (Ekanayake and Bambaradeniya, 2001). The ground was covered with small amounts of wet leaf litter and the soil was soft. There was approximately 10% canopy cover and the undergrowth consisted primarily of grasses. Observations of the lizard was made by the unaided eye from 2 m away between the hours of 1420 and 1600 hrs. The animal was not disturbed during observation. All measurements were taken to the nearest 0.1 mm using dial calipers. Observations A mature female Calotes liocephalus (snout to vent length: 54.0 mm, head length: 19.4 mm, head width: 11.9 mm, tail length: 156.0 mm, axilla to groin length: 26.5 mm) lying on the ground, approximately 50 cm from the road, was observed on 21 June 2006 at about 1420 hr. The temperature was 23.6°C and the humidity 93%. The weather was gloomy and the cloud cover was 8/8. Digging the nest hole.- First, the lizard lifted the anteri- or part of its body using its forelimbs. It then looked around for ~10 min. During this period it repeatedly turned its head 1 80° five times, without moving its body (Fig. 1). The female then began digging into the ground while scraping the soil with its forelimbs, which was thrown backward under its body through its raised hind limbs. This continued for approximately 5 minutes (Fig. 2). After that it stopped digging and looked around for approximately 5 min. while repeatedly turning its head 180° three times, without moving its body (Fig. 3). Again, it continued digging and this time the female dug the hole continuously for approximately 10 min. It stopped and looked around for about 5 min. while turn- ing its head 1 80° around three times, without moving its body as in Figure 3. After that, it continued to dig the © 2008 by Asiatic Herpetological Research 14 Asiatic Herpetological Research, Vol. 11 2008 Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. Figure 7. Figure 8. 2008 Asiatic Herpetological Research, Vol. 1 1 15 Figure 9a. Figure 9b. Figure 10. Figure 12. hole for another half an hour, stopping three more times for 5 min. each, to rest. The hole was dug into the ground at a 45° angle. The final hole was 92.6 mm deep and 79.1 mm in diameter (Fig. 4). During the rest intervals the body was coiled inside the hole with the anterior half bent at an angle of 90° to looking around (Fig. 5). There was a drizzle for ~15 minutes, but the female continued digging. Laying the eggs. - After half an hour of digging, the female turned its body 180° clockwise, placing the pos- terior part of its body inside the hole. It then looked around again (Fig. 6). The significance of this egg laying Figure 13. behavior was that the female removed herself slowly from the hole without lifting her limbs while it was lay- ing its eggs (Figs. 7-8). Eight eggs were laid at a rate of one per minute. The eggs were pure white and elliptical, with a mean length of 14.8 mm and a mean width 8.6 mm. After the eggs were laid, the female came out of the hole completely and started looking around (Fig. 9a, b). Then the female crept back into the hole for 1 5 min. to pack and place the eggs below ground level using the anterior part of its lower jaw (Fig. 10). Buiying the eggs and camouflaging the nest.- After coming out of the hole, the female turned 180° clock- 16 Asiatic Herpetological Research, Vol. 1 1 2008 wise and began to drag the soil towards the hole using its lorelimbs. The dragged soil was thrown backwards under its body while it lifted its hind limbs (Fig. 11). After dragging the soil for about 5 min., it turned 180° counter-clockwise and began pressing the soil with the anterior half of its lower jaw for half an hour. The hole was filled up to 18.4 mm below ground level (Fig. 12). After looking around, it dragged the surrounding Albizia saman (Family: Fabaceae) leaves over the nest site for camouflage (Fig. 13). It remained motionless for 2 min. and then ran towards the forest, during which time it was caught for measurement and then released. Discussion The oviposition behavior of this species varies from the oviposition behavior of Calotes versicolor. According to Amarasinghe and Karunarathna (2007), C. versicolor places its cloacal aperture over the opening of the hole while laying its eggs, but C. liocephalus places the pos- terior part of the body inside the hole while laying eggs. C. versicolor also lifts the anterior part of the body with its forelimbs while turning its head to look around, but C. liocephalus coils its entire body inside the hole while bending the anterior part of its body to look around. C. versicolor makes a knocking noise while packing and placing the eggs in the hole using its lower jaw while the C. liocephalus places them softly without making any noise. After the observation the eggs were removed from the hole and the hole was subsequently examined. The bottom was conical and the soil was soft, dark and wet. Finally the eggs were buried in a home garden to hatch. After approximately two and half months we observed five small holes where the hatchlings had come out. Unfortunately we could not observe the hatchlings. A Few diagrams, brief descriptions and notes of Calotes liocephalus are available in popular journals, books and magazines but almost nothing exists on the pre and post mating behavior, egg laying behavior, cap- tive breeding and their hatchlings. In addition Calotes liocephalus is an endemic, rare and threatened species and therefore it may become extinct if their population does not increase. For such a situation to be achieved, captive breeding methods may be needed for ex-situ conservation of this species. In addition further observa- tions are also needed for the conservation of Calotes lio- cephalus. Acknowledgments We wish to thank Mr. Kelum Manamendra-Arachchi (WHT - Wildlife Heritage Trust) for reviewing the man- uscript and Mrs. Zeenia Nissam of the Department of Zoology, Faculty of Natural Sciences of the Open University, Sri Lanka, for her generous support for the field visit. We also thank Mr. Niranjan Karunarathna and Mr. Gayan Wijeytunga (YZA - The Young Zoologists’ Association) for assisting the fieldwork. Finally, Ms. Debbie McCormick and Mr. F. S. Abeywickrama are acknowledged for their help in preparing this paper. Literature Cited Amarasinghe, A. A. T. and D. M. S. S. Karunarathna. 2007. Beobachtungen zum Eiablageverhalten der Indischen Schonechse Calotes versicolor (Daudin, 1802) (Reptilia: Agamidae) in einem anthropogenen Biotop in Sri Lanka. Sauria, Berlin, 29(3): 27-30. Bahir, M. M. and T. D. Surasinghe. 2005. A conservation assessment of the agamid lizards of Sri Lanka. The Raffles Bulletin of Zoology, Suppl. 12: 381-392. Deraniyagala, R E. P. 1953. A colored atlas of some ver- tebrates from Ceylon, Tetrapod Reptilia. National museums of Sri Lanka. Ekanayake, S. and C. N. B. Bambaradeniya. 2001. Trekking in the Knuckles forest - A trekking guide to Alugallena, Dekinda and Nitre cave nature trails. IUCN Sri Lanka. Manamendra-Arachchi, K. 1990. A guide to the Agamids in Sri Lanka. Occ. Papers of the Young Zoologist Association of Sri Lanka. 5: 1-8. Manamendra-Arachchi, K. and S. Liyanage. 1994. Conservation and distributions of the agamid lizards of Sri Lanka with illustrations of the extant species. Journal of South Asian Natural History 1(1): 77-96. Manamendra-Arachchi, K., A. de Silva and T. Amarasinghe. 2006. Description of a second species of Cophotis (Reptilia:Agamidae) from the highlands of Sri Lanka. Lyriocephalus 6(1): 1-8. Samarawickrama, V. A. M. P. K., K. B. Ranawana, D. R. N. S. Rajapaksha, N. B. Ananjeva, N. L. Orlov, J. M. A. S. Ranasinghe and V. A. P. Samarawickrama. 2006. A new species of the Genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology 13(3): 207-214. Submitted: 10 October 2006 Accepted: 23 September 2007 2008 Asiatic Herpetological Research, Vol. 11 pp. 17-23 On the Status of the Chinese Pitviper Ceratrimeresurus shenlii Liang and Liu in Liang, 2003 (Serpentes, Viperidae), with the Addition of Protobothrops cornutus (Smith, 1930) to the Chinese Snake Fauna Patrick David1’*, Haiyan Tong2, Gernot Vogel3 and Mingyi Tian4 1 Departement Systematique et Evolution, USM 602 Taxonomie-collection - Reptiles et Amphibiens, CP 30, Museum National d’Histoire Naturelle, 57 rue Cuvier, F-7523J Paris Cedex 05, France, 2 16, cour du Liegat, F -7 5013 Paris, France, dm Sand 3, D-6911 5 Fleidelberg, Germany, 4 College of Natural Resources & Environment, South China Agricultural University, Wushan, Guangzhou 510640, People’s Republic of China. * Corresponding author E-mail: pdavid@mnhn.fr Abstract.- Ceratrimeresurus shenlii Liang and Liu in Liang, 2003 was described as a new genus and a new species on the basis of the first “homed” specimen of pitviper recorded from the People’s Republic of China. This taxon has been overlooked in the literature. The original description is here translated verbatim into English. The holotype is compared with other homed pitvipers known from Asia. On the basis of its scalation and pattern, Ceratrimeresurus shenlii is synonymized with Protobothrops cornutus (Smith, 1930). The range of this latter species, previously endemic to Vietnam, is expanded northeastwards by approximately 780 airline km. A brief comment on the zoogeog- raphy of South China is given. Keywords.- Serpentes, Ceratrimeresurus, Ceratrimeresurus shenlii, Protobothrops cornutus, China, Vietnam, taxon- omy, zoogeography. Introduction A new Chinese genus and new species of Asian pitviper, Ceratrimeresurus shenlii Liang and Liu in Liang, 2003 has remained overlooked in the literature. Ceratrimeresurus shenlii was not included in the latest checklist of the Trimeresurus complex (Gumprecht et al., 2004), nor announced in Wolfgang Wiister’s invalu- able website “Venomous Snakes Systematic Alert” (http://biology.bangor.ac.uk/~bssl66/update.htm). Lastly, this taxon was not considered by Malhotra and Thorpe (2004, 2005) in their revision on the pitvipers of the Trimeresurus complex. The description of Ceratrimeresurus shenlii Liang and Liu in Liang (2003: 411; Plate 8: Fig. 13. Type local- ity: “Working site 02 at Wuzhishan forest, Ruyuan Xian”, Guangdong Province) appeared in a chapter of a book on the natural history of Nanling Nature Reserve, located in Nanling Mountains, in the north of Guangdong Province (Pang, 2003), a fact that may explain that the new taxon remained overlooked by the herpetological community. However, this new species was merely mentioned in September 2004 in the forum of a website dedicated to venomous snakes (http://www.venomdoc.com/fomms - last viewed on May 30th, 2005), where this species was regarded, with- out explanations, as a synonym of Protobothrops cornu- tus. It was also tentatively regarded as a synonym of Protobothrops cornutus by Vogel (2006). Only three other taxa of Asian pitvipers with more or less raised supraoculars were previously known from the mainland: Trimeresurus wiroti (see David et al., 2006), Protobothrops cornutus and Triceratolepidophis sieversorum. In this paper, we provide a translation of the original description. The new taxon is, according to the re-examination of the holotype, compared with the currently known homed species of mainland Asia and its status is discussed. Materials and Methods Body and tail lengths were measured to the nearest mm. The number of ventral scales is counted according to Dowling (1951). The terminal scute is not included in the number of subcaudals. The numbers of dorsal scale rows are given at one head length behind head, at mid- body (i.e. at the level of the ventral plate corresponding to a half of the total number of ventrals), and at one head length before vent, respectively. Values for symmetric head characters are given in left/right order. © 2008 by Asiatic Fferpetological Research 18 Asiatic Herpetological Research, Vol. 1 1 2008 Abbreviations of measurements and other meristic characters.- Measurements and ratios: HL: head length; SVL: snout-vent length; TaL: tail length; TL: total length; TaL/TL: ratio tail length /total length. Meristic characters: DSR: formula of dorsal scale rows; MSR: number of dorsal scale rows at midbody; IL: infralabials; SC: subcaudals; SL: supralabials; VEN: ventrals. Museum abbreviations.- BMNH: Natural History Museum, London, UK. - FMNH: Field Museum of Natural History, Chicago, USA. - MNHN: Museum National d’Histoire Naturelle, Paris, France. - ZFMK: Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. Results Translation of the original description of Ceratrimeresurus shenlii.- The original description appeared in Liang (2003: 411), but is credited to Liang and Liu. It was published in Chinese with a short English summary. A poor quality, small color photo- graph of the holotype appears on Plate 8, Figure 13 of the book. A verbatim translation of the original descrip- tion should read as follows (numbers placed in square brackets refer to our comments placed after this transla- tion): Jiao laotietou Ceratrimeresurus shenlii Liana |2| ° and Liu, gen. and sp. nov. (See colour plate 13 ) The upper eyelid forms on each side a raised triangle covered with small scales, the base of the triangle being slightly like a three-sided pyramid; the length of the tri- angular horn is 1 .5 times the diameter of the eye, and the height from the tip of the triangle to the edge of the upper eyelid is equal to the eye diameter. The tip of the snout is blunt, the upper snout surface is broad and flat- tened, with the lateral edge slightly upturned. The length of the snout is about a 1/4 of head length and 1/3 of head width. The nostril is placed near the tip of the snout, slightly directed sideward and almost rounded in shape, the opening being in the middle of a divided nasal scale; there is one upper nasal scale. There are 5 small scales between the two nostrils . The upper head surface is covered with granular scales, which become progres- sively larger and more keeled from the front to the back of the head. The cephalic surface is slightly convex and becomes progressively flat and broader backwards. There are 7 small interorbital scales . There are 11 small scales surrounding the base of the triangular horn. The tip of the triangular horn is one rather large scale, to which is adjacent a smaller scale which makes the horn looking as bifurcated. The eye is rather large and the third of its volume is out of the head. The iris is a verti- cally elongated oval. There is a row of small scales around the orbit and many scales between the snout side and temporal region. The forward sunken part of the occiput is roughly equal to the length of the snout, with the end of the neck plunged in it. The neck is slender, its diameter being less than half of the head width. There are 14 supralabial scales , among which the first and second ones are separated from the nasal scales by a row of 3 small scales, and the third to fifth ones do not enter the loreal pit161, but are separated from it by a row of small scales; the edge of the upper jaw is convex down- wards; 14 infralabials, the fourth being the largest, the first to the fourth ones in contact with the anterior chin shields, and the posterior chin shields being rather large. One large fang is located on the anterior part of the upper jaw, followed by 2 smaller teeth. No teeth on the palatine and pterygoid . Five small teeth are present on the lower jaw. The loreal pit is placed antero-inferiorly to the eye. The head is peach-shaped and slightly flattened, with a narrow and elongated neck; the body is rather wide but slender and long, with a long, pointed tail. The dorsal scales are strongly keeled, 23-23-21-17 rows, the rows of cervical scales are more oblique than those of the body scales. There are 187 ventral scales +77 pairs of subcaudals. The anal scale is single. The vertebral scales are normal. The back of the body is of a grey color, which becomes light grey on the sides. On the head upper sur- face, there is a ‘X’ shaped pattern, made of a blotch extending from the nasal scale to the front of the oppo- site triangular, and of a blackish brown elongated blotch extending from the posterior of the horn through the rearward supraorbital scale (behind the horn) backwards up to the posterior of the head. A white streak extends from the posterior margin of the eye through the tempo- ral up to the posterior of the head, followed by a blackish brown stripe from the comer of the mouth to the sides of the throat. The upper and lower lips are marked with blackish brown square shaped spots. Diffuse and irregu- lar spots cover the rest of the body throughout. The dor- sal side of the head and the body is brown in color. Dorsal blotches extend on each side from the middle of the body to the vent, the right and the left rows being off- set and these blotches being linked together by their inner comer; they are merged together in the middle of the body and linked each other again by the inner comer on the posterior part of the body, then merged together again from the vent to the end of the tail. Light grey- brown spots on the lower part on the sides of the body, roughly square in shape or almost rounded, oval or 2008 Asiatic Herpetological Research, Vol. 1 1 19 reduced to short crossbars. The venter is pale grayish- brown, without other spot in the middle, so that the ven- tral side of the body is light grey-brown color through- out. There is a pointed scale on the tip of the tail. The total length of the specimen: 362 + 78 = 440 mm, length of the tail/total length = 0.564, weight = 24 g. The specimen was collected in the thatch on the top ot a house near a forest, coiled with the head in the mid- dle ot its body. Not very active. It was knocked down with a little stick on the ground and captured. The local- ity is^the “Working site 02” at Wuzhishan forest, Ruyuan Xian , July 1996. The specimen is deposited in the Laboratory of Zoology of the Faculty of Biotechnology, Jinan University'91. Comments on the original description. - In this original description, no mention of the sex of the specimen is given, although it is a female according to the shape of the base of the tail. Other comments are: [1] : Translation: “Homed iron-head [snake]”. The ver- nacular name “ Laotietou ” is given in China to Protobothrops mucrosquamatus, due to the shape of its head, looking like an ancient iron. [2] : It should rather appear as Plate 8: fig. 13. [3] : This value includes the two supranasals plus the scales separating these latter ones. [4] : If one counts the cephalic scales on a line connect- ing the middle of the supraoculars, the value is 14 (see below). [5] : This value is obviously erroneous; we counted 8 SL. [6] : This is obviously a lapsus for the orbit. [7] : this lack of teeth on the palatine bones is surprising and may be an artifact. Teeth are very commonly miscounted, as they are masked by the tissues and one usually needs to peel back the gum tissues to count the sockets. All the tooth counts in this description may be unreliable (A. Malhotra, pers. comm, January 2006). [8] : Ruyuan County, in northern Guangdong Province, close to the Guangdong-Hunan border. The Nanling National Nature Reserve is located in the centre of Nanling Mountains, which extend from northeast Guangxi to southwest Fujian. [9] : Guangzhou, Guangdong Province. Redescription of the holotype of Ceratrimeresurus shenlii (Figs. 1-3).- The original description includes some imprecision or mistakes. In combining data of the original description and our own data, the morphology is as follows: Body moderately stout; head subtriangular, wide at its base, clearly distinct from the neck, thick and swollen when seen from the side, depressed between the uplifted areas of the supraoculars. Snout average in relative length, about one quarter of HL, bluntly rounded when seen from above, depressed in its center, obliquely trun- cated when seen from the side, with a distinct canthus rostralis. Eye relatively large. Tail long and tapering. The holotype is now in average condition and somewhat dessicated: SVL: 362 mm; TaL: 78 mm; TL: 440 mm; ratio TaL/TL: 0.177. VEN: 187; SC: 77, paired, plus one terminal scale; anal shield entire. DSR: 23-21-17 scales, rhomboid, distinctly keeled. Rostral visible from above, broader than high, trian- gular; nasals subrectangular, divided, with a round nos- tril opening near the tip of the snout, directed slightly sideward and almost rounded in shape; 2 intemasals on each side, separated by 2 small scales; 4/4 canthal scales, slightly larger than adjacent snout scales, border- ing the canthus rostralis ; 2 elongate upper preoculars above the loreal pit; lower preocular forming the lower margin of loreal pit; 3/3 small postoculars; 5/5 supraoc- ulars on an uplifted triangular area, of which the two central supraocular scales are strongly enlarged, triangu- lar and strongly obliquely erected (“horn like”) and extending out of the head margin, 1.5 times the diameter of the eye, convergent and originating from the same base covered with small scales; 8 or 9 slightly enlarged scales on upper snout surface on a line between the scales separating the intemasals and a line connecting the anterior margins of eyes, smooth, juxtaposed, irreg- ular in shape; 14 cephalic scales on a line between the base of the supraoculars (including the scales covering the uplifted basal area), smooth, flat and juxtaposed; occipital scales larger than cephalic scales, distinctly keeled; temporal scales small, obtusely keeled: 9 SL. 1st SL separated from nasal; 2nd SL bordering the anterior margin of the loreal pit, 3rd SL largest, separated from the subocular by 1 scale; 4th SL much smaller than 3rd SL, separated from the subocular by 1 scale; 5th and pos- terior SL much smaller; 14 IL. first four pairs in contact with anterior chin shields. The coloration and pattern agree with that of the original description. The light postocular streak is quite conspicuous, but the overall pattern of the body is rather faded. Discussion Nomenclatural considerations.- The combined descrip- tion of the genus and species was brief and may raise some questions about its validity. Two points need to be discussed. The first one relates to the combined descrip- tion of the genus Ceratrimeresurus and of its sole 20 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 1. Ceratrimeresurus shenlii, holotype. General view of body. Photograph by Tian Mingyi. Figure 3. Ceratrimeresurus shenlii, holotype. Lateral view of the head. Photograph by Tian Mingyi. included species, shenlii. According to Art. 13.4 of the International Code of Zoological Nomenclature (1999), in the present case, such a “description of a new nominal genus and a single included new nominal species is deemed to confer availability on each name under Article 13.1.1.” Art. 13.1.1 requires that every new name published be accompanied by a description purported to differentiate the taxon. This combined description is hence considered to be complying with the requirements of the Code. The second point relates to the identification of the holotype. No number was cited in the description, and this specimen was still deposited in Prof. Liang Qishen’s private collection in December 2005. Nevertheless, as the name of the collection into which the specimen will be eventually deposited was clearly indicated in the original description, and as it was confirmed to us that the specimen will eventually be deposited in this collec- tion, we consider that this point complies with Art. 16.4.2 of the Code and makes valid the description of Figure 2. Ceratrimeresurus shenlii, holotype. General view of body. Photograph by Tian Mingyi. Figure 4. Head of the second Chinese specimen of Protobothrops cornutus. Photograph by Fu Jie. the specific nomen. A comparison of Ceratrimeresurus shenlii with other horned species.- In the Asian mainland, only Protobothrops cornutus and Triceratolepidophis siever- sorum have erected supraoculars. Trimeresurus wiroti has only slightly raised supraoculars (see David et al., 2006). Another species occurring in western Indonesia, Trimeresurus brongersmai, has erected hom-like supraoculars. However, this latter species, related to Trimeresurus wiroti, differs from the other homed species by several characters, including the shape of its snout (David et al., 2006). On the basis of Ziegler et al. (2001), Herrmann et al. (2002), Herrmann and Ziegler (2002), Herrmann et al. (2004) and of specimens exam- ined by ourselves, a comparison between horned species and Ceratrimeresurus shenlii is given in Table 1 . Triceratolepidophis sieversorum differs by its greater number of ventral scales and a different structure of horns, free and strongly divergent. This species also 2008 Asiatic Herpetological Research, Vol. 1 1 21 *'*38 ... MUli ASO VII KAM RA-wucmufXA ' mm*. •**„ ' *!!«►! - mm hx-im >, Figure 5. Distribution map of of Protobothrops cornutus. shows a peculiar structure of keels of dorsal scales, which is not found in other homed pitvipers (Ziegler et al., 2001). The dorsal keels of Ceratrimeresurus shenlii are narrow and made of a single ridge. In contrast, Ceratrimeresurus shenlii cannot be dis- tinguished from Protobothrops cornutus otherwise than by minor characters. In both species, the horns stem from the same base and are free only in their outermost part. All scalation characters are similar, including the keeling of the dorsal scales (see Table 1). Other charac- ters include the keeled occipital scales, the large 3rd SL separated from the subocular by 1 scale row, 4th and 5th separated from the subocular by 1 or 2 scales, the num- ber of intemasals and of supraoculars. The sole differ- ence bears on the numbers of pairs of infralabials in con- tact with the anterior chin shield, first 3 pairs in P. cor- nutus vs. first 4 pairs in Ceratrimeresurus shenlii. The pattern is also similar, especially the dorsal blotches, the upper head dark pattern and the postocular streak. On the basis of the similarities in morphological characters, we synonymize Ceratrimeresurus shenlii Liang and Liu in Liang (2003) with Trimeresurus cornu- tus Smith, 1930, now Protobothrops cornutus (see Herrmann et al. [2004] for the generic position of this species). At the generic level, the point to be now discussed is if the homed species Trimeresurus cornutus Smith, 1930 should be referred to a genus distinct from Protobothrops, in which all other included species are hornless. In this case, the generic nomen Ceratrimeresurus is available. Pending molecular analy- ses that should clarify the relationships between the Chinese and the Vietnamese populations, we have to rely only on morphology. Only two currently recognized genera of mainland Asia include species with truly erect- ed supraoculars (“horns”), namely Protobothrops (only for P cornutus ) and Triceratolepidophis (T. sieverso- nim). On the basis of the similarities between P. cornu- tus and C. shenlii, we adopt a conservative approach in considering that erected supraoculars have been conver- gently evolved in several lineages. Consequently, we synonymize the genus Ceratrimeresurus Liang and Liu in Liang (2003) with Protobothrops Hoge and Romano Hoge, 1983. A second Chinese specimen of Protobothrops cor- nutus has appeared on Internet in summer 2005 (Fig. 4), from Shimentai Nature Reserve (24° 22'-24° 3 1 ' N, 113° 05'- 113° 31' E), Nanling Mountains, Yingde County, Guangdong Province (see Jim and Xu [2002] for a description of the area). The characters visible on this picture of a freshly killed specimen are identical with Table 1. A comparison between known specimens of Asian horned pitvipers. Taxon Sex Tal/TL MSR Ven Sc Cep SL IL Sup/Oc SupOc “C. shenlii1’ F 0.177 21 187 77 14 9/7 14/14 5/5 convergent R cornutus (1) M 0.203 21 189 78 16 9/9 14/14 6/6 convergent P. cornutus (2) F 0.184 21 192 71 13 9/9 13/14 6/6 convergent P. cornutus (3) F 0.182 21 193 72 13 9/9 12/13 7/7 convergent T. sieversorum (4) M 0.167 23 228 82 15 8/9 13/14 2/2 divergent T. sieversorum (5) F 0.157 22 235 79 15 8/9 13/14 2/2 divergent T. sieversorum (6) ? — 21 — — 16 10/10 13/14 — divergent List of cited specimens. Protobothrops cornutus. (1) ZFMK 75067, Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam (not seen; after Herrmann et al., 2004); (2) BMNH 1946.1 .19.25, “Fan-si-pan Mts., Tonking", now Mt. Phang Si Pang, Lai Chau Province, Vietnam; (3) MNHN 1937.35, “Tonkin”, northern Vietnam. - Triceratolepidophis sieversorum. (4) ZFMK 71262 (holotype), Phong Nha village, Phong Nha-Ke Bang Nature Reserve, Quang Binh Province, Vietnam; (5) ZFMK 75066, Phong Nha-Ke Bang Nature Reserve, Quang Binh Province, Vietnam; (6) FMNH 255258, Hin Nammo NBCA, Boualapha District, Khammouan Province, Laos (specimens (5) and (6) not seen; after Herrmann et at, 2002). 22 Asiatic Herpetological Research, Vol. 1 1 2008 those ot Ceratrimeresurus shenlii. The pattern is much similar to that of Protobothrops cornutus depicted in Herrmann et al. (2004). According to the data kindly communicated to us by Mr. Fu Jie, the author of the pho- tograph, this second specimen was seen in May 2005. It was held in alcohol in the home of a member of the Yao minority, to be most likely used as a medicinal beverage. The locality of this second specimen lies in a hilly area at approximately 7 5 airline km southeastwards from the type locality of Ceratrimeresurus shenlii in Nanling Nature Reserve. According to the Yao owner of this specimen, this snake was caught by himself while he was acting as a guide to a scientific survey of the local herpetological fauna. Several specimens were collected, but all died in captivity within some days. This species is there considered very rare. Conclusions The occurrence of Protobothrops cornutus in China makes a considerable northeastward range extension for this latter species, previously considered endemic to Vietnam (Nguyen et al., 2005) (see map on Fig. 5). Ziegler et al. (2006) recorded a specimen of Protobothrops cornutus from Ha Giang Province, in extreme northern Vietnam, close to the border with Yunnan and Guangxi Zhuang Autonomous Provinces. The previously northernmost known locality, Mt. Phang Si Pang (formerly Mt. Fan Si Pan) and Nanling Mountains are separated by about 980 airline kilome- ters. Chinese specimens are separated from the new locality cited in Ziegler et al. (2006) by about 760 airline km. We also consider that the wide gap between the Chinese population and its Vietnamese relatives may most likely reflect a lack of appropriate collecting effort in elevated areas of Guangxi Zhuang Autonomous and Guangdong Provinces than a real geographical gap. Protobothrops cornutus should be searched for in forests of mountain or hill ranges such as Darning Shan and Dayao Shan (Guangxi Zhuang Autonomous Province) and in various hills of northern Guangdong located between the Nanling Mountains and the mountain ranges of Guangxi Autonomous Region. The herpeto- fauna of this latter province is still quite poorly known. Besides the provinces of Guangxi and Guangdong, this species should be searched for in forested areas of south- ern Yunnan (see Herrmann et al., 2004). Herrmann et al. (2004) also showed that, in Central Vietnam, P. cornutus also occurs in the lowlands. However, lowland areas of southern China are quite dry (Anonymous, 1998). These lowlands separate the hill or mountain ranges of south- ern China which share a subtropical humid climate with high annual amounts of rainfall (above 2000 mm), a sit- uation which leads to the isolation of the populations of P. cornutus in this region. A discussion on the zoogeographical affinities between North Vietnam and various regions of South China are outside the scope of this paper, but a mere comparison between the snake faunas of northern Vietnam and the hills of Guangxi and Guangdong sug- gests strong similarities. The occurrence of Protobothrops cornutus in northern Guangdong, as well as of Shinisaurus crocodilurus and Amphiesma bitaenia- tum in northern Vietnam (David et al., 2005; respective- ly Le and Ziegler, 2003) reinforces the zoogeographical relationships of these areas connected by more or less contiguous hill or mountain ranges receiving high annu- al amounts of rainfall. Acknowledgments We are indebted to Q. Liang (Guangzhou, China), who was instrumental in letting TM re-examine the holotype of Ceratrimeresurus shenlii. We are grateful to A. Malhotra (Bangor, United Kingdom), M. W. Lau (Hong Kong, China) and T. Ziegler (Koln, Germany) for their comments on the draft of this paper; we especially thank T. Ziegler who made available and permitted us to cite his paper in press. Lastly, we thank J. Fu (Guangzhou, China) and N. Wu (Beijing, China) for their kind permis- sion to use in our paper the picture of the second Chinese specimen of Protobothrops cornutus. Literature Cited Anonymous (1998) Physiogeographical Atlas of China. 2nd Edition. China Cartographic Publishing House, Beijing. 51 + 112 pp. David, P., G. Vogel and O. S. G. Pauwels. 2005. On the occurrence of Amphiesma bitaeniatum (Wall, 1925) in Vietnam, with preliminary remarks on the group of Amphiesma parallelum (Boulenger, 1890) (Serpentes, Colubridae, Natricinae). Salamandra 41(4): 167-178. David, P., G. Vogel, S. P. Vijayakumar and N. Vidal. 2006. A revision of the Trimeresurus puniceus- complex (Serpentes: Viperidae: Crotalinae) based on morphological and molecular data. Zootaxa 1293:1-80. Dowling, H. G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of 2008 Asiatic Herpetological Research, Vol. 1 1 23 Herpetology 1(5): 97-99. Gumprecht, A., F. Tillack, N. Orlov, A. Captain and S. Ryabov. 2004. Asian Pitvipers. Geitje Books, Berlin. 368 pp. Herrmann, H.-W., T. Ziegler, B. L. Stuart and N. L. Orlov. 2002. New findings on the distribution, mor- phology, and natural history of Triceratolepidophis sieversorum (Serpentes: Viperidae). Herpetological Natural History 9(1): 89-94. Herrmann, H.-W., T. Ziegler, A. Malhotra, R. S. Thorpe and C. L. Parkinson. 2004. Redescription and sys- tematics of Trimeresurus cornutus (Serpentes: Viperidae) based on morphology and molecular data. Herpetologica 60(2): 211-221. International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature. Fourth Edition. International Trust of Zoological Nomenclature, London, xxix +306 pp. Jim, C. Y. and S. S. W. Xu. 2002. Perception of local inhabitants towards nature reserve establishment in a remote mountain area in South China. Proceedings of IUCN/WCPA-EA-4 Taipei Conference, March 18-23, 2002, Taipei, Taiwan: 459^167. Le, K. Q. and T. Ziegler. 2003. First record of the Chinese crocodile lizard from outside of China: Report on a population of Shinisaurus crocodilurus Ahl, 1930 from north-eastern Vietnam. Hamadryad 27: 193-199. Liang, Q. 2003. Reptiles of Guangdong Nanling Nature Reserve. In: Pang, X. (Ed.), Studies on Biodiversity of the Guangdong Nanling National Nature Reserve. Guangdong Science and Technology Press, Guangzhou: 408^117, PI. 8. (In Chinese). Malhotra, A. and R. S. Thorpe. 2004. A phylogeny of four mitochondrial gene regions suggests a revised taxonomy for Asian pitvipers ( Trimeresurus and Ovophis). Molecular Phylogenetics and Evolution 32: 83-100. Malhotra, A. and R. S. Thorpe. 2005. Erratum to “A phylogeny of four mitochondrial gene regions sug- gests a revised taxonomy for Asian pit vipers (; Trimeresurus and Ovophis)". [Mol. Phylogen. Evol., 32 (2004) 83-100], Molecular Phylogenetics and Evolution 34: 680-681. Nguyen, V. S, T. C. Ho and Q. T. Nguyen. 2005. Danh luc ech nhai va bo sat Viet Nam. A checklist list of Amphibians and Reptiles of Vietnam. Nha xuat ban Nong Nghiep, Hanoi. 180 pp. (In Vietnamese). Pang, X. (Ed.). 2003. Studies on Biodiversity of the Guangdong Nanling National Nature Reserve. Guangdong Science and Technology Press, Guangzhou. 474 pp., 8 pis. (In Chinese). Vogel, G. 2006. Terralog. Venomous snakes of Asia - Giftschlangen Asiens. Edition Chimaira, Frankfurt am Main & Aqualog Verlag ACS, Rodgau. 148 pp. Zhao, E. M., M. H. Huang, Y. Zong, J. Zheng, Z. J. Huang, D. T. Yang and D. J. Li (Eds.). 1998. Fauna Sinica. Reptilia Vol. 3. Squamata Serpentes. Science Press, Beijing, xvii + 522 pp., 8 pis., 4 col. Pis. (In Chinese). Ziegler, T., H.-W. Herrmann, P. David, N. Orlov and O. S. G. Pauwels. 2001 [2000], Triceratolepidophis sieversorum, a new genus and species of pitviper (Reptilia: Serpentes: Viperidae: Crotalinae) from Vietnam. Russian Journal of Herpetology 7(3): 199-214. Ziegler, T. and H.-W. Herrmann. 2002. Wiederentdeckung zweier gehornter Grubenotterarten in Vietnam. Mitteilungen Zoologische Gesellschaft fur Arten- und Populationsschutze e.V. 18(2): 24—26. Ziegler, T., N. T. Vu, K. Q. Le, Q. T. Nguyen, J. Hallermann, V. K. Le & M. H. Thach. 2006. Neue Verbreitungsnachweise einiger vennig bekannter Vietnamesischer Amphibien und Reptilien. Sauria 28(2): 29-40. Submitted 13 June 2006 Accepted: 04 April 2007 pp. 24-30 Asiatic Herpetological Research, Vol. 1 1 20081 The Effects of Incubation Temperature On Hatching Success, Embryonic Use of Energy and Hatchling Morphology in the Stripe-tailed Ratsnake Elaphe taeniura Wei Guo Du1’* and Xiang Ji1’2 1 Hangzhou Key Laboratory for Animal Sciences and Technology, Hangzhou Normal University, 310036, Hangzhou, Zhejiang, R R. China 2 Jiangsu Key Laboratory for Biodiversity and Biotechnology, College of Life Sciences, Nanjing Normal University, Nanjing 210097, Jiangsu, P R. China * Corresponding author E-mail: dwghz@126.com Abstract .- We incubated eggs of Elaphe taeniura at 22, 24, 27, 30 and 32°C to examine the effects of incubation tem- perature on hatching success, embryonic use of energy and hatchling morphology. Incubation temperature affected incubation length and most hatchling traits examined in this study. Incubation length increased nonlinearly as tem- perature decreased, with the mean incubation length being 101.7 d at 22°C, 86.0 d at 24°C, 66.3 d at 27°C, 53.9 d at 30°C, and 50.5 d at 32°C. Hatching successes were lower at the two extreme temperatures (41.2% at 32°C, and 50.0% at 22°C) than at the other three moderate temperatures (78.1-79.3%). Hatchlings from the extreme high incu- bation temperature (32°C) were smaller in body size and wet body mass. High incubation temperatures resulted in production of less developed hatchlings that characteristically had less developed carcasses but contained more unuti- lized yolks. The proportion of energy transferred from the egg contents to the hatchling was 71.1% at 22°C, 80.2% at 24°C, 81 .5% at 27°C, 82.6% at 30°C and 83.9% at 32°C. Taking the lowest hatching success at 32°C and the sub- stantially prolonged incubation lengths at 22°C into account, we conclude that these temperatures are not suitable for embryonic development in E. taeniura. Our data confirm the prediction that there are some thresholds over which incubation temperatures can affect hatching success, embryonic use of energy and hatchling morphology. Keywords.- Reptilia, Colubridae, Elaphe taeniura, egg, incubation, temperature, hatchling phenotype. Introduction As in other vertebrate and invertebrate taxa, temperature may profoundly influence embryonic development in reptiles. Compared to avian embryos, reptilian embryos can develop under a relatively wide range of tempera- tures (Birchard, 2004, Booth, 2004). Low temperatures slow embryogenesis but usually have little lethal effect on embryos; high temperatures result in faster embryon- ic development (and thus, shortened incubation or gesta- tion length) but often increase embryonic abnormality or mortality (e.g. Andrews and Rose, 1994; Andrews et al., 1997; Deeming and Ferguson, 1991; Sexton and Marion, 1974; Shine and Harlow, 1996). Apart from the effects on the rate of embryonic development and embryonic abnormality or mortality, thermal environ- ments experienced by developing embryos also affect a number of phenotypic attributes of the hatchling, includ- ing morphology (Du and Ji, 2002; Ji and Du, 2001a, b; Overall, 1994), energy reserves (Du and Ji, 2001), behavior (Burger, 1991; 1998), post-hatching growth (Brana and Ji, 2000; Du and Ji, 2003; Rhen and Lang, 1995), and gender in species with temperature-depen- dent sex determination (Janzen and Paukstis, 1991). It has been repeatedly reported for oviparous reptiles that eggs incubated at optimal temperatures not only exhibit high hatching success but also produce good-quality hatchlings. The range of optimal temperatures for reptilian embryos is often narrow and varies not only among but also within species. For example, the optimal incubation temperatures fall within the range from 24°C to 26°C in Xenochrophis piscator (checkered keelback; Ji et al., 2001) and Deinagkistrodon acutus (five-paced pit-viper; Lin et al., 2005) but, in Elaphe carinata (king ratsnake; Ji and Du, 2001b), Naja atra (Chinese cobra; Ji and Du, 2001a), Rhabdophis tigrinus lateralis (red-necked keel- back; Chen and Ji, 2002), Dinodon rufozonatum (red- banded wolf snake; Ji et al., 1999b; Zhang and Ji, 2002), Ptyas korros (gray ratsnake; Du and Ji, 2002) and P. mucosus (mucous ratsnake; Lin and Ji, 2004), generally within the range from 26°C to 30°C. Pelodiscus sinensis (Chinese soft-shelled turtle), however, has a wider range of optimal incubation temperatures, because tempera- tures exert no important effects on hatching success and hatchling phenotypes within the range from 24°C to 34°C (Du and Ji, 2003; Ji et al., 2003). In Eumeces chi- nensis (Chinese skink), eggs from a lower latitudinal population have a narrower range of optimal incubation © 2008 by Asiatic Herpetological Research 2008 Asiatic Herpetological Research, Vol. 1 1 25 temperatures than do those from a higher latitudinal population, primarily because of more stable thermal environments in the former population (Ji et ah, 2002). Overall, previous studies suggest that optimal tempera- tures for developing embryos differ among reptiles dif- fering in habitat use and/or distributional range. The stripe-tailed ratsnake Elaphe taeniura is a large sized (to 1800 mm SVL [snout-vent length]) oviparous colubrid snake that ranges from the central and southern provinces of China to Korea, Burma, Laos, Vietnam and India (Zhao, 1993). Wild population of this snake have declined dramatically due to habitat loss and over-har- vesting over the past two decades (Zhao, 1998). Fecundity, reproductive output and embryonic mobiliza- tion of energy and material during incubation have been reported for snakes from Zhoushan Islands (Ji et al., 1999a; 2000). Because eggs were never incubated at multiple temperatures, the range of incubation tempera- tures optimal for developing embryos of E. taeniura remains unknown. To fill this gap, we incubated eggs produced by females from a southern population (Guangxi, China) at five constant temperatures ranging from 22°C to 32°C. Specifically, our aims are to (1) examine the effects of incubation temperatures on hatch- ing success, embryonic use of energy and hatchling mor- phology, and (2) determine the range of optimal temper- atures for embryos of E. taeniura. Materials and Methods We obtained 13 gravid E. taeniura (SVL: 1220-1690 mm; postoviposition body mass: 277.9-755.0 g) in June 1998 from a private hatchery in Guilin (Guangxi, south- ern China), and brought them to our laboratory in Hangzhou, where they were maintained in a 2000 x 800 X 800 (length X width X height) mm wire cage placed in a room inside which air temperatures were never higher than 30°C. Food (eggs of Coturnix coturnix) and water were provided ad libitum. The snakes laid eggs between 23 June and 3 July (clutch size: 8.8±0.4, range: 7-11). We collected the eggs within a few hours after being laid. Each egg was measured (to the nearest 0.01 mm) for length and width with a Mitutoyo digital caliper and weighed (to the nearest 1 mg) on a Mettler balance. One freshly laid egg from each clutch was dissected to deter- mine the composition of eggs. Egg contents (yolk plus embryo) were placed in pre-weighed glass dishes, and weighed. Shells were briefly rinsed, dried by blotting with paper towels and weighed. Egg contents and shells were weighed again after oven drying to constant mass at 65°C. The remaining eggs were incubated systemati- cally at five constant temperatures (22, 24, 27, 30, and 32 [± 0.3]°C); such that eggs from single clutches were distributed almost equally among the five temperature treatments. Eggs were individually incubated in covered plastic jars containing known amounts of vermiculite and dis- tilled water at approximately -12 kPa water potential (vermiculite: water = 1:2). One-third of the egg was buried lengthwise in the incubating substrate, with the surface near the embryo exposed to air inside the jar. Jars were equally assigned to five incubators (Guanzhou medical instrument, China), with incubation tempera- tures set at 22, 24, 27, 30, and 32 (± 0.3)°C, respectively. We moved jars among the shelves in the incubator daily according to a predetermined schedule to minimize any effects of thermal gradients inside the incubator. Jars were weighed on alternate days, and distilled water was added evenly into substrates when necessary to compen- sate for evaporative losses and water absorbed by eggs, thereby maintaining the substrate water potential con- stant. The duration of incubation, measured as the number of days to pipping, was recorded for each egg. Hatchlings were collected, measured (for SVL and tail length), and weighed a few hours after hatching, and then euthanized by freezing to -15°C for determination of composition and sex. The killed hatchlings were sep- arated into carcass, residual yolk and fat bodies. The Table 1. The effect of temperature on incubation duration, hatching success, and sex ratio in Elaphe taeniura. Data on incubation duration are expressed as mean±1 SE. Temperature (°C) Duration of incubation (d) Hatching success (%) Sex ratio ($$/ 0.25). Hatching successes were apparently lower at the two extreme temperatures (22°C and 32°C) than at the other three moderate temperatures (24, 27, and 30°C) (Table 1). There were no between-sex differences in all exam- ined hatchling variables (all p > 0.05), so we pooled data for both sexes. All examined hatchling variables, except yolk sac, were positively correlated with initial egg mass). ANCOVAs with initial egg mass as the covariate showed that incubation temperatures significantly affected wet body mass (F474 = 3.67, p <0.01; Fig. la), SVL (F4 j4 = 4.21, p < 0.01; Fig. lc) and tail length ( F474 = 4.31,/? < 0.01; Fig. Id) of hatchlings, but not hatchling dry body mass ( F 474 = 0.38, p > 0.05; Fig. lb). Mean values for wet body mass, SVL and tail length were all smaller in hatchlings incubated at 32°C than in hatch- lings incubated at the other four temperatures (Fig. la, c, d). Hatchlings from different incubation temperatures differed in carcass mass (ANCOVA with the initial egg mass as the covariate: F474 = 6.36, p < 0.001; Fig. 2a) and yolk sac dry mass (ANOVA: F475 - 4.79, p < 0.01; Fig. 2b), but not fatbody dry mass (F474 = 0.37, p > 0.05; Fig. 2c). Hatchlings from 32°C had lighter car- cass dry mass but larger residual yolk sac, whereas hatchlings from 22°C had heavier carcass but smaller residual yolk sac (Fig. 2a, b). Energy contents (F4 74 = 2.78, p < 0.05) and non- polar lipids (F474 = 2.80, p < 0.05) differed among hatchlings from different incubation temperatures, with hatchlings incubated at 22°C containing lower quantities of energy and non-polar lipid than did those incubated at other four temperatures (Table 2). Conversion efficiency of energy during incubation at 22°C (71.1%) was thus lower than those at 32, 30, 27, and 24°C (83.9%, 82.6%, 81.5%, and 80.2%). Similarly, conversion effi- ciency of lipid at 22°C (55.3%) was lower than those at 32, 30, 27, and 24°C (70.9%, 69. %, 67.3% and 64.9 %). Additionally, shells of hatched eggs were significantly lighter than shells of freshly laid eggs (F5 84 = 3.19, p < 0.05; Table 2). Discussion As in numerous other reptiles, thermal environments experienced by developing embryos affect hatching suc- cess, incubation length, embryonic expenditure of ener- gy, and linear dimensions (SVL and tail length) and body composition of hatchlings in Elaphe taeniura. Our results provide support for the previous conclusion that reptilian embryos developing at relatively low or moder- ate temperatures produce well developed and thus, larg- er hatchlings (e.g. Deeming and Ferguson, 1991; Du and Ji, 2002; Ji and Du, 2001a, b). The larger hatchling size has an association with the greater carcass dry mass (Chen and Ji, 2002; Du and Ji, 2002; Ji et al., 1997; Ji and Sun, 2000). Data from the current study proved that this conclusion is also true in E. taeniura (Fig. 1; Fig. 2). The finding that more yolks remain unutilized at hatch- ing when eggs are incubated at high temperatures has been reported for nearly all reptiles studied to-date (e.g., Beuchat, 1988; Du and Ji, 2002; Ji and Du, 2001a, b; Lin et al., 2005; Phillips et al., 1990; Phillips and Packard, 1994). In E. carinata (Ji et al., 1997), E. taeniura (Ji et al., 1999a), D. rufozonatum (Ji et al., 1999b) and P. kor- ros (Ji and Sun, 2000), hatchlings exhibit a substantial increase in size (SVL) during the first post-hatching days due to the transfer of resources in the residual yolk into carcass. The proportion of lipids transferred from the egg contents into the hatchling was noticeably lower than those of energy. Given that mass-specific energy density is much higher in lipids than in proteins and carbohy- drates, this result provides evidence that lipids are the main source of energy for embryonic development. 28 Asiatic Herpetological Research, Vol. 11 2008 Hatchling size and mass were primarily determined by the embryonic expenditure of energy during incubation after removing the influence of variation in initial egg mass (Du and Ji, 2001, 2003; Ji and Du, 2001a, b). Given that the total energy invested in an egg is fixed, any increase in embryonic expenditure of energy during incubation may inevitably result in production of small sized or lighter hatchlings. Incubating eggs at low tem- peratures or high temperatures may increase energy expenditure for embryonic development due to the increased incubation length and/or embryonic metabo- lism (Booth, 1998; Booth and Astill, 2001; Packard and Packard, 1988). Consequently, eggs incubated at moder- ate temperatures usually produce larger and heavier hatchlings than did those at low or high temperatures. A prolonged exposure of eggs of E. taeniura to tem- peratures lower than 24°C or higher than 30°C may have a detrimental effect on embryonic development, as indi- cated by the fact that hatching success decreases dramat- ically at temperatures outside this temperature range (Table 1). The mean incubation length 30°C is 53.9 days, approximately 3.4 days longer than that at 32°C, so the ecological disadvantage of the increased incuba- tion length (and thus, decreased growth period prior to the onset of the first winter) due to a decrease in incuba- tion temperature from 32°C to 30°C is less pronounced. Considering that less developed hatchlings are produced at 32°C and that hatching success is low at this temper- ature, we conclude that the temperature of 32°C is out- side the range of optimal temperatures for incubating eggs of E. taeniura. Eggs incubated at 24°C exhibit high hatching success and produce well developed hatch- lings. However, the majority of hatchlings from eggs incubated at 24°C appear between late September and mid-October, so the growth prior to the onset (late November) of the first winter for these hatchlings is about 1.5-2 months. As incubation length increasingly increases as temperature decreases in E. taeniura (Table 1 ), we expect that the disadvantage of incubating eggs of E. taeniura at temperatures lower than 24°C can be very pronounced due to the increasingly prolonged incuba- tion length. For eggs of reptiles incubated under natural conditions, the prolonged incubation length at low tem- peratures also increases exposure of eggs to the effects of adverse biotic (increased microbial contamination) and abiotic factors(extreme thermal and hydric condi- tions) in the incubation environment of the eggs, which potentially reduces hatching success. Thus, the tempera- ture of 24°C is suitable but not optimal for incubating eggs of E. taeniura. Taking the energy expenditure dur- ing incubation, the rate of embryonic development and hatchling phenotypes into account, we consider that temperatures around 27°C are optimal for incubating eggs of E. taeniura. Acknowledgments This work was supported by a grant from Zhejiang Provincial Natural Science Foundation. Many thanks were given to X.-Z. Hu, W.-Q. Xu, and J.-Q. Du for their assistant in the laboratory. Literature Cited Andrews, R. M. and B. R. Rose. 1994. Evolution of viviparity: constraints on egg retention. Physiological Zoology 67: 1006-1024. Andrews, R. M., C. P. Qualls and B. R. Rose. 1997. Effects of low temperature on embryonic develop- ment of Sceloporus lizards. Copeia 1997: 827-833. Beuchat, C. A. 1988. Temperature effects during gesta- tion in a viviparous lizard. Journal of Thermodynamic Biology 13: 135-142. Birchard, G. F. 2004. Effects of incubation temperature. 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Further studies of egg incu- bation on red-banded wolf snakes Dinodon rufo- zonaturn , with comments on influence of hydric environments. Acta Zoologica Sinica 48:51-59. Zhao, E. M. and K. Adler. 1993. Herpetology of China, Ohio: Oxford. 235 pp. Zhao, E. M. 1998. China red data book of endangered animals: Amphibia and Reptilia. Beijing: Science Press. 245-247 pp. Submitted: 1 7 November 2006 Accepted: 22 September 2007 2008 Asiatic Herpetological Research, Vol. 1 1 pp.31-38 Behavioral Observations and Descriptions of the Endangered Knobby Newt Tylototriton wenxianensis and Their Application in Conservation Da-Jie Gong* and Mai Mu Department of Biology, College of Life Science, Northwest Normal University Lanzhou, Gansu, China. * Corresponding author E-mail: gongdj@nwnu.edu.cn Abstract.- Tylototriton wenxianensis is an endangered species at the status of VU. This paper introduces observations and studies including breeding, territorial, communication and antipredatory behaviors. The rules and mechanisms of the behaviors are recorded separately and described in the paper. Relationships between the behaviors and environ- ment are also explained tor the purpose of describing how the environment acted on the behaviors and how the behav- iors adapted to the environment. The paper ends with conservation plans in connection with behavioral ecology: A) protect the particular habitat and avoid anthropogenic threats, B) artificial construction for natural migration and gene communication and C) artificial breeding and re-introduction into nature. Keywords.- Observations, Tylototriton wenxianensis , behaviors, habitats, conservation. Introduction Tylototriton wenxianensis (Caudata: Salamandridae) is peculiar in China and poorly known. It is only found along the boundary between Gansu and Sichuan Province, narrowly distributed in Wenxian, Qingchuan and Pingwu (details see Table 1 and Fig. 1). It was defined as a threatened species with the status of VU (IUCN, 2006). A representative of the Family Salamandridae, T. wenxianensis has rough and toxic skin with seasonal colors. The bilateral warts stay longitudinal and clus- tered in the same size and the boundaries between them are not clear enough. It is dark all over apart from the red-orange fingers, toes and the venter of the tail (Fei, 1993). The newt lives in the heavily forested mountains at approximately 940 m. The adult is terricolous and usu- ally wanders about the pool. It stays hidden under wet gravel or small muddy caves covered by fallen leaves in the daytime and appears at night for preying (Gong and Mou, 2006). Behavioral studies on tailed amphibians have recently been reported. The studies were mainly focused in the following fields: breeding behaviors, including sexual recognition and sexual selection (Dawley, 1984), modes of courtship and mating (Arnold et al., 1972, 1977; Salthe, 1967, 1974), sperm competition (Arnold, 1977; Halliday, 1998; Massey, 1988; Sparreboom, 1995; Verrel et al., 1998), reproductive behavior (Fang, 1984; Harris et al., 2002; Park et al., 2000), parental care (Cramp, 1995; Nussbaum, 1985; Peterson, 2000), evolu- tion of reproduction (Arnold, 1977; Veith, 1998; Verrell and Krenz, 1998,); migratory behavior (Amtzen, 2002; Douglas, 1979; Griffiths, 1996; Serbiolova, 1995; Twitty, 1966); territorial behavior (Mathis, 1991; Mathis et al.,1998; Mathis et al.,2000; Simons et al., 1997); communications (Holliday, 1997; Houck, 1988; Houck and Sever, 1994; Rollmann, 1999; Verrell, 1989,1989a;); antipredatory behavior (Brodie, 1990; Graves and Quinn, 2000; Maerz et al.,2001; Sih et al., 2000, 2003; Storfer and Sih, 1998; Sullivan, 2002; Woody and Mathis, 1997). All the above provide a source of refer- ence for the behavioral study on T. wenxianensis. Methods Migratory behavior.- We chose the habitat in Qingchuan as the site to observe, where a large population was found (Gong and Mou, 2006). The newts are not active year round except for the breeding season, so their migratory behaviours seem quite obvious and clear. We © 2008 by Asiatic Herpetological Research 32 Asiatic Herpetological Research, Vol. 1 1 2008 Date Figure 2. Reproductive migration of Tylototriton wenxianensis. The polyline represents temperature, columns repre- sent number of the migratory newts and “ * ” rainy days. made continuous observations when they migrated for breeding towards the pools nearby. We marked each migratory group with varisized loops and observed by radio tracking and GPS. Finally we made records of the time, route length, size of groups and sex ratio in the migration. Reproductive behavior.- Reproductive behaviors included mating and spawning. We observed the course and took down the data of sex ratio, time, envioronment and quantity of the spawns etc. Conmmunication.- Communication happens along with courtship. We tried to understand how the newts com- municate with each other and why. The communications included chemical signals and body-contacts (Jiang, 2004). Mechanisms of chemical signals from glands are expected to be understood through dissection. Types of the body-contacts and their effects on courtship were studied. Territorial behaviors - We measured the size of the ter- ritory occupied by 6 female newts and size of the tails and bodies in contrast. Aggressive behaviors related to the available prey in the environment were also observed. Relationships between the data and phenome- non were discussed and concluded. Antipredatory behavior.- The newts were less aggres- sive without structures for aggressivity (Jiang, 2004). They moved too slowly to escape attacks from enemies and to defend themselves. We made some model ene- mies and demonstrated the predation and antipredatory behaviors artificially. Results and Analysis Behaviors.- We made continuous observations on the migratory behaviors and found 186 adults including 170 females and only 16 males, the ratio was 170:16 ($:$) = 10.625:1. Breeding lasted from early April to late May with a peak from April 8th to May 6th. The average tem- perature was 15.5±4°C (Fig. 2). Like many other newts, T .wenxianensis does not seem to have significant secondary sexual characters. It distinguishes and selects the opposite sex primarily by sight and sense of smell. The male seemed more active. It courted the female by wagging its tail and dancing round. Meanwhile, the female was secreting chemicals and emitting a strong smell to attract the male. Soon the male bit the female on the tail (or hind limb), then they circled around both with their tails wagging (Fig. 3). Experiments show that the male prefers to choose a big- ger one with more ovums and a stronger smell. Mating did not accompany spawning in the mean- time. It was the female that selected where to spawn. The male moved to the pool and settled down earlier than the female. However, the female left soon after spawning, while the male stayed untill the end of the Figure 3. Model of mating behavior. Shows the male (black) wagging its tail and dancing round. 2008 Asiatic Herpetological Research, Vol. 1 1 33 Table 1 . Distribution and flora of T. wenxianensis along the boundary between Gansu and Sichuan Province. Province City Country (town), county Village Latitude and longitude Altitude (m) Flora Gansu Longnan Bikou, Wenxian Bifenggou 103. 7°E 33.95°N 927 semitropics Gansu Gansu Longnan Longnan Liziba, Wenxian Liziba, Wenxian Moziping Hanjialiang 104.6°E 33.2°N 104.4°E 33.1°N 940 1100 semitropics- intersemitropics semitropics- intersemitropics Sichuan Guangyuan Daba, Qingchuan Wuxing 105. 1°E 32.5°N 1160 semitropics- intersemitropics Sichuan Guangyuan Daba, Qingchuan Dawuji 105.1 °E 32.46°N 998 intersemitropics Sichuan Guangyuan Daba, Qingchuan Laowuji 105°E 32.68°N 1092 semitropics- intersemitropics Sichuan Guangyuan Daba, Qingchuan Baishuling 105.14°E 32.32°N 1210 intersemitropics Sichuan Mianyang Shuitian, Pingwu Dafenling 104.23°E 32.58°N 1134 semitropics- intersemitropics Sichuan Mianyang Bazi, Pingwu Yinshanli 104.3°E 32.44°N 962 intersemitropics Sichuan Mianyang Bazi, Pingwu Miaoshanli 104.34°E 32.45°N 970 intersemitropics Figure 4. The sperm transmission. Swells show the col- loidal secretion. Figure 6. Rigid body: the reaction to anti-predator. Shows ribs jutting through the side warts. Figure 5. The courtship behavior. Shows the transmis- sion of pheromones in the fantail. Figure 7 The raised tail and warning color on the ventral surface of the tail. 34 Asiatic Herpetological Research, Vol. 1 1 2008 Figure 8. The protuberant glands and warning color on the back. Shows harmful and toxic skin secretion, glan- diform warts, spine glands, caudal glands, glandiform warts, spine glands, caudal glands, tergal glands, etc. Figure 10. Mimicry: newt playing dead, with venter upward and body rigid. v Figure 9. Playing dead. Shows the newts played dead with upward bottom and rigid body. Figure 11. The breeding pool (after spawning). Vicinity of Wuxing Village, Sichuan Province, southwest China. Table 2. Relation between spawning territory, food and size of six female newts. No. Areas of spawning territory (m2) Percent of total (%) TL (mm) SVL (mm) Food environment i? 0.64 10.39 58 71 poor 2? 0.81 13.15 59.2 72.2 poor 3? 0.55 8.93 55.7 70.1 poor 4? 1.26 20.45 65.9 73.2 medium 5? 2.3 37.34 67.4 74.6 high 69 0.6 9.74 56.1 71.4 poor Table 3. The antipredatory behaviors of T. wenxianertsis, grouped by different types. The observations were made in the field and in artificial conditions, a. harmful skin secretion; b. toxic skin secretion; c. parotids; d. glandiform warts; e. spine glands; f. caudal glands; g. tergal glands; h. tergal colors; i. ventral colors; j. rigid body; k. roll of body; I. exposed abdomen; m. raised tail; n. raised jaw; o. arched body; p. larger head; q. swagging tail; r. curved forehead; s. extended nerve. Types Imitative toxicant body Warning colors Warning posture S Pc s articular