t HARVARD UNIVERSITY Ernst Mayr Library of the Museum of Comparative Zoology Volume 9 • 2001 . MCZ LIBRARY JUL. 2 jj 2002 HARVARD UNIVERSITY I i mi Zhao Chengdu Institute of Biology. Academia Sinica, Chengdu. Sichuan. China Associate Editors Ki ll.ii Autumn Lewis & Clark College, Portland. Oregon. USA Theodore J. Papenfuss Museum of Vertebrate Zoology. University of California. Berkeley. California. USA J. Robert Macey James Ford Parham Department of Biology. Washington University. St. Louis. Museum of Paleontology. University of California. Missouri. USA Berkeley. California. USA Editorial Board Kraig Adler Cornell University. Ithaca. New York. USA Natalia B. Ananjeva Zoological Institute, Si. 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 Bihui Chen Anhui Normal University, Wuhu. Anhui. China I .|iiiiiii Cheng Xiang Ji Hangzhou Normal College. Hangzhou. Zhejiang, China Pi-peng Li Yantai Normal College, Yantai. Shandong. China Ronald Marlow University of Nevada. Las Vegas. Nevada. USA Robert W. Murphy Royal ( )ntario Museum. Toronto. Ontario. Canada Giiren NiLson University ol Goteborg. Goteborg. Sweden Nikolai Orlov Zoological Institute, St Petersburg, Russia Hidetoshi Ota Department of Biology, University of the Ryukyus. Nishihara, Okinawa. Japan Soheila Shafii Institute of Marine Biology. National Taiwan Ocean University. University of Shahid Bahonar. Kerman. Iran Keelung, Taiwan. China Ilya Darevsky Zoological Institute. St. Petersburg. Russia Indraneil Das Madras Crocodile Bank, Vadanemmeli Perur. Madras. India William K. Duellman University of Kansas. Lawrence. Kansas. USA Hajime Fukada Sennyuji Sannaicho. Higashiyamaku. Kyoto. Japan Carl Gans University of Michigan. Ann Arbor. Michigan. USA Robert F. Inger Field Museum. Chicago, Illinois, USA Hai-tao Shi Hainan Normal University. Haikou. Hainan. China \iu-ling Wang Xinjiang Normal University. Urumqi. Xinjiang. China Yue-zhao Wang Chengdu Institute of Biology. Academia Sinica. Chengdu. Sichuan. China Yehudah Werner Hebrew University, Jerusalem. Israel Ken-tang Zhao Su/hou Railway Teacher's College, Suzhou. Jiangsu. China Asiatic Herpetological Research is published by the Asiatic Herpetological Research Society (AHRS) and the Chinese Society for the Study of Amphibians and Reptiles (CSSAR) 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 outside of China and requests for subscription should be sent to AHR. Museum of Vertebrate Zoology. University of California. Berkeley. California. USA 94720. or by email to asiaherp@uclink.berkeley.edu. All correspondence within China should be sent to Ermi Zhao. Editor. Chengdu Institute of Biology. P.O. Box 416, Chengdu. Sichuan Province. China. Authors should consult Guidelines for Manuscript Preparation and Submission at the end of this issue. Subscription and membership are $25 per year ($45 for libraries). Postage outside of the USA and China, please add $5 per issue for surface mail or $10 per issue for air mail. Make checks or money orders payable in US currency to AHRS. If you do not have access to US currency, please notify us. and we will make other arrangements. Asiatic Herpetological Research Volume 9 succeeds Volume 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- 1 989 as the journal for the Chinese Society for the Study of Amphibians and Reptiles. Volume 2 succeeded 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. Cover: Turtles of Turkey. Upper left: Trionyx triunguis. Dalyan. 6/16/00. Upper right: Tesnuio graeca anamurensis. Anamurium. Anamur, 6/1 1/00. Lower left: Mauremys caspica rivulata Mamure Kalesi. Anamur. 6/10/00. Emys orbicularis. Lower right: Mamure Kalesi. Anamur. 6/10/00. All photographs by Tonya Van Leuvan-Smith. 2001 Asiatic Herpetological Research Vol.9, pp. 1-5 The Reproductive Biology of Rana boulengeri Wen-Jian Li Laboratory of Economic Frogs, Changde Teachers College, Changde, Hunan, 415000 China Abstract.- Maturation of Rana boulengeri Giinther occurs at the age of 24 to 36 months. After reproduction the gonads are in an intermittent period. Gonads begin to develop rapidly from next February to April. After May, the ovocytes grow mature in batches. In the wild, the adult frogs often spawn under waterfalls or in shallow pools where the water flows slowly, with high dissolved oxygen. The reproductive period lasts from May to August. The ovulation time of females and fertilization time are generally from 5:00 a.m. to 8:00 a.m. The Water temperature for deposition at the beginning is 15.5°C and the appropriate temperature for most deposition is 17.7-22.5°C. The total duration under artificial constant temperature of 23±0.5°C lasts 261 hours. The appropriate temperature for development is 22-24°C. This species can not develop below 4°C, and it dies at 30°C. The development of the ovary is divided into 6 stages and the development of the testis is divided into 5 stages. The morphological and histological studies at different stages of sex cells are described. Key words.- Amphibia, Rana boulengeri, reproductive biology, China Introduction Rana boulengeri is a large frog that is found in the Wuling Mountains of Hunan Province. It is an impor- tant species for maintaining the ecological balance of the region (Yuan and Wen, 1990). Besides, it can be used as a kind of medicine (Li et al., 1993). In order to help protect this natural resource, these studies on its reproductive biology have been carried out. Material and Methods Laboratory experiment and artificial breeding The studies on the natural condition of habitat, growth and ages, characteristics of reproduction, the develop- ment of genital gland and development of the embryo of Rana boulengeri was conducted for four years from 1989 to 1993. Field investigation The field sites chosen for investigation were places where Rana boulengeri are common. Field observa- tions were made two or three times a month. Results and Discussion Habitat Natural habitat of Rana boulengeri. The Xiang Xi mountains are a part of the Wuling Mountains, which are located on the border of Hunan, Hubei, Sichuan and Guizhou; between north Latitude 27°44'- 29°48' and east longitude 109°10'-1 H"20'; with a mean ele- vation of 686 meters (the highest, 1900 meters above sea level; the lowest, 75 meters above sea level). The mountains are covered with evergreen broad-leaf sub- tropical forest. The mean air temperatures for a whole year are 16.10 degree-days. The temperature summa- tion above 10°C is 50320; the mean sun hours per year are 1292. 7h; frostless period is 270.6 days; the mean rainfall per year is 1397.2mm; the mean humid- ity per year is above 82% (R.H). These conditions constitute an ideal environment for Rana boulengeri. Rana boulengeri frequently live in brooks or ponds in mountain forests where there are few people. The water in the brooks or ponds is very clear. During the day, R. boulengeri often hide at the edge of small caves and during the night, they go out to feed. From July to September of 1987, the physical factors in 35 observation points were measured. Their values are: the elevation of the habitat is 204-675 meters above sea level; air temperature 17.2-24°C; water tempera- ture 14.9-20.5°C; the depth of water 0.2-1.0 meters; the transparency of water 0.2-1.0 m; the humidity 92.0-97.5%. Activities of Rana boulengeri and temperatures of habitat. Because the air temperature and water tem- perature play important roles in the development and growth of R. boulengeri; we observed their relation- ship in a cave (450m above sea level). In autumn, when the water temperature fall lower than 12.5°C, R. boulengeri begins hibernation. In spring, when water temperature is above 12.5°C, they start to feed, and when water temperature is higher than 15.5°C, they enter the reproductive period. The air temperatures and water temperatures in all months are listed in Table 1. Vol. 9, p. 2 Asiatic Herpetological Research 2001 Table 1 . Air and water temperatures of the habitat of Plana boulengen (450m above sea level). Month Mean air temperature Mean water temperature 1 8.1 11.0 2 10.0 12.2 3 10.8 12.8 4 13.0 14.4 5 14.6 14.9 6 16.3 16.1 7 23.9 16.5 8 22.1 16.3 9 18.1 16.1 10 16.1 12.9 11 13.1 11.5 12 8.2 10.0 Chemical properties of the water. The chemical characteristics of the water inhabited by R. boulengen were measured and analyzed. The results are listed in Table 2. On the whole, the water is clear, thin, neutral and acidulous, with some contents of Ca, Mg and other minerals. Vertical distribution. Rana boulengeri are distrib- uted vertically from 200 to 700 meters above sea level, but most of them are found from 450 to 650 meters. Older, large adult frogs are mostly found above 600 meters, while young frogs and tadpoles are found lower. Growth and age When bred in artificial pools, the mean body length and the mean body weight of the frogs just complet- ing metamorphosis are 19.8 mm and 0.95 g respec- tively. When feeding lasts to the end of the year, their body weight increases to 3.07 g and their body length to 31.2 mm. After feeding for 2 years, their body weight grows to 20.50 g, and the body length to 58.1 mm. The growth curves of body weight and body length are logistic (i.e. "S" shaped) (Li, et al., 1993). In the field, it was found that the largest male was 261 g with a body length of 134 mm. The largest female was 202 g with body length of 120 mm. Reproductive characteristics Environmental conditions of breeding areas. In the wild, the adult frogs often spawn under waterfalls or in shallow pools. The mean water area and mean water depth of natural spawn areas are 2.16 m" and 0.35 m respectively. The water flows slowly with high dissolved oxygen. The mean value of pH is 6.3. Gen- erally, there are small stones, sand, ratty plant and leaves, or humus on the bottom. There are bryophytes and algae on the bottom as well as on the sides of the pools. Reproductive period. In west Hunan at 563 m above sea level, R boulengeri begin reproduction in May and end in August. The reproductive peak is in June and July. Female frogs may have three clutches, the egg number of the second and third clutches depend on the availability of food and environmental conditions. Temperature. When water temperature reaches 15.5°C R. boulengeri begins to spawn. The optimum temperatures for large numbers of females to spawn are 17.7-22.5°C. From July to August, the mean air temperature of natural spawning sites is 21.4°C, and the optimum humidity is above 95%. Reproductive behavior. In middle or late March, R. boulengeri end their hibernation and feed for a month. Then the male and female frogs aggregate at spawn- ing fields. Before mating, male frogs call for about 15 days as a courting period. The ovulation time of females and fertilization times are generally from 5:00 a.m. to 8:00 a.m. When the female spawns, she creeps slowly and her cloaca is near the walls of pool or brooks. Table 2. Analysis of the water inhabited by R. bou/engeri(mglL). Date of analysis: 5, January, 1990; elevation where water was sampled: 204m above sea level. PH total hardness Ca Mg Cu Zn Fe Hg Mn Cd 6.71 12.92»C 58.36 13.66 0.013 0.183 0.793 0.001 0.036 0.003 Pb Ag Co Cr Do Sulfate N(N02) N(HN03) N(NH4) Water type 0.0148 0.002 0.017 0.0199 4.45 9.89 0.001 0.026 0.075 Thin acidulous 2001 Asiatic Herpetological Research Vol. 9, p. 3 > o •B 7 9? o e a> o o _ * $&■* Figure 4. (A) Dorsal and (B) lateral view of the head of Hologerrhum dermali :(CMNH 5075); (C) dorsal and (D) lat- eral view of the head of Hologerrhum phi/ipp/num (PUM 2490). 2001 Asiatic Herpetological Research Vol. 9, p. 13 Table 1 . Summary of diagnostic characters distinguishing Hologerrhum dermal! {new species; from Panay Island) from Hologerrhum philippinum (Gunther, 1858; from the Luzon Aggregate Island Complex). Characters H. philippinum H. dermali Pretemporals Ventral nuchal blotches enlarged -, + reduced or absent Lateral black ventral spots -, + +, fused into line Dorsal nuchal spots 12-30 7-10 Midlabial stripe - + Dorsal live color tan to reddish brown purplish brown Ventral live color orange to salmon bright yellow Adult throat color tan to orange (some spotted white) purplish brown w ocelli Midventral dark stripe - + Parietal suture notched unnotched Ventral scale at 2nd dorsal reduction 97-112* 84-97 Subcaudals (females) 42-53 57-61 * Excluding apparently aberrant counts of CAS 61554 (Table 2). suture between pre- and postnasal; together, nasal scales square to triangular; dorsal border of nares formed by thick shelf of prenasal, ventral border formed by extension of postnasal; internasals as long as broad, slightly shorter than prefrontals, laterally contacting both pre-and postnasals, forming a vague right triangle with 45° face oriented anterolaterally; loreal single, as large as or only slightly smaller than ventral preocular, half as high as postnasal, pentago- nal, surrounded by postnasal, lateral edge of prefron- tal, dorsal and ventral preoculars, and second supralabial; prefrontals longer than internasals, with irregular lateral extensions caused by presence of con- cave curved suture with preoculars (concave surface oriented posterolaterally); frontal twice as long as broad, longer than to equal to its distance to the end of the snout, shorter than parietals; anterolateral corner of frontal barely contacting medial point of preocular (e.g., CAS 31553, 60951, MCZ R-25693-94, PNM 6505), or with substantive contact between frontal and preocular squeezed off by contact between supraocu- lar and prefrontal (e.g., CAS 60950, 61554, 62430, 134075; PNM 2120, 2490; USNM 498718, MCZ R- 25695); posteromedial point of frontal extends past posterior margin of supraoculars for distance shorter than or equal to length of internasals; supraoculars very large, nearly as long as and slightly narrower than frontal; parietals very large, laterally contacting dorsal postocular, pretemporal, and highly enlarged secondary temporal, together bordered posteriorly by three (CAS 61554, 134075), four (CAS 60950, 61553; USNM 498718, MCZ R-25695) or five (CAS 60951, 62430; PNM 2490, MCZ R-25693-94) undif- ferentiated nuchals; posterior tips of parietals extend caudally, parietal suture forming a distinct medially inverted V-shaped cleft, in which a single slightly enlarged to undifferentiated nuchal (Fig. 6) lies; tem- porals arranged in three to four irregular vertical rows with enlarged posttemporals extending caudally beyond posterior ends of parietals; temporal formu- lae: (L) 2/1 + 1/1+2, (R) 1 + 1 + 1/1+2 (PNM 2490), (L, R) 1+2+1/1+2 (CAS 60950), (R, L) 1 + 1+2+3 (MCZ R-25695), (L) 1 + 1+2+3, (R) 1 + 1+2+2 (CAS 60951), (L) 1+1+2+3, (R) 1 + 1+2+4 (MCZ R-25694) (L) 1 + 1 + 1/1+2, (R) 1 + 1+2+3 (CAS 61554, PNM 6505), (L, R) 1+1+1/1+2 (CAS 62430, 61558, 134075; USNM 318363, 498718; TNHC 60114, MCZ R- 25693, PNM 2120), (L) 1 + 1+2+3, (R) 2+1/1+2 (USNM 319037), (L); pretemporal relatively enlarged, its length much more than half that of sec- ondary temporal. Orbit surrounded by supraocular, two preoculars (dorsal larger than ventral), two postoculars, and supralabials 3-5; supralabials eight, fifth largest; Vol. 9, p. 14 Asiatic Herpetological Research 2001 Table 2. Scale row reduction formulae (Dowling, 1951b) variation in H. philippinum philippinum (Gunther, 1858; from the Luzon Aggregate Island Complex) and the type series of H. derma// (new species; from Panay Island). Specimen (sex) Reduction 1 Reduction 2 Hologerrhum phillippinum 3+4=3(4) CAS 6095 l(juv.) 19 17 4+5=4(5) 3+4=3(107) 17 15 3+4=3(106) 4+5=4(5) CAS 60950 (juv.) 19 17 3+4=3(4) 3+4=3(103) 17 15 3+4=3(105) 3+4=3(7) CAS 61553 (0 19 ' 17 4+5=4(5) 3+4=3(109) 17 15 3+4=3(112) 3+4=3(7) CAS 61554(0 19 17 4+5=4(6) 3+4=3(93) 17 15 3+4=3(95) +4(96) 3+4=3(100) 3+4=3(7) CAS 62430 (juv.) 19 17 3+4=3(6) 3+4=3(101) 17 15 3+4=3(103) 3+4=3(16) PNM 2490(0 19 17 5+6=5(5) 3+4=3(110) 17 15 4+5=4(109) 5+6=5(5) PNM 2120(0 19 17 5+6=5(6) 3+4=3(95) 17 15 -4(99) 3+4=3(4) PNM 2120 (juv) 19 n 3+4=3(4) 3+4=3(6) USNM 3 19037 (0 19 17 3+4=3(5) 3+4=3(97) 17 15 -4(96) 3+4=3(105) 17- 3+4=3(109) -15 3+4=3(5) USNM 318363(0 19 17 3+4=3(7) 3+4=3(100) 17 15 3+4=3(99) 3+4=3(5) USNM 498718 (m) 19 17 3+4=3(5) 3+4=3(5) TNHC 60114(0 19 17 3+4=3(8) -4(100) 17 15 ^4(105) 3+4=3(102) 17- -4(100) -15 4+5=4(4) MCZ R-25693 (0 19 17 3+4=3(4) 3+4=3(100) 17 15 3+4=3(100) 2001 Asiatic Herpetological Research Vol. 9, p. 15 Table 2. (continued) Specimen (sex) Reduction 1 Reduction 2 4+5=4(4) MCZR-25694(0 19 17 4+5=4(4) 3+4=3(102) 17 15 3+4=3(103) 5+6=5(8) MCZ R-25695 (juv) 19 : 17 5+6=5(6) 3+4=3(97) 17 15 3+4=3(100) Mean x = 6.3 ± 3.2 SD; n = 12 x =5.5±1.2SD;n=12 x = 102.2 ± 4.9 SD; n = 12 x =103.6 ±4.9SD;n=12 Hologerrhum dermali 3+4=3(5) PNM2711H(f) 19 17 3+4=3(6) 3+4=3(94) 17 15 3+4=3(92) -4(3) PNM 6505 p (f) 19 17 3+4=3(6) -4(97) 17 15 -4(92) 3+4=3(5) CMNH 5075 p (f) 19 17 3+4=3(7) 3+4=3(84) 17 15 3+4=3(84) Mean x =4.3±1.2SD;n = 3 x =6.3 ±0.6SD;n=12 x =91.7±6.8SD;n=12 x. =89.3±4.6SD;n = 12 " = Holotype; p = Paratype infralabials eight; mental subtriangular, with highly pointed posterior tip caused by medially concave curved suture with first infralabial; first infralabials differentiated, elongate, with curved medial points nearly contacting anterior to genials; second infralabi- als reduced, squarish; infralabials 2-5 increasing dra- matically in size (fifth largest in verntral aspect), then decreasing sharply to infralabial 8; infralabials 1^ in contact with anterior genials, 4—5 in contact with pos- terior genials; mental groove distinct and broad; sub- labials thin, followed medially by 3^4 similarly-sized, longitudinal rows of gulars, medial two pairs (anterior to first ventral) slightly enlarged; number of gular pairs between posterior genials and first ventral two (CAS 61553) or three (remaining specimens). Dorsals smooth, without apical pits, vertebrals undifferentiated from paravertebrals, in 146-176 (X- 157.4 ± 8.1 SD; n = 12) transverse rows on body, 42- Vol. 9, p. 16 Asiatic Herpetological Research 2001 Figure 5. (A) Dorsal and (B) ventral view of a paratype of Hologerrhum dermalt '(CMNH 5075); (C) dorsal and (D) ventral view of Hologerrhum philippinum (PNM 2490). 56 (JC = 49.3 ± 5.3 SD; n = 12) on tail; first scale row reduction (i.e., reduction of 19 to 17 scales around body; Table 2) occurring at point on body correspond- ing to ventrals 4-16 (left: X = 5.5 ± 1.2 SD; n = 12; right; X = 6.3 ± 3.2 SD; n = 12), second (17 to 15) occurring between ventrals 93-1 10 (left: X - 103.6 ± 4.9 SD; n = 12; right X = 102.2 ± 4.9 SD; n = 12); ventrals broad, each slightly angulated laterally, 136— 158 (X= 146.7± 6.4 SD; n = 12); subcaudals 42-56 (x = 47.5 ± 4.6 SD; n = 12). The single adult male specimen (USNM 498718) has 156 vertebrals, 149 ventrals, 56 caudals, and 55 subcaudals. Anal undi- vided; tail with enlarged vertebral row (dorsocaudals) formed by fusion of midvertebral row with both flank- ing paravertebral rows. Hemipenes of USNM 498718 are extremely narrow and elongate, and are covered with uniformly minute spines; hemipenes extend in situ to the 14th subcaudal plate. Measurements (in mm): SVL 251-347 mm (X = 280.8 ± 47.1 SD) for ten mature females; tail length X = 56-96 (X = 73.8 ± 15.0 SD) for eight mature females with complete tails. Coloration in preservative: Dorsum tan, orang- ish-tan to brown, with 12-30 (X = 21.8 ± 7.5 SD; n = 12) alternating dark brown to black spots (Figs. 1A, 3, 5C), each with three associated small white spots (Fig. 3), fading in intensity posteriorly, where they are replaced on scale rows 4-5 by a pair of dorsolateral black lines, gaining intensity posteriorly and continu- ing to tip of tail; faint vertebral thick gray stripe (1-3 scales in width) becoming increasingly apparent pos- teriorly from midbody; a pair of light cream lines dor- sal (medial) to black lines; posterior (distal) portions of each dorsal scale slightly to markedly darker than remainder of scale; dorsal occiput colored as body (PNM 2490) or slightly darker (CAS 60950) to mark- edly darker (CAS 134075); melanic pigment congre- gated on medial suture between parietals, on posterior portion of frontal, and on lateral edge of head; distinct longitudinal dark midnuchal stripe evident from pos- terior edge of parietals to second pair of nuchal spots (Fig. 1A), occasionally (e.g., juveniles CAS 134075 and MCZ R-25695, adult female USNM 318363) very dark and forming a distinct nuchal cross (Fig. IB); one specimen with a pair of bright white nuchal spots immediately anterior to nuchal cross (USNM 319037; Fig. 1A); lateral aspects of head colored as dorsal, with distinct thin black line dorsally bordering supralabials (Fig. 4D) and stretching from tip of snout to just beyond supralabial 8; labials creamy yellow to tan, occasionally with a few black flecks (CAS 60950) or with ventral half of labials dark gray (USNM 319037); venter immaculate cream to pale yellow or orange; each ventral with dark lateral pigment in the form of a small black spot or brown to black longitu- dinal bar (Figs. 1A, 3), becoming a confluent black ventrolateral stripe on posterior portions of body and tail; some specimens with subtriangular black mark- 2001 Asiatic Herpetological Research Vol. 9. p. 17 > \ — ^ ' / \ / \ I \^ / / / Figure 6. Comparison of the posterior portions of the parietal suture in (A) Hologerrhum philippinum (PNM 2490) and (B) Hologerrhum dermah '(CMNH 5075). ings on anterior 20 ventrals (USNM 319037. 498718; TNHC 60114); underside of head lighter (PNM 2490). to distinctly darker (USNM 319037) than remainder of venter, especially in juvenile specimens (MCZ R-25694-95; CAS 134075) where ventral head coloration resembles that of adult Hologerrhum n. sp. (see below); tongue black with pale gray tips of bifur- cated portions. Coloration in life: (Fig. 3) Dorsum described as "bright reddish to orange brown, darker anteriorly" or "reddish brown" (Taylor 1922b:200) or "grayish brown on neck, fading to orangish brown posteriorly" (Brown et al.. 1996: 13); labials dirty white to creamy yellow; venter "uniform pale, orangish tan" (Brown et al.. 1996:13) to "bright reddish salmon, lighter anteri- orly" or "uniform coral to red" (Taylor. 1922b:200); underside of head pale orange to "dusky with milk white spots" (Taylor, 1922b:200). Hologerrhum derma//, n. sp. Figures 4-6 Holotype: PNM 271 1. an adult female, collected at 0900 hr on 9 April, 1 992 by Rafe M. Brown at 1 5 1 0 m above sea level in the area known locally as "Hang- gud Tubig" ("Big Water"), on the western face of Mt. Madja-as. Barangay Alojipan. Municipality of Culasi. Antique Province, Panay Island. Philippines (11°23' N. 122 09'E). Paratypes: CMNH 5075. an adult female, same data as the holotype except collected at 1030 hr on 6 April 1992 by Rogelio V. Sison; PNM 3704. an adult female, collected February-March 1994 by Rogelio V. Sison at 750 m above sea level on Mt. Ranges, Sitio Banagon. Barangay Aningalan. Municipality of San Remegio, Antique Province, Panay Island, Philip- pines. Etymology: The specific epithet is chosen to honor Ronald "Dermal" Crombie, in recognition of his numerous contributions to Philippine herpetology and in thanks for the guidance he has provided RMB and JWF during the past several years of our work with Philippine amphibians and reptiles. Diagnosis: Hologerrhum dermali can be readily distinguished from its congener. H. philippinum. by (1) the presence of 7-10 (vs. 12-30) pairs of dark spots in nuchal and dorsal regions. (2) a bright yellow (vs. pale orange to reddish salmon) venter, (3) pres- ence (vs. absence) of a black midventral stripe, (4) bright white labials with midlabial black stripe (vs. cream or yellow labials; midlabial stripe absent), (5) dorsum dark purplish brown (vs. tan to dark orangish brown or dark brick red). (6) chin and throat of adults dark purplish brown with black and while ocelli (vs. pale tan to orange, immaculate or with faint white spots in some specimens). (7) pretemporal absent or much reduced (length less than half that of secondary temporal: vs. pretemporal invariably present and enlarged), and (8) posterior portions of parietals squared oft. with no medial cleft at parietal suture (vs. posterior tips of parietals pointed, extending caudally to form a medial inverted V-shaped cleft). Description of the Holotype: An adult female. Body cylindrical, ventrals convex, head slightly dis- tinct from neck, not flattened; eyes small, pupil round; vertebral ridge nonevideni. Vol. 9, p. 18 Asiatic Herpetological Research 2001 Figure 7. Habitat of Hologerrhum dermalisX the type locality (following heavy rain). Rostral much broader than deep, barely visible from above, subtriangular with ventromedial groove in dorsal aspect: nasal divided, naris piercing suture between pre- and postnasal: majority of dorsal border of nares formed by prenasal. majority of ventral formed by postnasal: internasals as long as broad, slightly shorter than prefrontals, laterally contacting both pre- and postnasals, forming a vague right trian- gle with 45° face oriented anterolaterally; loreal sin- gle, distinctly smaller than ventral preocular. half as high as postnasal, pentagonal, surrounded by postna- sal, lateral edge of prefrontal, dorsal and ventral pre- oculars and second supralabial: prefrontals longer than internasals. with irregular lateral extensions caused by presence of concave curved suture with preoculars (concave surface oriented posterolater- ally): frontal twice as long as broad, longer than its distance to the end of the snout, a little shorter than parietals: frontal-preocular contact squeezed off by substantive contact between posteriolateral corners of prefrontals and anteromedial corner of supraocular: posteromedial point of frontal extends past posterior margin of supraoculars for distance shorter than length of internasals; supraoculars very large, nearly as long as and narrower than frontal: parietals very large, laterally contacting dorsal postocular, pretem- poral. and highly enlarged secondary temporal, together bordered posteriorly by five undifferentiated nuchals; posterior ends of parietals squared off. with no medially inverted V-shaped cleft at parietal suture (Fig. 6); enlarged posttemporals extend posteriorly only slightly beyond caudal margin of parietals; tem- porals (L) 1 + 1+2+3. (R) 1 + 1/1+2 (pretemporal much reduced, its length much less than half that of suture between parietal secondary temporal). Orbit surrounded by supraocular, two preoculars (dorsal larger than ventral), two postoculars, and supralabials 3-5; supralabials eight, fifth largest; infralabials eight; mental subtriangular. with highly pointed posterior tip caused by medially concave curved suture with first infralabial; first infralabials differentiated, elongate, with curved medial points nearly contacting anterior to genials; second infralabi- als reduced, squarish; infralabials 2-5 increasing dra- matically in size (fifth largest in ventral aspect), then decreasing sharply to infralabial 8; infralabials 1—4 in contact with anterior genials, 4-5 in contact with pos- terior genials; mental groove distinct; sublabials thin, followed medially by four similarly-sized, longitudi- nal rows of gulars, medial two pairs (anterior to first ventral) enlarged; two pairs of gulars between poste- rior genials and first ventral. 2001 Asiatic Herpetological Research Vol. 9, p. 19 Dorsals smooth, without apical pits, vertebrals undifferentiated from paravertebrals, in 140 trans- verse vertebral rows on body, 64 on tail; scale row reduction from 19 to 17 in nuchal region and from 17 to 15 posterior to midbody (Table 2); ventrals 143, broad, each slightly angulated laterally; subcaudals 61; anal undivided; tail with enlarged vertebrals formed by fusion of midvertebral row with both flank- ing paravertebral rows; SVL 220 mm; tail length 68 mm. Coloration in preservative: Dorsum dark pur- plish brown with 10 tightly paired black spots, decreasing in size posteriorly (Fig. 5A) on anterior one third of body; caudal third of body with a pair of dorsolateral black lines (on scale rows 4-5) gaining intensity posteriorly and continuing to tip of tail; ver- tebral stripe absent; a pair of light, bright yellowish lines dorsal (medial) to black lines, especially bright on tail; posterior (distal) portions of each dorsal scale markedly darker than remainder; dorsal occiput col- ored as body; melanic pigment congregated on medial suture between parietals, on posterior half of frontal, and on lateral edge of head; supralabials bright white, dorsal border composed of thin black stripe (Fig 4B), from tip of rostrum to beyond angle of jaw; white labial coloration continues in the form of a broad white stripe to point opposite fifth ventral; midlabial thin black stripe (Fig 4B) continues posteriorly as ventral border of the white stripe in nuchal region; distinct dark brown midnuchal stripe evident from posterior edge of parietals to first pair of nuchal spots, very dark and confluent with nuchal spots, forming a distinct nuchal cross; chin and throat purplish brown with white circular spots encircled in black (ocelli) much like juvenile coloration in H. philippinum speci- mens; venter pale yellow with midventral thin black stripe, becoming interrupted on caudal portions of body, nearly obliterated by vent and continuing again caudal to vent for five ventrals; each ventral with dark lateral pigment in the form of a small black spot (ante- riorly) or black longitudinal bar (caudally), becoming a confluent black ventrolateral stripe on caudal por- tions of body and tail; tongue black with bright white tips on forked portions. Coloration in life: Dorsum and ventral surfaces of head light purplish brown, light areas dorsal (medial) to dorsolateral caudal lines medium yellow; labials bright milky white; venter very bright yellow with distinct black midventral stripe. Iris dark brown to brick red. Variation: One paratype (female, CMNH 5075, SVL 268 mm; tail 91 mm) has seven pair of dark dor- sal spots, slightly lighter midcephalic coloration and less yellow above the dorsolateral caudal black lines. The midventral black stripe continues to the tenth subcaudal. This specimen lacks the small pretempo- rals found in the holotype; temporals (R, L) 1 + 1/1+2; ventrals 143, subcaudals 60, vertebrals 156, dorsocau- dals 64 . The other paratype (female, PNM 3704, SVL 327 mm; tail 93 mm) has nine pairs of nuchal spots, a faint midlabial line, and lacks midventral stripes on the subcaudals (present on anterior 2/3 of body). PNM 3704 has the following counts (R, L) 1 + 1+2+3, 1 + 1 + 1/1+2; ventrals 149; subcaudals 58; vertebrals 155; dorsocaudals 57. Scale row reduction formula presented in Table 2. Ecology and habitat: The type of habitat in which H. dermali (Fig. 7) was collected on Mt. Madja-as has been classified as the transition zone between mixed dipterocarp (submontane) and mossy (upper montane) forests (Whitmore, 1984; Ferner et al., 1997). The forest consisted of two strata (a canopy of 10 m, and a subcanopy of 3-4 m with emergent trees as high as 18 m); herb and shrub layer vegetation was also abundant. The forest near the collection site was mossy and contained high densities of epiphytic ferns and orchids. Topography was qualitatively char- acterized as steep, with numerous valleys bordered by sheer rock escarpments and forest-covered ridges. The holotype was collected in a sun spot in the early after- noon in a rocky stream bed (10 m wide) with a central 4 m wide channel of rapidly running water. The speci- men was basking 1 .5 m from water on the top of a flat rock. The Mt. Madja-as paratype was collected in the mid-morning and was crawling through leaf litter on the forest floor (30 m from the same stream) when captured. Paratype PNM 3704, collected in San Remegio, was found on the floor of secondary forest near a small dry stream bed. The circumstances of collection are very similar to those reported for H. philippinum on Luzon (Taylor, 1922b; Brown et al., 1996; A. Diesmos, personal communication). Discussion The endemic Philippine genus Hologerrhum is now known to contain two species distributed on the Luzon and Visayan aggregate island complexes (Fig. 2). There are no known Hologerrhum from the Pala- wan, Mindoro, Mindanao, Sulu Archipelago, or the Batanes faunal subregions (Fig. 2). The absence of any clear close relatives of Holo- gerrhum (Leviton, 1963, 1965) among SE Asian colu- brids renders speculations regarding the genus' affinities somewhat moot. However, we note that both Hologerrhum and Cyclocorus share characteristics Vol. 9, p. 20 Asiatic Herpetological Research 2001 unique among Asian snakes, most notably, an unusual, presumably derived pattern of reduction in caudodorsal scale rows. In all species of Cyclocorus and Hologerrhum, caudodorsal reduction takes place by fusion of vertebral and paravertebral scale rows, resulting in an odd-numbered series of longitudinal rows of caudodorsals rather than an even number (characteristic of all other SE Asian colubrine snake genera known to us). The systematic affinities of the genus Hologerrhum are in need of further study. The description of Hologerrhum dermali brings the number of new species of vertebrates recently described the by the PNM/CMNH PBI team in the coastal Madja-as mountain range to six (Sison et al., 1995; Gonzales and Kennedy, 1990, 1996; Brown et al., 1997; Ferner et al., 1997; Brown et al., 1999). Other collections from Panay contain at least three probable undescribed species of frogs and many other species of amphibians and reptiles of uncertain taxo- nomic status (many of which are, doubtlessly, unde- scribed species; Ferner et al., this issue). Most of these species presumably are reliant on the closed-canopy rain forests of the western portions of Panay. Accord- ingly, most should be considered severely threatened by deforestation (see Ferner et al., 1997:fig. 2). Recent survey work in the northwestern portions of Panay indicates that Hologerrhum dermali occurs in forested habitats at lower elevations as well as the montane localities reported here (M. Gaulke, personal communication). Unfortunately, the low elevation for- ests of Panay Island have nearly all been removed by an aggressive timber industry in the central Visayas. We expect that numerous other undescribed popula- tions of amphibians and reptiles will be discovered in Panay and the remainder of the Visayas if biologists are permitted access to these forests in order to cata- log and describe Philippine biodiversity. Specimens Examined Hologerrhum philippinum: Philippines, Luzon Island. Zambales Province, Municipality of Masinloc, Barangay Coto, 4.3 km N, 0.5 km E of Mt. High Peak, elevation 1550 m (15° 31' N, 120° 07' E): PNM 2490; Bataan Prov., Mt. Mariveles: CAS 60950-51; Isabela Prov., Municipality of Palanan, Barangay Didian, Sitio Natapdukan, elevation 50 m: PNM 6505; Kal- inga Prov., Municipality of Balbalan, Barangay Bal- balan: CAS 61553-54, MCZ R-25694; Caminares Sur Prov., Municipality of Naga City, Mt. Isarog, eleva- tion 900 m: USNM 31863; Cagayan Prov., Municipal- ity of Baggao, Barrio Santa Margarita, elevation 150 m: USNM 319037, 498718; CAS 134075; Quezon Prov., Municipality of Tayabas, Barangay Camaysa, Mt. Banahaw, 1 150 m above sea level: TNHC 601 14 Camarines Norte Prov., Municipality of Ruis, Baran- gay San Lorenzo, Mt. Labo Range: PNM 2120; Mountain Prov., Mt. Polis: PNM 67; Laguna Prov., Mt. Makiling: MCZ R-25695; Polillo Island, Polillo Prov., near town of Polillo: CAS 62430, MCZ R- 25693; Catafiduanes Island, Municipality of Gigmoto, Barangay Summit Bordan, elevation 200 m: USNM 319037. Hologerrhum dermali: See Holotype and Paratypes sections for this species. Acknowledgments For logistical assistance in the Philippines, we thank the Department of the Environment and Natural Resources (DENR), A. Alcala (Silliman University), P. Gonzales and R. Caberoy (PNM), R. Kennedy (CMNH) and the provincial DENR authorities of Antique Province. The Protected Areas and Wildlife Bureau of the DENR facilitated collecting and export permits necessary for the field portions of this study. For the loans of specimens or assistance while vis- iting museum collections, we thank the following individuals and their respective institutions (museum acronyms follow Duellman et al., 1978 and Leviton et al., 1985): J. Vindum, R. Drewes and J. Slowinski (CAS), R. Crombie, K. de Queiroz, and G. Zug (USNM), R. Kennedy (CMNH), R. Caberoy (PNM), A. Diesmos (De La Salle University), and D. Canna- tella (TNHC). Financial support for RMB's travel to CAS while working on this project was provided by a C. Stearns Fellowship of the California Academy of Sciences. We owe particular thanks to R. Crombie and M. Gaulke for their help and assistance and to D. Wechlser for providing live photographs of H. philip- pinum. Support for field work (by RMB, JWF) was pro- vided in part by the Zoology and Botany Departments and the College of Arts and Sciences of Miami Uni- versity (Oxford, Ohio), the Society for the Study of Amphibians and Reptiles, The Explorer's Club, the Department of Biology of Thomas More College, and the Cincinnati Museum of Natural History. The PNM/ CMNH PBI was supported by a grant (to R. Kennedy and P. Gonzales) from the John D. and Catherine T. MacArthur Foundation and by the benefactors of Cin- cinnati Musuem of Natural History. We thank L. Bockstanz, T. LaDuc, A. Gluesenkamp, T Devitt, A. Diesmos, and D. Cannatella for comments on earlier drafts of this manuscript. The description of Hologer- rhum dermali constitutes contribution No. 24 to the 2001 Asiatic Herpetological Research Vol. 9, p. 21 results of the PNM/CMNH Philippine Biodiversity Inventory. Literature Cited Alcala, A. C. 1986. Guide to Philippine flora and fauna, Vol. X, amphibians and reptiles. Natural Resource Management Center, Ministry of Natural Resources, and University of the Philippines, Manila. 194 pp. Boettger, O. 1886. Aufzahlung der von den Philip- pinen bekannten Reptilien und Batrachier. Bericht liber die Senckenbergische Naturforschende Gesell- schaft in Frankfurt am Main 1 15:91-134. Boulenger, G. A. 1896. Catalogue of the Snakes in the British Museum (Natural History), Vol. Ill, containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae and Viperidae. Trustees of the Brit- ish Museum [Natural History], London, xiv + 727 pp. Brown, R. M., J. W. Ferner. and R. V. Sison. 1995. Rediscovery and redescription of Sphenomorphus beyeri (Reptilia: Lacertilia: Scincidae) from the Zam- bales Mountains, Luzon, Philippines. Proceedings of the Biological Society of Washington 108:6-17. Brown. R. M., J. W. Ferner, R. V. Sison, P. C. Gonza- les, and R. S. Kennedy. 1996. Amphibians and reptiles of the Zambales Mountains of Luzon Island, Republic of the Philippines. Herpetological Natural History 4:1-22. Brown, R. M., A. E. Leviton, and R. V. Sison. 1999 Description of a new species of Pseudorabdion (Ser- pentes: Colubridae) from Panay Island, Philippines with a revised key to the genus. Asiatic Herpetologi- cal Research 8:7-12. Brown, R. M., J. A. McGuire, J. W. Ferner, N. Icaran- gal, and R. S. Kennedy. 2000. Amphibians and reptiles of Luzon Island, II: preliminary report on the herpeto- fauna of Aurora Memorial National Park, Philippines. Hamadryad- 25: 175-195 Brown, W. C, R. M. Brown, and A. C. Alcala. 1997. Species of the hazelae group of Platymantis (Amphibia: Ranidae) from the Philippines, with descriptions of two new species. Proceedings of the California Academy of Sciences 49:405-421. Casto de Elera, R. P. Fr. 1895. Catalogo sistematico de toda la Fauna de Filipinas conocida hasta al pre- sente, y a la vez et de la Colleccion zoologica del Museo de PP. Domfnicos del Colegio-Universidad de Santo Tomas de Manila, vol. 1. Manila, viii + 701 pp. [Ophidiens, pp. 423-445.] Dowling, H. G. 1951a. A proposed standard system of counting ventrals in snakes. British Journal of Herpe- tology 1:97-99. Dowling, H. G. 1951b. A proposed method of expressing scale reduction in snakes. Copeia 1951:131-134. Duellman, W. E., T. Fritts, and A. E. Leviton. 1978. Museum acronyms. Herpetological Review 9: 5-9. Ferner J. W., R. M. Brown, and A. E. Greer. 1997. A new genus and species of moist closed canopy forest skinks from the Philippines. Journal of Herpetology 31: 187-192. Ferner, J. W., R. M. Brown, R. V. Sison, and Robert S. Kennedy. 2000. The amphibians and reptiles of Panay Island, Republic of the Philippines. Asiatic Herpetological Research 9: 34-70. Fischer, J. G. 1885. Ichthyologische und herpetolo- gische Bemerkungen Part IV: Ueber eine Kollection von Amphibien und Reptilien von Mindanao. Jahr- buch Hamburgischen Wissenschaftlichen Anstalten 2:80-81. Frost, D. R., and D. M. Hillis. 1990. Species in con- cept and practice: herpetological applications. Herpe- tologica 46:87-104. Gonzales, P. C, and R. S. Kennedy. 1990. A new spe- cies of Stachyris babbler (Aves: Timaliidae) from the island of Panay, Philippines. Wilson Bulletin 102:367- 379. Gonzales, P. C, and R. S. Kennedy. 1996. A new spe- cies of Cratewmys (Rodentia: Muridae) from Panay, Philippines. Journal of Mammology 77:25-40. Griffin, L. E. 1910. A list of snakes from Polillo, P.I., with descriptions of a new genus and two new species. Philippine Journal of Science 5:21 1-218. Griffin, L. E. 1911. A check-list and key of Philippine snakes. Philippine Journal of Science 6:253-268. Gunther, A. 1858. Catologue of colubrine snakes in the collection of the British Museum. Trustees of the British Museum of Natural History, London, xvi + 281 pp. Gunther, A. 1873. Notes on some reptiles and batra- chians obtained by Dr. Adolf Bernhard Meyer in Celebes and the Philippine islands. Proceedings of the Zoological Society of London 1873:165-172. Gunther, A. 1879. List of the mammals, reptiles, and batrachians sent by Mr. Everett from the Philippine Islands. Proceedings of the Zoological Society of London 1879:74-79. Hall, R. 1996. Reconstructing Cenozoic southeast Asia. Pp. 153-184. In R. Hall and D. Blundell (eds.), Vol. 9, p. 22 Asiatic Herpetological Research 2001 Tectonic Evolution of Southeast Asia. Geological Society, London, iv + 368 pp. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. Pp. 99-132. in Biogeography and geological evolution of South- east Asia. R. Hall and J. D. Holloway (eds.). Brack- huys, Leiden, The Netherlands, ii + 417 pp. Heaney, L. R. 1986. Biogeography of small mammals in SE Asia: estimates of rates of colonization, extinc- tion and speciation. Biological Journal of the Linnean Society 28:127-165. Jan, G., and F. Sordelli. 1870. Iconographie generates des ophidiens, Tome Troisieme. J. B. Bailiere et Fils, Paris. 188 pp. Lee, J. C. 1990. Sources of extraneous variation in the study of meristic characters: the effect of size and inter-observer variability. Systematic Zoology 39:31- 39. Leviton, A. E. 1963. Remarks on the zoogeography of Philippine terrestrial snakes. Proceedings of the Cali- fornia Academy of Sciences 31:369-416. Leviton, A. E. 1965. Contributions to a review of Phil- ippine snakes, IX. The snakes of the genus Cyclo- corus. Philippine Journal of Science 94:519-533. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthy- ology: Part I. Standard symbolic codes for institu- tional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Ross, C. A., and P. C. Gonzales. 1991. Amphibians and reptiles of Catanduanes Island, Philippines. National Museum Papers 2:49-76. Ruedas, L. A., J. R. Demboski, and R. V. Sison. 1994. Morphological and ecological variation in Otoptero- pus cartilagonodus Kock, 1969 (Mammalia: Chi- roptera: Pteropodidae) from Luzon, Philippines. Proceedings of the Biological Society of Washington 107:1-16. Simmons, J. E. 1987. Herpetological collecting and collections management. Society for the Study of Amphibians and Reptiles Herpetological Circular 16:1-70. Simpson, G. G. 1961. Principles of Animal Taxon- omy. Columbia University Press, New York, NY. 164 pp. Sison, R. V., P. C. Gonzales and J. W. Ferner. 1995. New island records from Panay, Philippines. Herpeto- logical Review 26: 48-49. Taylor, E. H. 1922a. The Snakes of the Philippine Islands. Monograph of the Bureau of Science, Manila, No. 16.312 pp. Taylor, E. H. 1922b. Additions to the herpetological fauna of the Philippine Islands, I. Philippine Journal of Science 21:161-206. Taylor, E. H. 1922c. Additions to the herpetological fauna of the Philippine Islands, III. Philippine Journal of Science 22:51 5-557. Taylor, E. H. 1922d. Herpetological fauna of Mount Makiling. Philippine Agriculturist 11:127-139. Wiley, E. O. 1978. The evolutionary species concept reconsidered. Systematic Zoology 21:17-26. Whitmore, T. C. 1984. Tropical Rain Forests of the Far East. Clarendon Press, Oxford. 209 pp. 2001 Asiatic Herpetological Research Vol. 9. pp. 23-24 The First Record of Ptyas korros (Co\ubr\dae) from Bangladesh M. Farid Ahsan1 and Shayla Parvin Department of Zoology, University of Chittagong, Chittagong 4331, Bangladesh, e-mail: mfahsan@ctgu.edu Abstract.- This is the first report of Ptyas korros from Bangladesh. The specimens reported here were collected from the Cittagong University campus between the years of 1985 and 1996. They match previously described specimens of Ptyas korros except in the number of loreal scales. Ptyas korros is known to occur in the Assam region of India and so its presence in adjacent regions of Bangladesh is expected, but unconfirmed until now. Key words.- Serpentes, Colubridae, Ptyas korros, Bangladesh, distribution, biogeography The number of species of snakes occurring in Bang- ladesh is still debatable because of the lack of confir- mation of some species. However, lists have been compiled based on a number of field studies con- ducted in different parts of the country. Sarker (1975) recorded 20 species, Montaquim (1979) reported 18 species, Montaquim et al. (1980) recorded 28 species and Rashid (1982) recorded 22 species from Bang- ladesh. Whereas Khan (1982) estimated the snakes species of the country as 78 and he later reported 79 species in 1987 and 81 species in 1992 (of which the occurrence of two are doubtful). Sarker and Sarker (1988) reported 88 species, although some are uncer- tain. This report established the presence of one spe- cies, Ptyas korros (Schlegel, 1837). During the identification of the snake species pre- served in the Departmental Museum of Zoology, Chit- tagong University (CU), four specimens (three 5 cm Figure 1. Specimen of Ptyas korros from Chittagong University Campus. collected from CU Campus [CUC] between 1985 and 1996, and one not known but possibly from CUC) of P. korros were found (Fig. 1). Until now, nobody has reported the occurrence of P. korros in Bangladesh, although Husain (1977) predicted it. Its known range extends from Assam (India) through Myanmar (Burma) to Indo-China, Thailand, throughout the Ten- asserim to the Malay Peninsula, Indonesia (Sumatra, Java, Borneo), Southern China including Hainan, Tai- wan and Hong Kong (cf. Giinther, 1864; Smith, 1943; Zhao & Adler 1993). The occurrence of P. korros in the CUC, Chittagong, extends its distribution to Bang- ladesh. It might also occur in nearby parts of the country such as Jamalpur, Greater Sylhet, Chittagong, Chittagong Hill Tracts (Rangamati, Khagracheri and Bandarbans Districts). The specimens from the CUC are olive brown above and yellowish white below. The scales on the posterior part of the dorsal side of the body are edged with a dark brown pattern (looking black) producing a regular network pattern. The outer margins of the ven- trals and caudals are also sometimes edged with dark brown (less dark than the dorsal color). The coloration of the CUC specimens resembles the P. korros speci- mens described by Smith (1943). Smith (1943) diag- noses P. korros from Ptyas muscosus by the number of dorsal scale rows (15 in P. korros vs. 16 or 17 in P. muscosus) and the number of ventral scales (160-187 in P. korros vs. 190-213 in P. muscosus). The CUC specimens have 15 dorsal scale rows and 177-188 ventral scales. They differ from P. korros specimens described by Giinther (1864) in the number of loreal scales. Giinther reported two loreals, but three of the four CUC specimens have only one loreal. Literature Cited Giinther. A. C. L. G. 1864. The reptiles of British India. Oxford & IBH Publishing Co., New Delhi. 452 pp + XXVI plates. Vol. 9, p. 24 Asiatic Herpetological Research 2001 Husain, K. Z. 1977. Bangladesher banayajonju sham- pad O tar sangrakshan (Wildlife resources of Bang- ladesh and their conservation). Bangla Academy Bijnan Patrika. 3(3): 1-1 1. (in Benglai). Khan, M. A. R. 1982. Wildlife of Bangladesh -a checklist. Dhaka University Press, Dhaka. 173 pp. Khan, M. A. R. 1987. Bangladesher banayaprani (Wildlife of Bangladesh) vol. 1. Bangla Academy, Dhaka. 169 pp. (in Benglai). Khan, M. A. R. 1992. Bangladesher shap (Snakes of Bangladesh). Bangla Academy, Dhaka. 227 pp. (in Benglai). Montaquim, M. A. 1979. Snakes of several districts of Bangladesh. Unpublished M. Sc. Thesis, University of Dhaka, Dhaka. 63 pp + v plates. Montaquim, M. A., Sarker, A. H.; Khan, M. A. R., and K. Z. Husain. 1980. List of the snakes of Bangladesh. Bangladesh Journal of Zoology. 8(2): 127-130. Rashid, S. M. A. 1982. On some snakes of Bang- ladesh with notes on their habit, habitat, status and distribution. Unpublished M. Sc. Thesis, University of Dhaka, Dhaka. 108 pp + xv plates. Sarker, A. H. 1975. Snakes of Bangladesh. Unpub- lished M. Sc. Thesis, University of Dhaka, Dhaka. Sarker, M. S. U. and Sarker, N. J. 1988. Wildlife of Bangladesh (a systematic list with status, distribution and habitat). The Rico Printers, Dhaka. 69 pp. Smith, M. A. 1943. The fauna of British India, includ- ing Ceylon and Burma, Reptilia and Amphibia, vol. 3 Serpentes. Taylor and Francis, London. 583 pp. Zhao, E. and Adler, K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, New York. 522 pp + 48 plates. 2001 Asiatic Herpetological Research Vol. 9, pp. 25-29 Influence of Temperature on Burrow Use by the Monitor Lizard Varanus panoptes o\ the Coastal Dunes at Fog Bay, Northern Australia Sean J. Blamires Morning Bell Language School, 671-3 Jukdo 2 Dong, Pohang City, Kyongsangbuk Do 791-052, South Korea. email: s_blamires@hotmail.com Abstract.- An increase in the number of Varanus panoptes burrows appearing among the sand dunes at Fog Bay in northern Australia was noticed during the dry season (June to August). Entrances of marked burrows were smoothed, and their interiors investigated using a burrowscope, to determine the monthly number of foraging and retreat burrows appearing over a 12 month period. In the dry season, increased numbers of both types of burrows were found. A temperature data logger was used to record temperatures of the sand surface, and at a depth of 50 cm. Burrow depths were measured as how far the burrowscope could be lowered into each burrow. There was a positive correlation between the number of retreat burrows and burrow depths. There was a negative correlation between the number and depth of retreat burrows and the minimum sand-surface temperature and the maximum sand temperature at 50 cm. The number of foraging burrows was independent of sand temperatures. The results indicate a likely thermoregulatory use of retreat burrows with more and deeper burrows prevalent when overnight surface and subterranean sand temperatures decrease. Key words.- Reptilia, Varanidae, Varanus panoptes, Australia, Northern Territory, fossorial, thermoregulation Introduction Burrows serve many ecologically important purposes to many animals (Hansell, 1993). Lizards expend less energy throughout the day if a large portion of time is spent in burrows (Bennett and Nagy, 1977). Monitor lizards (Varanus spp.) utilise burrows for a variety of purposes such as thermoregulation (Cowles and Bogert, 1944), reducing water loss (Green, 1972), finding prey (Pianka, 1969), and for oviposition and retreat (Cowles and Bogert, 1944; Auffenberg, 1983). In a recent study of the monitor lizard Varanus panoptes (Fig. 1 ) inhabiting the coastal dunes at Fog Bay, Northern Territory, Australia (12°42'S; 130°20'E), I detected a seasonal change in the num- ber of burrows present. A greater number of burrows appeared among the dunes in June-August (dry sea- son) compared to the rest of the year. These burrows also seemed quite deep compared to those seen in the wet season. Use of these burrows for oviposition is unlikely since Varanus panoptes produces eggs during the wet season. Free access to other water sources might rule out conservation of water loss. Overnight tempera- tures in the area often drop below 15°C in the dry sea- son. A likely explanation may be that the monitors retreat into burrows at a depth where sand tempera- tures remain high to conserve overnight body temper- Figure 1. Varanus panoptes'm the coastal dunes at Fog Bay, Northern Australia. ature, as has been found for some other monitors (Cowles and Bogert, 1944; Auffenberg, 1983). The monitors at Fog Bay inhabit the dunes and forage on the beach and dunes (Blamires, 1999). Increased for- aging activities may be another explanation for the number of burrows seen in the dry season. The aim of this investigation was to determine whether changes in the number of burrows between the wet and dry Vol. 9, p. 26 Asiatic Herpetological Research 2001 seasons is a result of foraging, thermoregulation, retreat or a combination of these, or other, influences. Material and Methods The area of the investigation is approximately 5 km of beach along the northern-most mainland beach at Fog Bay (12°42'S, 130°20'E), approximately 80 km from Darwin, Northern Territory, Australia. The area is pri- vately owned and closed to the public. The sandy dunes are immediately backed by grassland domi- nated by spinifex vegetation and dispersed Pandanus trees. Black soil plains, mangroves and salt flats back the grassland. The dunes of the southern-most 1 km stretch of beach are backed by monsoon forest. The entire 5 km of beach, and the grassland and monsoon forest backing the dunes, was walked twice monthly over a 12 month period. All burrows encountered were marked with a depth of surveyor's tape tied to nearby vegetation (Fig. 2). To estimate visitation rates the entrances were smoothed and checked, on revisiting, for varanid tracks leading into it. A small video-camera device, called a "burrow- scope" (Dyer and Hill, 1991; Dyer and Aldworth, 1998) was used to investigate the inside of burrows. The model I used was a modified version of that of Dyer and Aldworth, (1998). A small black and white CCD, 38mm x 38mm, camera (Samsung MOD-BW 204), now popular for home security systems (Capel, 1993) encased in a 375 ml jar was used. Infrared light- ing was used to illuminate the burrow for the camera, provided by 10 high intensity light emitting diodes fit- ted on a piece of Vero board. A small black and white video monitor (260 mm; 5.8 kg) was used for view- ing, operated directly from a 12V battery. The wiring from the camera to the monitor was approximately 2 m long and encased in a garden hose. The depth of each burrow was determined by measuring the depth of hose that fed into the burrow to reach its end. Burrows were assumed to be for foraging when were noted to intersect the burrow of a potential prey item (crab, skink or bird). The number of foraging burrows was totalled each month. Occupied burrows, and those with tracks appearing from the entrance after smoothing, not identified as foraging burrows, were classified as retreat burrows. The number of retreat burrows observed was totalled monthly. A temperature data logger (model 6003 A, Unidata Australia, Perth) was placed on top of one of the dunes with two probes extruding from the logger's case. One of the probes was placed 1 cm below the sand surface, while the other was buried at 50 cm ■y ( - ' ~/' gpj I *% Figure 2. Burrow of Varanus panoptes . below the surface. A digital temperature reading was taken by the logger every 30 minutes. The data were downloaded and compiled at the end of every month. Correlation analysis was done between the num- ber of foraging and retreat burrows counted each month and the mean burrow depth. Correlation analy- sis was also done between the number of foraging burrows, retreat burrows and burrow depths and monthly maximum and minimum sand temperature at the surface and at 50 cm depth. Results Overall 93 burrows were identified as either retreat or foraging burrows. Fifty-one were identified as forag- ing burrows and 42 as retreat burrows, three of the retreat burrows were identified as such because they were occupied by a monitor lizard. The numbers of retreat and foraging burrows observed each month are shown in Figure 3. There was a peak in the number of retreat burrows in July with a steep drop in August and September. The number of foraging burrows observed peaked in April, although the number stays high until July when the number drops. The number of retreat burrows exceeded the number of foraging burrows between May and September. There was a positive correlation between the number of retreat burrows and the burrow depths recorded each month (r = 0.67; P = 0.017; Table 1). Table 1 . Correlation coefficients between the number of retreat and foraging burrows and burrow depths. * denotes a significant correlation Retreat Burrows 0.67 0.017* Foraging Burrows 0.097 0.763 2001 Asiatic Herpetological Research Vol. 9, p. 27 12 10 O 3 SQ - o 6 z -2 O RETREAT BURROWS -Q FORAGING BURROWS JAN SEP MAR MAY JUL MONTH Figure 3. Monthly number of foraging and retreat burrows of V. panoptes at Fog Bay. NOV However, the number of foraging burrows is not correlated to burrow depths (Table 1 ). The number of retreat burrows are negatively correlated with the min- imum sand-surface temperatures (r = 0.463; P = 0.024) and the maximum sand temperature at 50 cm (r = -0.616; P = 0.033; Table 2). Burrow depths were also negatively correlated to the minimum sand sur- face temperature (r = -0.789; P = 0.002) and the maxi- mum sand temperature at 50 cm (r = -0.64; P = 0.033; Table 2). The number of foraging burrows was unaf- fected by sand surface temperature and sand tempera- ture at 50 cm (Table 2). This equates to more and deeper retreat burrows being dug, the cooler the sand temperature at the surface and at 50 cm. Discussion The field experiments herein were the first attempt with this modified design of burrow viewing equip- ment and an assessment of its effectiveness is war- ranted to justify the results. The only problem encountered was when the original silicon glue (a metal sealant), used to hold the camera to the jar cas- ing, was suspected of having a corrosive effect on the camera. The camera stopped working within a short time and, when inspected, small white spots were seen on the solder of the circuit board. The camera worked properly when the spots were cleaned off and a non-corrosive sealant (glass/window sealant) was used. From then on, the system provided clear images, with effective viewing of monitor lizard presence in burrows and the depth, width and curvature of all bur- rows. The narrower, rounder burrows of other animals were easily identifiable at the bottom of foraging bur- rows. The number of foraging burrows was independent of depth or temperature influences. However, the greater the number of retreat burrows dug, the greater their depth. The number and depth of retreat burrows was also negatively correlated to minimum sand-sur- face temperature each month. This is indicative of an increased need for the lizards to burrow to an increas- ing depth as the overnight temperatures cooled. Cooler sand temperatures at 50 cm also caused an increase in the number and depth of retreat burrows. Since thermal diffusion in sand is slow, sand tempera- ture at 50 cm heats and cools slowly (Packard and Packard, 1988). The sand temperature at 50 cm at Fog Bay was the highest overnight when the sand surface is lowest (Guinea, 1994). Thus, monitors may be bur- rowing more in the dry season to exploit the warmer overnight internal sand temperatures. Maximum inter- nal sand temperatures fall during the dry season and the monitors therefore dig further to find even warmer temperatures. Foraging burrows were also responsible for increasing the number of burrows seen in the dry Vol. 9, p. 28 Asiatic Herpetological Research 2001 Table 2. Correlation coefficients between the number of retreat and foraging burrows and burrow depth and sand surface (Tso) minimum and maximum temperatures and sand at 50 cm depth (TS50) minimum and maximum tem- peratures. * denotes a significant correlation. Retreat burrows r P Foraging r burrows P Burrow r depth P Min. Tso Max. Tso -0.643 0.316 0.024* 0.317 0.18 0.227 0.576 0.478 -0.789 0.722 0.002* 0.008* Min. TS50 -0.296 0.351 -0.145 0.653 -0.086 0.79 Max. TS50 -0.616 0.033* 0.026 0.937 -0.646 0.033* season, but their depths were constant, and indepen- dent of temperature. Of other possible reasons for an increase in moni- tor lizard burrowing in the dry season, egg ovipositing is unlikely, as no eggs were ever observed within bur- rows and V. panoptes. Burrow use to conserve water loss has been proven important for some arid zone monitor lizards (Green, 1972, Vernet et al., 1988). This has not been demonstrated for tropical monitor lizards, although there is a considerable difference in humidity and rainfall between the wet and dry seasons in Australia's wet-dry tropical region (Bureau of Meteorology, 1989). Water loss in lizards increases with decreasing ambient humidity (Hillman and Gor- man 1977) and without physiological adaptations to prevent water loss, burrowing to moist sand may be utilised to prevent desiccation by V. panoptes when ambient humidity decreases. More needs to be investi- gated on the water economies, and uses of other water sources, in this monitor to determine if water loss is influential in increasing burrowing during the dry sea- son at Fog Bay. Acknowledgments I thank Bill Milne, Faculty of Technology, Northern Territory University for constructing the burrowscope. Greg Hill and Pam Dyer provided feedback on its design. Equipment was purchased from Oatley Elec- tronics. Project funding was by Australian Geo- graphic, The Queen's Trust for Young Achievers and The Centre for Tropical Wetlands Management, Northern Territory University. Literature Cited Auffenberg, W. 1983. The burrows of Varanus benga- lensis: characteristics and use. Records of the Zoolog- ical Survey of India. 80:375-385. Bennett, A. F. and K. A. Nagy. 1977. Energy expendi- ture in free ranging lizards. Ecology 58:697-700. Blamires, S. J. 1999. Quantifying predation on sea turtle nests by varanids at Fog Bay. MSc Thesis. Northern Territory University. Bureau of Meteorology. 1989. Climate of Australia. Morphea Press, Canberra. Capel, V. 1993. Security Systems and Intruder Alarms. Newnes, Oxford. 267 pp. Cowles, R. B., and Bogert, C. M. 1944. A preliminary study of the thermal requirements of desert reptiles. Bulletin of the American Natural History Museum 83:261-296. Dyer, P. K., and G.J.E. Hill. 1991. A solution to the problem of determining the occupancy status of wedge-tailed shearwater Puffinus pacificus burrows. Emu 91:20-25. Dyer, P. K., and K. Aldworth.. 1998. The "burrow- scope": modifications to burrow viewing equipment. Emu 98:143-146. Green, B. 1972. Water loss of the sand monitor lizard (Varanus gouldii) in its natural environment. Ecology 53:452-457. Guinea, M. L. 1994. A possible model to explain win- ter nesting by the flatback turtle, Natator depressus at Fog Bay, Northern Territory. Pp. 154- 155. In R. James (ed.), Australian Marine Turtle Conservation Work- shop. Queensland Department of Environment and 2001 Asiatic Herpetological Research Vol. 9, p. 29 Heritage, Australian Nature Conservation Agency. Gold Coast. Hansell, M. H. 1993. The ecological impact of animal nests and burrows. Functional Ecology 7:5-12. Hillman, S. S., and G. C. Gorman. 1977. Water loss, desiccation tolerance and survival under desiccating conditions in two species of Carribean Anolis. Oeco- logia 29:105-1 16. Packard, G. C, and M. J. Packard. 1988. The physio- logical ecology of reptilian eggs and embryos. Pp. 523-605. In C. Gans and R. B. Huey (eds.). Biology of the Replilia, Vol. 16. Alan Liss, New York. Pianka, E. R. 1969. Habitat specificity, speciation and species density in Australian desert lizards. Ecology 50:498-502. Vernet, R.. M. Lemire. and C. Grenot 1988. Field studies on activity and water balance of a desert mon- itor Varanus griseus (Reptilia: Varanidae). Journal of Arid Environments. 15:81-90. 2001 Asiatic Herpetological Research Vol. 9, pp. 30-33 Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx aegyptius microlepis from the United Arab Emirates Peter L. Cunningham P.O. Box 17258, AlAin. United Arab Emirates, email: plc@ emirates. net.ae Abstract.- Uromastyx aegyptius microlepis are almost exclusively herbivorous and feed on a variety of plant species with Pennisetum divisum being extensively utilized in the United Arab Emirates. The survival rate, as determined over a one year period without any rainfall is <50%. Burrow openings are mainly aligned in a southerly and easterly direction with the average burrow entrance size being 60x155 cm. Temperatures measured at 30 cm down the burrow entrance are on average 6°C lower than the ambient temperature. Key words.- Reptilia. Squamata. Agamidae, Uromastyx aegyptius microlepis. United Arab Emirates, ecology Figure I.The Spiny-tailed lizard, Uromastyx aegyptius microlepis. Introduction Uromastyx aegyptius microlepis (Spiny-tail Lizards; Fig. 1) belong to the Family Agamidae and occur throughout the Arabian Peninsula, Iran, Iraq and Jor- dan (Arnold 1986. Leviton et al. 1992). They are diurnal ground dwelling lizards that may reach sizes of up to 60 cm and are generally yellowish-grey in colour with an impressive spiny club-like tail. Adult U. a. microlepis are documented as being mainly her- bivorous, although insects form part of their diet, while juvenile lizards are thought to be mainly insec- tivorous (Arnold 1984, Baha El Din 1996, Brown Figure 2. The study site and habitat of U. a. microlepis in Abu Dhabi. 982, Highveld and Slimani 1998, Jongbloed 1997, Manthey and Shuster 1996). Material and Methods Twenty mature Uromastyx aegyptius microlepis indi- viduals in a scattered population, ±35 km northwest of Al Ain (24°25'07"N and 55°35'01"E; Fig. 2) in the Abu Dhabi Emirate of the United Arab Emirates (UAE), were observed for one full day each (sunrise to sunset). These observations took place during sum- mer (May and June 1999) and winter (December 1999 and January 2000), respectively. I spent 480 observa- tion hours studying these lizards. Direct observations of plant species utilized during their feeding bouts was conducted using an 8x40 binocular over a dis- tance of 100m so as not to influence the feeding behavior. Tracks were also followed once the lizards had retreated underground to confirm sightings. Fecal pellets were collected for later analysis. Burrow 2001 Asiatic Herpetological Research Vol. 9, p. 31 information (orientation, burrow entrance height and width, temperature above ground and 30 cm down burrow) was collected from 25 active burrows once the lizards had retreated. Survival rate was determined for the 20 study individ- uals by investigating their burrows for any signs of activity in June 2000, one year after the first observa- tions were made. Results Diet. Uwmastyx aegyptius microlepis utilized the fol- lowing 10 plant species: Class Monocotyledonae Gramineae (Grass family): Pennisetum divisum, Stipagrostis plumosa Class Dicotyledonae Amaranthaceae (Cockscomb family): Aerva jav- anica Asclepiadaceae (Milkweed family): Leptadenia pyrotechnica Boraginaceae (Borage family): Moltkiopsis cili- ata, Heliotropium kotschyi Chenopodiaceae (Goosefoot family): Haloxylon salicomicum Cucurbitaceae (Gourd family): Citrullus colocyn- this Leguminosae (Pea family): Tavemiera cuneifolia Polygalaceae (Milkwort family): Polygala eri- optera Survival Rate. Of the 20 study individuals first observed in May 1999, only 9 were still present in June 2000. Table 1 . Vegetation selected by Uromastyx aegyptius microlepis as documented by different authors from the Arabian Peninsula. * Pulicaria glutinosa observed being utilized in other areas although not in the present study area. This Study UAE- Abu Dhabi Aerva javanica Citrullus colocynthis Haloxylon salicomicum Heliotropium kotschyi Leptadenia pyrotechnica Moltkiopsis ciliata Pennisetum divisum capitata Polygala erioptera *Pulicaria glutinosa Stipagrostis plumosa Tavemiera cuneifolia Jongbloed(1997) UAE - Sharjah Fagonia sp. Pennisetum divisum Mandevile(1965) Saudi Arabia Aristida plumosa Astragalus gyzensis Citrullus colocxnthis Horwoodia dicks on eae Launaea capitata Moltkiopsis ciliata Neurada procumbens Plantago boissieri Zygophyllum sp. Vol. 9, p. 32 Asiatic Herpetological Research 2001 Burrow specifics (n=25). Burrow openings were ori- entated as follows: North 2, West 3, South 12 (S/West: 4, South: 5, S/ East: 3) and East 8. Mean average burrow opening height and width was 61 cm and 155 cm (n=25), respectively. Mean average temperature at a depth of 30 cm down the burrow was 38°C (n=25). The ambient tem- perature measured at 1.5 m above ground at 13h00 was 44°C. Discussion Diet. Ten perennial plant species are selected by Uro- mastyx aegyptius microlepis as observed during the study period, with the coarse desert grasses Pennise- tum division and Stipagrostis plumosa being favored. Jongbloed (1997) identified 3 plant species selected by the lizards from the Sharjah area in the UAE while Mandaville (1965) identified 8 plant species from the stomach contents of six Uromastyx aegyptius speci- mens from Saudi Arabia (See Table 1). Foley et al. (1992) identified 23 plant species, mainly annuals, selected by the same species in Israel. As no rain had fallen during the study period no annuals were observed and therefor the exclusion from the diet. It would therefor be expected that more plant species, especially flowering annuals, would be utilized after rains. Rainfall is highly variable (± 100mm p. a.) and unpredictable in the UAE (Bot- tomley 1996) forcing the lizards to rely on the avail- able perennials. According to Zari (1996 and 1998) Uromastyx philbyi feed on a variety of annual grasses and perennial shrubs in western Saudi Arabia. High- field and Slimani (1998) indicate Uromastyx acan- thinurus's preference for the family Chenopodia (fat hen/spinach) as well as the salt tolerant Artiplex genus and a wide variety of ephemeral plants after rains in Morocco. An analysis of 170 fresh adult fecal pellets (85 summer and winter, respectively) confirmed the her- bivorous nature of U. a.microlepis as only one pellet (<1%) contained insect remains, that of a Tennebri- onid beetle. According to Highfield and Slimani (1998) adult fecal pellets of U. acanthinurus in Morocco contained 6% insect matter. Brown (1982) also mentions U. a.microlepis feeding on locusts from the UAE. It still has to be investigated if the lizards would show a preference for insects during autumn and spring although it is expected that the lack of insects in the diet could be contributed to the fact that most insects, Tenebrionidae at least, are crepuscular or nocturnal (Tiger 1996). Other items consumed by Uromastyx as observed during the fecal analysis of the pellets include: sheep droppings (17 pellets), date kernels (4 pellets) and feathers (3 pellets). It is expected that above men- tioned items are consumed erroneously by the lizards as insect material although Brown (1982) suggests that sheep droppings be consumed to extract undi- gested plant material. If the sheep droppings, date kernels and feathers were consumed as insect mate- rial, it would suggest that the lizards would prey on insects more often if they were more readily available. Survival rate. Nine (45%) of the 20 study individuals survived a period of one year from May 1999 to June 2000. Three of the 1 1 missing individuals were found as severely malnourished carcasses. The other 8 indi- viduals not accounted for are presumed to have suc- cumbed below ground in their burrows, some of which were covered by sand. No signs of emigration were determined. No rainfall occurred during this period in the study area. The area is also frequented by large numbers of domestic camels and sheep, which compete directly with the lizards for the avail- able food. It is suggested that the lack of rainfall and lack of annual plants as a result of this, together with the extra pressure of domesticated stock, resulted in the high mortality observed. Burrow specfics. Burrow openings were mainly aligned to the south and east with 80% of the burrow entranced facing this way. During summer, tempera- tures consistently reach between 40°C and 47°C between lOhOO and 16h00. North and west facing burrow entrances would be hotter than burrows facing in other directions, especially during midday. During winter the lizards would also benefit from the south- erly orientation warming the burrows quicker and more effectively. Wind direction during summer is predominantly from the northwest and can cause severe sandstorms locally known as "shamal" (El- Ghonemy 1985). This causes lizard whose burrows face that direction to spend more time on den clearing, a time consuming an energetically costly affair espe- 2001 Asiatic Herpetological Research Vol. 9, p. 33 cially during summer when plant availability and pal- atability are low. Burrow size is often related to the size of the resi- dent lizard. The mean average size of 61 x 155 cm (n=25) is indicative of an adult Uromastyx colony. Bigger burrow openings have higher temperatures at a depth of 30 cm, especially when facing to the north or west. The mean average temperature of 38°C, during the heat of the day, at a depth of 30 cm down the burrow is on average 6°C lower than the ambient temperature. This gradient is imperative to the survival of the liz- ards. During the fieldwork it was often found that the lizards were resting in the burrows at a depth of just over 30 cm, scurrying deeper once the thermometer was inserted down the burrow. According to Brown (1982), the burrows can be more than 6 feet long and 3 to 4 feet deep in the form of a spiral. Jongbloed (1997) confirms this depth even indicating reports of up to 12 feet deep burrows. The same author men- tions that an excavated burrow of 5 feet in depth was 2 feet across and 1 foot high and still not at its end. Burrows do not only serve as thermoregulatory havens for Uromastyx aegyptius microlepis, but also for the Desert Monitor (Varanus griseus) (Pers.obs.) and certain Larks (Cunningham 2000, Williams et al. 1999). Williams et al. (1999), states that Uromastyx burrows can potentially reduce Hoopoe Lark (Alae- mon alaudipes) water loss by as much as 81% during the hottest periods during summer. Temperatures at a depth of 30 cm are relatively stable therefor being exploited by many desert dwelling animals (Love- grove and Knight-Eloff 1988). Burrows are thus an effective way of escaping predators, the harsh desert environment as well as controlling water loss. Literature Cited Arnold, E. N. 1984. Ecology of lowland lizards in the eastern United Arab Emirates. Journal of Zoology, London 204:329-354. Arnold, E. N. 1986. A key and annotated checklist to the lizards and amphisbaenians of Arabia. Fauna of Saudi Arabia 8:385-435. Baha El Din, S. 1996. Terrestrial reptiles of Abu Dhabi. Pp. 124-147. In P.E. Osborne (ed.), Desert Ecology of Abu Dhabi. Pisces Publications, Newbury, UK. Bottomley, N. 1996. Recent climate of Abu Dhabi. Pp. 36-49. In: P.E. Osborne (Editor), Desert Ecology of Abu Dhabi. Pisces Publications, Newbury, UK. Brown, J. N. B. 1982. Spiny-tailed Agamid - Uro- mastyx microlepis (Arabic "Dhub"). Emirates Natural History Group Bulletin Nol6:20. Cunningham. P. L. 2000. The use of burrows by Hoo- poe Lark (Alaemon alaudipes). Tribulus 10. 1 :2 1 . El-Ghonemy, A. A. 1985. Ecology and flora of Al Ain Region. University of the United Arab Emirates, Al Ain. Foley. W. J., A. Bouskila, A. Schkolnik and I. Chosh- niak. 1992. Microbial digestion in the herbivorous liz- ard Uromastyx aegyptius (Agamidae). Journal of Zoology, London. 226:387-398. Highveld, A. C. and T. Slimani. 1998. The Spiny- Tailed Lizard at home: Uromastyx acanthinurus in Southern Morocco. Reptiles - Guide to keeping rep- tiles and amphibians, July 1998: 76-87. Jongbloed, M. 1997. Observations in a Dhub colony. Tribulus 7.2:23-24. Leviton, A. E., S. C. Anderson, K. Adler and S.A. Minton. 1992. Handbook to Middle East Amphibians and Reptiles. Scociety for the Study of Amphibians and Reptiles, Oxford, Ohio. 252 pp. Lovegrove, B. G. and A. Knight-Eloff. 1988. Soil and burrow temperatures, and the resource characteristics of the social mole-rat Cryptomys damarensis (Bathy- ergidae) in the Kalahari Desert. Journal of Zoology, London 216:403-413. Mandeville, J. 1965. Plants eaten by Uromastyx microlepis Blanford and other notes on this lizard in Eastern Arabia. Journal Bombay Natural History Society 62(1): 161 -163. Manthey, U. and N. Schuster. 1996. Agamid Lizards. T.F.H. Publications Inc., USA. Tiger, B. J. 1996. A preliminary assessment of the arthropods of Abu Dhabi. Pp. 172-195. In P.E. Osborne (ed.). Desert Ecology of Abu Dhabi. Pisces Publications, Newbury, UK. Williams, J. B., B. I. Tielman and M. Shobrak. 1999. Lizard burrows provide thermal refugia for larks in the Arabian Desert. The Condor 101 (3):7 14-7 1 7. Zari, T. A. 1996. Effects of body mass and tempera- ture on standard metabolic rate of the herbivorous desert lizard, Uromastyx philbyi. Journal of Arid Envi- ronment 33:457-461. Zari, T. A. 1998. Effects of sexual condition on food consumption and temperature selection in the herbiv- orous desert lizard, Uromastyx philbyi. Journal of Arid Environment 38:371-377. 2001 Asiatic Herpetological Research Vol. 9, pp. 34-70 The Amphibians and Reptiles of Panay Island, Philippines John W. Ferner1, 2, Rafe M. Brown2, 3, Rogelio V. Sison4 and Robert S. Kennedy2 Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, USA. email: fernerj@thomasmore.edu; Geier Collections and Research Center, Museum of Natural History and Science, 1301 Western Avenue, Cincinnati, Ohio 45203, USA; Section of Integrative Biology and Texas Memorial Museum, University of Texas, Austin, Texas 78712, USA. email: rafe@mail.utexas.edu;4 Zoology Division, National Museum of the Philippines, Executive House, P. Burgos Street, Manila, Philippines, email: nmzoo@fastmail.i-next.net. Abstract.- We provide species accounts for 20 amphibians (frogs and toads) and 72 reptiles (one turtle, 36 lizards, and 35 snakes) from the central Philippine island of Panay and its associated land-bridge islets. Panay is a moderately-sized island (123,000 km-) that currently is separated from the nearby islands of Negros, Masbate, and Cebu by shallow seas, indicating that dry land connections once allowed free exchange of flora and fauna between these land masses at several periods during the Pleistocene. This fact, coupled with the wealth of knowledge on herpetological communities of Negros and Cebu, has led biologists to assume that the amphibians and reptiles of Panay are reasonably well known. Our data suggest that this is far from true. Our recent survey work (1989 until present) has resulted in the discovery of at least six (and probably 12-15) new species of vertebrates, most of which appear to be high elevation rainforest Panay endemics. In this paper we note numerous taxonomic problems that await the attention of biologists; these surely will result in an increase of the known species from Panay. We expect that many widespread species complexes currently of uncertain taxonomic status will eventually be recognized as Panay endemics, further bolstering the total diversity and degree of endemism on Panay. The status of Panay herpetofauna warrants particular attention because many of the suspected new and endemic species appear to be forest obligates. With the near complete removal of Panay 's low elevation forests and increased pressure on its mid- and upper montane forests, we suspect that many of Panay's endemics are threatened by extinction. Panay should be regarded as a tropical island with a unique geological history, a spectacular compliment of habitat types, and a diverse set of species communities - all of which are beset with a conservation crisis of deforestation and probable extinction. Panay may be a particularly appropriate model island for large scale conservation efforts aimed at sustainable management of forest resources. However, before informed conservation and management plans can be enacted, additional biodiversity survey work must be conducted on Panay. Key words.- Reptiles, amphibians, SE Asia, Philippines, Panay, Visayan Islands, conservation crisis, herpetofauna. Introduction The central (or "Visayan") Philippine island of Panay (Figs. 1 , 2), is part of a Pleistocene aggregate island complex that includes Negros, Cebu, Guimaras, Mas- bate, Ticao, and several small land-bridge islands associated with each of these larger land masses. Although amphibian and reptile species similarity among Visayan Islands was noted much earlier (Tay- lor, 1920, 1922a-d, 1928; Inger, 1954; Leviton, 1963c; Brown and Alcala, 1970a), it was not until Heaney (1985, 1986) explicitly illustrated the position of the 120 m underwater bathy metric contour in the Philippines that the land bridge implications of chan- nel depth became fully appreciated by biologists. It is now understood that five major Philippine island groups, (complexes of islands separated by shallow channels) intermittently formed much larger land mass amalgamations at various times during the mid- to late-Pleistocene (Fig. 1). It is presumed that these events allowed free exchange of fauna and flora via land-positive connections between the otherwise iso- lated islands of today. Each of the Philippine Pleis- tocene aggregate island platforms (Fig. 1) are now recognized by biogeographers as subprovinces (Tay- lor, 1928) due to the fact that each supports highly- celebrated suites of endemic taxa (Taylor, 1928; Dick- inson, 1991; Heaney and Regalado, 1998; Heaney et al., 1998; Alcala and Brown, 1998; Kennedy et al., 2000). 2001 Asiatic Herpetological Research Vol. 9, p. 35 Figure I.The Philippines (darkly shaded islands), with the major Pleistocene aggregate island platforms indi- cated by tracing of the 120 m submarine bathymetric contour (following Heaney, 1985, 1986). Star = Manilla. One unfortunate result of the prevailing perspec- tive since Heaney 's (1985) landmark paper, has been herpetologists' lack of attention to islands that are land-bridge (once connected by dry land) to well sur- veyed members of the same Pleistocene island com- plex. A case in point is Panay, a central Philippine island of 123,000 km" with several mountain peaks of more than 2000 m in elevation (Figs. 1, 2). Perhaps not unreasonably, numerous biologists over the past century have assumed or suggested that a species' presence on the well-surveyed Negros implied its undocumented presence on Panay as well (see Inger, 1954; Leviton, 1963; Brown and Alcala, 1970, 1978, 1980, 1986; Brown and Rabor, 1967; Alcala, 1986). Nevertheless, cases of low-level island endemism have been documented on individual islands within Pleistocene island complexes (Alcala, 1958, 1962; Taylor, 1920; 1922a-d; 1923; 1925; Inger, 1954; 0-150 150-400 400-800 800-1200 1200-1600 > 1600 Figure 2. Panay Island, its position in the Philippines indicated by darkened arrow (inset). Discrete shading indi- cates increasing elevational increments (see key), enclosed numbers indicate collection localities (see Table 1 ), underlined bold text indicates provinces (boundaries indicated with darkened dashed lines), and darkened circles indicate major cities. Vol. 9, p. 36 Asiatic Herpetological Research 2001 Table 1 . Study sites and collection localities for museum specimens of amphibians and reptiles from Panay Island. "No further data available" indicates specimens for which municipalities were the only available collection data in museum catalogs. The term "municipality" is synonymous with "town" but municipalities are also the political entity surrounding towns by these names. Thus, municipalities contain small surrounding villages (Barrios [="Barangays "] which may contain smaller "Sitios"). It is reasonable to assume that most collections localized to the municipality level were also collected in the town of the same name. Entries including "and vicinity" indicates specimens that may have been collected in the municipal town, or they may be from smaller surrounding Baran- gays. * Combined into a single locality due to the close proximity of collection sites at the common borders of these municipalities. Site Province Municipality Notes Ibajay Barangay Bugtong-bato Libacao Libacao town "and vicinity" Makato Makato town "and vicinity" Malinao no specific locality data Malay Nogas Point Nabas Nabas town "and vicinity" Tangalan Tangalan town "and vicinity" Malay Borocay Isl., no specific locality data Barbaza Barbaza town "and vicinity" Bugasong Bugasong town "and vicinity" Culasi Barangay Alojipan (Mt. Madja-as; PNM/CMNH site) Tobias Fornier Barangay Tobias, Villaflor Valderrama Barangay San Agustin, (Mt. Baloy and vicinity; PNM/CMNH site) San Remegio Barangay Aningalan, Aningalan mountain range Tibiao no specific locality data Caluya Caluya Isl., no specific locality data Sibay Sibay Isl., no specific locality data Caluya Semirara Isl., no specific locality data Hamtik Barangay Gunisang-an Sibalom Barangay Egaiia San Jose Barangay Bagumbayan Capiz no specific locality data Marubusao Burias lloilo no specific locality data lloilo City lloilo City "and vicinity" Ajuy no specific locality data Calinog Mt. Baloy and vicinity 1. Aklan 2. Aklan 3. Aklan 4. Aklan 5. Aklan 6. Aklan 7. Aklan 8. Aklan 9. Antique 10. Antique 11. Antique 12. Antique 13. Antique 14. Antique 15. Antique 16. Antique 17. Antique 18. Antique 19.* Antique 20. Capiz 21. Capiz 22. lloilo 23. lloilo 24. lloilo 25. lloilo 2001 Asiatic Herpetological Research Vol. 9, p. 37 Site Province Municipality Notes 26. lloilo Barotac Viejo Barangay San Francisco 27. lloilo Carles no specific locality data 28. lloilo Carles Sicogon Island; including Buaya area 29. lloilo Dingle Bulabog-Putian National Park; PNM/ CMNH Site 30. lloilo Estancia no specific locality data 31. lloilo Lambunao no specific locality data 32. lloilo Mandurriao no specific locality data 33. lloilo Pototan Pototan town "and vicinity" 34. lloilo Cabatuan no specific locality data 35. lloilo Estancia Gigante North Isl., no specific locality data 36. lloilo Estancia Gigante South Isl., Gabi area 37. lloilo Estancia Gigante South Isl., no specific locality data 38. lloilo Calinog E. Catalbac ("Calinog town") 39. lloilo Estancia Calagna-an Isl.; Barangkalan and vicinity 40. lloilo Concepcion ca 1 km NW SBS lyang Beach Resort 41. lloilo Concepcion Pan de Azucar Isl., no specific locality data 42. Negros Occidental Pulupandan Inampulagan Isl., 8 km W Munic. W. Pulupandan, Negros Occ. Prov. 43. lloilo, Subprov. Guimaras Unknown Guimaras Isl., no specific locality data 44. lloilo, Subprov. Guimaras Jordan Guimaras Isl.; Jordan town "and vicinity" 45. lloilo, Subprov. Guimaras Unknown Guimaras Isl.; 2.1 km SW Daan Banwa 46. lloilo, Subprov. Guimaras Nueva Valencia Guimaras Isl.; Nueva Valencia town "and vicinity" 47. lloilo, Subprov. Guimaras Buenavista Guimaras Isl.; Buenavista town 48. lloilo, Subprov. Guimaras Nueva Valencia Panobolon Isl.; Nueva Valencia town. 49. Unknown Unknown "Masaya" (not on available maps) 50. Unknown Unknown Panay Island: no Further Data Vol. 9, p. 38 Asiatic Herpetological Research 2001 Brown and Alcala, 1961, 1970a-b, 1978, 1980, 1982a, 1986, 1994; Brown et al, 1997a-b; Brown et al., 1999), suggesting that the two large islands war- rant individual attention by biogeographers. Despite the assumption that many widespread Philippine spe- cies were present on Panay (e.g., see Alcala, 1986), biogeographical summaries and taxonomic reviews historically have listed only nine vouchered (based on museum specimens) species of snakes, six geckos, six skinks, and seven frogs (Inger, 1954; Leviton, 1963c; Brown and Alcala, 1970, 1978, 1980) - 30% of the herpetofauna considered in the present report. Until very recently the higher elevation forested regions of Panay have not been explored by biolo- gists. At present, the results of only a few preliminary surveys in a few high elevations localities are avail- able (Gonzales and Kennedy, 1990, 1996; Sison et al., 1995; Ferner et al., 1997; Brown et al., 1997a; 1999). The purpose of this paper is to synthesize museum records that document the diversity, endemism, status, and distribution of the amphibians and reptiles from Panay and to report on several recent collections resulting from the National Museum of the Philip- pines/Cincinnati Museum of Natural History Philip- pine Biodiversity Inventory (1989-1994). Another primary goal is to draw attention to recently-discov- ered endemic species of amphibians and reptiles from Panay and to stress the need for immediate faunal sur- veys on this tropical island beset by a conservation crisis of deforestation. Material and Methods Study sites and collection locations are presented in Table 1 and shown in Figure 2. Additional site and habitat information for many TNHC, PNM, CAS, FMNH, and CMNH specimens may be found in the museum catalogs. The PNM/CMNH Philippine Biodiversity Inventory team conducted field studies at sites 11 and 13 (Figs. 3-9) and various members of the team (particularly RVS and RMB), visited other locations (Figs. 10-14) to do more limited surveying. We established elevational transects in a variety of habitat types (Ruedas et al., 1994, as modified by Brown et al. 1995a, 1996, 2000) and utilized standard collection and specimen preservation techniques (Simmons, 1987; Heyer et al., 1994). Detailed exami- nation of all material was conducted by RMB and JWF. We follow the taxonomy of Taylor (1922c), Brown and Alcala (1978, 1980) and Brown et al. (1995a; 1995b) for gekkonid and scincid lizards. The taxon- omy of Inger (1954, 1996), Frost (1985), Duellman (1993), Brown and Alcala (1994), Inger and Tan (1996), Alcala and Brown (1998) and Emerson et al. (2000), was consulted for amphibians and we include (in parentheses) the unsubstantiated taxonomic hypotheses of Dubois (1992) for reference. While no suitable taxonomy currently is available for Philippine agamids (see Taylor, 1922c), we consulted Musters (1983) and McGuire and Alcala (2000) for identifica- tion of our Draco specimens. Snake taxonomy was based on Taylor (1922a), Leviton's "Contribution to a Review of Philippine Snakes" series (Leviton 1961, 1962, 1963a-c, 1964a-d, 1965, 1967, 1968, 1970a-b, 1979, 1983, 1985), Leviton and Brown (1959), Inger and Marx (1965), Malnate and Underwood (1988), Wynn and Leviton (1993), McDiarmid et al. (1999), and Brown et al. (1999). Scale counts on snakes were taken using the methods of Dowling (1951a, 1951b). Field techniques in our surveys have been described in Ruedas et al. (1994) and Brown et al. (1995a; 1996; 2000). Museum abbreviations follow Duellman et al. (1978) and Leviton et al. (1985). In order to examine large scale faunal similarities between Panay and other large island of the Philip- pines, we assessed overall herpetofaunal simmilarities by calculating coefficients of similarity using a simple index (C - 2W/a+b; see Brown and Lomolino, 1998, for review) for the amphibians and reptiles of Panay and other islands. Our data for these calculations were based on all available literature (above) and updated conservative estimates of the taxonomy of Philippine amphibians and reptiles (Brown, Crombie, Diesmos, unpublished data). Results We present records for 20 amphibians (anurans; Figs. 15-25), and 72 reptiles (one turtle, 36 lizards, and 35 snakes; Figs 26-52) from Panay and its associated land-bridge islets. The results of faunal similarity cal- culations are presented in Figure 53. Individual spe- cies accounts, with comments on their status, are presented below. 2001 Asiatic Herpetological Research Vol. 9, p. 39 Species Accounts Amphibia Anura Bufonidae Bufo marinus (Linnaeus) (Fig. 15) Introduced on most inhabited islands of the Philip- pines (Inger, 1954; Diesmos, 1998; Alcala and Brown, 1998), this species is particularly common in agricul- tural areas and near human habitation. It is wide- spread and abundant in disturbed and agricultural areas on Panay. Localities and specimens: (Site 1) PNM 1 144 — 47, 1228-33; (Site 11) CMNH 4958-59, PNM 1289-96, 2552-54; (Site 12) PNM 1232-33; (Site 19) CMNH 5197; (Site 23) USNM 339985-86; (Site 24) FMNH 61482-84; (Site 31) PNM 1 144-47; (Site 33) USNM 339987-88, 340062. Microhylidae Kaloula conjuncta negrosensis Taylor (Fig. 16) Taylor (1920, 1922a) first collected this form on Negros and Guimaras islands. He (Taylor, 1922a) rec- ognized it as a full species but Inger (1954) placed K. negrosensis in the synonymy of Luzon's Kaloula con- juncta; no further taxonomic studies have been forth- coming. Kaloula conjuncta negrosensis is both a burrower and a climber (Taylor, 1920; Alcala, 1962, 1986); it has been found in the detritus of the forest floor (750-1075 m on Negros; Alcala, 1962) and in the leaf axils and holes of trees (Alcala 1962). They also emerge and congregate around water that collects in pools and ditches in the rainy season (Taylor, 1920, 1922a). Our Mt. Madja-as specimen was found in a tree hole 10 cm from the ground (tree < 0.5 m dbh). Specimens from near the base of Mt. Baloy were found in breeding aggregations around water buffalo wallows near a large river. We suspect that the Visayan populations represent a distinct evolutionary lineage and that they will eventually once again be recognized as a full species once new data become available (Brown and Diesmos, unpublished data). Localities and specimens: (Site 8) CAS 127890; (Site 1 1) PNM 2555; (Site 13)TNHC 56340-46; (Site 16) CAS 127510-11, 127591; (Site 18) CAS 127815; (Site 47) CAS 124446. Kaloula picta (Dumeril and Bibron) (Fig. 17) This frog is found at low elevations (100-200 m) in open areas near human habitations (Boulenger, 1882; Alcala, 1956, 1958). It is a burrowing species that conceals itself under leaf litter and topsoil until the beginning of the rainy season. Choruses may contain hundreds of individuals (Alcala, 1962; Alcala and Brown, 1998). This appears to be the first published account of this species on Panay. Localities and specimens: . (Site 16) CAS 127617- 37; (Site 18) CAS 127816, 127827; (Site 23) CAS- SU 14219-20; USNM 78079-80, 78842. Kaloula sp. In 1992, while conducting survey work at Barangay Alojipan (Site 11), we heard the distinctive honking calls of a forest species of Kaloula. Although we were unable to locate and collect specimens, we noted that the calls were superficially similar to Kaloula kalin- gensis (Taylor, 1922a; single honk, delivered approxi- mately every 15 to 20 minutes) from Luzon. Neither Kaloula kalingensis nor the related K. kokacii (Ross and Gonzales, 1991) have been reported from the nearby island of Visayas, although recent field work in the last remaining low elevation forests of Negros (Municipality of Ayungon, Negros Oriental Prov.; C. N. Dolino and A. C. Diesmos, personal communica- tion) reveals the presence of a forest species (also pre- viously unrecorded) with single honking calls there as well. Gaulke (in press) recently has discovered a pop- ulation of Kaloula in NW Panay that may be the same as that heard (but not collected) previously on Panay and Negros. Localities and specimens: none. Ranidae Limnonectesc\. leytensis (Boettger) (Fig. 18) This species is widely distributed in patches and con- sidered common in some localities on the nearby Negros (Alcala, 1986; Alcala and Brown, 1998). However, it has been collected from only one locality on Panay (Sison et al., 1995). On Negros, this species inhabits coolor high elevation mountain streams between 150 and 900 m (Alcala, 1962) and probably breeds terrestrially but deposits eggs in close proxim- ity of water (Alcala and Brown, 1956; Alcala, 1962; Brown and Alcala, 1982b). When hatching, terrestrial embryos of these and related species fall, are carried by males, or are washed into water (Alcala, 1962; see also Inger et al., 1986; Brown and Iskandar, 2000). We find it doubtful that Visayan specimens identified Vol. 9, p. 40 Asiatic Herpetological Research 2001 Table 2. List of known species from Panay and smaller, nearby, land-bridge islands and other islands within the political boundaries of major Panay Island provinces. Entries include Panay species with vouchered specimens in major museum collections (see text and species accounts for discussion of taxonomy used) and the first published accounts by authorities utilizing museum specimens. IR = island record or first published record from within major Panay island provinces. (1 Previously reported from Panay, nearby islets, or Visayan sea by Alcala (1986) but with- out specific reference to specimens. 2 See also Gaulke (in press). 3 Recorded from land-bridge islets but currently not recorded from the mainland of Panay. 4 A record from Semirara Isl., within the political boundary of Panay's Antique Province, but land-bridge to Mindoro Isl.; not likely to be discovered on Panay in the future. 5 Apparent major range extension, in need of verification or based on specimens with locality data probably in error.) Bufo marinus Kaloula conjuncta negrosensis Kaloula picta Kaloula sp. Limnonectes d leytensis Limnonectes visayanus Occidozyga laevis Platymantis corrugatus Platymantis dorsalis Platymantis negrosensis Platymantis insula tus Platymantis panay en sis Platymantis sp. 1 Platymantis sp. 2 Platymantis sp. 3 Rana cancrivora cancrivora Rana erythraea Ranad. everetti Rana vittigera Po/ypedates leucomystax Cuora amboinensis amboinensis Bronchocela sp. Draco spilopterus Hydrosaurus pus tula tus Gonocephalus sp. Cosymbotus platyurus Cyrtodactylus annulatus Inger, 1954 IR (see Taylor, 1920; 1922a)1 IR IR2 IR Inger, 1954 Inger, 1954 IR2 IR2 Sisonet al., 1995 Brown and Alcala, 1970b3 Taylor, 1920; Inger, 1954; Brown et al., 1997a IR IR IR Inger, 1954 Inger, 1954 Sison etal., 1995 IR Inger, 1954 Gaulke and Fritz, 1998 Taylor, 1922c Musters, 1983 IR2 Sison etal., 1995 Brown and Alcala, 1978 IR2 2001 Asiatic Herpetological Research Vol.9, p. 41 Cyrtodactylus philippinicus Gehyra mutilata Gekko gecko Gekko gigante Gekko mindorensis Hemidactylus frenatus Hemidactylus stejnegeri Hemiphyllodactylus insularis Lepidodactylus lugubris L epidodactylus planicaudus Brachymeles boulengeri taylori Brachymeles talinis Brachymeles tridactylus Dasia grisea Das/a semicincta Emoia atrocostata Lamprolepis smaragdina philippinica Lipinia pulchella taylori Mabuya indeprensa Mabuya multicarinata borealis Mabuya multifasciata Parvoscincus sisoni Sphenomorphus arborens Sphenomorphus coxi divergens Sphenomorphus cumingi Sphenomorphus jagori grandis Sphenomorphus steerei Tropidophorus grayi Varanus sal va tor nuchal is Acrochordus granulatus Python reticulatus Ahaetulla prasina preocularis Boiga angulata IR Brown and Alcala, 1978 Taylor, 1922c; Brown and Alcala, 1978 Brown and Alcala, 19783 Sison et al., 1995 Brown and Alcala, 1978 Sison etal., 1995 Brown and Alcala, 1978 Brown and Alcala, 19783 Brown and Alcala, 1978 IR Brown and Alcala, 1980 Brown and Alcala, 1980 Brown and Alcala, 19804 IR5 IR3 Brown and Alcala, 1980 IR Sison etal., 1995 IR IR2 Ferneretal., 1997 Brown and Alcala, 1980 IR5 IR5 Brown and Alcala, 1980 Brown and Alcala, 1980 Sison etal., 1995 Gaulke, 1991a, 1991b, 1992 IR Leviton, 1963c Leviton, 1963c, 1968 IR2 Vol. 9, p. 42 Asiatic Herpetological Research 2001 Bo/gad cynodon Bo/ga ci. dendrophila Calamaria gervaisi Cerberus rynchops Chrysopelea paradisi Cyclocorus lineatus alcalai Dendrelaphis caudolineatus terrificus Dendrelaphis pictus p ictus Elaphe erythrura psephenoura Gonyosoma oxycephala Hologerrhum dermali Lycodon aulicus capucinus Oligodon modestum Psammodynastes pulverulentus Pseudorabdion mcnamarae Pseudorabdion oxycephalum Pseudorabdion talonuran Tropidonophis negrosensis Zaocys luzonensis Calliophis calligaster gemianulis Hydrophis belcheri Hydrophis cyanocinctus Hydrophis elegans Hydrophis inornatus Lapemis hardwickii Laticauda colubrina Ramphotyphlops braminus Rhamphotyphlops cumingii Typhlops castanotus Typhlops hypogius ( = T. ruber ? ) Tropidolaemusc\ wagleri Trimereserus flavomaculatus IRZ IR Leviton, 1963c; Ingerand Marx, 1965 Gyi, 1970 Sisonetal., 1995 IR Leviton, 1970b Leviton, 1963c, 1970b Leviton, 1979 IR2 IR2 Leviton, 1965 Sisonietal., 1995 IR2 Sisonietal., 1995 IR Brown etal., 1999 Leviton, 1963c; Malnate and Underwood, 1988 Ross etal., 1987 Leviton 1963b, 1963c IR1 IR1 IR IR IR1 IR IR IR Wynn and Leviton, 1 993 IR? (see McDiarmid et al., 1999) IR Gaulke (in press) 2001 Asiatic Herpetological Research Vol. 9, p. 43 as Rana cf leytensis are conspecific with specimens referred to this species from the Mindanao Aggregate Island Complex (Leyte, Samar, Bohol, and Mind- anao). Locality and specimens: . (Site 25) PNM 1 1 14-15. Limnonectes visayanus (Inger) (Fig. 19) This large fanged frog is found in clear forest streams; it is known to hide in rock crevices during the day and call from rocks and stream banks above water at night (Alcala, 1962; Alcala and Brown, 1998). We found L. visayanus at low elevations on rocks in large rivers near Mt. Madja-as. This species may breed and lay eggs outside of water (Alcala, 1962). Localities and specimens: (Site 2) PNM 1715-20; (Site 3) CAS 137592-95, 137590-91, 139164-66, USNM 305671-76; (Site 4) PNM 1613-20, 1623-27; (Site 6) CAS 137596-98; (Site 7) PNM 1799-800, 1828-31, 1836-39, 1845, 1855-60, 1865-77; (Site 1 1 ) CMNH 4894-98, 4899, PNM 1 302-06, 26 17-21; (Site 13) TNHC 56337; (Site 14) PNM 3710-12, 3732, 3764-68, 3805; (Site 25) 1085-92, 1133-38, 1140; (Site 28) CAS 124093-106, 124442-14, 124950-58; (Site 39) CAS 124121, 124293-97; (Site 44) CAS 125308-309, 125312; (Site 47) CAS 125302-307; (Site 50) USNM 78072-78. Occidozyga laevis (Gunther) (Fig. 20) Occidozyga laevis is found in flooded fields in agri- cultural areas, in road-side ditches and open sewers, and in streams and rivers from lowlands to high eleva- tion forested sites (Inger, 1954; Alcala, 1962; pers. obs.). On Negros this species has an altitudnal range of sea level to 1 150 m (Alcala, 1962). Specimens are common in stream-side pools along larger rivers in forested areas near Mt. Madja-as and Mt. Baloy. Localities and specimens: . (Site 1 ) PNM 1 1 10-11, 1113, 1116-29, 1141, 1184-99; (Site 2) PNM 1116- 29, 1690-1714, 1721-23, 1731, 1757-58; (Site 3) CAS 137586-88, 139148, 139167-68, USNM 305647-48; (Site 4) PNM 1600-02, 1611, 1621-22; (Site 6) CAS 137614-15, USNM 305649, 305650- 54; (Site 7) PNM 1832-35, 1841-44, 1848-54, 1862- 64, 1880; (Site 9) PNM 1156, 1165-68, 1170-71; (Site 10) PNM 1110-11, 1113; (Site 11) CMNH 4951-57, PNM 1329, 2655-61; (Site 12) PNM 1 163- 64; (Site 14) PNM 3730-31, 3782, 3804; (Site 22) CAS-SU 9813; (Site 23) CAS-SU 14224-25, 14373; (Site 24) FMNH 61478-81; (Site 25) PNM 1141; (Site 26) PNM 1160-62; (Site 27) CAS-SU 14049; (Site 28) CAS 124959-70, 124059-76, 124426, 124432-33, 124439; (Site 30) CAS-SU 14223; (Site 32) PNM 1172-73; (Site 38) CAS 132880, 132887- 901, 134089-96; (Site 39) CAS 124171, 124190-91; (Site 40) USNM 339989; (Site 41) CAS 125001, 124177, 124184-85, 124194-96; (Site 42) CAS-SU 23946-49. 23952-58, 23961-63; (Site 44) CAS 125361-62; (Site 47) CAS 125311, 125344-59. Platymantis corrugatus (Dumeril) (Fig. 21) This widely distributed terrestrial frog inhabits the forest floor from sea level to above 1300 m (Alcala, 1986). On Mt. Madja-as we found P. corrugatus in leaf litter and in limestone crevices. This account and that of Gaulke (in press) appear to be the first pub- lished records of this species from Panay. Localities and specimens: (Site 6) CAS 137616-19, 139149, 185494; (Site 11) CMNH 4960-63, 5118, PNM 2556-59; (Site 13) CMNH 3160-65; (Site 25) PNM 1 103-06; (Site 28) CAS 124058. Platymantis dorsalis (Dumeril) This common forest frog is found in the detritus of the forest floor as well as in tree cavities and low tree ferns (Alcala, 1962; Alcala and Brown, 1998). Although recent studies (Brown et al., 1999) indicate the presence of numerous cryptic species in the P. dorsalis complex on Luzon (and we suspect that fur- ther new species await discovery in the Visayan islands), the calls of some P. dorsalis have been heard on Mt. Baloy and Mt. Madja-as are, at least superfi- cially, similar to the short, whistling, ascending fre- quency sweep of true P. dorsalis from Luzon Island (Brown et al., 1997c). Thus, while we expect that more species in the dorsalis complex will soon be dis- covered in the Visayas, we can confidently assert that at least some Panay populations are indistinguishable from P. dorsalis of Luzon Island (Brown et al., 1997c; 1999). This account and that of Gaulke (in press) appear to be the first published records of this species from Panay. Localities and specimens: . (Site 2) PNM 1734—56; (Site 6) CAS 137620-40, 137649-51, 139150-63, USNM 305655-70; (Site 1 1 ) CMNH 4964-98, 5206, PNM 2559, 2562-88; (Site 13) TNHC 56347-50; (Site 14) PNM 3713-19, 3729, 3733^45, 3756-62, 3772-74, 3783-85, 3788-89, 3791-96, 3830-31, 3857, 3860, 3862-65, 3882-83, 3886-88, 3893-95, 3906-08; (Site 25) PNM 1093-102; (Site 28) CAS 124419, 124428-31, 124440-41, 124689-91; (Site 39) CAS 124123-33, 124146-47; (Site 41) CAS 124041^12, 124122, 124186-89, 125014-19. Vol. 9, p. 44 Asiatic Herpetological Research 2001 Platymantis negrosensis Brown, Alcala, Alcala, and Diesmos This recently-described forest frog (Brown et al, 1997b) has only been documented from two sites on Panay but is also known from localities on the nearby island of Negros, from approximately 300 to 1625 m in elevation (Alcala, 1958; Alcala and Brown, 1957; Alcala, 1962). On that island, this species occupies arboreal microhabitats in primary forest (Brown and Alcala, 1961; Alcala, 1962; Brown et al, 1997b). We documented this island record for Panay (Sison et al., 1995; then recognized as P. guentheri) from speci- mens collected in forest on Mt. Baloy at about 950 m. This species is related to P. luzonensis but differs in characteristics of the advertisement call an external morphology (Alcala and Brown, 1998). Localities and specimens: . (Site 13) CMNH 3166; (Site 14) PNM 3889. Platymantis insulatus Brown and Alcala (Fig. 22) A frog known from primary and secondary forest situ- ated on karst limestone outcrops, this species was originally discovered (Brown and Alcala 1970b) on the forest floor and in the open mouths of small caves at low elevations (Alcala and Brown, 1998) on the island of Gigante South off Panay's northeast coast. A recent (June 2000) visit to the type locality by R. Brown and A. Alcala confirmed this species' persis- tence despite the complete removal of the original for- est. The presence of an endemic species on such a small, land-bridge island is puzzling and suggests that it may still be (or at least, may have once been) present on eastern Panay. Unsurveyed limestone for- mations along the northeastern coast are the most promising possibility for locating this species on Panay. Localities and specimens: . (Site 35) CAS 157235- 39; (Site 36) CAS 137641-12; (Site 37) CAS 117440-41; 119967-69, MCZA-72946. Platymantis panayensis Brown, Brown and Alcala Only recently described (Brown et al., 1997a) from our collections from Mt. Madja-as, this species is closely related to P. hazelae from Negros and occu- pies similar microhabitats in high elevation cloud for- ests. Taylor (1920:101) apparently had a specimen (collected by R. McGregor) of this species on hand during the description of Philautus (= Platymantis) hazelae and he considered it conspecific with the Negros population. On the basis of Taylor's (1920) account, Inger also (1954) included P. hazelae in his list of species from Panay . We collected the majority of the type specimens of this species from leaf axils, leaves on shrubs, and the leaf litter on the forest floor. The call has not yet been formally described, but consists of a pure, ringing, tonal note with no frequency or amplitude modula- tion; it sounds to the human ear like the sound pro- duced by the ringing of a small bell (1-2 notes/s). Localities and specimens: (Site 6) CAS 1 37641^42; (Site 11) PNM 2314-18, 2495, 2589-90; CMNH 4113-15,4116-20,4868-69. Platymantis sp. 1 Several immature specimens of a tiny, dark brown, tuberculate (1 1-15 mm SVL) species of Platymantis were taken at high elevations on Mt. Madja-as and on Mt. Baloy. They appear most similar to Platymantis pygmaeus of the Sierra Madre mountains of Luzon's east coast. Due to the fact that the available specimens are all sexually immature, we cannot recognize them taxonomically until further material and recordings of advertisements calls become available. Localities and specimens: (Site 11) CMNH 8132 (Site 1 3) CMNH 3 1 73-74, 3177. Platymantis sp. 2 Two immature specimens of a diminutive, black, smooth-skinned (12-13 mm SVL) species of Platy- mantis were collected at 1450 m from under leaf litter. The small sample size and immaturity of the speci- mens necessitates that taxonomic recognition of the species must await the collection of further material. Locality and specimens: (Site 1 1) CMNH 8133-34. Platymantis sp. 3 This unidentified species is represented by a single, very large, black specimen with two dorsolateral light lines; it is appears possibly related to P. pseudodorsa- lis from Luzon (Brown et al., 1999). Locality and specimen: (Site 6) CAS 185495. Rana (= " Fejervarya") cancrivora cancrivora Gravenhorst This common frog is found in swamps, ponds, flooded rice fields and ditches (Inger, 1954). It is found in almost any pool of water at low elevations (Alcala and Brown, 1998). Inger (1954) first reported it on Panay Island. Localities and specimens: . (Site 1) PNM 1178-82; (Site 8) CAS 127893-95, 127899-904; (Site 9) PNM 2001 Asiatic Herpetological Research Vol. 9, p. 45 1156; (Site 16) CAS 127509, 127559-81, 127611, 127678-79; (Site 17) CAS 127800; (Site 18) CAS 127801-803; (Site 19) PNM 2607-16; (Site 22) USNM 78862-902; (Site 23) CAS-SU 14259-60, 14452-79, 15486-514, USNM 77984-96, 77988- 78042, 78048, 78062-65; (Site 24) FMNH 61398- 402; (Site 26) PNM 1160-62; (Site 27) CAS-SU 14513-25; (Site 28) CAS 124262-71, 185665-69; (Site 30) CAS-SU 14252-57; (Site 34) CAS-SU 9762-63; (Site 36) CAS 124343; (Site 37) CAS 124564-76; (Site 38) CAS 132878-79; (Site 39) CAS 124135, 124175; (Site 40) USNM 339990, 340059- 60; (Site 41) CAS 124178-79, 124320-28, 125020- 28; (Site 42) CAS-SU 23966; (Site 44) CAS 125 194- 96; (Site 46) CAS 125183-93; (Site 47) CAS 125174-82; (Site 48) CAS 124791-92; (Site 50) USNM 78066-71. Rana(= " Hylarana") e/>tf/vae,3(Schlegel) (Fig. 23) This widely-distributed and common frog is believed to have been introduced to the Philippines (Diesmos, 1998; Alcala and Brown, 1998), originally on Negros (Inger, 1954; Alcala, 1962; Alcala and Brown, 1998). It's presence on Panay has been previously docu- mented (Taylor, 1920; Inger, 1954). We found our specimens along the grassy boarders of flooded rice fields. Localities and specimens: (Site 2) PNM 1725-26; (Site 3) CAS 137589; (Site 4) PNM 1603-10, 1612; (Site 7) PNM 1840; (Site 8) CAS 127891-92, 127905-912; (Site 10) PNM 1174-76; (Site 11) CMNH 4870, PNM 1309-28, 2591-92; (Site 14) PNM 3816; (Site 19) PNM 2622-54; (Site 22) CAS- SU 9744; (Site 23) CAS-SU 14537-47, USNM 77730-983, 78043-47, 78049-61; (Site 24) FMNH 61391-97, 61449—52; (Site 27) CAS-SU 14578-84; (Site 30) CAS-SU 11120-24, 14526-36, FMNH 40527; (Site 33) USNM 38650-54; (Site 38) CAS 132881-86,' 134086-88; (Site 39) CAS 124209-212, 124214; (Site 44) CAS 125158-69; (Site 47) CAS 124143-57, 125310; (Site 50) USNM 77617-729, 78413-38. Rana{- " Chalcorana") cf. et-weff/Boulenger Species of the Rana everetti complex are found in and along streams from 300 to about 1300 m (Inger, 1954; Brown and Alcala, 1955; Alcala, 1962; Alcala, 1986; Brown et al., 2000). Our Mt. Baloy expedition in 1989 first recorded the presence of this frog on Panay (Sison et al., 1995). Negros populations (expected to be conspecific with those on Panay) are only found near water during breeding; this species has most often been collected in overhanging, streamside vege- tation (Brown and Alcala, 1955; Alcala, 1967). As noted by Brown et al. (2000), the taxonomic status of the Visayan populations referred to Rana everetti is in need of further study. Localities and specimens: . (Site 2) PNM 1732-33; (Site 14) PNM 3771, 3800-03, 3806-14, 3817-24, 3896, 3913; (Site 39) CAS 124213, 124215-16. Rana (= " Fejervarya") vittigera Wiegmann (Fig. 24) This species occurs in open, agricultural areas near sources of water (ponds, flooded rice fields). The dis- tinctive, rapid honking call of this species can be heard in choruses of up to hundreds of individuals. This record appears to be the first published account of this species from Panay. Localities and specimens: . (Site 11) CMNH 4871 — 72, PNM 2593-94; (Site 19) PNM 2596-606; (Site 40) USNM 339991; (Site 41) CAS 124197; (Site 46) CAS 125360. Family Rhacophoridae Polypedates leucomystax (Gravenhorst) (Fig. 25) This common tree frog is widely distributed in Panay (Inger, 1954; Alcala, 1986) in agricultural areas, for- est edges, and disturbed forests. It ranges from near sea level to 1000 m on Negros (Alcala. 1962) and was first documented on Panay by Inger ( 1954). Our spec- imens were collected in banana plantations and rice fields near the base of Mt. Madja-as and Mt. Baloy. Localities and specimens: . (Site 1) PNM 1107-09, 1209-27; (Site 2) PNM 1729-30; (Site 6) CAS 137599; (Site 7) PNM 1797-98; (Site 11) CMNH 4997-98, PNM 2560-61; (Site 13) TNHC 56336, 56338; (Site 14) PNM 3763, 3799; (Site 16) CAS 127512-14, 127540-58, 127592-95, 127608-10, 127647^19, 127657, 127670-76, 127680-97, 127701; (Site 17) CAS 127721-26; (Site 18) CAS 127841; (Site 24) FMNH 61485; (Site 25) PNM 1107-09; (Site 28) CAS 124110-13, 124420, 124422-25, 124581; (Site 29) USNM 339992-93; (Site 30) CAS-SU 11113-16, 14764, FMNH 40569, 44263; (Site 37) CAS 124114, 125034-37; (Site 39) CAS 124158, 124192, 124204-07; (Site 40) USNM 339994; (Site 41) CAS 124176, 125031-32; (Site 42) CAS-SU 23950-51, 23959-60, 23964-65; (Site 47) CAS 125342-43. Vol. 9, p. 46 Asiatic Herpetological Research 2001 Reptilia Testudines Bataguridae Cuora amboinensis amboinensis (Daudin) (Fig. 26) This common species is found throughout the Philip- pines (Gaulke and Fritz. 1998) and on Panay (Gaulke, in press) in low elevation forests, agricultural areas, and near streams and swampy areas; it is generally considered nocturnal (Alcala, 1986). We have col- lected this species in stream-side habitats at low ele- vations on Panay. Localities and specimens: (Site 7) PNM 1888-89, 1891-95; (Site 11) CMNH 5500, 5501, 5502; PNM 1288, 5657-58; (Site 23) USNM 78103-04, 78746- 49; (Site 41) CAS 153872; (Site 45) CAS 185507; (Site 50) USNM 78081-102. Squamata (Lizards) Agamidae Bronchocela cristatella (Kuhl) and B. marmo- rata (Gray) These arboreal lizards (Alcala, 1986) are found from lowland cultivated areas to lower midmontane pri- mary and secondary forests; they are most often encountered sleeping at night in stream-side vegeta- tion. No suitable taxonomic reference is available for Philippine Bronchocela and so the true identities of Panay specimens referred to B. cristatella and B. mar- morata are uncertain. There is little consensus regard- ing the identity of Bronchocela throughout the Visayas. Although specimens from Negros and Panay key out to earlier descriptions of both Bronchocela marmorata and B. cristatella (Taylor, 1922c; Alcala, 1986), both "species" appear to be highly variable and diagnostic characters vary ontogenetically. We con- sider it unlikely that two independent lineages occur in sympatry on Panay and, at present, we hold in abeyance the identity of these populations until a thor- ough taxonomic revision is available. Localities and specimens: . (Site 6) CAS 137605; (Site 22) CAS-SU 10948; (Site 23) USNM 77133- 38, 78105-107; (Site 41) CAS 124333; (Site 47) CAS 125337-38. Draco spilopterus (Weigmann) (Figs 27, 28) This species is common at lower to mid-montane ele- vations and often is found in coconut groves and for- est edges (Alcala, 1986; McGuire and Alcala, 2000). Draco spilopterus is the only Draco species currently recognized from the Visayan and Luzon aggregate island complexes, despite biogeographic and morpho- logical evidence suggesting that Luzon and Visayan populations constitute independent evolutionary lin- eages (Taylor, 1922c; Heaney, 1985, 1986; McGuire and Alcala, 2000). Draco spilopterus may warrant further taxonomic attention once biochemical studies of species boundaries become available (Taylor, 1922c; McGuire and Alcala, 2000). Localities and specimens: . (Site 3) CAS 137578, 185504; (Site 4) PNM 1628-38; (Site 6) CAS 137608-609, 185505; (Site 7) PNM 1759-79; (Site 8) CAS 127886, 127916, 127961, 128031; (Site 11) PNM 1275-82, 2720-21; (Site 14) PNM 3769-70, 3878; (Site 18) CAS 127851-52; (Site 19) TNHC 58465-67, 58471-80, 58482-90, 58850; (Site 43) CAS 39686; (Site 44) CAS 125295, USNM 38990- 96; (Site 47) CAS 125277-94. Hydrosaurus pustulatus (Eschscholtz) (Figs. 29, 30) The Mt. Madja-as specimens were collected in over- hanging stream-side vegetation. On Mt. Baloy we also collected a specimen in similar riparian habitats. These large omnivorous agamids can be found on trees and shrub-layer vegetation, overhanging streams and rivers (Alcala, 1986). The taxonomy of Philippine Hydrosaurus is in need further taxonomic studies. Localities and specimens: (Site 11) CMNH 5043; (Site 13) TNHC 56762; (Site 22) USNM 77091-103, 85073-74; (Site 44) CAS 125336, USNM 38988-89; (Site 50) 77104-28, 78168-87. Gonocephalus sp. This genus was reported as an island record for Panay from site 2 by Sison, et al. (1995). The name G. sophiae has been applied to Negros populations (Tay- lor, 1922c; Alcala, 1986) and we might expect that if Negros populations are indeed G. sophiae (and this name applies to a distinct lineage that is independent from G. semperi), specimens from Panay might be referable to this species as well (see Gaulke, in press). At present, no suitable taxonomic reference exists, and species boundaries are unclear. Due to this fact, we do not apply a specific epithet to this population. Philippine populations of the genus Gonocephalus are greatly in need of taxonomic review. 2001 Asiatic Herpetological Research Vol. 9, p. 47 Localities and specimens: (Site 2) PNM 1130-32; (Site 14) PNM 3858. Gekkonidae Cosymbotus platyurus (Schneider) We found specimens of this common house gecko species in both the city and in agricultural areas; they are widely distributed on Panay (Brown and Alcala, 1978) and are usually encountered under lights. Localities and specimens: (Site 7) PNM 1803, 1815-16; (Site 8) CAS 128020; (Site 11) CMNH 5137, PNM 1261-74; (Site 19) CMNH 5089-96, 5098-104, PNM 2722-36; (Site 22) USNM 78776- 832, 103480; (Site 23) USNM 77144-57, 339998- 340010; CAS-SU 9613; (Site 26) PNM 1234-14, 1247, 1249; (Site 29) USNM 339995-97; (Site 34) CAS-SU 9612, 12021; (Site 40) USNM 34011-12; (Site 50) FMNH 41302. Cyrtodactylus annulatus (Taylor) (Fig. 32) Brown and Alcala (1978) and Alcala (1986) report this species in forested areas ranging from sea level to 1200 m on the nearby land-bridge islands of Negros, Cebu, and Inampulugan (Site 42) and its presence on Panay is not surprising. Nevertheless, this report and that of Gaulke (in press) appear to be the first records of this species from Panay. Brown and Alcala (1978) reported that this species has been collected on the forest floor, from under logs, beneath bark, and on the trunks of trees. We found Panay specimens in second- ary forest on root masses of trees protruding through eroding banks along a large river at the base of Mt. Madja-as. Predation on this species by flying snakes (Chrysopelea paradisi) was observed in these same microhabitats. Locality and specimens: (Site 28) CAS 124614-15, 124768-78 (Site 42) CAS-SU 28009-10, 28013-14, 28016-19, 28031, 28036-41, 28044-46,28050-60. Cyrtodactylus philippinicus (Steindachner) (Fig. 33) This species of Cyrtodactylus is found in a variety of habitats in the forest, particularly in rotting logs (Brown and Alcala, 1978; Alcala, 1986) and has been collected from sea level to nearly 1200 m (Brown and Alcala, 1978). In the Visayas, this species also has been collected on the nearby land-bridge islands of Negros, Pan de Azucar, and Boracay but never before on Panay. The specimens from Mt. Madja-as were collected in primary forest, during the day, under loose tree bark. Localities and specimens: .(Site 6) CAS 137607; USNM 496868; (Site 8) CAS 127883; (Site II) CMNH 5125-28, PNM 2751-53; (Site 13) TNHC 56339; (Site 18) 127864, 127869; (Site 28) CAS 124783-84; (Site 41) CAS 124046, 124780-82. Gehyra mutilata (Weigmann) (Fig. 34) This common and widely-distributed lizard is consis- tently found not only around human habitation (in darker areas, away from lights), but also in gardens and forested areas on trees (Brown and Alcala, 1978; Alcala, 1986). Localities and specimens: (Site 2) PNM 1682-86; (Site 3) CAS 137579; (Site 4) PNM 1649-53; (Site 7) PNM 1824; (Site 8) CAS 127888, 127922-29, 127948-52, 127964, 127975-76, 127999-128000, 128054-55; (Site 11) CMNH 5105-16, 5198, PNM 1250-58, 2737-46; (Site 16) CAS 127504, 127607; (Site 18) CAS 127804, 127866; (Site 19) PNM 2737- 46; (Site 23) USNM 77158, 78834-35; (Site 26) PNM 1245-49; (Site 28) CAS 124434-36, 124616- 19; (Site 29) USNM 340013-14; (Site 37) CAS 125029; (Site 39) CAS 124118-120, 124161-170; (Site 40) USNM 340015; (Site 41) CAS 124180-83; (Site 42) CAS-SU 28012, 28047-49; (Site 44) CAS 124505-510; (Site 46) CAS 124687-88; (Site 47) CAS 124682-86, 125129-31; (Site 48) CAS 124511- 12; (Site 50) FMNH 41383. Gekko gecko (Linnaeus) This common species is found around human habita- tion and in forest adjacent to disturbed areas. We col- lected specimens in secondary forest near the base of Mt. Madja-as. Localities and specimens: (Site 1) PNM 1056-63; (Site 2) PNM 2667; (Site 7) PNM 1062-1063, 1792- 96; (Site 8) CAS 128004; (Site 11) CMNH 5018-25, PNM 1282, 1330, 2662-69; (Site 16) CAS 127582; (Site 17) CAS 127745; (Site 19) PNM 2665; (Site 23) CAS-SU 9585-88; USNM 77142-43, 340018-19; (Site 28) CAS 124979; (Site 29) USNM 340016-17; (Site 31) PNM 1 143; (Site 35) CAS 124393, 124318, 124866-75; (Site 37) CAS 124315-17, 124929^19; (Site 39) CAS 124389-92; (Site 41) CAS 124319, 124580; (Site 42) CAS-SU 27929; (Site 44) CAS 125251; (Site 46) CAS 125249-50; (Site 47) CAS 125247-48; (Site 49) CAS-SU 9589; (Site 50) FMNH 41377-81, 41376. Gekko gigante Brown and Alcala (Figs. 35, 36) Gekko gigante was described by Brown and Alcala (1978) from the tiny land-bridge islands of Gigante Vol. 9, p. 48 Asiatic Herpetological Research 2001 North and Gigante South, off the northeast coast of Panay (Fig. 2). This species has not been studied since its 1968 discovery until a recent visit to the type locality by RMB and A. Alcala in June 2000. We found G. gigante in small sea-side caves in karst lime- stone outcrops along the south coast of Gigante South. The northeast coast is the best place to survey for this species on Panay if suitable limestone habitat can be located. Localities and specimens: (Site 35) CAS 124318, 124866-75 (Site 37) CAS 124315-17, 124929-49. Gekko mindorensisTay\or (Fig. 37) Brown and Alcala (1978), and Alcala (1986) report that individuals of this species are common on walls of caves, on tree trunks, and in leaves around the but- tresses of trees. Elsewhere in the Visayas this species has been reported from Negros, Cebu, and Caluya (Brown and Alcala, 1978). We have caught this spe- cies on cement walls, abandoned buildings in the for- est, in road-cut culverts, and on large dead tree trunks and stumps. Sison et al. (1995) reported this as an island record from Site 29. The taxonomic distinctive- ness of this species requires verification; recent data suggest the widespread G. mindorensis may be con- specific with G. kikuchii from Taiwan (Crombie and Ota, unpublished data; see also comment by Taylor, 1922c). Localities and specimens: (Site 8) CAS 127882, 127884-85, 128021; (Site 1 1) One uncataloged speci- men, deposited in PNM; (Site 16) CAS 127700; (Site 18) CAS 127817; (Site 28) CAS 124767; (Site 29) PNM 2500; (Site 39) CAS 124136. Hemidactylus frenatus Schlegel in Dumeril and Bibron This common house gecko is widespread on Panay and often is associated with Cosymbotus platyurus (Alcala, 1986) in well lighted areas in human habita- tion. We collected several specimens near the base of Mt. Madja-as on walls of houses in agricultural areas. Localities and specimens: (Site 3) CAS 137580, USNM 496869-70; (Site 7) PNM 1801-02, 1804-14; (Site 8) CAS 127878-81, 127897, 127913-14, 127930-45, 127953-58, 127963, 127977-78, 128001-03, 128022-26, 128034-35, 128056-57, 136742^14; (Site 11) CMNH 5148-54, PNM 1259- 60, 1646-48, 2771-76; (Site 16) CAS 127505-07, 127515-16, 127583, 127615-16, 127642^*6, 127652-56, 127658-59, 127699; (Site 17) CAS 127710-20, 127727-M, 127748-65, 127795-99, 136741; (Site 18) CAS 127805-14, 127818-26, 127829-10, 127848-50, 127865; (Site 19) CMNH 5129-36, 5139^17, PNM 2754-70; (Site 23) USNM 78833, 78836-39; (Site 26) PNM 1245-1246; (Site 28) CAS 124081-92, 124427, 124663, 125002-12; (Site 29) USNM 340021-27; (Site 33) USNM 340028-32; (Site 35) CAS 124363, 124504; (Site 37) 124364-72, 124357-62, 124582-610, 124662; (Site 39) CAS 124137^3, 124217^2; (Site 40) 340034- 41; (Site 41) CAS 124373-85, 124664-81, 125013; (Site 42) CAS-SU 28007-08, 2801 1, 28021, 28034- 35, 28042-43; (Site 44) CAS 124466, 124526-58 [Nadulao Island]; (Site 46) CAS 124458-65, 125315- 30, 124736-66; (Site 47) CAS 124448-57, 125128, 125313-14; (Site 50) FMNH 41384-86. Hemidactylus ste/negeri 0\a and Hikida Previously referred to H. garnoti (Brown and Alcala, 1978). the status of Philippine populations was clari- fied by Ota and Hikida (1989) and Ota et al. (1993). Sison et al. (1995) reported this species as an island record for Panay. Locality and specimen: (Site 25) CMNH 3225. Hemiphyllodactylus insu/arisJay\or While Brown and Alcala (1978) referred Philippine (and Panay Island) populations of Hemiphyllodacty- lus to H. typus, collections in USNM contain males and are not, therefore, the widespread unisexual H. typus, but probably are better referred to Taylor's H. insularis (R. Crombie, personal communication.). Localities and specimens: . (Site 3) CAS 137581- 83; (Site 8) CAS 127889, 127965-71; (Site 18) CAS 127855-57. Lepidodactylus lugubris (Dumeril and Bibron) This lowland gecko is common in patchy distributions and found in leaf axils, under bark, in tree holes, in coconut groves, and associated with large rocks in coastal areas (Brown and Alcala, 1978; Alcala, 1986). Though reported from Negros, Cebu, Inampulugan, Boracay, Gigante, and Pan de Azucar (Brown and Alcala, 1978), this species has not yet been reported from Panay. Localities and specimens: (Site 8) CAS 127947, 127959, 127979, 127982-87, 127989, 128005-07, 128010, 128012-14, 128017, 128043-44, 128046- 50, 128059, 154716; (Site 28) CAS 124579; (Site 37) CAS 124200, 125030, 125033; (Site 39) CAS 1241 16-17; (Site 41) CAS 124981-125000; (Site 42) CAS-SU 28015, 28061; (Site 44) CAS 124561-63; (Site 46) CAS 124492. 2001 Asiatic Herpetological Research Vol. 9, p. 49 Lepidodactylus planicaudus Stej neger Alcala and Brown (1978) recorded this species in coconut trees, mangroves, and in fern axils; they have also been taken on tree trunks in forests from sea level to 700 m. Elsewhere in the Visayan Aggregate Island Complex it is known from Cebu, Guimaras, Caluya, Masbate and Boracay (Brown and Alcala. 1978). Localities and specimens: (Site 3) CAS 139939; (Site 8) CAS 127921, 127980-81, 127988, 127990, 128008-09, 128011, 128015-16, 128018-19, 128045, 128051-53; (Site 16) CAS 127698; (Site 17) CAS 127709; (Site 39) CAS 124115, 124134; (Site 46) CAS 124823-26. Scincidae Brachymeles boulengeri taylon 'Brown (Figs. 38, 39) Usually encountered in agricultural areas (especially coconut plantations) adjacent to forest, this species most often is collected from its preferred microhabi- tat, inside rotting coconut logs. It has also been col- lected in mature and secondary forest, from sea level to 1200 m on numerous islands in the Visayas (Brown and Alcala, 1980) but this is the first record from Panay. Localities and specimens: (Site 1) PNM 1 148; (Site 25) PNM 1 148; (Site 39) CAS 124157; (Site 41) CAS 124044 (Site 42) CAS 27930-31, 27946-51, 27953. 27973-84, 27987-93. Brachymeles talinis Brown This skink occupies decaying vegetation and humus on the floor of primary and, occasionally, secondary forest (Brown and Alcala, 1980; Alcala, 1986). Alcala and Brown (1980) reported its elevational range from sea level to approximately 1200 m. Localities and specimens: (Site 6) 137603; (Site 8) CAS 127870-75, 127962, 128029-30, 154689; (Site 14) PNM 3852, 3856, 3859, 3909-10; (Site 16) CAS 127517-39, 127584-90, 127606, 127663-68; (Site 28) 154692, 200522-31; (Site 39) CAS 124148, 200521; (Site 42) CAS-SU 27972, 27996-97; (Site 50) CAS 137603. Brachymeles tridactylus Brown (Fig. 40) This species has been collected under logs, humus, and other debris in forested regions from sea level to approximately 900 m (Brown and Alcala, 1980; Alcala, 1986). On Mt. Madja-as we found them under logs in well-regenerated second growth forest below 200 m. Localities and specimens: (Site 3) CAS 137566-75; (Site 6) CAS 137602, 137604; (Site 8) CAS 127876- 77, 127915, 127974, 128027-28, 128037^12; (Site 11) PNM 5514-15; (Site 42) CAS-SU 27950, 27952, 27985-86, 28456. Das/a grisea (Gray) In the Philippines, this species has been recorded from the islands of Mindoro, Luzon, and Marinduque (Brown and Alcala, 1980). Its inclusion in this paper is based on a single record from Semirara Island, a small island south of Mindoro that is included in the political boundary of Antique province but is on the land-bridge to Mindoro. We do not expect this species to be discovered on Panay and we do not regard it as a Visayan Aggregate Island complex resident. Locality and specimen: (Site 18) CAS 134218. Dasia semicincta (Peters) A large lowland species, previously reported only from Mindanao in the Philippines (Brown and Alcala, 1980; Alcala. 1986). Outside of the Philippines D. semicincta is known from Borneo (Brown and Alcala, 1980). This is the first record of this species from Panay; it also is a substantial range extension in need of verification. Locality and specimen: (Site 23) USNM 78840. Emoia a trocostata (Lesson) This species of skink is common in mangroves and can be found active on tree trunks, in tree holes, and in rock crevices (Alcala and Brown, 1967; Brown and Alcala, 1980; Alcala, 1986). Taylor (1922c) reported that this species swims in brackish water, burrows in sand, and feeds on small crabs. On Panay and Negros, mangrove forests have nearly all been cleared, possi- bly rendering this species rare in recent collections. Localities and specimens: (Site 16) CAS 127638- 39; (Site 18) CAS 127842-45; (Site 22) CAS-SU 13585; (Site 23) USNM 77139, 7841 1, 78841, 80934; (Site 37) CAS 125040; (Site 41) CAS 124329-32, 124620-24; (Site 42) CAS-SU 27920; (Site 47) CAS 125335; (Site 50) USNM 77141. smaragdina philippinica Lamprolepis (Mertens) This skink is common in coconut groves and gardens and a variety of natural vegetation types (Alcala, 1986). Near Mt. Madja-as and Mt. Baloy we observed them on coconut trunks at low elevations. Localities and specimens: (Site 4) PNM 1639; (Site 7) PNM 1780. 1782, 1786; (Site 8) CAS 127887, Vol. 9, p. 50 Asiatic Herpetological Research 2001 127917-20, 127946, 127991-98, 128058, 128060- 61; (Site 11) CMNH 5038^0, PNM 1283, 2681-82; (Site 14) PNM 3798; (Site 16) CAS 127508, 127612- 14, 127640-41, 127669; (Site 17) CAS 127747, 127794; (Site 18) CAS 127853-54; (Site 19) TNHC 56472-73; (Site 28) 124053-57; (Site 35) CAS 124198, 12471 1-13; (Site 39) CAS 124201-02; (Site 40) USNM 340061; (Site 42) CAS-SU 27954-62, 27969; (Site 44) CAS 125301; (Site 46) 125298-300; (Site 47) CAS 124447, 125296-97. Lipinia pulchella taylon '(Brown and Alcala) This widely distributed arboreal skink had not been reported from Panay prior to our visits to Mt. Majda- as. Previously, L. p. taylori had only been known from Negros (Brown and Alcala, 1980). Brown et al. (1996; 2000) have collected specimens from Luzon that key out to this subspecies, potentially indicating the need for a review of Brown and Alcala's (1980) taxonomy of this species. This is a new island record for Panay. Localities and specimens: (Site 11) CMNH 5083; (Site 25) PNM 1156. Mabuya indeprensa Brown and Alcala Sison et al. (1995) reported this species as an island record, collected at about 200 m on Mt. Baloy. Well known from forested, montane habitats on Negros and Cebu (Brown and Alcala, 1980), this species is found in leaf litter, stumps and fallen logs, and around tree buttresses. Localities and specimens: (Site 1)PNM 1149; (Site 9) PNM 1 153-55; (Site 11) CMNH 51 19-22, 5199- 202, PNM 2747-49, 5511-13; (Site 13) one uncata- loged specimen deposited in PNM; (Site 16) CAS 127596-97, 127599, 127601-02, 127604-05; (Site 18) CAS 127863; (Site 25) CMNH 3247, PNM 1 151- 1152. Mabuya multicarinata borealis Brown and Alcala (Fig. 41) This subspecies commonly is found under tree bark, logs and piles of vegetation (Brown and Alcala, 1980; Alcala, 1986); in the Visayas it has been collected on Negros, Caluya, Semirara, Gigante, Pan de Azucar, and Cebu, but this is the first record from Panay. Brown and Alcala (1980) reported finding one speci- men as high as 1500 m on Luzon island. Localities and specimens: . (Site 2) PNM 1657-60, 1668-80; (Site 6) USNM 496871; (Site 16) CAS 127598, 127600, 127603, 127650-51; (Site 18) CAS 127862. 127867-68; (Site 28) CAS 124107-109, 124971-78; (Site 35) CAS 124493-97; (Site 36) CAS 124199, 125039; (Site 37) CAS 125038; (Site 41) CAS 124050. Mabuya multifasciata (Kuhl) This species was found in a variety of habitats from beaches and low elevation agricultural areas, to dis- turbed forest adjacent to primary forest on Mt. Madja- as. Though known to occur throughout the Philip- pines, in the Visayas, this species has only been reported from Negros (Brown and Alcala, 1980); this is the first published account from Panay. Localities and specimens: (Site 2) PNM 1661-65; (Site 3) CAS 137585, USNM 496872; (Site 4) PNM 1640-45, 1655-56; (Site 6) CAS 137610-12, 139147; (Site 7) 1781, 1783; (Site 1 1 ) CMNH 5203-04, PNM 5510; (Site 14) PNM 3845-46, 3853; (Site 22) CAS- SU 13632; (Site 25) PNM 1149-55; (Site 42) CAS- SU 27968, 27971, 27998; (Site 44) CAS 125264-66; (Site 46) CAS 125334; (Site 47) CAS 125262-63, 125333; (Site 50) FMNH 4 1 389^104. Parvoscincus s/son/Femer, Brown and Greer All specimens of this recently-described species were collected during the day from beneath loose soil and leaf litter in the forest between 900 and 1125 m (Ferner et al., 1997). At present, this species is known only from Mt. Madja-as; the only other member of the genus is from Palawan Island (P. palawanensis). Locality and specimens: (Site 11) CMNH 3797-99, PNM 2308-10. Sphenomorphus arbore/?sTay\or This forest species is found under logs and leaves and occasionally on buttresses at the base of tree trunks (Brown and Alcala, 1980; Alcala, 1986). Our speci- mens were collected in primary forest between 1400 and 1600 m in pit-fall traps. Localities and specimens: . (Site 6) CAS 137644; (Site 1 1) CMNH 5062-74, PNM 2684-710; (Site 14) PNM 3720, 3746-48, 3753, 3867-69, 3871, 3875, 3898, 3901-03; (Site 41) CAS 124048-49. Sphenomorphus coxi divergens~\ay\ox This species is often found at the forest edge and in forest gaps. It is widely-distributed in the archipelago (Brown and Alcala, 1980; Alcala, 1986). Sphenomor- phus coxi divergens previously has been reported from Luzon, Marinduque, and Mindoro but not the Visayas (Brown and Alcala, 1980). This is the first published record of a specimen from Panay. Locality and specimens: (Site 11) CMNH 5123, PNM 2750. 2()()l Asiatic Herpetological Research Vol. 9, p. 51 Sphenomorphus cumingi '(Gray) This large forest species is found around fallen logs and tree buttresses and is believed to be limited to low to medium elevations (Brown and Alcala, 1980; Alcala, 1986). Sphenomorphus cumingi previously has been reported from islands in the Luzon and Min- danao aggregate island complexes; this is the first published record of a specimen from Panay. Locality and specimen: (Site 28) CAS 124779. Sphenomorphus jagori grandis Taylor (Fig. 42) This large skink is found under rotting logs, in sun spots in forested and disturbed areas, and along streams (Brown and Alcala, 1980; Alcala, 1986). Localities and specimens: .(Site 6) 137645^8; (Site 14) PNM 3724-5, 3755; (Site 16) CAS 127660- 62, 127677; (Site 28) CAS 125267-76, 154659-68; (Site 37) CAS 124835, 185508; (Site 41) CAS 185509-516. Sphenomorphus 5/eere/Stejneger We found this species on Mt. Madja-as in leaf litter in montane forest. Brown and Alcala (1980) report that this species is common in leaf litter of primary forest and in secondary growth. Localities and specimens: (Site 3) CAS 137600- 601, 139169-70, USNM 496873-74; (Site 11) CMNH 5026-37, PNM 2670-80; (Site 14) PNM 3708-09, 3721-23, 3726, 3749-52, 3754, 3775-81, 3786-87, 3797, 3826-29, 3836-42, 3847-51, 3854- 55, 3861, 3870, 3872-74, 3876-77, 3879-81, 3890- 92, 3914; (Site 25) PNM 1159; (Site 28) CAS 124437-38, 124577-78; (Site 36) CAS 124830; (Site 37) CAS 124827-29, 124839; (Site 39) CAS 125091; (Site 47) CAS 124625-61, 125064, 125090. Tropidophorus grayi Giinther (Fig. 43) Sison et al.'s (1995) Mt. Baloy specimen was the first record of this species from Panay. We also found a specimen at the base of Mt. Madja-as underneath a rock on the edge of a small stream in second growth forest. Localities and specimens: (Site 11) CMNH 5117; (Site 13) One uncataloged specimen deposited in PNM. Varanidae Varanus salvator nuchalis (Giinther) (Figs. 44, 45) This monitor lizard is found in agricultural and dis- turbed areas, mangrove swamps, and forested areas from sea level to about 1200 m (Alcala, 1986; Gaulke, 1991a, 1991b, 1992). On Mt. Madja-as we collected one specimen during the day in second growth forest where it was active near a small stream. Localities and specimens: (Site 1 ) PNM 1 142; (Site 11) PNM 5660; (Site 22) CAS 1 1018; (Site 31) PNM 1142; (Site 37) CAS 124879; (Site 44) CAS 124881; (Site 47) CAS 124880; (Site 50) USNM 77129-32, 80115-19; FMNH 41417-18. Squamata (Snakes) Acrochordidae Acrochordus granulatus (Schneider) Found in mangroves and at river mouths, this species feeds exclusively on fish and is widely distributed and common (Alcala, 1986). We are not aware of other published accounts of this species from Panay. Locality and specimens: (Site 23) USNM 78412, 78744-45, 78906, CAS-SU 8695, 8769. Boidae Python reticulatus (Schneider) (Fig. 46) While this snake is considered widely distributed and common in the tropical rain forests as well as near human habitation (Alcala, 1986), only one specimen has been collected on Panay Island (Leviton, 1963c; see also Gaulke, in press). Locality and specimen: (Site 44) CAS 124916. Colubridae Ahaetulla prasina preocularis (Taylor) (Fig. 47) This snake may be found in low trees and shrubs from sea level to about 800 m on Panay (Leviton, 1963c, 1968; Alcala, 1986). The specimens from Mt. Madja- as were collected from vegetation overhanging a small stream at low elevation. Localities and specimens: (Site 2) PNM 1689; (Site 7) PNM 1787; (Site 1 1) CMNH 5084-85, PNM 2716; (Site 14) PNM 3825; (Site 29) USNM 340042; (Site 44) CAS 125339; (Site 50) FMNH 41 108. Vol. 9, p. 52 Asiatic Herpetological Research 2001 Boiga angulata (Peters) We found our specimen in a coconut tree in a clearing adjacent to virgin forest at 900 m on Mt. Madja-as. While this species is known from Negros (Leviton, 1970a), until this report and that of Gaulke (in press) it has never been recorded on Panay. Locality and specimen: (Site 1 1) CMNH 5504 Boiga xf cynodon (Cuvier inV. Boie) This species has been found in forested areas at low altitudes on Palawan, Mindanao, and Luzon (Leviton, 1963c, 1970a; Alcala, 1986) but previously it has not been reported in the Visayas (see also Gaulke, in press). Locality and specimen: (Site 41) CAS 125173. Boiga ci. dendrophila (Boie) Mangrove snakes usually are found in branches of low trees and bushes in forested areas (Leviton 1968; Alcala, 1986) at low elevations. This is the first record of a species in this complex from Panay. Localities and specimens: (Site 28) CAS 124388; (Site 39) CAS 124386-87. Calamaria geravaisi DumerW and Bibron A burrowing snake, this species is commonly found in the humus under rotting logs and feeds on earthworms (Leviton, 1963c; Inger and Marx, 1965; Alcala, 1986). On Mt. Madja-as we found specimens under rotten logs and large flat rocks. Localities and specimens: (Site 2) PNM 1087; (Site 11) CMNH 5081-82, PNM 2714-15; (Site 23) CAS- SU 15953-57, 15962-65; (Site 44) CAS 124612. Cerberus rynchops (Schneider) This aquatic snake has been collected in brackish swamps, mangroves, fish ponds, and river estuaries in coastal areas (Gyi, 1970; Alcala, 1986) Localities and specimens: (Site 1) PNM 1053-55, 1077-83; (Site 20) CAS-SU 13079; (Site 22) CAS- SU 8696-97, 8719; CAS-SU 12380, CM R2423, R2426; (Site 23) USNM 77159-178, 78907-19; (Site 40) 340043; (Site 50) FMNH 41115-17. Chrysopelea paradis/'Boie This species was not known from Panay (Leviton, 1963, 1964a) until Sison, et al. (1995) reported it as an island record from Site 1. On Mt. Madja-as we found this arboreal snake during the day in root masses overhanging a river bank; one specimen was preying on a Cyrtodactylus annulatus when captured. We have also observed this species in coconut groves at sea level near the town of San Jose (site 19). Localities and specimens: (Site DPNM 1050; (Site 8) CAS 128032; (Site 11) CMNH 5041-5042, PNM 2683; (Site 19 TMM 56474; (Site 27) CAS 185-672; (Site 36) CAS 125172; (Site 46) CAS 125331-32. Cyclocorus lineatus alcalai Leviton We found our specimens in habitats ranging from dis- turbed second growth at sea level to first growth for- ested riparian sites at 1400 m. At the time of its description (Leviton, 1967), this subspecies was known only from the nearby islands of Negros and Cebu (Fig. 2). This is the first published account of this species from Panay. Localities and specimens: (Site 1) PNM 1047, 1065; (Site 2) PNM 1065, 1688; (Site 3) CAS 137576; (Site 6) CAS 137606; (Site 11) CMNH 5086-87, PNM 2717-18; (Site 14) 3884-85, 3912; (Site 16) CAS 127702, 127706; (Site 28) CAS 124051-52, 124421; (Site 37) CAS 125171; (Site 47) CAS 124445. Dendrelaphis caudolineatus terrificus (Peters) This subspecies is known from Panay and Negros and usually is found in forested and cultivated areas in or along swamps and streams (Leviton, 1970b; Alcala, 1986). Sison et al. (1995) reported this as an island record from 200 m on Mt. Baloy. The specimen from Mt. Madja-as was sleeping in bushes near a river when captured. Localities and specimens: (Site 4) CAS 1654; (Site 8) CAS 127896, 128033; (Site 1 1) CMNH 5080; (Site 13) CMNH 3254; (Site 16) CAS 127703-705, 127707-708; (Site 18) CAS 127828, 127847, 127859; (Site 28) CAS 125170; (Site 39) CAS 124203, 185673-74; (Site 42) CAS-SU 28004; (Site 50) FMNH 41093-96. Dendrelaphis pictus pictus (Gmelin) (Fig. 48) This common snake is arboreal and usually found near streams, in vegetation surrounding flooded rice fields, and in swampy areas (Leviton, 1963c, 1970b; Alcala, 1986). The Mt. Madja-as specimens were col- lected at night where they slept in stream-side vegeta- tion. Localities and specimens: (Site 1) PNM 1049, 1071; (Site 3) CAS 137577; (Site 7) PNM 1788-90; (Site 9) PNM 1071; (Site 11) CMNH 5078-79, PNM 2713; (Site 14) PNM 3843-14; (Site 22) CAS-SU 8660-70, 8718, 14932, 14936-37, CMNH 2408-13, 200! Asiatic Herpetological Research Vol. 9, p. 53 CM R2226, R2408-13; (Site 23) CAS-SU 8698-99, 8708-710; USNM 77419-591, 340044-51; (Site 30) CAS-SU 14931 ; (Site 38) CAS 200256; (Site 44) CAS 125255-61; (Site 46) CAS 125254; (Site 47) CAS 125252-53; (Site 48) CAS 124725; (Site 50) FMNH 4109-104, 41106, USNM 77592-609. Elaphe erythrura psephenoura Leviton This is a common lowland snake, often found near human habitations (Leviton, 1979; Alcala, 1986). On Mt. Madja-as we collected one specimen from the for- est floor where it was active at midday. Localities and specimens: (Site 1) PNM 1048, 1051; (Site 1 1) PNM 5662; (Site 22) CAS-SU 12389, 13212-13; (Site 23) USNM 340052; (Site 30) CAS- SU 13217; (Site 38) CAS 131700; (Site 42) CAS-SU 28001; (Site 47) CAS 125141-^12, 125340. Gonyosoma oxycephala (Reinwardt in F. Boie) This is an arboreal snake that is found in disturbed and primary forests (Alcala, 1986). On Mt. Madja-as we collected one specimen that was active at mid day in a tree above a large river (4 m from the ground). This record, and that of Gaulke (in press) are the first published accounts of this species from Panay. Localities and specimens: (Site 1)PNM 1084; (Site 9) PNM 1084; (Site 1 1) CMNH 5503; PNM 5659. Hologerrhum derma// Brown, Leviton, Ferner, and Sison We first collected this newly-described species (Brown et al., this issue) between 1030 and 1510 m in climax forest on Mt. Madja-as. Specimens were col- lected in a dry stream bed and in leaf litter on the for- est floor 30 m from a large river. One specimen from the Municipality of San Remegio was collected at approximately 700 m above sea level. Recent survey work in NW Panay suggests that this species also occurs in forested areas at lower elevations (Gaulke, in press). Localities and specimens: (Site 1 1 ) CMNH 5075, PNM 2711 (14) PNM 3704. Lycodon aulicus capucinus (H. Boie in F. Boie) (Fig. 49) This common, widespread, nocturnal snake is often found in gardens, agricultural areas, and around houses (Leviton, 1965; Alcala, 1986). Our specimens were collected near rice fields away from forested areas. Localities and specimens: (Site 1) PNM 1072-75; (Site 3) CAS 137584; (Site 8) CAS 127960; (Site 12) PNM 1072; (Site 13) PNM 1380; (Site 22) CAS-SU 8671, 8700-01, CMNH 2443, CM R2443; (Site 23) USNM 77616, 340053; (Site 26) PNM 1073-75; (Site 47) CAS 125341. Oligodon modes turn (G un ther ) This species is found under rotting logs and forest floor debris; it is known from sea level to 400 m (Alcala, 1986). In the Visayas, this species was only known from Negros (Leviton, 1963a) before Sison et al. (1995) first reported it on Panay. Localities and specimens: (Site 1) PNM 1066; (Site 2) PNM 1067; (Site 14) PNM 3790, 3866. Psammodynastes pu/veruientus(H. Boie in?. Boie) Leviton (1963c; 1983) and Alcala (1986) report this snake as common up to elevations of 1000 m or more in moist forests on Negros Island; Leviton's (1963c) listing of this species from Panay was not repeated in his 1983 review of the genus in the Philippines and no specimens were reported from Panay in the later paper (Leviton, 1983). This account and that of Gaulke (in press) appears to be the first vouchered records of this species from Panay. Localities and specimens: (Site 1) PNM 1068; (Site 2) PNM 1069; (Site 25) PNM 1067, 1070. Pseudorabdion /ncnan?araeTay\or Prior to Sison et al. (1995) this species was known only from Negros and Luzon (Brown and Leviton, 1959; Alcala, 1986). Specimens from Mt. Baloy were collected at 950 m under rotting logs in original for- est. Locality and specimens: (Site 13) Two uncataloged specimens in PNM (PNM Field Numbers 163 and 209). Pseudorabdion oxycephalum (Gunther) Previously considered a rare snake endemic to Negros Island, this species is now known from other localities in the Luzon and Mindanao aggregate island com- plexes (Brown and Leviton, 1959; Leviton, 1963c; Alcala, 1986; Brown et al., 1999). This is the first published record from Panay. Pseudorabdion oxycephalum is been found in humus and under rot- ting logs from sea level to about 750 m. Localities and specimens: (Site 6) CAS 137643; (Site 39) CAS 124174, 124193; (Site 41) CAS 124043. Vol. 9, p. 54 Asiatic Herpetological Research 2001 Pseudorabdion talonuran Brown, Leviton and Sison The discovery of this new species at high elevations was surprising. Both specimens were found under logs on Mt. Madja-as in forest classified as the transi- tion zone between mixed dipterocarp (submontane) and mossy (upper montane; Whitmore, 1984). The holotype was found at 1500 m and the paratype at 1410 m. Locality and specimens: (Site 1 1 ) CMNH 5076, PNM2712. Tropidonophis negrosensis (Taylor) This species of water snake is common along forest streams from sea level to about 700 m (Leviton, 1963c; Alcala, 1986; Malnate and Underwood, 1988). The specimen from Mt. Madja-as was found on a river bank at midday. Localities and specimens: . (Site 3) CAS 185749; (Site 6) CAS 137613; (Site 11) CMNH 5124; (Site 14) PNM 3911; (Site 22) CAS-SU 15971; (Site 28) CAS 12461 1; (Site 41) CAS 124047. Zaocys luzonensis Gunther A common tropical forest snake, this species ranges from sea level to over 1 100 m (Leviton, 1983; Alcala, 1986; Ross et al., 1987). On Mt. Madja-as, we found two specimens active at midday in disturbed forest at low elevation. Localities and specimens: (Site 1)PNM 1052; (Site 2) USNM 269078; (Site 7) PNM 1791; (Site 11) CMNH 5505; PNM 5663. Elapidae Calliophis calligaster gemianulis (Peters) We collected one specimen on Mt. Madja-as at approximately 800 m on a mountain path away from water. They have previously have been found in a variety of semifossorial habitats associated with regenerated and climax forest (Leviton 1963b, 1963c; Alcala, 1986) Localities and specimens: (Site 11) CMNH 5088, PNM 2719; (Site 13) PNM 1379; (Site 21) UPLB 2184; (Site 22) CM R2581; (Site 23) CAS-SU 12966- 68; (Site 44) CAS 125364; (Site 47) CAS 125363. Hydrophis belcheri Gray This species has been recorded from the Visayan sea and is thought to be rare (Alcala, 1986). Localities and specimens: (Site 5) USNM 38588; (Site 37) FMNH 202832-36, 202838, 202840-42. Hydrophis cyanocinctus Daudin Alcala (1986) mentioned records from the Visayan Sea. Locality and specimens: (Site 37) FMNH 202843- 54. Hydrophis elegans Gray Locality and specimen: (Site 5) USNM 38589. Hydrophis inornatus Gray Locality and specimen: (Site 23) CAS-SU 8778. Lapemis hardwickii Gray Alcala (1986) reported this species from the Visayan sea; he noted that specimens were collected by trawl- ing. Localities and specimens: (Site 23) USNM 77610- 15. Laticauda colubrina Schneider This sea snake is found among rocks and in coral reef areas near shore; it is commercially used for its meat and hide (Alcala, 1986). Locality and specimens: (Site 37) FMNH 202797- 801,202804-808,202810. Typhlopidae Ramphotyphlops braminus (Daudin) This species is common under rocks and other debris in agricultural areas. It can also be found in similar microhabitats along the edges of forested areas (Alcala, 1986). Taylor (1922d) reported catching many specimens under rocks immediately following rains. Localities and specimens: . (Site 1) PNM 1044-46; (Site 8) CAS 127972, 128036; (Site 17) CAS 127746; (Site 18) CAS 127846, 127858, 127860; (Site 30) CAS-SU 12544-49; (Site 37) CAS 125041; (Site 46) CAS 124503. Ramphotyphlops cumingii (Gray) Alcala (1986) reports this blind snake is found in epi- phytic ferns in the trees of moist forests at low to medium elevations. In all of his field work, Taylor (1922d) was unable to capture this species, suggesting he may have overlooked its preferred microhabitat. 2001 Asiatic Herpetological Research Vol. 9, p. 55 Localities and specimens: (Site 28) CAS 169877; (Site 35) CAS 125092. Typhlops castanotusWynn and Leviton Described very recently (Wynn and Leviton. 1993). this distinctive bicolored species is known from Negros. Panay (Makato). Boracay. and Inampulugan islands. It has been collected under debris in bamboo and coconut groves, hardwood forests and forest rem- nants. Localities and specimens: (Site 3) CAS 139171; (Site 8) CAS 127973; (Site 42) CAS-SU 27934-16. Typhlops hypogius Savage, Typhlops luzon- ms/s Taylor, and Typhlops ruberBoettger Found in detritus under rotting logs, these snakes have been collected in forests and disturbed areas near for- est from sea level to about 800 m (Alcala. 1986). Wynn and Leviton (1993) followed McDowell ( 1974) in referring Philippine T. luzonensis and T. hypogius to T. ruber. Recently, McDiarmid et al. (1999) asserted that T. luzonensis and T. hypogius should be recognized until more compelling evidence is pre- sented that would suggest they are conspecific with T. ruber. We agree, noting that the type localities for T. hypogius (Cebu), T. ruber (Samar), and T. luzonensis (Luzon) are each located on separate Pleistocene aggregate island complexes that might be expected to support independent lineages of blind snakes. If this is so, and the Visayan islands contain a single distinct lineage in this species complex, the name Typhlops hypogius would most likely apply to specimens from Cebu. Negros. Panay. Masbate. and smaller, land- bridge islands. Localities and specimens: (Site 18) CAS 127861. Family Viperidae Tropidolaemus cf wagleri (H. Boie in F. Boie) (Figs. 51, 52) This arboreal viper is common in forest bushes and small trees as well as mangroves (Taylor. 1922d; Lev- iton. 1964b; Alcala, 1986). The Mt. Madja-as speci- men was found in a banana plantation at 800 m near primary forest. We find it unlikely that all SE Asian populations currently referred to T. wagleri will prove to be a single species. This is the first account of a specimen referable to this species from Panay. Localities and specimens: (Site 1 1 ) CMNH 5076. Discussion The primary goal of this report has been to provide a comprehensive and synthetic review of the amphibian and reptiles species known from Panay Island and is surrounding land-bridge islets. As noted, over the past century, biologists have expected that the documented presence of a species on the neighboring island of Negros implied its undocumented presence on Panay as well (see Inger, 1954; Brown and Alcala. 1970. 1978, 1980. 1986; Brown and Rabor, 1967: Alcala. 1986). However, Panay (and its smaller satellite islets) supports low levels of endemicity, primarily as exem- plified by populations at higher montane elevations. Panay has several species of amphibians and reptiles that have not yet been reported on Negros or Cebu, is known to support species that so far have not been documented on Panay. The presence on Panay of numerous new and undescribed species suggests that calculations of Panay 's herpetological endemicity will continue to rise with continued survey work. Addi- tionally, basic taxonomic studies of species com- plexes with representatives on Panay will no doubt further contribute to the total number of Panay endemics. There are only a few reports of amphibians and reptiles of high elevation forests of Panay (Sison et al., 1995; W Brown et al.. 1997a; Ferner et al., 1997; R. Brown et al. 1999, this issue). Essentially, on each occasion that herpetologists have surveyed higher ele- vation forests of Panay, they have discovered new spe- cies. Other surveys conducted in forested regions of Panay include survey efforts of A. Diesmos, R. Crom- bie. and M.GauIke (in press). Further high elevation surveys in well forested regions of Panay are greatly needed to gain an understanding and appreciation of these presumably relictal faunal elements. There are numerous records that are included in this report that were not at all unexpected. These include widespread SE Asian and Philippine species that biologists have expected or assumed were present on Panay. Others have even been listed as known from Panay, but without specific reference to reliable local- ity data or museum specimens (see Alcala. 1986). These include the frogs Bufo marinus, Kaloula picta, Occidozyga laevis, Platymantis corrugatus, Platy- mantis dorsalis, Rami vittigera, the skink Mabuya multifasciata, and the snakes Ramphotyphlops brami- nus, Acrochordus granulatus, Psammodynastes put- verulenius. Gonyosoma oxycephala, and Tropidolaemus cf wagleri. Another group of species includes forms known from other well-surveyed islands in the Visayas Vol. 9, p. 56 Asiatic Herpetological Research 2001 r u Cfl ^^ (T, ■; ■a o o i ID _Q cc "O _i c m o E o c o n Ifl 1^ 1_ CO CO co ■? CO o t> CD 0) S I/I CD > i_ o O CD 01 >~ IT, ni i— 1 , — ^ <~ HI LU in CD CD 5 CT CD LL .£_ Mp^dtiL*. 2001 Asiatic Herpetological Research Vol. 9. p. 57 Vol. 9, p. 58 Asiatic Herpetological Research 2001 Fig. 1 1 . Flooded mangrove forest on northeast coast of Negros Island (Photo: RMB). Fig. 12. South coast of Gigante North Island (Site 35; Photo: RMB). Fig. 13. Limstone cave on Gigante South Island; habitat of Gekko gigante (Site 37; Photo: RMB). Fig. 14. Jagged karst limestone habitat of Platymantis insulatus on Gigante South Island (Site 36; Photo: RMB). 2001 Asiatic Herpetological Research Vol. 9. p. 59 -ig. 15. Bufo marinus (Photo: RMB). Fig. 16. Kaloula corijuncta negrosensis (Photo: RMB). ig. 17. Kaloula picta (Photo: RMB). Fig. 18. Limnonectes cf leytensis (Photo: RMB). :ig. 19. Limnonectes visayanus (Photo: RMB). Fig. 20. Occidozyga laevis (Photo: RMB). :ig. 21 . Platymantis corrugatus (Photo: RMB). Fig. 22. Platymantis insulatus (Photo: RMB). Vol. 9, p. 60 Asiatic Herpetological Research 2001 Fig. 23. Rana erythraea (Photo: RMB) Fig. 24. Rana vittigera (Photo: RMB). Fig. 25. Polypedates leucomystax (Photo: RMB). Fig. 26. Cuora amboinensis (Photo: RMB). m Fig. 27. Draco spilopterus (Photo: J. McGuire). Fig. 28. Draco spilopterus with extended patagium (Photo: J. McGuire). Fig. 29. Mature female Hydrosaurus pustulatus Fig. 30. Immature female Hydrosaurus (Site 23; Photo C. Banks). pustulatus (Site 1 1 ; Photo; J. McGuire). 2001 Asiatic Herpetological Research Vol. 9. p. 61 <~ ~*-t ~X.4 Fig. 31 . Mature male Gonocephalus sp (Photo: RMB). Fig. 32. Cyrtodactylus annulatus (Photo: RMB). Fig. 33. Cyrfodacfy/us philippinicus (Photo: RMB). Fig. 34. Gehyra mutilata (Photo: RMB). ^r v4 A. ^^k ■ w :5^£>*rr. .^dMk .>\ Fig. 35. gigante (Site 37: Photo RMB) Fig. 36. Gekko gigante eggs in cave crevice (Site 37; Photo: RMB). ~» - «* - > V~M^^; Fig. 37. Gekko mindorensis (Photo: RMB) Fig. 38. Brachymeles boulengehtaylon (Photo: RMB). Vol. 9, p. 62 Asiatic Herpetological Research 2001 -■• " -«p Fig. 39. Brachymeles boulengeri taylori (Photo: Fig. 40. Brachymeles tridactylus (Photo: RMB). RMB). Fig. 41 . Mabuya multicarinata borealis (Photo: Fig. 42. Sphenomorphus jagori grandis (Photo: J. McGuire). ' RMB). Fig. 43. Tropidophorus grayi (Photo: RMB). Fig. 44. Varanus salvator nuchalis, dorsal view (Photo: J. McGuire). Fig. 45. Varanus salvator nuchalis, ventral view Fig. 46. Python reticulatus (Photo: RMB). (Photo: J. McGuire). 2001 Asiatic Herpetological Research Vol. 9. p. 63 Fig. 47. Ahaetulla prasina preocularis (Photo RMB I ■■ Fig. 48 Dendrelaphis pictus pictus (Photo: RMB). Fig. 49. Lycodon aulicus capucinus (Photo: RMB). Fig. 50. Pseudorabdion mcnamarae (Photo: RMB). Fig. 51 . Typical male Tropidolaemus cf wagleri (Photo: RMB). Fig. 52. Typical female Tropidolaemus cf wagleri (Photo: RMB). Vol. 9, p. 64 Asiatic Herpetological Research 2001 (Negros. Cebu) that we expected to find on Panay as well. Their presence on Panay was almost a certainty based on known biogeographic relationships of these islands. These species include the frogs Kaloula c. negrosensis, Limnonectes cf leytensis. the skinks Lipinia pulchella taylori, Brachymeles boulengeri taylori, Mabuya multicarinata borealis, Tropidopho- rus grayi. Emoia atrocostata, the geckos Cyrtodacty- lus annulatus, Cyrtodactylm philippinicus, the sail-tin agamid Hydro saurus pustulatus, and the snakes Boiga angulata. Boiga cf cynodon, Boiga cf dendophilia, Cyclocorus lineatus alcalai, Pseudorabdion mcnama- rae, Oligodon modestum, and Trimereserus flavomac- ulatus (Gaulke, in press). Several other records are major range extensions and real surprises. The skink Dasia semisincta is oth- erwise known only from Borneo and Mindanao Island and its presence on Panay is based on a well vouchered specimen (USNM 78840) that can not be discounted. The presence of a new species in the genus Hologerrhum was surprising in that this genus previously was considered a monotypic Luzon Aggre- gate Island Complex endemic (Leviton, 1963c; Brown et al., this issue; Gaulke. in press). A forest species of Kaloula related to K. kalingensis and K. kokacii is the first record of this species group outside the bound- aries of the Luzon Aggregate Island Complex (Inger, 1954; Brown and Alcala. 1970; Alcala and Brown, 1998; Brown and Diesmos, unpublished data; Gaulke. in press). Some records (based on few specimens or dubious locality data) may be in error: these include Sphenomorphus coxi divergens, Sphenomorphus cum- ingi, and S. fasciatus (of Sison et al., 1995); other- wise, if accurate, these records represent major range extensions beyond the confines of the Mindanao. Mindoro. and Luzon aggregate island platforms. As mentioned, a number of new species recently have been described as Panay endemics; the presence of endemics in high elevation habitats of Panay was not unexpected. These species Hologerrhum dermali (Brown et al.. this issue; see also Gaulke, in press). Parvoscinus sisoni (Ferner et al., 1997). Platymantis panaxensis (Brown et al., 1997a), Pseudorabdion tal- onuran (Brown et al., 1999). Kaloula sp., Platymantis sp. 1, Platymantis sp. 2. and Platymantis sp. 3 (this report, see also Gaulke, in press). Finally, there are numerous "subspecies", species, or members of widespread Visayan. Philippine, or SE Asian species complex members on Panay that are of uncertain taxonomic status and in need of basic taxo- nomic review (see individual species accounts). Many species currently listed from Panay are. we expect, distinct lineages that eventually will be recognized as Panay or Visayan endemics. These species are too numerous to list and extend from common, low eleva- tion forms to rare high elevation forest obligates. There is much basic taxonomic work still to be con- ducted on Panay. in the Visayas. and in the rest of the Philippines and we expect our estimates of Panay 's endemicity to generally rise with further systematic studies of the herpetofauna of the Philippines. Faunal similarity calculations (Fig. 53) indicated that, as expected, the Visayan islands of Negros and Cebu (situated on the same Pleistocene aggregate island platform as Panay) were among the islands fau- nistically most similar to Panay (Masbate was not considered due to the absence of substantial records from this island). We interpret this as evidence for mid- to late-Pleistocene land bridge connections between these islands (see Heaney, 1985. 1986). Within the Visayan Aggregate Island Complex, esti- mates of amphibian faunal similarities exceeded those of reptiles but when Panay is compared to islands out- side the Visayan Aggregate Island Complex, reptile faunal similarities exceed those of amphibians. As expected, amphibian faunal similarity between Luzon and Panay is much lower than estimates calculated for Cebu or Negros and Panay. However, surprisingly, Luzon and Panay had a higher reptile coefficient of similarity than did Cebu and Panay. A portion of this unusual finding may be the result of sampling error reflecting the degree to which survey data for Negros. Panay. and Luzon are available. Calculations of simi- larities between Negros and Luzon (not shown) are higher in reptiles (0.58) than they are for amphibians (0.45). In comparisons with islands on other (non- Visayan) major Philippine aggregate island platforms, reptilian species similarity was consistently higher than that of amphibians. This may in part be due to antiquated taxonomy. Recently, disproportionate amounts of taxonomic work has been conducted on amphibian groups, resulting in the recognition of more amphibian endemics, with fewer "shared" spe- cies among islands. In contrast, many reptiles "spe- cies" are shared between Panay and Luzon. Some of these may represent species complexes in need of tax- onomic resolution. Alternatively, these calculations may reflect the relatively greater dispersal abilities presumed for reptiles when crossing ocean barriers. It is tempting to consider that more reptile species may be shared between these islands because reptiles may be more tolerant of exposure to heat and salt water, and may have a higher probability of surviving dis- persal events (e.g.. via rafting) than would be expected for amphibians. In general, these results sup- 2(K)1 Asiatic Herpetological Research Vol. 9. p. 65 Figure 53. Faunal similarity coeffi cients, calculated bet ween Panay and other major islands in the Philippines. See text for formula and discussion. port the suggestion that the herpetofaunal communi- ties of islands within the Visayan Aggregate Island Complex are very similar but also that they have their own degree of endemism and are far from being iden- tical. Species that we might expect to soon be discov- ered on Panay include populations that are otherwise known from Negros and Cebu islands. However, we note that the high elevation endemics of Negros (e.g., Platymantis hazelae, Pseudorabdion montanum) are not expected to be discovered on Panay; in their place we expect Panay to support it's own high elevation populations of closely-related montane endemics (e.g., Platymantis panayensis, Pseudorabdion talonu- ran). These include the frogs Rhacophorus pardalis (Fig. 54). Rhacophorus appendiculatus (widespread on Mindanao and Luzon aggregate island complexes: both known from Negros), Platymantis spelaeus (known from Negros; Fig. 55). geckos such as Lepi- dodactylus lugubris (widespread in the Philippines), Lepidodactylus herrei (currently comprised of two subspecies: L. h. herrei on Negros and L. h. medianus on Cebu), Lepidodactylus christiani (known from Negros and Cebu). Pseudogekko brevipes (known from Negros and Cebu: P. compressicorpus has been collected on Masbate). a Luperosaurus species (L. cumingi has been collected on Negros), and skinks like Lipinia quadrivittata quadrivittata (from Negros and Cebu). Lipinia auhculata auriculata (from Negros and Masbate), and Lipinia rabori (from Negros). Snake species we expect will be found on Fig. 54. Rhacophorus pardalis, present on Nergos but not yet recorded on Panay (Photo: RMB). 1 Fig. 55. Platymantis spelaeus, known from caves in southern Negros: this species has not yet been recorded on Panay (Photo: RMB). Fig. 56. Oxyrhabdion leponnum visayanum. well-known and common throughout Nergos Island, but not yet recorded on Panay (Photo: RMB). Panay with continued survey efforts include forms possibly related to T. canlaonensis, T. hedraeus (Negros forms). Oxyrhabdion leponnum visayanum (from Negros and Cebu; Fig. 56). and Ophiophagus Hannah (recorded from numerous islands in the Phil- ippines). The Philippine endemic crocodile. Crocody- lus mindorensis. may have recently been rediscovered Vol. 9, p. 66 Asiatic Herpetological Research 2001 on Negros (E. Alcala, personal communication) and may be present on Panay if suitable habitat can be located. As noted, the Gigante Island endemics Plan- mantis insulatus and Gekko gigante might be expected to occur on karst limestone outcrops on Panay's northeast coast. The future of exploration on Panay guarantees continuation of the kind of discovery reported here. There is an immediate need for continued basic sur- vey efforts in the montane portions of Antique. Aklan. and Iloilo provinces, all of which contribute to the western coastal mountain range that supports so much of Panay's herpetological endemicity. Additionally, low elevation portions of Panay (principally Capiz and Iloilo provinces) are also herpetologically unknown. If areas of overlooked primary forest or well-regenerated secondary forest can be located, we have high expectations that these will support novel herpetological communities and generate continued discoveries of new taxa. Areas of particular interest include karst limestone outcrops along the northern and eastern coasts, mangrove fragments, cave habi- tats, and isolated outcrops of moderate elevation in eastern Panay (Fig. 2). As noted, the best place to search for the Gigante endemics Platymantis insula- tus and Gekko gigante is limestone outcrops support- ing caves along Panay's northeastern coast. Other rare Negros species (i.e.. Luperosaurus cumingi, Lepido- dactylus herrei, Platymantis spelaeus) may eventually be revealed on Panay as well, once adequate surveys in preferred microhabitats (forest canopies and karst limestone caves; C. N. Dolino, personal communica- tion) become available. Like most islands in the Visayan Aggregate Island Complex. Panay should be regarded a priority for future conservation initiatives and programs aimed at sustainable resource management. Panay is a unique island (not at all identical to Negros) that deserves its own conservation efforts. The few remaining forests of Panay continue to be felled at an alarming rate, suggesting that its endemic flora and fauna may disappear before even being recorded by biologists. Low elevation forests and mangroves are all but gone, and even the most dis- turbed and negatively impacted sites warrant immedi- ate study of the kind that currently is underway on Cebu and Negros (A. Alcala. and E. Alcala, C. N. Dolino. J. C. Gonzales, and M. Pedregosa. personal communication). It is our hope that Panay will be rec- ognized as a model island ecosystem, ripe for collabo- rative efforts of conservation biologists, taxonomists. biogeographers, community organizers, and politi- cians. Conservation efforts targeted at the community level represent the best opportunity for foreign, gov- ernment, and non-government organizations in their effort to halt the destructive practices of non-sustain- able timber and mineral extraction industries that cur- rently operate unchecked in the central Visayan islands of the Philippines. Acknowledgements We thank the management and staff of the Protected Areas and Wildlife Bureau of the Philippines Depart- ment of the Environment and Natural Resources (DENR) for facilitating collecting and export permits necessary for portions of this study. For logistical assistance, we thank P. Gonzales (PNM) and M. Ebreo (DENR, Iloilo City), the officers and staff of the Regional DENR offices in Iloilo, and the provin- cial DENR authorities of the Municipalities of Valder- rama. Culasi. and San Jose. The PNM/CMNH Philippine Biodiversity Inventory (PBI) was funded in part by a grant from the John D. And Catherine T. MacArthur Foundation, with additional support from CMNH benefactors. Support for JWF's Panay field work (1989 and 1992) came from CMNH and the Faculty Development Fund and Department of Biol- ogy. Thomas More College. Support for RMB's par- ticipation in field work on Panay (1992. 1993, 1996) came from the Roschman Student Enrichment Fund, the Alumni and Friends of Miami University Under- graduate Research Grant. Zoology and Botany Departments, and the College of Arts and Sciences of Miami University, and CMNH. We thank the follow- ing individuals (and their respective institutions) for assistance, provision of working space, and help assembling museum records: J. Vindum (CAS). A. Resetar (FMNH). R. Crombie (USNM). and J Rosado (MCZ). Financial support for RMB's travel to CAS was provided by the Stearns Grant of the Cali- fornia Academy of Sciences. The dedicated assis- tance, untiring enthusiasm, and cheerful company of K. Auffenberg. J. Barcelona, J. Bulalacao. D. Burt, D. Busemeyer. J. Cabalquinto, E. Canada, P. Comintan. A. Diesmos, J. Demboski, M. Ebreo, R. Fernandez. J.Lasugas. M. Manuel. J. McGuire. H. Miranda, E. Mockford. L. Moores, K. Reis.L and D. Ruedas, J. Ruthven. E. Sagcal. V Samarita. R. Wacdisen. and V. Yngente is greatly appreciated. We owe particular thanks to A. Alcala for his effort to organize and logis- tically support a recent excursion to Gigante Island. We thank J. Barcelona, L. Bockstanz, D. Cannatella, R. Crombie. A. Diesmos, M. Gaulke, J. McGuire, and J. Slowinski for comments on portions of earlier drafts of this paper, and R. Crombie and M. Gaulke for sharing their ideas, time, and unpublished data. 2001 Asiatic Herpetological Research Vol. 9. p. 67 This paper is contribution no. 27 to the results of the PNM/CMNH PBI. Literature Cited Alcala, A. C. 1956. Kaloula picta on Negros Island. Silliman Journal 31:44-146. Alcala, A. C. 1958. Amphibians on Negros Island, including two new records. Silliman Journal 5:171- 174. 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Leviton. 1993. Two new spe- cies of blind snake, genus Typhlops (Reptilia: Squa- mata: Typhlopidae) from the Philippine archipelago. Proceedings of the Biological Society of Washington 106:34-45. 2001 Asiatic Herpetological Research Vol. 9. pp. 71-76 The Discovery of Mauremys iversonh\ ike Turtles at a Turtle Farm in Hainan Province, China: The Counterfeit Golden Coin James Ford Parham1 and Haitao Shi2 Department of Integrative Biology, University of California, Berkeley, California 94720-3140, USA, email: parham@socrates.berkeley.edu. Department of Biology, Hainan Normal University, Haikou City, Hainan Province 571 158, China Abstract.- During a visit to a turtle farm in Hainan Province, we discovered turtles that closely resemble Mauremys iversoni. The Mauremys iversoni-hke turtles at the turtle farm were intentionally produced hybrids of two commonly reared species, Mauremys mutica and Cuora trifasciata. According to the turtle farmer, the intentional production of hybrids between these two turtles is a common practice. The hybrids fetch extremely high prices (-1000 USD/kg) since they are sold as Cuora trifasciata. the "golden coin" turtle, to unsuspecting buyers. We suggest, but cannot prove, that all Mauremys iversoni-Mke turtles might be turtle farm hybrids. This hypothesis could explain all of the available evidence and could alleviate a lot of the confusion surrounding this species unusual geographical distribution, sudden appearance in the pet trade, as well as its variable morphology. Key words.- Turtles, Bataguridae, Geoemydidae. Mauremys. Cuora trifasciata. Mauremys iversoni. China. Hainan, hybrids, turtle farm Introduction Mauremys iversoni Pritchard and McCord 1991 is one of ten new Chinese chelonians described from speci- mens purchased through the pet trade since 1987. From the beginning, our knowledge of this species' geographical distribution has been fraught with confu- sion. The type specimen of Mauremxs iversoni (UF 71866: institutional abbreviations follow Leviton et aL 1985) and 19 other individuals are reported to have come from Fujian province (Fig. la), but eight additional specimens from a locality in Guizhou (over 1000 km away) were also reported (Fig. lb). Prit- chard and McCord ( 1991 ) propose two hypotheses to explain this disjunct distribution: 1) The distribution is (or was) continuous between these localities; 2) One of the localities (perhaps the type locality) is the result of turtles being relocated by traveling Bud- dhists. An additional wrinkle is provided by Iverson and McCord (1994) who suggest that the type series of Mauremys "guangxiensis" (-'Clemmys guangxien- sis" Qin 1992) from Guangxi (Fig. lc) is actually a composite of Mauremys mutica and Mauremxs iver- sow- like specimens. However, some differences between the M. "guangxiensis" iversoni-Mke speci- mens, the Guizhou M. iversoni specimens, and the Fujian M. iversoni specimens have been noted (Prit- chard and McCord. 1991; Iverson and McCord, 1994). The result is the occurrence of different-look- ing M. iversoni-\ikc turtles from three disparate Figure 1. The reported localities of Mauremys iversoni like turtles in China: A) Type locality, Nanping, Fujian Province (Pritchard and McCord, 1991): B) Kweiying, Guizhou Province (Pritchard and McCord, 1991); C) Nanning, Guangxi Province (Qin, 1992); D)Tunchang, Hainan Province (This study). regions in China between the years of 1986 and 1991 (Fig. 1 ). Based on observations from a breeding facil- ity located on Hainan Island (Fig. Id), we propose a third hypothesis that might explain all of the available data and possibly alleviate much of the confusion: All Mauremxs iversoni-\'\ke turtles are the result of the Vol. 9, p. 72 Asiatic Herpetological Research 2001 Figure 2. An intentionally produced hybrid trom a Chi- nese turtle farm (MVZ 230475) that closely resembles Mauremys iversoni. Photo by JFR hybridization of Mauremys mutica and Cuora trifasci- ata in captivity. Chinese turtle farms The role of turtle farms in the Asian turtle trade is often overlooked. The primary reason for this is the secrecy of the turtle farmers themselves. Despite our ignorance, the practice of farming turtles in China is a widespread, lucrative endeavor. As far back as 1991, Zhou and Zhou report that Cuora trifasciata is being bred "everywhere". In the past ten years, the number of turtle farms is rumored to have increased dramati- cally. Although it is not possible to provide exact numbers, there are estimated to be at least ten breed- ing facilities on Hainan alone. In an attempt to assess the impact of this burgeoning business to the Asian turtle trade, one of us (HS) has gained access to a large turtle farm in Tunchang. Hainan province (Fig. Id) from 1996 to the present. This paper represents the second report of this effort (see also Shi and Par- ham, 2001). According to the owner of the turtle farm, the Tun- chang breeding facility was first established in 1983. The foundation of its breeding stock came from doz- ens of wild collected Cuora trifasciata. Mauremys mutica, and Ocadia sinensis from Hainan as well as additional specimens from a farm in Guangdong that was established in 1978. It currently houses an esti- mated 15.000+ geoemydids. Early on. the number of breeding ponds and turtles was limited and the estab- lishment of breeding stocks proceeded without clear aim. Almost all the local species of turtles were col- lected from the field or bought from villagers and often kept in the same pond or enclosure. In November of 1999, the authors visited a smaller facility that included several indoor breeding ponds. Figure 3. A comparison of Mauremys iversonip\as\ta from the type description with turtle farm hybrids and putative parental species: A) Mauremys mutica Uom a turtle farm in Tunchang, Hainan Province (MVZ 230477); B-D) Mau- remys tVerson/Uom the type description. They are either from Fujian or Guizhou Province; E-F) Intentionally pro- duced hybrids from a turtle farm in Tunchang, Hainan Province; G) Mauremys iverson/'Uom the type description. It is either from Fujian or Guizhou Province; H) An intentionally produced hybrid from a turtle farm in Tunchang, Hainan Province (MVZ 130475); I) Cuora trifasciata 'from Tai Pin, central Hainan Province (MVZ 23932); J) Cuora trifasciata Atom the pet trade (MVZ 230636). Photos by JFP [a,e-f,h-j] and Pritchard and McCord (1991 )[b-d,g]. 2001 Asiatic Herpetological Research Vol. 9, p. 73 Although numerous species were observed, Maure- mys mutica and Cuora trifasciata were the most abun- dant. Many of the C. trifasciata were kept separately. but one pond included C. trifasciata. M. mutica. and even a Chelydra serpentina. When asked if any of the turtles ever hybridized, the workers of the farm pro- duced several animals that closely resembled Maure- mys iversoni. One of these hybrids, a subadult male, was procured as a voucher (MVZ 230475; Fig. 2. 3h). JFP was given permission to photograph two other hybrids (Fig. 3e,f). The turtle farm hybrids As with the three previously reported incidences of M. iversoni (Fujian, Guizhou. and Guangxi). there are some differences in the Hainan farm specimens. Unfortunately, it is difficult to understand these differ- ences because of inconsistencies in the reporting of previous measurements, small sample sizes, and the fact the most important specimens remain in private hands. For example, Pritchard and McCord (1991) present bivariate plots of measurements for adult Mauremys iversoni. but inexplicably include only five of the adult males. It is not stated from which locality (Fujian or Guizhou) these specimens are from or why the measurements of all 10 of the adult males from Fujian (the type locality) are not included. Table 1 of Pritchard and McCord includes 10 shell measure- ments for all the specimens, but excludes the two measurements (the interpectoral length [IPLj and the interanal length [IAN]) that would have allowed the type series to be included in the bivariate plot. Iver- son and McCord (1994) perform a more detailed study of variation within East Asian Mauremys. but do not present the raw data and only include one speci- men of Guizhou Mauremys iversoni (no voucher). The relevant shell measurements of MVZ 230475 are as follows: IAN= 1.4 cm, IPL= 1.1 cm. CL= 1 1.4 cm. IAN/CL= .1228, IPL/CL=.096. MVZ 230475 is a juvenile and smaller than three specimens excluded from the adult category by Pritchard and McCord (1991). Bearing this consideration, a comparison shows that the morphology of MVZ 230475 is unlike the type series of Mauremys iversoni. and apparently unlike the other Mauremys. It is uncertain what the variation between and among the Mauremys iversoni series (from Guizhou and Fujian) is fully represented by the polygon in Pritchard and McCord (1991). Since the relevant scale ratios from their entire series, including the subadult specimens, are not reported, we are forced to rely on the close phenetic similarity between the turtle farm hybrids and M. iversoni for our conclusions. It is important to emphasize that if Figure 4. Cuora trifasciata Uom a Chinese turtle farm showing light head coloration. Photo by HS. all M. iversoni are turtle farm hybrids, then we should expect them to have an extremely variable morphol- ogy, especially if they are the result of separate hybri- dogenic events. This pattern is borne out by the previously reported M. iversoni-\\kc turtles (Pritchard and McCord, 1991; Qin. 1992; Iverson and McCord, 1994). In terms of coloration, MVZ 230475 and the hybrids photographed at the Tunchang turtle farm closely resemble Mauremys iversoni. The head color- ation of MVZ 230475 is lighter than that of the speci- men figured by Pritchard and McCord (1991 ). but a variation in head coloration in Mauremys iversoni was noted by Fritz and Obst (1999). MVZ 230475 is almost identical to the lighter individual figured in that work. The variable head coloration in M. iversoni can be linked to the variation in C. trifasciata. For, while most C. trifasciata have dark postorbital mark- ings. C. trifasciata with light head coloration are known (Fig. 4), especially in turtle farms. Pritchard and McCord ( 1991 ) state that M. iversoni can be dis- tinguished from M. mutica by a horseshoe-shaped coalescence of blotches from the pectorals to the anals (with the open end anterior). Although they state that this pattern is not found in Mauremys mutica. Zhou and Zhou ( 1991, p. 38) illustrate a M. mutica (locality not stated) with this this plastral pattern. Later. Iver- son and McCord (1994) illustrate a M. mutica from Taiwan (FMNH 127181 ) that also has a clear horse- shoe-shaped coalescence of blotches. Even without this character, M. iversoni can be distinguished from M. mutica by its more rounded anterior lobe that lacks a prominent gular projection. Furthermore, many M. iversoni specimens, including some of the type series Vol. 9, p. 74 Asiatic Herpetological Research 2001 of M. iversoni and the Hainan farm hybrids (Fig. 3b- f) have transverse trending blotches on the pectorals, a feature not known to occur in Mauremys, but common in juveniles of C. trifasciata. The variation among the figured specimens in the type description is extreme (Fig. 3b-d,g). It is not stated whether these specimens are from Fujian or Guizhou. Comparisons of the plastral figures from Pritchard and McCord (1991) with the specimens from the Tunchang farm show that the latter speci- mens lie within the range of variation of M. iversoni. The most notable differences are that one (Fig. 3e) has more irregularly shaped plastral blotches and MVZ 230475 (Fig. 3h) has only a narrow, light, central fig- ure on the plastron. Perhaps the most important fea- ture to be noted is that no two specimens look alike. In the type description, Pritchard and McCord ( 1991 ) describe the plastral pigmentation as "very variable in intensity", but it is unclear whether the figured speci- mens represent the most typical patterns of Mauremys iversoni or the entire range of variation. Luckily, there are three characters of M. iversoni that clearly set it apart from other East Asian Maure- mys: 1) its olive or yellow head, 2) rounded end of anterior lobe of the plastron and 3) red coloration on the underside of the limbs. Until the description of Mauremys iversoni, the red coloration of the limbs was a diagnostic characteristic of C. trifasciata. The light head coloration and rounded lobe are also remi- niscent of C. trifasciata. Furthermore, in Mauremys iversoni the sulcus between the humeral and abdomi- nal scutes lies closer to the hyo/hypoplastral suture than it does in other Mauremys. An alignment of this sulcus with the junction of these bones is a character shared by C. trifasciata and other taxa with a kinetic plastron. In other words, the morphology of Maure- mys iversoni is intermediate between C. trifasciata and M. mutica, and therefore consistent with a hybrid origin. A detailed comparison of the mitochondrial DNA and allozymes of this specimen and other pet trade "species" is currently under way (Parham et al., 2000; in prep.). Discussion During our joint visit to the Tunchang turtle farm, the workers stated that production of Mauremys iversoni- like turtles was the result of infrequent, accidental hybridization events. Since that time, the owner of the farm has confided that the intentional production of C. trifasciata X M. mutica hybrids is a common practice undertaken by several turtle farmers. Because practitioners of Chinese traditional medicine claim that C. trifasciata has many medicinal proper- ties, and recently it has also been suggested that it has the ability to cure cancer, it is highly valued. As a result, it is often called the "golden coin" turtle. The price of turtles is determined by their weight. In the year 2000. 1 kg of M. mutica sold for approximately 100-120 yuan (-15 USD) whereas 1 kg of C. trifasci- ata sold for 6,000-8,000 yuan (-1000 USD). Accord- ing to the turtle farmer, it is possible to sell the hybrids of C. trifasciata and M. mutica as pure C. tri- fasciata since, to the untrained eye, these species closely resemble one another, especially in the yellow coloration of the head. The confusion of these two forms should not be surprising since Timmins and Khounboline (1999) report that even people familiar with C. trifasciata have mistakenly identified juvenile M. mutica as this species. Therefore, it is possible to produce and sell large numbers of counterfeit "golden coins" to unsuspecting buyers without having to obtain large numbers of adult Cuora trifasciata. The high degree of phenetic similarity between the C. trifasciata X M. mutica hybrids from Hainan and the M. iversoni-tokc turtles reported from Fujian, Guizhou and Guangxi strongly suggests that all of these animals may be the product of accidental or intentional hybridization in turtle farms. This hypoth- esis explains the unusual morphology, its sudden appearance in the pet trade, absence in historical col- lections, and the confusion surrounding the distribu- tion of this taxon. The timing of the discovery of these turtles shortly follows the increased demand for Cuora trifasciata as a reputed cure for cancer (van Dijk et al., 2000). This increased demand may have initiated the establishment of numerous turtle breed- ing facilities that led to the production of Mauremys iversoni-Yikc turtles. We stress that the practice of housing and breeding several species of chelonians in a single enclosure or pond is probably not restricted to the Tunchang farm alone. If the establishment of other facilities mirrors the one on Hainan, the sudden appearance of new and unusual turtles in the pet trade would be expected. The ability of turtles to hybridize is well documented (Fritz and Baur, 1994; Fritz, 1995). Certain species, such as Mauremys pritchardi McCord 1997. have already been implicated as possible hybrids (Artner et al.. 1998). Others, such as Sacalia pseudocellata Iverson and McCord 1992. Ocadia glyphistoma McCord and Iverson 1994. and Ocadia phillipeni McCord and Iverson 1992 are known from a paltry number of specimens which might be indicative of an accidental production in a turtle farm (van Dijk, 2000; Lau and Shi. 2000). It is probably not a coincidence that many of the newly described species (O. phil- 2001 Asiatic Herpetological Research Vol. 9, p. 75 lipeni, S. pseudocellata, and M. iversoni) can be dis- tinguished from their congeners by characters found in the commonly reared and highly valued C. thfasci- ata. Studies of the DNA of the holotypes of these species, compared with multiple, field-collected, vouchered specimens of established species from known localities, could easily determine the validity of these taxa. Conclusions It is uncertain whether all of the reported collection data from the pet trade, such as the distributional data for Mauremys iversoni are actual or fabricated. The rest of the available evidence strongly suggests, but does not prove, that all M. iversoni-Wke turtles could have a turtle farm origin. Parham and Li (1999) openly question the Yunnan locality of Cuora pani that was provided by the same pet dealer credited with collecting the original Mauremys iversoni series. The high prices that unusual turtles with locality data can fetch from turtle fanciers could serve as incentive to fabricate collection data. Furthermore, there is also incentive to hide the true locality as a trade secret, especially if the locality is a Chinese turtle farm. Of course, given the current scarcity of turtles in Asia it is difficult to prove that a locality is in error or whether the turtles have been extirpated. Verified distribution data for rare chelonians (e.g., Iverson, 1992) are criti- cal to understanding their current and historical status. False localities unnecessarily complicate our meager understanding of turtle distributions. In light of the confusion surrounding pet trade data, we recommend that workers should treat them with utmost caution until they are supported by evidence that is more reli- able. Given the seemingly uncontrollable Asian Turtle Crisis, determining which species are valid evolution- ary lineages and which are turtle farm hybrids is important. . Otherwise, crucial resources could be wasted on invalid taxa. For example, the Red data Book of Endangered Animals (Wang, 1998) suggests that Fujian and Guizhou provinces protect M. iversoni and that surveys and ecological studies should be per- formed in order to determine the proper conservation strategies. We concur with van Dijk's (2000) asser- tion that researchers should make the investigations on the validity of the pet trade species a priority so that the limited resources for conservation can be directed towards established taxa that are undergoing unimagined, precipitous declines. Acknowledgments The authors would like to thank John Iverson for prompting a comparison between the turtle farm hybrids and the type description of Mauremys iver- soni. We would also like to thank Dr. Jaeger, the edi- tor of Herpetologica, for granting us permission to reproduce part of Figure 1 from Pritchard and McCord (1991). Ted Papenfuss and Jim Buskirk pro- vided helpful comments. JFP is also grateful to Carl Gillies, Kevin Padian, and Tonya Van Leuvan-Smith. This project was funded by the National Science Foundation (JFP), the University of California Museum of Paleontology (JFP), The National Natural Science Foundation of China (HS), The Provincial Natural Science Foundation of Hainan (HS), and the Hainan Provincial Ecological Key Discipline Fund (HS). Literature Cited Artner, H., H. Becker, and U. Jost. 1998. Erstbericht liber Haltung und Nachzucht der Japanischen Sumpfschildkrote Mauremys japonica (Temminck and Schlegel, 1835). Emys 5:5-22. Fritz, U. 1995. Schildkroten-Hybriden 2. Halsberger- Schildkroten (Cryptodira). Herpetofauna 95:19-34. Fritz, U. and M. Baur. 1994. Schildkroten-Hybriden. 1. Halswender- Schildkroten (Pleurodira). Herpeto- fauna 94:28-34. Fritz, U. and F J. Obst. 1999. Neue Schildkroten aus Siidostasien Teil II. Bataguridae {Cyclemys, Heose- mys, Mauremys, Ocadia, Pyxidea, Sacalia) und Tri- onychidae. Sauria 2 1 ( 1 ): 1 1 -26. 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Mauremys pritchardi, a new bat- agurid turtle from Myanmar and Yunnan, China. Che- Ionian Conservation and Biology 2:555-562. McCord, W. P. and J. B. Iverson. 1992. A new species of Ocadia (Testudines:Bataguridae) from Hainan Island, China. Proceedings of the Biological Society ofWashington 105:13-18. McCord, W. P. and J. B. Iverson. 1994. A new species of Ocadia (Testudines:Batagurinae) from southwest- ern China. Proceedings of the Biological Society of Washington 107:52-59. Parham J. F. and D. Li. 1999. A new locality for Cuora pani Song 1984 with comments on its known range. Asiatic Herpetological Research 8:1 1 1-1 13. Parham J. F., K. H. Kozak, W. B. Simison, and C. R. Feldman. 2000. New species of turtles from China: Endangered species or infrequent hybrids'? Fourth Asiatic Herpetological Conference: July 16-20, 2000, Chengdu, China. [Abstr.] Pritchard, P. C. H. and W. P. McCord. 1991. A new emydid turtle from China. Herpetologica 47(2): 139- 147. Shi, H. and Parham, J.F 2001. Preliminary observa- tions of a large turtle farm in Hainan Province, Peo- ple's Republic of China. Turtle and Tortoise Newsletter 3:2-4. Timmins, R. J. and K. Khounboline. 1999. Occur- rence and trade of the golden turtle, Cuora trifasciata, in Laos. Chelonian Conservation and Biology 3(3):44 1-447. Qin, J. 1992. A new species of genus Clemmys: C. guangxiensis. Pp. 60-62. In Y. Qian, E. Zhao, & K. Zhao (eds.). Animal Science Research. A Volume issued to celebrate the 90* birthday of Mangven L.-Y Chang. China Forestry Press, Beijing. van Dijk, P. P. 2000. The status of turtles in Asia. Che- lonian Research Monographs 2:15-23. van Dijk, P. P., B. Stuart and G. J. Rhodin. 2000. Asian Turtle Trade: Proceedings of a workshop on conservation and trade of freshwater turtles and tor- toises in Asia. Chelonian Research Monographs 2:164 pp. Wang, S. (ed.) 1998. China Red Data Book of Endan- gered Animals, Amphibia & Reptilia. Science Press, New York. 332 pp. Zhou J. and Zhou T. 1992. Chinese Chelonians Illus- trated. Jiangsu Science and Technology Publishing House, Nanjing. 89 pp. 2001 Asiatic Herpetological Research Vol.9, pp. 77-100 A Review of the Distribution of Diploid, Triploid andTetraploid Green Toads {Bufo viridis Complex) in Asia Including New Data from Iran and Pakistan1 Matthias Stock1, Daniel Frynta2, Wolf-Rudiger Grosse1, Claus Steinlein3 and Michael Schmid3 Martin-Luther-Universitdt Halle-Wittenberg, Institutfiir Zoologie, Domplatz 4, D - 06099 Halle/ Saale, Germany, e-mail: stoeck@zoologie.uni-halle.de. ' Charles University, Department of Zoology, ViniCnd 7, 128 44 Praha, Czech Republic. * Universitat Wiirzburg, Institutfiir Humangenetik, Biozentrum, Am Hubland, D - 97074 Wiirzburg, Germany. Abstract.- A summary of the distribution of green toads containing most of the data published after the discovery of polyploid forms (1976) including a map, an index and a bibliography are presented and discussed. 21 Asian type localities of hitherto described nominal green toad taxa are shown. The tetraploids are distributed in high mountains and extremely continental regions with strong climatic shifts. Records of triploid specimens are situated in supposed contact zones between the parapatric diploid and tetraploid toads at foothills of Middle Asian high mountains, and triploid bisexual populations occur in the Karakoram and West-Himalayas. Habitats of diploids appear to be restricted to lowlands and valley grounds. Polyploids seem to be more resistant. The methods hitherto used for the determination of the ploidy level and their applicability are evaluated. We show new data on this species complex from Iran including cytometric, karyological, bioacoustic and morphological data and we draw taxonomic conclusions for tetraploid Bufo oblongus, diploid Bufo viridis kermanensis, and probably diploid Bufo kavirensis. New information on the distribution of triploids in northwestern Pakistan based on flow cytometric measurements is presented. The ploidy level of Bufo latastii is revealed to be diploid. Key words.- Amphibia, Bufonidae, Bufo viridis complex, Bufo oblongus lectotype, Bufo viridis kermanensis, Bufo kavirensis, Bufo latastii, Bufo pseudoraddei pseudoraddei, Bufo pseudoraddei baturae, distribution, type localities, Asia, Iran, Pakistan, chromosomes, ploidy determination, calls, systematics This paper is a chapter of the doctoral dissertation of M. Stock. Introduction Since the discovery of tetraploid forms of the Bufo viridis complex in the northern Tien Shan (Bachmann et al„ 1978; Mazik et al., 1976) many records of dip- loid, triploid and tetraploid green toads have been made, especially on the territory of the former Soviet Union and some few in other countries. New findings as the detection of possibly all-triploid, gonochoristic populations of this complex in the Karakorum range of Pakistan (Stock et al., 1998, 1999) provide arguments that the number of investigations in this species com- plex will increase during the next years because not only zoogeographic and phylogenetic but also cytoge- netic, bioacoustic and biochemical questions will be studied. Since a fundamental summary of the data on the territory of the former Soviet Union was done by Borkin et al. (1986a), the development of knowledge in this species complex has been dramatically increased. Hence, the present paper was aimed to summarize the recent information about the distribution which is one of the preconditions for the understanding of the natu- ral history of these toads. The data on chorology will also be an important prerequisite for a revision of sys- tematics and nomenclature in this species complex. We discuss the hitherto applied techniques of ploidy determination, the size of the (known) range of polyploid green toads, any detectable correlation of their occurrence with climatic/ecological factors, and we try to enlarge the knowledge about the ploidy level of taxa hitherto described from various type localities. Because the knowledge on the ploidy of green toads from the territory of Iran and Pakistan is very scarce, we also present and discuss some new, sporad- ically collected data from different sources, and first results of a field excursion to Pakistan. Vol. 9, p. 78 Asiatic Herpetological Research 2001 Figure 1 . Map of Middle and Central Asia and parts of the Middle East with records of diploid, triploid and tetraploid green toads after 1976. For code numbers see appendix. 2001 Asiatic Herpetological Research Vol. 9, p. 79 A *loi A a&£.3^ A '^^AA □ A ^jS A 1 ■Is ,A m D bonoaa-e-r\avus (aipioia; i r A Kashgar (tetraploid) ■ Kapkan (diploid) 27.5 I 25 » E 22.5 o 1 20 B S 17.5 « 15 c 9 12.5 co «> 10 3 . 9 10 11 Figure 5. Chromosomes of diploid male Bufo viridis kermanensis Uom Kerman. a - Giemsa-stained metaphase, b - Giemsa-stained karyotype, c - Q- banded metaphase, d - Q-banded karyotype. Iran near Baghestan (Fig. 1: 13; Fig. 8F) in the moun- tainous region north-eastern of the Central Iranian Plateau (abstract by Stock et al.. 1998). A sample of five toads from Birjand (Fig. 1: 13a), the type locality (Fig. 1: E) of Bufo oblongus, was found to be tetrap- loid (Fig. 4). Table I outlines the results of the flow cytometric measurements in specimens from Kerman and Birjand. Table 2 summarizes the results of eryth- rocyte size measurements. The erythrocyte sizes were typical of diploid and tetraploid green toads, respec- tively, according to Stock and Grosse (1997a). The mean DNA content of B. oblongus (17.02 pg) rather corresponds to that of tetraploids from Kashgar (17.5 pg; Stock, 1998b) measured with the same technique, but during another session. The Giemsa-staining (Fig. 5a, b) revealed a diploid karyotype of B. viridis kermanensis. These convention- ally stained chromosomes posses the characters of those from some other Eurasian diploid Bufo viridis which have been examined until now (Birstein. 1981; Bogart. 1972; Roth and Rab. 1987; Schmid. 1978: Ullerich. 1966). The pair 6 terminally exhibits in its long arms a secondary constriction which is caused by the telomeric position of the nucleolus organizer regions (NORs) in B. viridis (see also Roth and Rab, 1987; Schmid 1978). The Quinacrine-banding in chromosomes of B. viridis kermanensis (Fig. 5c. d) shows differently intensive fluorescing chromosomal regions in pairs 6 to II, but distinct bright fluorescence »» . 1 h m 1 2 3 Mi mi in* 4 5 6 XXXX MAI MM 7 8 9 1 * UJVMM 4 5 6 ■vkX «MI xxk» Figure 6. Chromosomes of a tetraploid female Bufo oblongus Uom Birjand. a - Giemsa-stained metaphase, b - Giemsa-stained karyotype, c - Q-banded metaphase, d - Q-banded karyotype. was only found in short and long arms of pairs 6 and 8. Birstein (1981) detected Q-bands in a B. viridis from the Crimea in pairs 6 to II (the most distinct ones in both arms of pairs 7, 10, and II). In a triploid taxon (Bufo pseudoraddei balurae) of the B. viridis complex from the Karakoram. we found Q-bands in the long arms close to the centromere of triplet 1, in the short arms of triplets 6 and 7 and in both arms of triplets 8 to 11 (Stock etal., 1999). Similarly, the karyotype of the tetraploid Bufo oblongus exhibits Q-bands in chromosomes of the quartets 1, 6 to 11 (Fig. 6c. d). The most interesting linding is the occurrence of Q-bands in the short arms of only two out of the chromosomes of quartet 6. These two chromosomes have also larger long arms than the two remaining Q-negative chromosomes, and there- fore, we conclude that the Q-positive pair also repre- sents the only one which carries the telomeric NORs as Roth and Rab (1987) found in tetraploid toads from Kyrgyzstan. We are preparing a detailed cytogenetic study. Like in the triplet 1 of triploids from Karakoram. one or two chromosomes of quartet I in B. oblongus posses Q-bands in their long arms. Furthermore, addi- tional differences are visible in the occurrence and position of Q-bands among the chromosomes in each of the quartets 7. 8 and 10 (Fig. 6c. d). These observa- tions in B. oblongus provide arguments for considering this tetraploid form allopolyploid. The question of autoploidy or alloploidy of polyploids from various Vol. 9. p. 82 Asiatic Herpetological Research 2001 Table 1 . DNA content in diploid toads from Kerman (Bufo viridis kermanensis), diploid toads from Choqua Zanbil (B. wridisssp.), tetraploid toads from Birjand [Bufo ob/ongus), and diploid B. iatastiiUom Skardu measured by flow cytometry of DAPI stained erythrocytes with chicken nuclei (= 2.34 pg DNA/nucleus) as a standard. SD = standard deviation. Kerman, diploid C. Zanbil, diploid Bi rjand, tetraploid Skardu, diploid (N = 7) (N = 2) (N = 5) (N = 15) . . . , ,~ *~ 1~ *,~ ii f . ii ^^ ii , , n ^^ s en Q. ■o CO Q. ■o Q. "D D. co — CO CO CO ■d c 3 C 0) c CO C CD CO C CD ■o c CO c CD s? o 5? O s? O S? O c o c O c CJ c O 2 c < 20 v> 3 a. 15 10 / <► <> ,- <> <> / , * ' A" A A - 'A r ■■ ■■ H * 1 1 «.^i 0 s M A M A ▲ D Tetraploid. Middle and Central Asia (Stock, 1998a, b) Birjand Diploid, Middle Asia (Stock, 1998a) Lar Valley (Andren and Nilson, 1979) Cheshmeh-ye-Sefied-Ab (Andren and Nilson, 1979) Fasham Polur Linear (Tetraploid, Middle and Central Asia (Stock, 1998a. b)) • Linear (Diploid, Middle Asia (Stock, 1998a)) 10 15 20 25 Water temperature (°C) 30 Figure 7. Pulse rate of mating calls of green toads from Iran in comparison with pulse rate of diploid and tetraploid toads from Middle and Central Asia. Birjand: type locality of Bufo oblongus Nikolsky, 1 896. Toads from Birjand were found to be tetraploid (Fig. 4). Cheshmeh-ye-Sefid-Ab: type locality of Bufo kavirensis, Andren and Nilson, 1979: the call data appear to elucidate that the taxon is diploid. Call data from Lar valley (Andren and Nilson, 1979), Fasham and Polur (present study) provide arguments that these toads are diploid. green toads in the Elburz mountains and confirm the evolutionary conservatism of the mating calls (disc, in Stock, 1998b; Stock et al., 1999). The single mating call of a tetraploid male ( 1 3a) from the type locality, Birjand (E), of Bufo oblongus confirmed previous data on tetraploids from various regions of Middle and Cen- tral Asia (Castellano et al., 1998; Stock, 1998a, b). Morphology and Taxonomy (Fig. 8). Although the locality (1) is relatively close to the type locality (A) of Bufo luristanicus (Schmidt, 1952), the morphology of the toads from Gholaman (Fig. 8C) and Choqa Zanbil differs completely from that of the B. luristani- cus holotype (Fig. 8H). This confirmed once more the occurrence of at least two different green toad taxa in the north-western Zagros mountains of Iran as already Schmidtler and Schmidtler (1969) as well as Eiselt and Schmidtler (1973) stated. First, the form which was previously (Schmidtler and Schmidtler, 1969; Mertens, 1971b; Eiselt and Schmidtler, 1973) called B. viridis arabicus and to which the diploid toads from Gholaman and Choqa Zanbil belong to. Now, the name "arabicus" is no longer applicable since Bal- letto et al. (1985) restricted it to B. arabicus Heyden, 1827 from the Arabian Peninsula. Therefore, we pre- liminarily use the term Bufo viridis ssp. for them. The second taxon occurring in the north-western Zagros mountains until eastern Iraq (Afrasiab and Ali, 1988) is B. (surdus) luristanicus. As to be seen in the appen- dix, the ploidy level of all three subspecies of B. sur- dus distinguished by Schmidtler and Schmidtler (1969) and Eiselt and Schmidtler (1973) is still unknown (cp. Baloutch and Kami, 1995). The seven topotypic specimens of Bufo viridis kermanensis from Kerman (13b) are very similar to the holotype (NMW 19647) shown and described by Eiselt & Schmidtler (1971, 1973). This diploid form is different from the large sized diploid B. viridis turanensis as already Hemmer et al. (1978) stated and which was confirmed by our data. The diagnostic differences between Bufo kaviren- sis and B. viridis kermanensis consist according to Andren and Nilson (1979) in only two morphometric traits (I: ratio "distance between posterior border of nostril and anterior corner of eye/internasal distance" 0.80 - 1.06 in B. kavirensis and 1.21-1.55 in B. v. ker- manensis; II: ratio "width of upper eyelid /interorbital distance" 1.1 1 - 1.61 in B. kavirensis and 1.51 - 2.00 in B. v. kermanensis). Both ratios were not confirmed because the first ranged for our seven B. viridis ker- manensis from 0.9 - 1.18 (and 0.93 - 1 .05 in two spec- imens, Fig. 1: 13c, by Borkin et al., 2000); the second ratio varied also strongly from 0.8 - 2.92 (and 1.83 - Vol. 9, p. 84 Asiatic Herpetological Research 2001 2.31, Borkin et al., 2000). Including the data shown above, we preliminarily consider (diploid) B. kaviren- sis to be a junior synonym of diploid B. viridis ker- manensis. This also agrees with the range presumed by Eiselt and Schmidtler (1973) for their taxon. The relationships between diploid B. viridis kermanensis and diploid B. viridis ssp. (previously called B. viridis arabicus, see above) require additional investigations. The morphological differences between Bufo oblongus (Fig. 8A, B, D) and B. viridis kermanensis (Fig. 8E) which were considered to be "very sharp regarding neighboring races" (Eiselt and Schmidtler. 1973) were now explained by the different ploidy lev- els. The females of B. oblongus (e.g., Fig. 8B) exam- ined from Birjand exhibited a coloration pattern which is very similar to that of the specimen ZISP 1952.1 from the type series (Fig. 8A). We therefore designate it here as the lectotype of B. oblongus, Nikolsky, 1896, and we consider this taxon according to the topotypes to be tetraploid. This demonstration of tetraploidy for B. oblongus is important for the tax- onomy of green toads in Asia because the name is El .§ «a of "5 > I CO E" * S ? -D .o .9- LU (0 E 2 ^ Q. O , If) oi M !<§ CD " 5 S E « •a co E 5. o >- ^= x: e/> £ "S E O 0) £?. O) "O °! 2 ^ Es *1 ° o Source Intercall interval (s) [N of intervals measured] Call duration (s) [N of calls measured] Fundamental frequency (Hz) Pulse rate (s'1) Pulse duration (ms) Interpulse interval (ms) WaterT ("C) AirT CO Male Locality (code in Fig. 1) en o to •w v-/ ^ -a- <5 CM a in CM C\J o CM o co rn O) o> en en t^ f^ t~- h- f~ a en cn en en r c c c c r> o o o o B> co co CO CO -a c < CD CD CD 7 7 7 Li ffl O. CO Q. CO T3 ■o TJ a. ex Q. CO CO S C c C CD CD CD •CD •CD ■o CD CD CD ■o c T3 c c a. o. Q. < < < C CO •D C < T3 C CO CO CD Cvl_ „ — , t CD CO 2, o CD CD C\i CM CO CD C\i o o in o o C\J N ^ CO CD 2_ CM CD eg CD o en cvi ■w- CM t- co co T- Y- f- ^ in in Cvi C\i CO CD TJ- o CO o co f- cm co ■* in §. co i .o CO CM- E CD E ■o CD — ' n CO m 3 CO CD CO 4! ■2 O CD < O CD in r~- i- co CD "O- in en to t- i- cm co 2001 Asiatic Herpetological Research Vol. 9. p. 85 Table 4. Mating call data of triploid Bufo pseudoraddei baturae'xn comparison with Bufo /atasti/Uom India and Pakistan. *■ S u. a *-* a E £ 3 ■o ■ 1 i. 1 c o 'to 51 28 o t O CO — cb ra.E o ^_ CD o CZ o a c o X |2 o, I t 2 3P a a 3 £ c 3 ■ W 3 a. 22 to a « 3 a. | o E c ro ffi ■a 3 c o- 3 S> Li- — IS CO CO = ^ CO = o CO-, o£ a a 3 O 2600 h, see Zlotin, 1997) might have caused a genetically caused selective advantage of the polyploid forms. This may also account for the missing of diploid toads in the east of the range where only tetraploids have been detected. The eastern boundary of the range of tetraploid green toads and the obviously parapatric occurrence of B. raddei in Northern China and Western Mongolia was discussed by Peters (1971: "B. viridis" for tetraploids), Borkin and Kuzmin (1988), and Stock (1998b). The most western known records of tetraploids (Kopet Dagh, Khorasan) seem to correspond with the region of tran- sition from winter-mild, summer-dry steppe climates of Middle Asia and Afghanistan to the winter-humid, summer-dry climates of Mediterranean type in Iran (Walter and Lieth, 1967; Miiller, 1996). The deserts of Central Iran with fewer than 100 mm rainfall/year and large sand or salt areas seem to separate the polyploids in the East of Central Iran (and Afghanistan?) from the diploids in the West. Although B. surdus may provide unexpected karyological data, at the moment, in spite of only few studies, records of polyploid green toads in the West of Iran appear to some extent improbable. This may rather result from the history of distribution than from the possible relationship between polyploidy and environmental selective pressure. The present image of the distribution of diploid and tetraploid toads possibly also corresponds to the experi- ence that (especially allopolyploid organisms may have advantages in regions with extreme or changing environments (Futuyama 1990: 69 - "polyploids are more resistant"), when colonizing new habitats (Bretagnolle et al. 1998) or were considered to have a higher competitive ability (e.g., Lumaret et al., 1997). However, particularly the distribution of diploid plant species and their polyploid relatives probably more strongly reflects the influence of historical factors than of their current ecological requirements (Bretagnolle et al. 1998) or adaptedness. Morescalchi (1990) stated: "In amphibians, changes in genome size (...), far from being random, they are related to metabolic and onto- genetic factors which are of crucial importance in the adaptive strategies of these animals". 2001 Asiatic Herpetological Research Vol. 9, p. 91 Acknowledgments We thank Prof. Dr. J. Martens, Mainz, for mating call recordings from northern Iran, and Dr. T. Klapperstiick, Halle, for using the CYDOK image analyses system. Prof. Dr. N. Ananyeva, St. Peters- burg, Dr. R. Giinther, Berlin, Dr. C. McCarthy, Lon- don, Dr. G. Nilson, Goteborg, Dr. J. B. Rasmussen, K0benhavn, for loaning preserved specimens. Chro- mosome photographs were kindly prepared by G. Hesse, Wurzburg. The basic computer-map was cre- ated by H. Nagel, Halle, according to our concept. Many thanks to Dr. K. J. Baig, Islamabad, Dr. L. Ya. Borkin, St. Petersburg, Dr. S. Castellano, Torino, Prof. Dr. A. Dubois, Dr. A. Ohler, Paris, Dr. T Papenfuss, Berkeley, J. F. Schmidtler, Munich, and Prof. Dr. E. Zhao, Chengdu, for help with getting literature. Dr. S. L. Kuzmin, Moscow, kindly allowed to use his data- base "Amphibians of the former USSR". M. Stock is very grateful to the family of M. and M. Khatiri, Gorgan, for their kind hospitality, R. Dressel, Dresden, for optimism and assistance during the field work in Pakistan, Dr. D. Lamatsch, Wurzburg, for help with flow cytometry, and T. Diet- erich, Greifswald, for a toad from Tengiz Lake (Kaza- khstan). Literature Cited Afrasiab, S. R. and H.A. Ali. 1988. A new record of toad Bufo surdus Boulenger (Amphibia, Bufonidae) from Iraq, with preliminary key for Iraqi Amphibia. Bulletin Iraq National Museum 8 (1): 115-123. Andren, C. and G. Nilson. 1979. A new species of toad (Amphibia, Anura, Bufonidae) from the Kavir Desert, Iran. Journal of Herpetology 13 (1):93-100. Anonymous. 1993. The Times Atlas of the World. Comprehensive Edition. London, Bartholomew and Times Books, Harper Collins:pl. 1-123 + l-222pp. Ataev, E. A. 1987. K ekologii danatinskoii zhaby na Kyurendage. Izvestija Akademii Nauk Turkmenskoi SSR. Seriya biologiceskikh nauk 3:67-69. (In Rus- sian). Bachmann, K., A. Konrad, E. Oeldorf and H. Hem- mer. 1978. Genome size in the green toad (Bufo viri- dis group). 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Diagnoses reptilium et amphib- iorum in Persia orientali a N. Zarudny collectorum. Annuaire du Musee Zoologique de l'Academie Imperial des Science, St. Petersburg 1:369-372. Nikolsky, A. M. 1897. Presmykajushchiyasya, amfibii i ryby, sobrannyya N. Zarudnym" v' vostochnoi Per- sii. Annuaire du Musee Zoologique de l'Academie Imperial des Science, St. Petersburg 2:306-361. Nikolsky, A. M. "1899", 1900. Reptiles, amphibiens et poissons recueillis pendant le voyage de Mr. N. Zarudny en 1 898 dans la Perse. Annuaire du Musee Vol. 9, p. 94 Asiatic Herpetological Research 2001 Zoologique de l'Academie Imperial des Science, St. Petersburg 4:375-417. Orlova, V. F. and V. K. Uteshev. 1986. The tetraploid toad of the Bufo viridis group from Dzungarian Gobi. Mongolia, pp. 137-141. In Rocek, Z. (ed.), Studies in Herpetology. Proceedings Ord. General. Meet. Societ. Europ. Herpetol.: Prague. Peters, G. (1971): Materialien zur Okologie und Ver- breitung der Amphibien in der Mongolei. Mitteilun- gen Zoologisches Museum Berlin 47: 315-348. Pisanets, E. M. 1978. O novom poliploidnom vide zhab Bufo danatensis Pisanets. sp. n. iz Turkmenii. Doklady Akademii Nauk Ukrainskoj SSR, Ser. B, Geol., chim. i biolog. Nauki 3:280-284. (In Russian). Pisanets, E. M. 1987. Razlichiya v kladke ikpry zele- noi i danatinskoi zhab. Vestnik Zoologii 6:80-81. (In Russian). Pisanets, E. M. 1991. Novye dannye po kariologii Bufo viridis - Complex (Amphibia, Bufonidae) i voprocy proischozhdeniya aziatskikh tetraploidnykh zhab. Herpetological Researches 1:41-50. Leningrad. (In Russian). Pisanets, E. M. 1992a. Evidence of hybridization between di- and tetraploid toads (Bufo viridis - com- plex) from Eurasia (Anura:Bufonidae). Pp. 357-360. In Korsos, Z. and I. Kiss (eds.). Proceedings 6th Ord. General. Meet. Societ. Europ. Herpetol.: Budapest. Pisanets, E. M. 1992b. Poliploidiya i otkloneniya v morfologii gonad u tetraploidnikh zhab Bufo danaten- sis (Amphibia: Anura: Bufonidae). Citologija i gene- tika 26:35-39. (In Russian). Pisanets, E. M. and N. N. Shcherbak. 1979. Sistema- tika zelenykh zhab (Amphibia, Anura) fauny SSSR. Vestnik Zoologii 4:1 1-16. (In Russian). Pisanets, E. M. and O. I. Vasilenko. 1995. Reproduc- tive age of di- and tetraploid toads (Bufo viridis turan- ensis and B. danatensis) in South Turkmenia. pp. 46- 47 In Ananjeva, N. B. (ed.) Abstracts of the 2nd Asian Hepetol. Meet. Ashgabad. Pisanets, E. M. , S. V Mezhzherin and N. N. Shcherbak. 1996. Issledovanya po gibridizacii i vneshei morfologii aziatskikh zhab (Amphibia: Bufonidae) i opisanie novogo vida Bufo shaartusien- sis sp. nov. Dopovidi nacional'noi akademii nauk Ukraini 6:147-151. (In Russian). Roth, P. 1986. An overview of the systematics of the Bufo viridis group in Middle and Central Asia. Pp. 127-130. In Rocek, Z. (ed.) Proceedings of the 6th Ordinary General Meeting of the Societas Europea Herpetologica. Prague. Roth, P. and P. Rab. 1986. Karyotype analysis of the Bufo viridis-Group: Systematic implications. Pp. 131- 136. In Rocek, Z. (ed.) Proceedings of the 6th Ordi- nary General Meeting of the Societas Europea Herpe- tologica. Prague. Roth, P. and P. Rab. 1987. Sequential chromosome banding studies in the natterjack toad, Bufo calamita. 335-338. In Van Gelder, J. J., H. Strijbosch and P. J. M. Bergers (eds.). Proceedings 4th Ordinary General Meeting of the Societas Europea Herpetologica. Nijmegen. Schmid, M. 1978. Chromosome banding in Amphibia. I. Constitutive heterochromatin and nucle- olus organizer regions in Bufo and Hyla. Chromo- soma 66:361-388. Schmidt, K. P. 1952. Diagnoses of new amphibians and reptiles from Iran. Nat. Hist. Misc. Chicago Ac. Sci. 93:2. Schmidt, K. P. 1955. Amphibians and reptiles from Iran. Vidensk. Medd. Dansk. Naturh. Foren 117:193- 207. Schmidtler, J. J. and J. F. Schmidtler. 1969. fiber Bufo surdus; mit einem Schlussel und Anmerkungen zu den ubrigen Kroten Irans und West-Pakistans. Salamandra 5 (3/4): 113-123. Schneider, H. and E. M. Egiasaryan. 1995. The mat- ing call of the green toad, Bufo viridis viridis (Anura, Amphibia), in Kazakhstan. Russian Journal Herpetol- ogy 2:1-4. Schroer, T 1996. Morphologie und Ploidiegrade von Wasserfroschen aus unterschiedlichen Populations- systemen in Nordost-Polen (Anura, Ranidae). Zeitschrift fur Feldherpetologie 3:133-150. Shcherbak, N. N. and M. L. Golubev. 1981. Novye nakhodki zemnovodnykh i presmykayushchikhsya v Srednei Azii i Kazakhstane. Vestnik Zoologii 1:70-72. (In Russian). Stock, M. 1995. Morphologische, ethologische und cytologische Untersuchungen zum taxonomischen Status di- und tetraploider Griinkroten (Bufo viridis - Komplex) in Mittelasien. Diploma Thesis, Martin- Luther-University Halle- Wittenberg, 107 pp. Stock, M. 1997a. Mating call differences between diploid and tetraploid green toads (Bufo viridis com- plex) in Middle Asia In Rocek, Z. and Hart, S. (eds.) Herpetology '97, Abstracts of the Third World Con- gress of Herpetology 2-10 August. 1997, Prague, Czech Republic. Stock, M. 1997b. Untersuchungen zur Morphologie und Morphometrie di- und tetraploider Griinkroten (Bufo viridis -Komplex) in Mittelasien (Amphibia: 2001 Asiatic Herpetological Research Vol. 9, p. 95 Anura: Bufonidae). Zoologische Abhandlungen Staatliches Museum Tierkunde Dresden 49:193-222. Stock, M. 1998a. Mating call differences between diploid and tetraploid green toads (Bufo viridis com- plex) in Middle Asia. Amphibia-Reptilia. 19:29-42. Stock, M. 1998b. Tetraploid toads (Bufo viridis com- plex) from north-western China and preliminary taxo- nomic conclusions for Bufo nouettei Mocquard, 1910. Zeitschrift fur Feldherpetologie (Bochum) 5:139-166. Stock, M. and W.-R. Grosse. 1997a. Erythrocyte size and ploidy determination in green toads {Bufo viridis complex) from Middle Asia. Alytes (Paris) 15:72-90. Stock, M. and W.-R. Grosse. 1997b. New data from the Bufo viridis complex in Central Asia. In Rocek, Z. and Hart, S. (eds.) Herpetology '97, Abstracts of the Third World Congress of Herpetolgy 2-10 August. 1997. Prague, Czech Republic. Stock, M., W.-R. Grosse, M. Schmid and C. Steinlein. 1998. Ploidy level diversity in the Bufo viridis com- plex of Central Asia Zoology (Jena). Suppl. 1:76. Stock, M., M. Schmid, C. Steinlein and W.-R. Grosse. 1999. Mosaicism in somatic triploid specimens of the Bufo viridis complex in the Karakoram with examina- tion of calls, morphology and taxonomic conclusions. Italian Journal of Zoology 66 (3): 215-232. Toimastov, S. S. 1989. Osobennosti morfologii zhab iz okrestnostei oz. Jashil'kul' na Pamire. Izvestiya Akademii Nauk Tadzhikskoi SSR, otdelenie biolog- icheskikh nauk 4 (1 17):43-48. (In Russian). Toktosunov, A. T. 1984. Ecological basis of altitude adaptation of the vertebrates of Tian-Shan. Nauka Publ. House. Leningrad. (In Russian). Toktosunov, A. T. and E. Yu. Mazik. 1977. 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Vol. 9, p. 96 Asiatic Herpetologieal Research 2001 Apppendix 1 L\£' Ploidy Topographic description Source Method 1 2n Iran. Zagros Mountains. Gholaman. 30 km W Khorram „„..„, _,„„ Abad. 33=25' N. 48° 1 2' E preSent paper erythr. size, microdens. la 2n Iran. SW-slope of Zagors mountains. Khuzestan Province. Choqa Zanbil. 32°31'N. 48°32' E. 560 m a.s.l. present paper low cytom. 2 2n iran. N-slope of Elburz mountains, valley 15 km S Gorgan, approx. 1 100 m a.s.l. Stock (1995) chrom count, of larvae and uveniles 2a 2n Iran. S-slope of Elburz mountains. N of Theran. near Polur. approx. 2350 m a.s.l. Martens unpublished, present paper calls 2b - Iran, S-slope of Elburz mountains, N of Theran, valley NE of Fasham. aporox. 2540 m a.s.l. Martens unpublished, present paper calls 2n Iran, S-slope of Elburz mountains. Lar valley NE Andren and Nilson (1979). present paper calls 2c 2n Iran. Tehran Province, N environs of Tehran, garden of Plant Pests and Diseases Research Institute Borkin et al. (2000) flow cytom. 2d 2n Iran. Tehran Province, approx. 70 km W of Karaj. Karpuz-Abad village Borkin et al. (2000) flow cytom. 3 2n Iran, NE. frontier zone near Turkmenistan, approx. 50 km NE Gonbad-e-Kavus, 250 m a.s.l.. 37°38' N, 55°29' E Stock (1997a), Stock and Grosse (1997a), Stock (1998a) chrom. count, of larvae, adults, erythr. size 4 4n Turkmenistan Nebil-Dagskii Rayon N-slope of Stflck ( , Qg7a) StSck wd Grosse ( 1997a), chrom. count, of larvae, adults, Bolshoi Balkhan. approx. 15 km S of Oglanly village, stock (1998a) erythr size, calls 500 m a.s.l., 39°43' N, 54°29' E 5 2n Turkmenistan. Ashgabadskava oblast near Danata Pisanets (1978), Borkinetal. (1986a) chrom. count 2n -"- Mezhzherin and Pisanets (1990), Pisanets (1992a) ext morph. 4n, 3n Pisanets (1978) chrom. count 4n Borkinetal (1986a) chrom. count. 4n Mezhzhenn and Pisanets (19901. Mezhzhenn and Pisanets ( 1991 ). Mezhzherin and Pisanets (1995a. b) chrom. count and/or ext morph. 4n Ataev (1987), Pisanets (1992a) ext morph. 4n 1 Pisanets (1992b) chrom count, ext morph. 4n Turkmenistan, Ashgabadskaya oblast, stream 2-4 km SE of Danata village and warm spring approx. 4 km SE of Danata. 200 m a s 1 Stock ( 1997a), Stock and Grosse (1997a), Stock (1998a) chrom. count, of larvae, adults, erythr. size, calls 7 4n "South-west Kopet-Dagh", not exactly localized Pisanets and Vasilenko (1995) method not described 6 4n Turkmenistan, Kuruchsudon-Reserve, Kopet-Dag (not exactly localized) Borkinetal (1986a) chrom. count, and/or flow cytom. 7 2n Turkmenistan, Kyzyl-Arvatskii Rayon. Kopet-Dag- Range, valley approx. 25 km SW of the station Bami. S of the pass. 750 m a.s.l., 38°37' N. 56°38' E Stock (1997a). Stock and Grosse ( 1 997a), Stock (1998a) chrom. count of larvae, erythr. size, microdens. 8 2n Turkmenistan. Aydere oasis, 1 Kara-Kala Borkinetal. (1986a) chrom count and/oi flow cytom 4n Turkmenistan. Adere-oasis. E Kara-Kala Borkinetal. (1986a) chrom. count and/or flow cytom. 4n i Turkmenistan, Kara- Kala Mezhzhenn and Pisanets (1995a. b) chrom. count and/or ext. morph. 9 2n Turkmenistan, lake Sarykamysh Borkinetal (1986a) chrom. count 9a 2n „ ., ._ — ~ o ut. Schneider and Egiasarvan (1995), Kazahkstan. Guryev Town. Chornaya Rechka Dujsebaveva et al. ( 1997) calls 10 2n Turkmenistan. Ashgabadskava oblast. Bacharden Pisanets ( 1992a) ext morph. 2n .". Mezhzhenn and Pisanets ( 1991 1 method not mentioned 2n Turkmenistan, Ashgabadskaya oblast, Bacharden Mezhzhenn and Pisanets (1995a. b) chrom. count.and/or ext. morph. 2n Turkmenistan. Ashgabadskaya oblast. S Bacharden, approx. 10 km W Kelyata, 500 m a.s.l., 38°14' N, 57°31'E Stock (1997a). Stock and Grosse (1997a), Stock (1998a) chrom. count, of larvae, erythr. size, microdens. 11 2n Turkmenistan. Ashgabadskaya oblast near Ashgabad Borkinetal. (1986a) chrom. count 2n Mezhzherin and Pisanets ( 1 990), Mezhzherin and Pisanets (1991 1 ext. morph. 2n Turkmenistan, Ashgabad, village Kodzh Mezhzhenn and Pisanets (1991) method not mentioned 2n Mezhzhenn and Pisanets (1995a. b) chrom count and/or ext morph. 2n Turkmenistan, Ashgabadskaya oblast near Ashgabad ! Pisanets and Vasilenko (1995) method not described 2n Pisanets (1991) chrom. count 2n Turkmenistan, Ashgabadskava oblast, Ashgabad Stock (1995) chrom. count, of larvae 4n Turkmenistan, Ashgabad Roth and Rab (1 9861 chrom. count. 12 4n Turkmenistan, village Kuruchaudan Pisanets (1987) clutch 12a 2n Iran, 20 km WNW Kapkan. Khorasan, 37°22' N, 58°32'E, 1670 m a.s.l. present paper erythr. size, microdens. 13 4n Iran. 10 km NE Baghestan. Khorasan. N of Kuh-e- Kalat 34°09' N. 58°25' E. 1900 m a.s.l. Stock et al. (1998a). present paper chrom. count, erythr. size, microdens. and flow cytom 13a 4n Iran, Khorasan, Birjand, 32°33' N, 59° 10' N, about 1500 m a.s.l. present paper chrom. count, flow cytom. 13b 2n Iran. Kerman Province, Kerman, 30° 18' N, 57°05' E, 1860 m a.s.l. present paper chrom. count, flow cytom. 13c 2n Iran. Kerman Province, Bahr-e-Aseman Mountains, Sarduiyeh village in Sarduiyeh District 2500 m a.s.I. Borkin et al. (2000) flow cytom. 14 , Turkmenistan, lake Shach-Senem (not exactly zn [localized, SE of 9) Borkinetal. (1986a) chrom. count.and/or flow cytom 15 2n Turkmenistan, Ashgabadskaya oblast near Iolotan Mezhzherin and Pisanets (1990), Mezhzherin and Pisanets (1991) ext. morph. 16 4n Badchyz-Reserve, AkarChechme Pisanets (1978) chrom. count 4n Borkin et al. (1986a) chrom. count 4n _ " . Mezhzhenn and Pisanets (1995a, b) chrom. count.and/or ext. morph. 17 2n Uzbekistan. Buchara Roth and Rab (1986) chrom. count 2001 Asiatic Herpetological Research Vol. 9, p. 97 IS 2n Uzbekistan. Samarkand Borkrn et al (1986a) chrom cuunl 19 4n Uzbekistan. Dzhisakskaya oblast Ravon Farislt Stock (1997a), Stock and Grossed 997a), chrom. count, of larvae, adults, Nurauu-Reserve N-slope r of Nuratau Range, 900 - | „ k , 9Qg ^ ■ ca,K 1600 ma.s.l.. 40 35 N. 66 30 E | ' 4n Uzbekistan, Dzhisakskaya oblast. Rayon Farish, S Stock (1997a), Stock and Grosse (1997a), chrom. count, of larvae, erythr. size, bank of Aidar-Kul near of Nuratau- Range, 300 m as. 1. Stock (1998a) microdens., calls 20 4n Tadzhikistan. 60 km E of mouth of Janob into Zeravshan river, right bank of Zeravshan river, near Pasucax Kryukovetal. (1985) chrom. count. 21 4n Tadzhikistan, Gissar- Range. Ziddi. 3000 m a.sl. Borkinetal. (1986a) chrom count 22 -In Tadzhikistan. Ramit-gorge, Chuhgaram Borkinetal. (1986a) chrom count 4n Tadzhikistan. Ramit-gorge. Chuligaram Mezhzhenn and Pisanets ( 1995a, b) chrom. count and/or ext morph 4n ■"- Pisanets (1992b) chrom. count., ext. morph , Tadzhikistan, 1 5 km SW of the mouth of Sardan- n Miena and Sorbo. gorge of Kafinugan, near Javroz Kryukovetal. (1985) chrom count. 23 2n and 4n Tadzhikistan, Dushanbe Roth and Rab (1986) chrom. count. 4n Tadzhikistan, N Dushanbe Kryukovetal. (1985) chrom. count 24 4n Uzbekistan, Gissar- Range, 20 km of mouth of Tupalang-Darya. 800 m a si Borkinetal. (1986a) chrom. count. 25 2n Tadzhikistan, 90 km SW Dushanbe. Kafirnigan-valley Borkinetal. (1986a) chrom. count. 26 4n 1 adzhikistan, 90 km S Dushanbe. Kafirnigan-valley. near lsambai Pisanets (1992b) chrom. count., ext. morph. 27 4n Tadzhikistan. S Pamir, near Lyangar Mezhzhenn and Pisanets (1990) ext. morph. 4n Tadzhikistan. S Pamir, near Lyangar Mezhzhenn and Pisanets (1991) method not mentioned 4n 1 -"-1800m a.sl Pisanets 1 1992b) chrom. count . ext. morph 27a 4n Tadzhikistan, near Faizabad Mezhzhenn and Pisanets (1991) method not mentioned 4n I ad/hikistan. near Faizabad Pisanets (1991) chrom count 4n Tadzhikistan. near Faizabad Mezhzhenn and Pisanets ( 1995a. b) chrom count. and/or ext. morph. 4n - " - .localization using the information:„I20 km N of Dushanbe" in: Pisanets (1992a) ext. morph. 4n . " _ Pisanets 1 1992b) chrom. count., ext. morph. 4n ladzhikistan. Vakh valley, environs of Obi-Garm Mezhzhenn and Pisanets 1 1991 1 method not mentioned 4n Tadzhikistan. Obi-Gann. 2300 m a.s.l. Pisanets (1992a) ext. morph. 4n ladzhikistan. Obi-Garm Mezhzhenn and Pisanets (1995a, b) chrom. count. and/or ext. morph 4n . ". Pisanets (1992b) chrom count . ext. morph 27b 4n fadzhikistan. mouth of Komarou-river. 2000 ma.s.l. (not exactly localized) Borkinetal. (1986a) chrom. count. 28 2n Tadzhikistan. 10-12 km SW of the mouth of Vakhsh and Javansu. near Kurgan-Tiube Kryukovetal. (1985) chrom count. 2n Tadzhikistan. 1 0- 1 2 km W of the mouth of Vakhsh and Javansu. left of Vachsh, Kuibyshevskii Rayon Kryukovetal. (1985) chrom. count. 2n Tadzhikistan, 50 km NE of mouth of Vakhsh into Pvandzh. near Dusti Kryukovetal. (1985) chrom. count. 29 2n Tadzhikistan. Chirik Borkinetal. (1986a) chrom. count. 2n environs of Chaartuz. Chirik Pisanets (1991 i chrom. count. 2n Tadzhikistan, Beshkenlskre peski, valley of Kafinugan nver. environs of Chaartuz Mezhzhenn and Pisanets ( 1991) method not mentioned 2n -"- Pisanets (1992a) ext morph. 30 4n Tadzhikistan, S of Aktau-Range, 80 km S Dushanbe Kryukovetal. (1985) chrom. count 31 4n Uzbekistan, Tashkent Borkinetal. (1986a) chrom count and/or flow cy torn. 4n . " . Kudryavcev et al. (1988) cytophotometry 4n . " . Roth and Rab (1986) chrom. count 4n Uzbekistan, Tashkent, 450 m as.l., 41°16' N. 69°13' E Stock [1997a), Stock and Grosse (1997a) Stock (1998a] chrom. count of larvae, adults, erythr. size, calls 31a 2n Kazakhstan, environs of Chimkent Mezhzherin and Pisanets (1995a, b) chrom count and/or ext. morph. 4n Kazakhstan, environs of Chimkent Mezhzhenn and Pisanets ( 1991) method not mentioned 31b 4n Kazakhstan. Kyzyl-kum desert, 50 km SW of Bairkurn DuJ,ehayeva e, al. ( 1997), Castellanoet al. village, Chimkent region. 250 ma.s.l. 67°25 N, 42 02 ,„-Ug ' chrom count 31c 4n Kazakhstan, Chimkent region. Jagbagly village, 1 100 mas.l..70i32,N,42°25'E Dujsebayeva et al. (1997), Castellano et al. (1998) chrom. count. 32 4n Uzbekistan. 80 km E Tashkent, entry of Chatkal- Reserve. 5 km SE Burchmulla, 900 m a.s.l., 41°35' N, 70°07' E Stock ( 1 997a), Stock and Grosse (1997a), Stock (1998a) chrom. count. and/or erythr. size, calls 33 4n Uzbekistan. Kuraminsky-Range, 3000 ma.s.l. Borkin et al. (1986a) chrom. count 34 4n Tadzhikistan. S Pamir, near Khorog Mezhzherin and Pisanets (1990) ext morph. 4n Tadzhikistan. Gorno-Badakhshan. S Pamir, near Khorog Mezhzherin and Pisanets (1991) method not mentioned 4n - " - Mezhzhenn and Pisanets (1995a. b) chrom count and/or ext morph 4n -"- Pisanets (1992a) ext morph 35 4n ladzhikistan, S Pamir, near Ishkashim Mezhzhenn and Pisanets ( 1990i ext morph. , Tadzhikistan, Gorno-Badakhshan, S Pamir, near Ishkashim Mezhzherin and Pisanets (1991) method not mentioned 4n Tadzhikistan. S Pamir, near Ishkashim Me/hzhenn and Pisanets ( 1995a, b) chrom. countanoVor ext. morpjt chrom count 36 2n | Kazakhstan, Dzhambul Roth and Rab ( 1 986) 36a 2n (outside the map ) Kazakhstan. Tengis Lake. 1 2 km W Dieterich leg., examined by Stock, ol Abaya village, about 50 40 N 69 4(1 1 unpublished flow cytometry (one specimen) 37 4n Kazakhstan, desert near Burubaital, S Balchash lake region Egembergdieva ( 1983) cited by Borkin et al. (1986a) unknown, original paper not available 4n ! Kazakhstan, desert near Burubaital,SBalchash lake Borkinetal (1995) flow cytometry region Vol. 9, p. 98 Asiatic Herpetological Research 2001 37a 4n Kazakhstan. Karaoi village. 350 m as 1 , 74 47' N. Dujsebayeva et al ( 1997). Castellano et al. 45°54'E (1998) :hrom. count., calls 37b 4n Kazakhstan. Zhidely Channel. Hi River Delta, 370 m Dujsebayeva et al. (1997), Castellano et al. a.s.i.,75°12'N.45°18'E (1998) :hrom. count., calls 38 2d _,. „ ,„„ , Mazik. Kadyrova and Toktosunov (1976) and Kyrgyzstan, Chu-valley. 650 m a.s.l. C|ted bv Borkin et al . ( 1 9Sh., 1 :hrom. count. 2n Kyrgyzstan, Chu-valley Toktosunov and Mazik (1977) method not mentioned 2n Kvfvzstan Tulek Castellano et al (1998) chrom count, calls 2n Kyrgyzstan, Tyulyok, not exactly localized | Borkin et al. (1986a) chrom. count.and/or flow cytom. 39 2n Kyrgyzstan, Frunze (Bishkek) Bachmann et al. (1978) cytophotometry 2n Borkin et al 1 1986a) chrom count, and/or flow cytom. 3n 4n Kyrgyzstan. S Bishkek Kuzmin(1995 187) Kyrgyzstan. Frunze (Bishkek) Borkin et al. (1986a) chrom. count. 4n Kvrgvzstan. environs of Frunze (Bishkek) Mezhzhenn and Pisanets ( 1991 ) jmefhod not mentioned 4n Bachmann et al. (1978) cytophotometrv 2n Kyrgyzstan. Bishkek (Frunze) Stock ( 1 997a ), Stock and Grosse ( 1 997a) chrom. count.and/or erythr. size ? 4n Kyrgyzstan. Kyrgyzskn Khrebet, 1200 m a.s.l. and 2500 m a.s.l.. not exactly localized Fikhtman(1989) method not mentioned 9 2n, 3n, 4n Kyrgyzstan. Kok-jar, 25 km S of Bishkek, approx. 1300 m a.s.l., artificial bassin about 6 km from the villare Castellano etal. (1998) chrom. count., calls 40 4n Kyrgyzstan, Toktogul-valley. 900 m a.s.l. Toktosunov (1984) and cited by Borkin et al. (1986a) chrom. count. 41 2n? Kyrgyzstan, SW slope of Fergana- Range, near Arslanbob, 1 800 m a.s.1. Pisanets and Shcherbak ( 1979) method not described 3n .". Stock (unpublished) chrom. count, (one juvenile animal) 4n _ " . Borkin et al. (2000) method nut mentioned 42 4n Kyrgyzstan, Arpa-valley. 3500 m a.s.l. Toktosunov (1984) and cited by Borkin et al. (1986a) chrom. count. 43 4n China, Kashgar, 39°29'N, 76°02'E. 1350 m a.s.l. Stock (1998b) chrom. counl . flow cytom . and/or ervlhr size, calls 44 4n China, E-Pamir. Taxkurgan, 37°47'N, 75° WE, 3350 m a.s.l. Stock et al. (1998a), Stock (1998b) chrom. count., calls 45 4n Tadzhikistan, Central Pamir, lake Jashilkul. 3734 m Toimastov (1989) ext. morph. 4n - " - (also 3n, 2n ?) Mezhzhenn and Pisanets ( 1 990) ext. morph. 4n _ " - Mezhzhenn and Pisanets (1991) method not mentioned 4n - " - (also 3n, 2n ?) Pisanets (1992a) est morph 4n - " - (also 3n, 2n ?) Pisanets (1992b) chrom. count., ext. morph. 4n - " - (also 3n, 2n ?) Mezhzhenn and Pisanets ( 1 995a, b) chrom. count.and/or ext morph. 46 3n Pakistan. Karakoram, Sust, Hunza-Valley, 36°46iN, 74°50'E, 2950 m a.s.l. Stock etal. (1998. 1999) chrom. count., flow cytom., and/or erythr. size, calls 47 , Pakistan. Karakoram. Pasu, Hunza- Valley. 36°30'N, ia 74°52'E. 2600-2800 m a.s.l. Stock et al. ( 1 998, 1 999 ). present paper chrom. count., flow cytom., and/or erythr. size, calls 47a 3n Pakistan. Northern Areas, Hunza-Valley, river bank. Karimabad near Ganesh, 36°I8'N, 74°4FE, 2060 m a.s.l. present paper flow cytom. 48 3n Pakistan, W-Himalayas, Upper Swat valley. Kulalai, 35° 1 9'N, 72°36'E, 1 750 ma.s.1. Stock etal. (1998, 1999) chrom. count, only one specimen), and/or erythr. size, calls 48a 3n Pakistan, Northern Areas, Gilgit, 35°54'N, 74°24'E, 1550 m a.s.l. Stock etal. (1998, 1999) chrom. count., flow cytom.. and/or erythr. size, calls 48b 3n Pakistan, Northern Areas, Gupis, 36°14'N, 73°27'E, opposite Yasin valley, 2160 m a.s.l. present paper flow cytom. 48c 3n Pakistan, Shandur Pass, Lake, border of NWFrontier Prov. and Northern Arears, 3720 m a.s.l. present paper flow cytom. 48d 3n Pakistan, NWFrontier Prov.. Hindu-Kush, Buni, approx. 36°20'N, 72°20'E, approx. 1900 m a.s.l. present paper flow cytom. 48e 3n Pakistan, NWFrontier Prov., Hindu-Kush, Chitral City, 35°53'N. 71°47'E, 1480 m a.s.1. present paper flow cytom. 48f 3n? Afghanistan. Kabul ?. 34°31iN, 69°12'E, sample not surely localized for Kabul Bachmann et al. (1978): "36% more DNA than diploid B. viridis ", Hemmer et al. (1978) microdens. 49 2n Pakistan, Northern Areas (Baltistan). Himalaya, Satpara river SW of Skardu, 35"17- N, 75°37'E. 2300 m a.s.1., see also M and N present paper flow cytom., chrom. count 50(?) 4n China, Xinjang; Hotan; localization in the present map not sure Whu Mm and Zhao Yajiang (1987) chrom. count. 51 4n Kyrgyzstan. Naryn Stock (unpublished) chrom. count. 52 4n Kyrgyzstan. Kara-Kudzhur-valley Toktosunov (1984) and cited by Borkin et al. (1986a) chrom. count. 52-57 4n Kyrgyzstan. whole environs of Issyk-Kul. 1670 m a.s.l Mazik etal. (1976) chrom. count 4n .-. Toktosunov (1984) and cited by Borkin et al. (1986a) chrom. count. 4n Roth and Rab ( 1 987b) , chrom. count. 4n Borkin ( 1 989) partly chrom count. 4n .". Mezhzhenn and Pisanets I !99Sa. b) Fikhtman(1989) chrom. count.and/or ext. morph. 4n .". method not mentioned 53 4n Kyrgyzstan, Chu-valley. approx. 20 km W Rybache, Stock (1997a), Stock and Grosse (1997a) erythr. size 4n Kyrgyzstan. SW bank of Issyk-kul Me/hzhenn and Pisanets ( 1991 ) method not mentioned 2001 Asiatic Herpetological Research Vol. 9, p. 99 ,, . Kyrgyzstan. Issyk-kul. N-bank near Sary-Kamysh. Stock ( 1997a), Stock and Grosse (1997a). —al- .:„ —n. 34 n 1670 m a.s.l.. 42°29'N,76°20'E ISlock (1998a) eryuir. size, cans 4n Kvrgvzstan. Issyk-kul [Castellano et al (1998) chrom. count., calls 55 4n Kyrgyzstan, Issyk-kul. S-bank nearTamga, 1670 m a.s.l Stock (1997a). Stock and Grosse (1997a). Stock (1998a) chrom. count, of adults, erythr. size, calls 56 4n Kyrgyzstan. Rayon Tyub. village I-runze Borkin (1989) partly chrom count 57 4n Kyrgyzstan. NO-bank of Issyk-Kul. Rayon Tyub. near Kuturga Borkin (1989) partly chrom. count. 4n Kyrgyzstan. Issyk-kul. N-bank near Chon-Oryuktu. 1670 m a.s.l. Stock (1997a). Stock and Grosse (1997a) chrom. count of adults, erythr. size 58 4n Kyrgyzstan. Kemin-valley. 2500 m a.s.l. Toktosunov (1984) and cited by Borkin et al. (1986a) chrom count. 59 2n Kazakhstan. Almaty Birstein (1981). Pisanets ( 1 99 1 ) chrom. count 2n Kazakhstan. Almaty ] Mezhzhenn and Pisanets ( 1 99 1 ) method not mentioned 4n Kazakhstan, Almaty Mezhzherin and Pisanets ( 1995a. b) chrom. count. and/or ext morph. 4n Kazakhstan. Almaty Borkin et al. (1995) flow cytometry 4n Kazakhstan. Almaty. 900 m a.s.l., 76°55' N, 43° 15' E Dujsebayeva et al. (1997). Castellano et al. (1998) chrom. count. , calls 59a 2n Kazakhstan, Kopa, 20 km S of Kopa station. 75"47' N. 4< 25' E Dujsebayeva et al. (1997), Castellano et al (1998) chrom. count, calls 59b 4n Big Almaty Lake. Zailiskii Alatau Range. 2300 m as 1 , 77°N. 43°04' E Dujsebayeva et al (1997), Castellano et al (1998) chrom. count., calls 60 4n Kazakhstan. Kapchagav. Ili-river Borkin etal. (1986a). Borkin et al (1995) chrom. count and/or flow cytom. 60a . Kazakhstan. Bashn, 1 km S Altyn-Emel Mountain Range 44' 10' N. 78 "45' E Borkin etal. (1995) flow cytometry 60b 4n S foothills of the Koyandytau Mountain Range Borkin etal. (1995) How cytometry , Ayan-Saz Point. Borokhudzir river valley, between Kovandvtau and Dsunganan Alatau Mountains Borkin etal. (1995) flow cytometry 61 -, ^ 4n Kazakhstan, valley of river Chingzhal, basin of lake ' ' Alakol. 6- 1 0 km near Andreevka Golubev(1990) method not mentioned 4n Kazakhstan. Andreevka Mezhzhenn and Pisanets (1995a. b) chrom count and/or ext. morph. 4m Kazakhstan. Taldy-Kurganskaya oblast Pisanets 1 1992b) chrom. count., ext. morph. 4n Kazakhstan. Ucharal Mezhzhenn and Pisanets (1995a, b) chrom. count and/or ext. morph. 62 4n China. Xinjang; Wusu (Usu ?) Whu Mm and Zhao Yajiang 1 1987) chrom. count. 63 4n China. E-Tian Shan (E-Narat Shan), Kunas 43°14'N, 84 40'E, :i45M, Stock (1998b) chrom. count., calls 65 4n Kazakhstan. Aksnr-Farm. Zaysan-lake Borkin etal. (1986a) chrom count 4n SW-part of Zaysan-gorge, 1 2 km NW of settlement Maikapchagai near Aksiir-Farm Shcherbak and Golubev ( 1981) method not mentioned 64 4n China, Xinjang, Wulumuqui (Urumqi) Whu Min and Zhao Yajiang (1987) chrom. count 66 4n Mongolia, river Ded-Nariin-Gol, S-slope of Mongolian Altai. 2000 m a.s.l., 1 2 of the detailed map in: Borkin and Kuzmin (1988) mostly chrom count. 67 4n Mongolia, oasis Chug-Bulag, Bulgan-Somon, Chovd- Aimak Borkin etal. (1986b) chrom. count. 4n corresponds to 1. 2 and 3 of the detailed map in Borkin and Kuzmin (1988) chrom. count. 68 4n Mongolia, Ml kin W of settlement Bulgan, ( :hovd Aimak. nver Bulgan-Gol Meyer (1991) ext. morph. 69 4n Mongolia, spring Chujten-Bulak. S-slope of Mongolian Altai, 2000 m a.s.l., spring Bayan-Mod, 15 km N of spring Icher-Tol, 1600 m spring Icher-Tol, 83 km NW of settlement Bulgan, 1 600 m a.s.l., spring Chavchig- Us or Chavchig-Bulag-spring Shara-Bulag Borkin and Kuzmin (1988) partly chrom. count. 70 4n Mongolia, river Uench-Gol, 46°n.B., 92°w. L., 1350 m a.s.l. ( 1 1 of the detailed map in: Borkin and Kuzmin (1988) partly chrom. count. 4n Mongolia, spring Jaman-Usny-Bulag or Jaman-Us. 30 km E of settlement Uench, 1 800 m a.s.l., 1 2 of the detailed map in: Borkin and Kuzmin (1988) partly chrom. count. 71 4n Mongolia, oasis Ushigiin-Us, Dsungarian Gobi, 90 km SW of settlement Bulgan. 9 of the detailed map in: Borkin and Kuzmin ( 1988), Orlova and Uteshev(l986) partly chrom. count 72 4n' Mongolia, spring Domdzhigiin-Us. Dsunganan Gobi. 85 km S of settlement Bulgan. 10 of the detailed map in: Borkin and Kuzmin ( 1 988), Orlova and Uteshev(1986) partly chrom. count 73 4n China, Xinjang; Hami prefecture, 1.' km M Koumenzi. 2090 m a.s.l. Zhao and Adler( 1993) ext morph. 74 4n China. Xinjang. Hami Whu Min and Zhao Yajiang (1987) chrom. count A ? Iran, l.unstan. Shah Bazan. km 324 of the Transiranian ^h™*' ' n o^m'' ^^q,,^- „ •, .„ , , , e D , , , ,. ■ Schmidt er (1969 , Mertens 1971b), Eiselt railway, type locality of Bufo {surdus ) lunsmmcus a[)d Schmidtler ( , 973 . see als0 0 ^ s no ploidy determination B 2n? Iran. NW pan of Central Iranian Plateau. Cheshmeh-ye Sefied-Ab, type locality of Bufo ku\ir?n.sis AndnSn and Nilson (1979), Stock (1998a): diploid'?, present pajper calls C 2n Iran, Kerman, type locality of Bufo viridis kermunensi* Eiselt and Schmidtler ( 1 97 1 ), Hemmer et al. (1978), present paper chrom. count, flow cytometry D 4n Turkmenistan, Achgabadskaya oblast, Danata village, type locality of Bufo danaiensis Pisanets (1978). Borkin and Kuzmin (1988), see also number 5 chrom count E 4n E-Iran, Ssaman Shakhi mountains near Birjand, type locality of Bufo oblongus Nikolsky (1896 1897), Eiselt and Schmidtler (1973). Roth (1986). Borkin and Kuzmin ( 1 988). present paper chrom. count, flow cytometry F 2n Hemmeret al. (1978). Pisanets and Shcherbak Tadzhikistan. Dushanbe, type locality of Bufo i in dis (1979), Roth (1986), Borkin etal. (1986b), no ploidy determination of the type turanensis Kuzmin (1995, 1999), Mezhzherin and series Pisanets (1995a. b). see also number 23 Vol. 9, p. 100 Asiatic Herpetological Research 2001 c -, Tadzhikistan, SW-part Beshkent desen, near Shaartus. PisanetM 1991 1, Pisanets el al (1996), •aasA type localitv of Bufo shaarmsiensis Kuzmin (1999). see also number 29 H ? „ .. . ~ .. , , ... rn_r / • -j- \ Eiselt and Schmidller ( 1 973 ). Borkin and Pakistan, Pishin, type locality of Bufo (vmdts ) Kuzmn ,98g S(6ck ■ a, 999 Borkl[) e( zugmayen 'al.(2000) no ploidy determination J 3n and/or 4n(?) Pisanets and Shcherbak (1979: diploid Kyrgyzstan. Arslanbob. type locality of Bufo viridis without determination), Borkin and Kuzmin asiomontanus (1988), Borkin et al. (1997), Kuzmin (1999), Borkin et al. (2000) chrom. count, (only one triploid specimen tested by Stock, unpubl.l. calls K 4n „ „ _. . . , „ . ,„, Kashchenko(1909). Stock (1997a), Stock and Kyrgyzstan, Kokmoinok. terra typica of Bufo Qws%e , 99? R mjn , 999 see ^ chrQm (vind«)iim<:otor number 52 to 57 L 4n China. Kashgar. one of three type localities of Bnjo Mocquard (1910). Borkin and Kuzmin (1988), nouenei sensu Mocquard; type locality of the B Stock ( 1 998b), additional papers are cited in nouettei lectotvDe the last one chrom. count., calls, flow cytometry M 2n Pakistan, Shinu village near Siachen glacier, type locality of Bufo siachinensis Khan ( 1 997 ). Baig ( 1 998 ), Stock et al. ( 1 998, 1999): junior synonym of B. latastii. see N and 49 chrom. count., flow cytometry N 2n N-India. Ladakh. a locality was not exactly described, terra typica of Bufo latastii Boulenger (1882: 295), Dubois and Martens (1977), Hemmer et al. (1978), Pisanets and Shcherbak (1979), Gruber (1981 ), Borkin et al. (1986a). Roth (1986). Borkin and Kuzmin (1988), Khan (1997). Baig (1998), Kuzmin ( 1999), Stock et al. (1999). see M and 49 chrom. count, flow cytometry O 3n Pakistan, Swat valley, Mingora province, terra typica of Bufo (viridis ) pseudoraddei pseudoraddei Mertens (1971a), Pisanets and Shcherbak (1979). Roth (1986), Borkin and Kuzmin (1988), Baig (1998), Kuzmin (1999), Stock et al. (1999), additional papers are cited in the last one chrom. count, (only one specimen). calls P 3n N-Pakistan, Karakoram, type locality of Bufo pseudoraddei baturae Stock etal. (1999) chrom. count., calls, flow cytometry Q ? SE-Iran, Bazman, Tamin in Sarhad, Duz-Ab. Ziaret in Sarhad. type locality of Bufo persicus Nikolsky ( 1 899). Carvsky ("1925". 1 926): "synonym of B surdus ", Schmidt (1955), Schmidtler and Schmidtler ( 1969). Eiselt and Schmidtler (1973), see also R and S no ploidy determination R ? Belutchistan (W-Pakistan?), not exactly localized, terra f^X'a'n^Schrnidt'ler ( 1969?. Eisllt^d typica of Bufo surdus Schmidtler ( 1 973). see also Q and S no ploidy determination S 7 SW-lran. Mehkuh. 70 km S of Shiraz, type locality of Schmidtler and Schmidtler (1969), Eiselt and Bufo surdus annulatus Schmidtler ( 1 973) no ploidy determination T 4n China, type locality of Bufo ssp. taxtorensis Fei et al. (1999), Stock (1998b) no ploidy determination in ssp. description, Stock (1998b): chrom count, (only one specimen), calls U 7 China, Kok-Far (= Kokyar). type locality of the depicted specimen of Bufo viridis var. pewzowi , three other type localities for the series Bedriaga (1898: 61 and Fig. 2. plate I), Hemmer et al ( 1 978). Borkin and Kuzmin (1988), Fei etal. (1999) no ploidy determination V 7 China. Tschik-Tym (= Qiktim). Turfan, type locality of Bufo viridis var. grum-grzinuiiloi ; "Turfan": one of 14 type localities of Bufo viridis var. strauchi Bedriaga (1898: 61). Hemmer etal. (1978). Borkin and Kuzmin ( 1 988 ) no ploidy determination 2001 Asiatic Herpetological Research Vol.9, pp. 101— 106| First Record of the Smooth-Backed Parachute Gecko Ptychozoon lionotum Annandale 1905 from the Indian Mainland Samraat Pa war' and Sayantan Biswas2 Wildlife Institute of India, Chandrabani, Dehradun- 248 001, India. Present addresses: '3/21, Mohanwadi, AlandiRoad, Yerawada, Pune-4 11006, India. 2 30/3 Jheel Road, Calcutta-70003 1 , India. Abstract.- The smooth-backed parachute gecko, Ptychozoon lionotum is reported from the mainland India for the first time. The nearest known previous record was from Pegu, Myanmar, about 700 km southeast of the previous location. The species was collected in Langtlai and seen in the Ngengpui Wildlife Sanctuary, both in south Mizoram. The collected individual was kept in captivity for four and a half months, during this time, opportunistic observations on activity pattern, food habits, escape and parachuting behavior were made. Both individuals showed slow, deliberate pre-escape movement previously unrecorded for Ptychozoon. Information on morphological characters and morphometric measurements is presented. Explanations for the disjunct distribution are discussed. Key words.- Reptilia, Gekkonidae, Ptychozoon, parachute gecko, Northeast India, Myanmar, distribution, biogeography, behavior Ptychozoon is a genus of arboreal geckos distributed over much of Southeast Asia, primarily in moist tropi- cal evergreen and semi-evergreen forests (Brown, 1999; Brown et al. 1997; Smith, 1935). At present, six species are recognized under the genus: Ptychozoon kuhli, P. horsfieldii, P. lionotum, P. intermedium, P. rhacophorus and P. trinotaterra (Brown et al., 1997; Brown, 1999). To date, the only species reported for India is Ptychozoon kuhli, from the Nicobar Islands (De Rooij, 1915; Smith, 1935; Tiwari, 1961). We present here the first record of the smooth-backed parachute gecko Ptychozoon lionotum based on two records from the state of Mizoram (21°56'N to 24°31'N and 92°16'E to 93°26'E) in Northeastern India (Fig. 1). The first individual was sighted on 29th June 1998 in Lawngtlai town of south Mizoram during a short survey. Subsequently, on 21st April 1999, a second individual was sighted by SP from the vicinity of Ngengpui Wildlife Sanctuary (NWLS; 22°21'24" N to 22°30'06" N and 92°45'12" E to 92o50'20•' E) in south Mizoram, during a herpetofaunal community study (Pawar, 1999). We could only collect the first individual, and although SP could get a superficial look at the second one before it escaped (see below), we presume that it was the same species as the straight-line distance between the two sites is only about 40 km. Both the localities lie in the low to mid- elevation region of south Mizoram. The vegetation is of the tropical (moist) evergreen type, corresponding to Northern Tropical Evergreen Forest (lb/c2; Cham- 20° N 18°N BAY OF BENGAL 16° N 0 50 100 150lrilom«OT 96° E Figure 1. Present record (1) of Ptychozoon lionotum (BNHM 1445) from south Mizoram (India), along with nearest previous record (2) from Pegu (Myanmar). Vol. 9, p. 102 Asiatic Herpetological Research 2001 Table 1 . Mensural (in mm) and meristic measurements of two specimens of Ptychozoon lionotum . The vouchered record reported here (BMNH 1445) is compared with one of the syntypes (ZSI 2601). The latter spec- imen was fully discolored and severely mutilated so many characters were not discernable (NA) and so measure- ments beyond mm were not possible. Character Head length BNHM 1445 16.8 ZSI 2601 15 Head width 16.8 16 Head height 12.7 11 Eye diameter 4.8 NA Eye to nostril distance 8.2 NA Eye to snout distance 11.3 11 Eye to ear distance 7.7 8 Inter orbital distance 10.3 NA Inter narial distance 3.7 NA Tympanum diameter 2.3 3 Neck length 11.9 NA Snout to forelimb length 36.0 NA Axilla to groin length 47.0 42 Body flap width (Greatest width from base of flap to tip) 8.1 9 Body flap length (From axilla to groin) 39.6 41 Fore arm length 18.7 12 Fore limb length 27.8 21 Femur length 13.6 NA Tibia length 12.7 NA Hind leg length 22.9 20 Hind foot length 39.6 37 Length of I st Toe 6.3 NA Length of IV th Toe 9.4 NA Snout to vent length 94.6 88 Tail length 93.0 NA Tail width 7.6 NA Tail depth 6.3 NA Terminal tail flap length 20.1 NA Terminal tail flap width 10.9 NA Supralabials 10/11 NA Infralabials 9/9 NA Transverse dorsal bands in the axilla-groin region 4 NA Number of lobes fused before straight flap 7 NA 2001 Asiatic Herpetological Research Vol. 9, p. 103 Character Supranasals in contact BNHM 1445 No ZSI 2601 NA No. of tail lappets 19 NA Subdigital lamellae (L/R) Finger 1 11/10 NA Finger II 11/13 NA Finger III 12/16 NA Finger IV 15/15 NA Finger V 14/14 NA Toel 11/11 NA Toe II 12/12 NA Toe 111 16/15 NA Toe IV 14/14 NA ToeV 14/14 NA pion and Seth. 1 968) and Chittagong Tropical Ever- green Forest ( lb/c4; Wikramanayake et al. 1998). Identification of the species as P. lionotum is based on the presence of the following combination of char- acters: absence of enlarged tubercle on the dorsum; denticulate tail lobes of the tail directed somewhat backwards; tail not tapering (Smith. 1935); presence of pre-digital notch on the forearm skin fold (Brown. 1999; Brown et al. 1997; Cox et al. 1998). We also compared our specimen with one of the syntypes of P. lionotum (ZSI 2601. from Pegu. Myanmar) and a specimen of P. kuhli (ZSI 2603. from Nicobar Islands) housed at the National Zoological Collection of Cal- cutta. Selected morphometric measurements (after Brown, 1999; Brown et al. 1997; Das. 1997; Ota. 1989; Zug and Moon. 1995) were recorded using Mitutoyo Digimatic callipers, with an accuracy of 0.1 mm (Table 1 ). The color pattern of the specimen (in life) was as follows (Fig 2): Dorsally medium to dark gray with darker markings; dermal appendages lighter in color, mottled: distinct wavy dark, grayish-brown transverse bands present, four between the axilla and groin: chin and gular region dirty white to yellowish, white on chest, belly and underside of tail heavily powdered with gray-brown: underside of thighs, arms and der- mal appendages was similar to gular region in color. The collected individual showed some degree of color change in captivity, ranging from light (bands dis- tinct) to dark gray (bands barely distinct). The individual was judged to be an adult female based on the absence of preanal and femoral pores and the lack of hemipenal swellings at the tail base. The sex was later confirmed by dissection. The individual was kept in captivity in a glass ter- rarium measuring 2x1x1.5 ft, for four and a half months. During this period, we frequently took the gecko out at different times of the day, which allowed us to make additional behavioral observations, includ- ing those on its escape and parachuting behavior. The specimen was later euthanized and preserved in 70 9c ethanol after fixing in 10% formalin, and deposited in the reptile collection at the Bombay Natural History Museum (BMNH fide Leviton et al., 1985) in Mum- bai (No. 1445). Natural History and Behavior A mosaic of bamboo-dominated patches, remnant mature forest, teak plantations and jhum fallows of varying ages surround the town of Lawngtlai (900- 1000 m elevation), where the first individual of Pty- chozoon was seen. At 1930 hrs on 29th June 1998. the parachute gecko was seen in a circuit house situated in the outskirts of the town. It was resting on the inside ledge of a window in the corridor, at a height of about 2 m from the floor. The corridor was enclosed, the only entry points being the windows and the doors al the ends of the passage. In the same passage, there were a few Hemidactylus frenatus, while a nearby corridor was occupied by Cosymbotus platyurus. While resting, the dermal appendages of the parachute gecko were closely apposed to the body, and it did not show any movement, except for a vigilant but slug- gish lateral movement of the body towards the outside of the ledge when attempts were made to capture it. Vol. 9, p. 104 Asiatic Herpetological Research 2001 Figure 2. Ptychozoon lionotum (Adult female, BNHM 1445) from Mizoram, northeast India. On 6th April 1999. SP, along with his field assis- tant, spotted the second Ptychozoon at 1820 hrs, next to a dirt track in a patch of mature evergreen forest south of NWLS houndary. -40 km (straight-line) south of the first locality. NWLS is the only remaining patch of unfragmented. mature primary forest in the area, characterized by a three-tiered structure, with towering, buttressed, deciduous emergents up to 50- 60m in height, followed by middle and tertiary can- opy trees (Pawar. 1999). This area, especially the Ngengpui valley, experiences five rainless months, but the effective dry period is much shorter, with humid- ity being consistently high during these months due to fine, localized precipitation from cloud and fog. This individual was smaller than the first one and was spot- ted at a height of 5 m on the trunk of a Sterculia scaphigera tree. The tree is characterized by a deeply fluted trunk and a smooth but slightly flaking bark, and occurs as a deciduous canopy-emergent in pri- mary evergreen forest above 500 m elevation. The patch of forest was on a slope at an altitude of approx- imately 450 m. and the tree (385 cm in girth at shoul- der height) was towards the edge of the patch, slightly down slope, about 3 m from the dirt track and the observers. The gecko was sitting on the outer ridge of one of the trunk folds with its head pointing down- ward, barely visible on the lichen-covered bark. It was twilight, and upon sighting it. SP observed the animal for about a minute with the aid of a torch and binocu- lars before attempting to capture it. Meanwhile, the gecko had apparently become wary and steadily started moving laterally in the manner of the first indi- vidual away from the two observers, towards the other side of the trunk and out of sight. When SP tried to dislodge the gecko with a bamboo pole, the animal rapidly moved further around the trunk. It then lumped onto some lianas which were 2.5 m from the trunk, landed 1.5 m lower than its previous position with its head up. ran further up and vanished into a mass of dry branches which were caught in the lianas. All further attempts to trace the gecko were futile, and we presume that either the gecko jumped on to another tree or liana, or fell to the ground somewhere down slope when the lianas were shaken to dislodge it. Hemidactylus frenatus, H. garnoti, Cosymbotus platyurus and Gekko gecko are four other gekkonines that were commonly seen in the same area. During its four months of captivity, the individual was offered a variety of insects, of which it took cock- roaches and moths most readily. In the daytime, it remained motionless, either on one of the branches in its terrarium. or on one of the tar strips at the corners of the enclosure. Towards evening however, it would become active, and was often observed moving around the terrarium. making audible leaps across the corners of the enclosure. When taken out. its behavior was very different during day and night. If kept on a branch or tree trunk in the daytime, it would remain motionless with its limbs closely apposed to the trunk, and move only if provoked. If not disturbed for a long time, it would start moving slowly with the same slow, deliberate movement that it had displayed dur- ing its capture, either out of sight to the other side of the branch/trunk, or run up the tree. In the evenings however, it showed much more alacrity in trying to escape, often with the same preliminary lateral move- ment of its body. On two occasions it also resorted to launching itself into the air, and when it fell to the ground, remained motionless. This escape behavior has been earlier observed in these geckos, and remain- ing immobile ostensibly makes it difficult to locate them (Brown et ai, 1997). However, the slow pre- 2001 Asiatic Herpetological Research Vol. 9, p. 105 escape movement that we observed in both the speci- mens has not been reported before, and we reason that this behavior probably aids the gecko to position itself for parachuting or simply move out of sight (such as the blind side of a tree trunk) inconspicuously, with- out disclosing its crypsis. To gain further insights into this escape behavior that we observed in both the specimens, we dropped the individual on seven occa- sions from heights of 3-5 m and observed its behav- ior. In all instances, the dermal appendages came into play apparently due to air resistance, and the gecko dropped relatively softly on the ground without any horizontal displacement. The ecology and behavior of species of Pty- chozoon is poorly known, and there has been much discussion about its alleged ability to "glide" (Gunther, 1864; Smith, 1935; Tiwari, 1961 and refer- ences therein). It has been argued that the dermal appendages do not help in gliding, but enhance its camouflage. However, experimental studies have now demonstrated that the dermal appendages do allow the gecko to take advantage of air resistance while mak- ing long sallies (Heyer and Pongsapipatana, 1970; Marcellini and Keefer, 1976) and may serve a dual function in crypsis and escape or locomotion (Marcel- lini and Keefer, 1976). Recently, Brown et al. (1997), based on their observations of the escape behavior of P. intermedium in the wild, have argued that "para- chuting" is a more appropriate term than "gliding" to describe this behavior in these geckoes. Our observa- tions apparently sustain the arguments of Brown et al. (1997). Further studies on the preflight behavior of Ptychozoon species may provide interesting insights into the escape behavior of this extraordinary group. Biogeographical Notes The syntype (ZSI 2601 ), collected by Major Beddome and W. Theobald from Pegu in south Myanmar, was previously the northwestern most distribution record of Ptychozoon (Annandale, 1905; Brown et al. 1997). The present record thus adds a crucial link to the dis- tributional information for the genus, and increases the known range ca.700 km towards the northwest. This also adds another case of range disjunction in a region that already has numerous examples of taxa showing dramatic discontinuities in their range (Mani, 1974). After the collision of the Indian plate with the Asian mainland in the Eocene (54-36 mybp) (Molnar and Tapponnier, 1975), Indo-Malayan faunal and flo- ral elements have colonized different parts of the India, resulting in more Indo-Malayan faunal repre- sentatives within India than vice versa (Das, 1996; Mani, 1974). Geckos are notorious for their penchant for waif-dispersal (Case et al. 1994). It has been observed that Ptychozoon species are not obligate for- est dwellers (Annandale, 1904; Brown et al. 1997), and probably are capable of dispersing through both forest and human inhabited areas (Annandale 1904; Brown et al. 1997). Mizoram is dominated by the Lushai Hills, a series of parallel hill ranges running from north to south and increasing in elevation from west to east (Pachuau, 1994). To the west of these hills lie the Chittagong Hill tracts of Bangladesh, and to the east lie the Chin Hills and the Arakan Yoma mountain ranges of Myanmar. The latter, also running in a north-south direction, lie between the lowland moist evergreen forests of south Mizoram and Pegu. Along the foothills of the Arakan Yomas, flanking the west- ern side, lie more or less contiguous rainforests, which forms a habitat bridge between these two areas (Collins et al. 1991). It is likely that this species has extended its range northwards along this route. This conjecture will get firmer footing if surveys in these forests reveal the presence of P. lionotum along these tracts. The areas beyond the Lushai Hills of Mizoram and the adjoining Chin Hills of Myanmar in contrast, are more arid with relatively dry forests. Moreover, recent surveys have not revealed the presence of Pty- chozoon species in these areas, and it is unlikely that it exists there (George Zug, pers. comm.). Our inquiries revealed that not many local people know of this gecko, but those who did, opined that it was rarely seen because it mostly "lived high up in the trees". There have also been unconfirmed reports of a parachute gecko from north Mizoram (Lai Ramthanga, pers. comm.). That this area has been inadequately surveyed is evident from the fact that the six-month herpetofaunal study conducted by SP yielded a number new taxa and distributional records (Pawar, 1999). Further exploration will probably reveal that Ptychozoon is present in other parts on this region, and its range may not be as disjunct as it appears now. Acknowledgments The Wildlife Preservation Society of India and Wild- life Institute of India supported our surveys in Mizo- ram. We are particularly grateful to the Mizoram Forest Department for permits and their support in the field. Zokhima was more than a field assistant to us. J.B. Alfred, S.K. Chanda and S.K. Talukdar at ZSI, Calcutta, kindly permitted us to access the collection and K. Deuti, I. Das and N.C. Gayen helped us to trace the specimens. R.M. Brown, I. Das, A. Sinha, N.M. Ishwar and K. Vasudevan provided us with criti- Vol. 9, p. 106 Asiatic Herpetological Research 2001 cal comments and literature. Aysegul Birand helped prepare the map for Fig. 1 . Literature Cited Annandale, N. 1904. Contribution to Oriental Herpe- tology I: The Lizards of the Andamans, with the description of a gecko and a note on the reproduced tail in Ptychozoon homalocephalum. Journal of Asi- atic Society of Bengal 73:12-22. Annandale, N. 1905. Notes on some Oriental Geckoes in the Indian Museum, Calcutta, with the Description of New Forms. Annual Magazine of Natural History 7:26-32. Brown, R. M. 1999. New species of parachute gecko (Squamata: Gekkonidae: Genus Ptychozoon) from northeastern Thailand and central Vietnam. Copeia 1999(4): 990-1001. Brown, R. M., J. W. Ferner, and A. C. Diesmos. 1997. Definition of the Philippine Parachute gecko, Pty- chozoon intermedium Taylor 1915 (Reptilia: Squa- mata: Gekkonidae): redescription, designation of a neotype and comparisions with related species. Her- petologica 53:357-373. Case, T J., D. T. Bolger, and K. Petren. 1994. Inva- sions and competitive displacement among house gec- kos in the tropical Pacific. Ecology 75(2):464-477. Champion, H. G. and S. K. Seth. 1968. A revised sur- vey of the forest types of India. Government of India Printing, Delhi. 404 pp. Collins, N. M., J. A. Sayer, and T C. Whitmore (eds.). 1991. The Conservation Atlas of Tropical Forests: Asia and the Pacific. IUCN, Macmillan Press Ltd., London. 256 pp. Cox, M. J., P. P. van Dijk, J. Nabhitabhata, and K. Thiraklupt .1998. A Photographic guide to snakes and other reptiles of Thailand and South Asia. Asia Books, Bangkok. 142 pp. Das, I. 1996. Biogeography of the reptiles of south Asia. Krieger Publishing Company, Florida. 87 pp. Das, I. 1997. Rediscovery of Lipinia macrotympamim (Stoliczka, 1873) from the Nicobar Islandsjndia. Asi- atic Herpetological Research 7:23-26. De Rooij, N. 1915. The reptiles of the IndoAustralian Archipelago, Vol. I and II. E.J.Brill, Leiden. 334 pp. Giinther, A. C. L. G. 1864. The Reptiles of British India. Ray Society, London. 452 pp. Reprinted ca. 1982, Oxford and IBH Publishing Co., New Delhi. Heyer, W. R. and S. Pongsapipatana,. 1970. Gliding speeds of Ptychozoon lionotum (Reptilia: Gekkonidae) and Chrysopelea ornata (Reptilia: Colu- bridae). Herpetologica 26: 317-319. Leviton, A. E., Gibbs Jr., R. H., and Dawson, C. E. 1985. Standards in herpetology and icthyology. part I. Standard symbolic codes for institutional resource collections in herpetology and icthyology. Copeia 1985(4):802-821. Mani, M. S. 1974. (eds.) Ecology and Biogeography in India. Dr. W. Junk b. v. Publishers, The Hague. 773 pp. Marcellini, D. L. and Keefer, T.E. 1976. Analysis of the gliding behavior of Ptychozoon lionotum (Rep- tilia: Gekkonidae). Herpetologica 32(3):362-366. Molnar, P. and P. Tapponnier. 1975. Cenozoic tecton- ics in Asia: Effects of a continental collision. Science 1 89(4201 ):419-426. Ota, H. 1989. Japalura breviceps Gressit (Agami- dae:Reptilia), A valid species from a high altitude area of Taiwan. Herpetologica 45(l):56-60. Pachuau, R. 1994. Geography of Mizoram; R.T Enterprise, Aizawl. 153 pp. Pawar, S. 1999. Effect of habitat alteration on her- petofaunal assemblages of evergreen forest in Mizo- ram, North-East India. M.Sc. Thesis. Saurashtra University. 64 pp. Smith, M. A. 1935. The Fauna of British India, Cey- lon and Burma: Amphibia and Reptilia, Vol.11. - Sau- na, Vol. II. Taylor and Francis Ltd., London. 240 pp. Tiwari, K. K. 1961. The eggs and flight of the gecko Ptychozoon kuhli from Car Nicobar. Journal of Bom- bay Natural History Society 58(2):523-527. Wikramanayake, E. E. Dinerstein, P. Hedeo and D. Olson. 1998. Terrestrial Eco-regions of the Indo- Pacific Map, Publ. by WWFAVCMC, Washington D.C., USA Zug, G. R. and B. R. Moon. 1995. Systematics of the Pacific slender-toed geckos, Nactus pelagicus com- plex: Oceania, Vanuatu and Solomon populations. Herpetologica 5 l(l):77-90. 2001 Asiatic Herpetological Research Vol.9, pp. 107-112 A New Species of Eremias (Sauna: Lacertidae) from Highlands of Kermanshah Province, Western Iran Nasrullah Rastegar-Pouyani1 and Eskandar Rastegar-Pouyani2 1 Department of Biology, Faculty of Science, Razi University, Kermanshah, Iran, e-mail: nasrullah® razi.ac.ir. 'Department of Biology, Faculty of Science, Sabzevar University, Sabzevar, Iran. Abstract.- A new and distinctive species of the genus and subgenus Eremias is described from the highlands of Kermanshah Province, western Iran at about 1800 m elevation. It is easily distinguished from all other species of the typical subgenus (E. velox, E. persica, E. strauchi, E. nigrolateralis, E. lalezharica, E. afghanistanica, E. regeli, E. suphani, and E. nikolskii) by a variable number of postmentals (4-5 pairs), smaller size, and distinctive color pattern. Furthermore, it can be distinguished by having a combination of characters against any of the species in the typical subgenus. Systematics of the genus and subgenus Eremias is shortly discussed. Key words.- Lacertidae, Eremias, Eremias (Eremias) montanus. Western Iran , Zagros Mountains, Kermanshah province, Siah-Darreh Figure 1. Location of Kermanshah province on the Ira- nian Plateau. Introduction The lacertid genus Eremias Fitzinger, 1834 encom- passes about 33 species of mostly sand, steppe, and desert dweller lizards which are distributed from northern China, Mongolia, Korea, Central and south- west Asia to southeastern Europe (Rastegar-Pouyani and Nilson, 1997). The genus is Central Asian in its relationships and affinities (Szczerbak, 1974). About 15 species of the genus Eremias occur on the Iranian Plateau mostly in northern, central, and eastern regions (Rastegar-Pouyani and Nilson, 1997; Ander- son, 1999). To date, no comprehensive study has been carried out on Eremias fauna of the Iranian Plateau and the systematic status of most taxa is in great need of a revisionary work. Szczerbak (1974), however, 45 46 .47 48 Iraq \_ Iran KERMANSHAH ■ J PROVINCE Ty locality CT Hamedan □ Kermanshah^ Lorestan"^_^ 50 km 35 34 Figure 2. The type locality of Eremias (Eremias) mon- tanus, vicinity of the Village of Siah-Darreh, about 60km northeast of the city of Kermanshah, Kerman- shah Province, western Iran. revised Eremias and divided it into two distinguished genera based on morphological characters: Mesalina Gray and Eremias Fitzinger (see under systematic dis- cussion). As the first record of Eremias, in this paper we describe and introduce a new species of this genus from the upland and mountainous regions of Kerman- shah province, western Iran at about 1800 m eleva- tion. This province is located on the western periphery of the Iranian Plateau (Fig. 1 ) and a major part of it is covered by the Zagros Mountains. The type locality of Eremias (Eremias) montanus (sp. nov.) is located in an upland area surrounded by the Zagros Mountains Vol. 9, p. 108 Asiatic Herpetological Research 2001 with steppe vegetation (e.g., Astragalus, Euphorbia, Zygophyllurn), about 60 km northeast of city of Ker- manshah, Kermanshah province, western Iran (Fig. 2). Eremias montanus, new species (Figs. 3-4) Holotype and type locality: An adult female, Field number P198, collected by the senior author on 14 August 1995 from the upland regions of the Zagros Mountains, 60 km northeast of city of Kermanshah (47° 5'E, 34° 52'N), Kermanshah Province, western Iran , at about 1800 m elevation Paratypes: Two adult females. Field number PI 99- P200, other information as for the holotype. Diagnosis and comparison: A small-sized lacertid, maximum snout-vent length (SVL)= 57.2mm, tail length = 95mm, with 13-14 longitudinal and 27-28 transverse rows of ventral plates, slightly converging posteriorly; with 63-67 small, granular scales across middle of dorsum. A species belonging to the subge- nus Eremias: subocular reaches mouth edge; one fron- tonasal; two supraoculars which are not completely separated from frontal and frontoparietals; distance between the femoral pores being narrow; color pattern "striped"; inhabitant of steppe and mountain-steppe landscapes (Szczerbak, 1974: 83). On the other hand, it differs from all other species of its relevant subgenus based on having several dis- tinguishing characters; the color pattern is distinctive and it is distinguishable from all other species in this character i. e., dorsum uniformly dark-brown without spots and ocelli, interrupted by five light longitudinal stripes; the vertebral stripe bifurcating on the nape, two paravertebrals on each side; a broad dorso-lateral stripe containing one or two regularly arranged light spots (different from the other Eremias species in this character); Furthermore, it differs from each species of the typical subgenus in the following character combinations (Bischoff and Bohme, 1980; Bohme and Szczerbak, 1991; Rastegar-Pouyani and Nilson, 1997; Szczerbak, 1974) : From Eremias nigrolateralis Rastegar-Pouyani and Nilson, 1997 in having a much smaller size, lack of separation of the third pair of submaxillary shields by granular scales (0% versus 100%), lower count of gulars (23-24 versus 41-42), variable number of sub- maxillary shields (33% versus 0%), reach of femoral Figure 3. Eremias (Eremias) montanus holotype. Figure 4. Eremias (Eremias) montanus paratypes. 2001 Asiatic Herpetological Research Vol. 9, p. 109 pores to the knee (100% versus 0%), and distinct dif- ferences in color pattern. From Eremias persica Blanford, 1874 in having a smaller size, lower count of gulars (23-24 versus 28- 38), the absence of distinctly keeled upper caudal scales (100% versus 75%), variable number of sub- maxillary shields (33% versus 4%) and distinct differ- ences in color pattern. From E. velox (Pallas, 1771) in having a smaller size, in the absence of distinctly keeled upper caudal scales (100% versus 0%), variable number of sub- maxillary shields (33% versus 5%) and in color pat- tern. From E. strauchi Kessler, 1878 in having a smaller size, the absence of distinctly keeled upper caudal scales (100% versus almost 0%), variable number of submaxillary shields (33% versus 9%) and in color pattern. From E. lalezharica Moravec, 1994 in having variable number of submaxillary shields (33% versus 0%), higher count of dorsals (63-68 versus 54-59), no contact of gulars with the second pair of submaxillary shields, lack of a small scale between prefrontals, and distinct differences in color pattern. From E. afghanistanica Bohme and Szczerbak, 1991 in a much higher count of dorsal scales (63-68 versus 44-46), lower count of gulars (23-24 versus 25- 28), variable number of submaxillary shields (33% versus 0%), and in color pattern. From E. nikolskii Bedriaga, 1905 in a higher count of dorsals (63-68 versus 45-59), variable num- ber of submaxillary shields (33% versus 8%), and in color pattern. From E, regeli Bedriaga, 1905 in a higher count of dorsals (63-68 versus 43-61), higher number of scales in the 9th- 10th caudal annulus (27-28 versus 17-25), the absence of distinctly keeled upper caudal scales (100% versus 0%), variable number of sub- maxillary shields (33% versus 3%), and in color pat- tern. Description of holotype: An adult female, preserved in 75% ethyl alcohol in good condition; body small and moderately depressed; a species of the subgenus Eremias (Szczerbak, 1974:83); five pairs of submaxil- lary shields, first three pairs in contact, the fourth and fifth pairs widely separated; first pair of submaxillary shileds as large as the fifth and in contact with mental anteriorly, with first and second infralabials laterally; the fifth sabmaxillary pair each in contact with the fourth pair anteriorly, being surrounded by 8 granular scales laterally and posteriorly; 7-8 supralabials, 4-5 of which anterior to subocular which borders the mouth; two large nasals in contact with rostral anteri- orly, with first and second supralabials laterally, and with frontonasal and first loreal posteriorly, the former being single, broader than long and laterally in contact with first loreal and posteriorly with prefrontals; two prefrontals each smaller than frontonasal and almost as long as broad and laterally in contact with second loreal and posteriorly with frontal and granules of supraocular region; only frontonasal with distinct concavity; frontal almost as long as prefrontal and frontonasal together, broadened and posteriorly and laterally partly in contact with supraoculars (and partly separated from the latter by 2-3 large scales, not by granules) and posteriorly with frontoparietals; two frontopariatals almost as large as a single supraocular, laterally being in contact with the second supraocular, and posteriorly with interparietal and parietals, the former being small and relatively lozenge-shaped, surrounded by frontoparietals and parietals; two vey large and plate-like parietals, almost as long as broad, being in contact behind interparietal; a vestigial occipital; two loreals, first one small, surrounded by first two supralabials, nasal, frontonasal, and the sec- ond loreal which is distinctly large with an evident keel; 6-6 supraciliaries, separated from supraocular by a series of 42-44 granules; postocular elongate, sur- rounded by granules anteriorly; temporal region- mostly covered by granular scales becoming large towards the orbit, more than 100 on each side; tym- panic scale distinct and elongated obliquely, almost the same size on both sides; tympanum vertically elongated, slightly larger than orbit; no distinct supratemporal; subocular huge, broader than long with a distinct ridge being extensively in contact with the lower edge of orbit; lower eyelids with a semi- translucent membrane made up of about 22 enlarged scales; collar well pronounced , not serratted, made up of 10 scales, the two medial ones the largest; gular fold weakly developed, 23-24 gulars from symphysis of chin shields to median gular, becoming enlarged posteriorly; 13-14 longitudinal and 27-28 transverse rows of almost squarish ventral plates from collar to hindlimbs; anterior series of ventrals to some extent irregular, median ventral longer than broad; dorsal scales juxtaposed, smooth, granular, becoming slighly larger posteriorly, 63-68 scales across the middle of dorsum, and about 160-164 scales in a single row from occiput to a point just above the vent; proximal caudals larger than posterior dorsals but the change being gradual; caudals becoming large, elongate, and slightly keeled distally, arranged in distinct whorls, 27-28 scales in the 10th whorl behind the vent; upper forearm covered dorsally by enlarged, juxtaposed, and almost lozenge-shaped scales; lower forearm covered Vol. 9, p. 110 Asiatic Herpetological Research 2001 with granules; upper hindlimbs covered externally by granules, externally by large shields; tibia covered dorsally by slightly pointed granules, ventrally by large plates which are slightly keeled, two plates in a transverse row; no fringes on the toes, 18-20 uni-and bi-carinate lamellae under fourth toe, proximal part of lower fourth toe containing two complete rows of lamellae, distal part with a single row (in this charac- ter it is quite different from all other species of its rel- evant subgenus); 18-19 femoral pores in each side, the two series separated anteriorly by a narrow space con- sisting of three scales; preanal region encompassing 24 large shields, the four median ones being the larg- est; 6 plates in longitudinal row from the space between femoral pores to anterior edge of the vent. Coloration and color pattern: Dorsum uniformly dark-brown without spots and ocelli, interrupted by five light, narrow longitudinal stripes: one vertebral bifurcating on the nape, two paravertebrals on each side; a broad dorso-lateral stripe containing one or two regularly arranged light spots; a ventro-lateral series of dark-brown spots, to some extent forming a stripe; upper surface of head uniformly olivish-brown; temporal and labial regions suffused by dark-brown; submaxillary region light-gray-cream; ventral region dirty white, suffused by bluish-brown; upper surface of limbs dark-brown containing numerous light spots; proximal upper surface of tail brown, distal part light brownish-gray; lower surfaces of limbs and tail yel- lowish-white. Description of paratypes: The paratypes, two adult specimens, one male (PI 99) and one female (P200) approximate the holotype in almost all pertinent details. However, there are some minor differences between paratypes and the holotype as follows: Male paratype: five submaxillary shields on the right side but only four on the left, the fifth vestigial; 13-14 longitudinal and 30-31 transverse ventral plates; 10- 1 1 collars, 3-4 median ones the largest; 23-24 gulars in a longitudinal row from chin shields to collar; 20- 20 femoral pores, separated by three small scales; 23- 24 lamellae under fourth toe; 23-27 scales around 10th tail annulus; 8-9 labials, 5 of them anterior to subocular; 6-6 lower labials; lower nasal resting on the first two supralabials; temporal scale vestigial; 62- 63 scales around widest part of dorsum; 162-167 scales in a single longitudinal row from occiput to vent. Measurements (mm): SVL = 58.5; TL = incomplete; Forelimb = 24; Hindlimb = 39; HL = 15.5. Female paratype: only four pairs of submaxillary shield, the fifth vestigial; 15-17 longitudinal and 31- 33 transverse rows of ventral plates; 9 collars, 4-5 median ones the largest; 24-25 gulars in a single lon- gitudinal row from chin shields to collar; 20-20 femo- ral pores, separated by a space corresponding to two scales; 25-26 scales on the 10th tail annulus, 24-25 subdigital lamellae under fourth toe; 8-9 supra- and 4- 6 infralabials; temporal scales relatively small; frontal separated from the first supraocular by two large scales (not by granules); 65-67 scales around widest part of dorsum; 163-167 scales in a single longitudi- nal row from occiput to vent. Measurements (mm): SVL - 52.9; TL = incomplete; Forelimb = 23; Hindlimb = 35.5; HL = 13.6. In color pattern they are quite similar to the holotype. Habitat: During field work on the western regions of the Iranian Plateau in 1995, we surveyed the Zagros Mountains and the nearby mountain steppes in the northern parts of Kermanshah province. 60 km north- east of Kermanshah city, in the highland steppes, in vicinity of the Village of Siah-Darreh in an area named Sarpal, we came across to three specimens of a new taxon of the genus Eremias, described here as a new species. The habitat, which is surrounded by the Zagros Mountains, is characterized by a mountain- steppe; the vegetation is luxuriant steppe association: mainly Astragalus, Euphorbia, Artemisia, and Amigdalus as well as various species of the families Graminaceae and Compositeae (Fig. 5). Since it is a mountainous region with relatively high elevation, the winter being harsh and cold, the summer being mild and rather short. All the speci- mens were foraging around the shrubs probably look- ing for prey. They were quite shy and wary and very difficult to capture. When alarmed, they took refuge under the shrubs or inside the underground holes. One of the most effective anti-predatory adaptations evolved in these lizards is the ability to lose the tail (autotomy) when being touched by the predators (or collectors). Hence, we could only collect one speci- men with a complete tail and the other specimens lost their tails during capturing. In September 1998, we re-surveyed the type local- ity in order to find more specimens of this species but without success. Whether it being a relictual and rare species, confined only to the type locality, or being distributed over a wider area in the western margin of the Iranian Plateau is yet to be established. With regard to the occurrence of Eremias monta- nus in the highlands of Kermanshah province, the Procter record of E. velox, as the westernmost record, from Kuretu (Iran-Iraq border) (Procter, 1921:252) should seriously be reconsidered. Asiatic Herpetological Research Vol. 9. p. I 1 1 <# js»*h»- "ym -*> •" E ilfc •r ■w ■-** Figure 5. Habitat and type locality of Eremias (Eremias) montanus. 60 km northeast of Kermanshah, vicinity of Siah-Darreh village, Kermanshah Province, Western Iran. Etymology: Eremias montanus is so named as it is apparently restricted in distribution to the upland and mountainous steppes of northeastern regions of Ker- manshah province, western Iran. Taxonomic account: As mentioned before, so far, the most complete work done on the complicated genus of Eremias (sensu lato) is of Szczerbak (1974) who studied almost all species and species complexes of this genus throughout the range. Based on morpholog- ical characters and geographic distribution, S/.czerbak (1974) subdivided the inclusive genus Eremias (s. 1.) into two distinct genera; the genus Mesalina as a north African and lowland southwest Asian clade, and the genus Eremias (sensu stricto) which is mainly occurring in Central and northeast Asia. (Szczerbak. 1974). Furthermore. Szczerbak (1974) subdivided Ere- mias (s.s) into five distinct subgenera: Eremias (Szcz- erbak, 1974: 83), Rhabderemias (Szczerbak, 1974: 201 ), Ommateremias (Szczerbak, 1974: 146). Parere- mias (Szczerbak, 1974: 22-23), and Scapteria ( Szcz- erbak, 1974:247). Except for the subgenus Pareremias, which is a Central and east Asian clade, all of the major species groups of the genus are presented on the Iranian Pla- teau (Anderson, 1999). Arnold ( 1986) who studied the hemipenes of lacertids supported the Szczerbak's sub- generic names. In a more recent study Arnold placed Eremias as the sister (axon of a clade including Acan- thodactylus, Mesalina, and Ophisops-Cabrita (Arnold. 1989:238, 240 ). But Mayer and Benir ( 1994) have proposed a different scenario. According to these authors, Eremias is the sister taxon of Mesalina and both of them belong to a larger clade also containing Omanosaura and Ophisops. They believe that Eremias is not closely related to Acantho- dactylus. The Czech Biological Expedition to Iran in 1996 collected 8 specimens of an undetermined species of Eremias related to E. persica from the Zagros Moun- tains in Esfahan province at about 2000-2200 m ele- vation (Frynta et al., 1997: 9-10). Whether it is a new taxon or just a variety off. persica is yet to be known. Material examined: Eremias montanus (n = 3): P 198-200 (Field number), from around the Siah-Darreh Village (about 1800 m elevation). 60 km northeast city of Kermanshah. Kermanshah province, western Iran. Vol. 9. p. l : Asiatic Herpetological Research 2001 Eremias nigrolateralis (n = 2): GNHM. Re. ex. 5147-5148, from 150 km northeast of Shiraz. Fars province, south-central Iran. Eremias persica (n = 4): GNHM. Re. ex. 5159- 5162. from 150 km northeast of Shiraz. Fars province, south-central Iran. Eremias persica (n = 28) : GNHM. Re. ex. 5163- 5190, from 45 km east of Arak on the road to Qum, Markazi province, north-central Iran. Eremias persica (n = 4) : GNHM. Re. ex. 5191 - 5194, from 65 km west of Tehran, between Eshtehard- Saveh, Tehran province, northern Iran. Eremias persica (n = 2) : GNHM. Re. ex. 5195- 5196. from 45 km east of Golpaygan, Esfahan prov- ince, central Iran. Eremias persica (n = 4) : GNHM. Re. ex. 5197- 5200. from 50 km north of Delijan on the road to Qum. Markazi province, north-central Iran. Eremias persica (n = 1) : GNHM. Re. ex. 5201, from 50 km north of Abadeh. Fars province, south- central Iran. Eremias persica (n = 1) : GNHM. Re. ex. 5202. from 50 km east of Hamedan on the road to Qazvin, Hamedan province, western Iran. Eremias persica (n = 3) : GNHM. Re. ex. 5203- 5205. from 5 km west of Takestan on the road to Zan- jan, Zanjan province, northwestern Iran. Eremias velox (n = 4) : GNHM. Re. ex. 5122(1- 4). from around the Carin River. 250 km E-SE Almaty (Alma Ata). Kazakhstan. Eremias velox (n = 2) : GNHM. Re. ex. 5120(1- 2), from Mulali Kurozek. eastern Kazakhstan. Eremias velox ( n = 2 ) : GNHM. Re. ex. 5121(1- 2), from the Taldi Korgau District, northeast Kazakh- stan. Eremias velox (n = 2) : GNHM. GK. 18881 (1-2). from Archenjan Village ( 1 ), and 30 km north of Mary (2), Turkmenistan. Eremias strauchi (n = 3) : GNHM. Re. ex. 441 1 (1-3), from Golestan National Park, Mazandaran province, northeastern Iran. Abbreviations : GNHM. Re. ex. = Gothenburg Natu- ral History Museum. Reptilia exotica; GNHM. GK. = Gothenburg Natural History Museum. General Kata- logue. Acknowledgements We wish to thank the Razi University authorities (Ker- manshah-Iran) for their generous help and support during field work in western Iran. We thank the Gothenburg Natural History Museum (Gothenburg-Sweden) for loan of Eremias specimens. Also we thank Dr. Michael Golubev for translation of the relevant Russian literature. Literature Cited Anderson, S. C. 1999. The Lizards of Iran. Society for the study of Amphibians and Reptiles 442 pp. Arnold, E. N. 1986. The hemipenis of lacertid lizards (Sauria : Lacertidae): structure, variation and system- atic implications. Journal of Natural History 20:1221- 1257. Arnold, E. N. 1989. Towards a phylogeny and bioge- ography of the Lacertidae : relationships within an Old- World family of lizards derived from morphol- ogy. Bulletin of the British Museum (Natural His- tory). Zoology 55(2):209-257. Bischoff, W. and W. Bohme. 1980. Der systematische Status der turkischen Wustenrenner des Subgenus Eremias (Sauria: Lacertidae). Bonner Zoolgische Beitrage, N. F. 26:297-306. Bohme. W. and N. N. Szczerbak. 1991. Ein neuer Wustenrenner aus dem Hochland Afghanistans. Ere- mias (Eremias ) afghanistanica sp. n. (Reptilia : Sau- ria : Lacertidae). Bonn. Zool. Beitr. 42:137-141. Frynta. D.. J. Moravec, J. Cihakova, J. Sadlo, Z. Hod- kova, M. Kaften. P. Kodym. D. Krai. V. Pitule. and L. Sejna. 1997. Results of the Czech Biological Expedi- tion to Iran. Part 1. Notes on the distribution of amphibians and reptiles Acta Societatis Zoologicae Bohemicae 61 :3-17. Mayer. W. and G. Benyr. 1994. Albumin-Evolution und Phylogenese in der Familie Lacertidae (Reptilia: Sauria). Annalen Naturhistorischen Museums in Wien 96B:62 1-648. Moravec. J. 1994. A new lizard from Iran. Eremias (Eremias ) lalezharica sp. n. (Reptilia: Lacertilia: Lacertidae). Bonner Zoolgische Beitrage 45(1 ):6 1-66. Procter. J. B. 1921. Further lizards and snakes from Persia and Mesopotamia. Journal of the Bombay Nat- ural History Society 28(1 ):25 1-253 Rastegar-Pouyani. N. and G. Nilson. 1997. A New Species of Eremias (Sauria: Lacertidae) from Fars Province, South-Central Iran. Russian Journal of Her- petology4(2):94-101. Szczerbak, N. N. 1974. Yashchurki Palearktiki (Pale- arctic species of Eremias ). Kiev. 295 pp. 2001 Asiatic Herpetological Research Vol.9, pp. 113-121 Lizards of the Northern Mongolian Deserts: Densities and Community Structure KONSTANTIN A. ROGOVIN1, DMITRY V. SEMENOV1, AND GEORGY I. SHENBROT2 'A.N.Severtzov Institute of Ecology and Evolution, Russian Academy of Science, Leninsky pr. 33 Moscow 1 17071, Russia, ~ Ramon Science Center, Ben-Gurion University of the Negev, P.O. Box 194, Mizpe Ramon, 80600, Israel Abstract.- Spatial organization and population densities of three-species lizard community was studied in the Gobi Desert, Mongolia. To evaluate the effect of habitat variables on the distribution and abundance of each species we used the stepwise procedure of factor selection with ANOVA on each step. To describe the distribution of species' spatial niches in the space of environmental variables, we used stepwise discriminant function analysis (DFA). The number of species in 1-ha grid areas varied from 0 to 4. Phrynocephalus versicolor was the only species distributed over the 91% of grids occupied. There was a positive relationship between distribution and local species abundance. A set of two to three habitat variables determined the abundance of each species. The result of DFA signify to the well pronounced segregation, but not even distribution of species spatial niches in the space of resources. Key words.- Lizards, Mongolia, community, ecology, density 91* 96* 101* 106° Fig 1. Map of Mongolia and location of sites where data were collected. Eastern Gobi, 4-Barun-Churay Basin. 111* 116' Southern Gobi, 2-Western Gobi, 3- Introduction The reptile communities of Mongolian deserts are characterized by several specific features discussed elsewhere (Ananjeva and Semenov, 1986; Borkin and Semenov, 1984; Munkhbajar, 1976; Semenov and Borkin, 1986; Semenov and Shenbrot, 1988). There are few species in the fauna with low species richness at sites, a low level of species turnover between habi- tats, low abundance of most species and high domi- nance of only one species, Phrynocephalus versicolor. A few common species have rather broad spatial niches, diverse behavioral and physiological charac- teristics (diverse range of thermobiological patterns, wide active search for food items, etc.). Vol. 9, p. 114 Asiatic Herpetological Research 2001 The degree of interest in comparative studies of structure and function of reptile communities increased dramatically after seminal papers written by E. Pianka (Pianka, 1973, 1975). Most of the ensuing studies were devoted to the species rich and diverse communities of desert lizards in Australia, south- western North America and southern Africa (Case, 1983; Fuentes, 1976; Henle, 1989; Inger, Colwell, 1977; Millado et al., 1975; Pianka,1986; Scheibe, 1987; Shenbrot et al., 1992; Simbotwe, 1984). At the same time study of species poor communities in the Central Asian desert can provide a significant infor- mation not only in comparison with other continents, but also can help us to understand better which factors rule in reality structure and dynamic of lizard commu- nities of many species. Up to now there were only two examples of such studies made in China (Chang et al., 1993;Luietal.,1992). The main objective of this paper was to study the specific features of spatial organization and popula- tion densities of three-species lizard community in the Gobi Desert, Mongolia. The study was conducted dur- ing a long-term research program on the biodiversity of the Mongolian desert biota, and was sponsored by the Permanent Soviet-Mongolian Biological Expedi- tion. Material and Methods Mongolian desert A map (Fig. 1) illustrates the location of desert regions of Mongolia. Three desert regions to the south of the Altai Mountains are partly separated from one another by chains of low mountains and hills. These three are Trans-Altai Gobi (South), Alashan Gobi (East) and Sungarian Gobi or Barun-Churay Basin (West) (Yunatov, 1950). Besides these deserts there are desert areas between the Altai and Hangai moun- tains, usually called Western cold deserts, and some arid lands in the Great Lakes and Ubsu-Nur Basins (northwest). Three arid subzones of the Mongolian desert are defined (Sokolov and Gunin, 1986): extra — arid desert (<50 mm of rainfall per year), real desert (50— 100 mm per year) and steppe — like desert (100—150 mm per year). However, the climatic border that restricts the distribution of plants (Kazantseva, 1986; Volkova et al., 1986) and animals (Podtyazhkin and Orlov, 1986; Semenov and Borkin, 1986) exists only between the southern part of Trans- Altai Gobi (< 50 mm per year) and the northern waste belt of desert lands with more predictable precipitation (100-200 mm per year). The narrow real desert subzone appears transitive with no specific features of vegetation. Thus, only the southern (extra-arid) and northern sub- zones are well pronounced. The southern desert occurs mostly in the Trans- Altai Gobi and is characterized by a few very dry, unproductive biotopes inhabited by five lizard species (two agamids, two gekkonids, and one Eremias spe- cies). Among this group only one species, Phryno- cephalus versicolor is common in the northern subzone (Semenov and Borkin, 1986). The northern deserts, which extend to the south- west, west, and south-east of Mongolia and along both slopes of the Mongolian Altai and Gobi-Altai mountains, are characterized by pronounced microre- lief and rich vegetation, although the main landscape types are the same as in the southern subzone. The vegetation in rock and gravel valleys consists of perennial grass (Stipa), forbs, onions and succulents, and a variety of annual plant species. Shrub vegetation is often associated with foothills and sand dunes, or is spread along the dry river beds (Lavrenko, 1978). The difference in climate between western and eastern parts of the northern Mongolian desert is not pro- nounced (Murzaev, 1952); some differences exists in the composition of the flora (Yunatov, 1950). Lizard species There are four lizard species inhabiting northern Mongolian deserts: Alsophylax pipiens, Phrynoceph- alus versicolor, Eremias przewalskii and E. multiocel- lata. Among these, only the three last mentioned species are abundant and relatively widespread. Data collection We collected data during three field trips to the Mon- golian northern deserts in June-August 1985, 1986 and 1988. Forty five 1-ha grids were established in the Northern desert subzone (see map, Fig 1 ). Grids were distributed so that they covered the whole range of habitats from the middle slopes of the mountains to the clay basins and sand dunes. Each habitat type was sampled equally. Two factors determined the number of grids at a desert region: diversity of habitat types and abundance of lizards. Each grid was divided into 25 smaller sample plots, 20 x 20 m. The centers of sample plots were marked with 50 cm aluminum stakes. Lizards were sampled by repeated, regular search of established grids during two to four consec- utive days during periods of their maximal diurnal activity. Nearly all encountered lizards were captured by hand. Most of the surveys accounting were con- ducted before the appearance of hatchlings. In the rest of the cases hatchlings were not counted. Each cap- 2001 Asiatic Herpetological Research Vol. 9, p. 115 Table 1. Density (no. ha ) of lizard species. 0) 'o a> a (A TO N s. O w c "(5 c 3 O W c o "35 $ ■_ (0 a> c a. 0) ■a ro a> a (0 c w 1 c '5 re (0 (0 Q. c 1- (0 O) E 3 (A O T3 a o v> O C o •— s ■^ a o £ !E >» c o 4* i* .c 10 c o •o c >> (0 (A 0) C CO n O re (0 CO O _l CO /? versicolor E przewalskii E. multiocellata 1.7±0.9 52.8+8.9 54.4±8.7 36.6±8.2 5.5±4.5 41.2+6.3 0.8±0.4 0-3 24-91 32-83 10-106 1-10 22-64 0-3 0 0 0 2.6±1.3 0 16.3±6.3 0.9±0.5 0-18 0-44 0-4 0 2.1+1.3 0.4±0.4 0.5±0.4 0 2.0±1.8 0 0-12 0-2 0-5 0-11 tured lizard was marked by toe-clipping (the most dis- tal phalange only) and/or by a number painted with permanent color marker on its back, and released. Densities of lizard species were estimated by the total number of animals caught on a grid. The reliability of density estimates was verified by recapture of marked individuals and by the observation of marked lizards on grids. Two days were sufficient to catch > 80% of lizards. A 0.5 kg soil sample was taken from the center of each plot for laboratory texture analysis. The number of shrubs (by species) in each plot was counted in a circle, 5 m in radius. To determine vegetation cover and volume by height layers within each plot, the height and diameter of crown of the shrubs (up to 30 shrubs of each species in the grid) were measured. The abundance of annuals was evaluated by clipping all annuals on 0.25 m" sample plots (4 sample plots placed at random in each of 25 plots in 1-ha grid area). Twenty three parameters were used in the sub- sequent analysis (Table 2). Data on 1 125 descriptions of sample plots and 1710 records of 3 lizard species were used in the analysis. In total, we recorded 1528 individuals of the agamid lizard P. versicolor, 142 of the lacertid lizard Eremias przewalskii and 40 E. mul- tiocellata. Statistical data processing To evaluate the effect of habitat variables on the distri- bution and abundance of each species we used the stepwise procedure of factor selection (Shenbrot, 1988) with ANOVA of each step. Before analysis, the original values of species' densities and habitat vari- ables were transformed to an interval scale with five intervals for habitat variables and three intervals for species' abundance variables. The influence of each variable was estimated separately by ANOVA and the variables with nonsignificant effect were omitted. In the next step, each of variables that was not omitted earlier was added separately to the most powerful variable to select the most powerful pair of variables. This procedure was repeated until all possible vari- ables were included in the analysis or until the addi- tion of the new variable does not increase the proportion of explained dispersion. To describe the distribution of species' spatial niches in the space of environmental variables (eco- logical space) and to reduce the dimensionality of this space, we used stepwise discriminant analysis. Dis- criminant axes were calculated based on the data set consisting of the values of habitat variables for each point of lizard's registrations and with lizard species as a grouping variable. Vol. 9, p. 116 Asiatic Herpetological Research 2001 Results Densities and distribution through habitat types The results of lizard density and diversity estimations on the 1-ha grids indicate the low local species diver- sity in Mongolian deserts (Table 1). The number of species in our grid areas varied from zero to three. Among 45 grids there were two grids with no lizards, 21 grids with only one species. 18 grids with two spe- cies and four grids with three species. Phrynoceph- alus versicolor was the only species distributed over the most number of grids (91% of grids occupied). Eremias przewalskii was found on 33%, E. multioce- lata on 24% and A. pipiens on 4% of grids. Regarding distribution through the main habitat types (Table 1), P. versicolor was also the most widely distributed spe- cies but with the pronounced habitat preference. The second was E. multeocellata, and the third was E. przewalskii. The last species had the most restricted habitat preference, namely sandy-loess hills in saline depressions with shrub vegetation of Nitraria sp. The abundance of P. versicolor was positively cor- related with its broad distribution. The density of this species varied from 1 to 106 individuals per 1-ha (Table 1). The second most abundant E. przewalskii (44 individuals per lha maximum), and the third was E. multiocelata (11 individuals per 1-ha maximum). All three species coexisted at rather high densities in saline depressions with sandy-loess hills covered with Nitraria sp. shrubs. There were no pronounced nega- tive correlation between the densities of two dominant species on grids situated within this habitat type (P. versicolor-E. przewalskii: R"=0.04, n=20, ns). Table 2. Designation and description for the 23 habitat variables included in the analysis. Mnemonic Variable Unit RCK GRW CLY SCS SCH WDS WDD NRB FRB AGR ANN ALL PGR MIC HAL sue HLX NIT SHC SV1 SV2 SV3 SV4 Content of rocks in the soil Content of gravel in the soil Content of clay in the soil Sand cover area Sand cover height Dry river bed area Dry river bed depth Number of rodent burrows Abundance of annual forbs Abundance of annual grasses Overall abundance of annual grasses and forbs Perennial Allium covet Perennial grass cover Cover of microphyllous shrubs Cover of halophytuos shrubs Cover of small succulent shrubs Cover of Haloxylon Cover of Nitraria Overall shrub cover Perennial plant crown volume at the level 0-25 cm Perennial plant crown volume at the level 25-50 cm Perennial plant crown volume at the level 0.5-1 m Perennial plant crown volume at the level 1-2 m %% %% %% %% cm %% cm no/sq.m. no/sq.m. no/sq.m. no/sq.m. %% %% %% %% %% %% /o /o o/ o/ /o /o %% %% 0/ O/ /o /o o/o. Vo% 2001 Asiatic Herpetological Research Vol. 9, p. 117 1 0.5 0 -0.5 -1 AGR 1 0.5 0 -0.5 -1 NIT SHV2 SHV2 1 0.5 0 -0.5 -1 GRV 1 0.5 0 -0.5 -1 1 Fig 2. Habitat use profiles of P. versicolor. For each habitat variable rank, mean use by species is plotted as a difference between capture frequency on plots of this rank and capture frequency of all plots. The size of histograms therefore denote the selectivity (positive or negative) of habitat use. For mnemonics see Table 2 Spatial niches There was a statistically significant influence of habi- tat variables on the distribution and abundance of all three studied lizard species (Table 3). For each of these species we extracted a set of two to three habitat variables, determining 6.6-28.7% of observed vari- ance in abundance. Densities of two species (P. versi- color, E. przewalskii) were moderately affected by habitat variables, whereas density of E. multiocellata was weakly affected. There was a statistically significant influence of habitat variables on the distribution and abundance of all three studied lizard species (Table 3). For each of these species we extracted a set of two to three habitat variables, determining 6.6-28.7% of observed vari- ance in abundance. Densities of two species (P. versi- color, E. przewalskii) were moderately affected by habitat variables, whereas density of E. multiocellata was weakly affected. CLY 1 0.5 0 -0.5 -1 Fig. 3. Habitat use profiles of E przewalskii. For expla- nation see Fig. 2. FRB MIC Fig. 4. Habitat use profiles of E multiocellata. For explanation see Fig.2. Patterns of habitat usage based on selected vari- able sets for each species are presented on Figures 2-4. Phrynocephalus versicolor clearly avoided microsites with low gravel content, very low and very high annual grass abundance, moderate and high shrub crown volume in the level 0.25-0.5 m and pre- ferred microsites with moderate gravel content, mod- erate annual grass abundance and very low shrub Vol. 9, p. 118 Asiatic Herpetological Research 2001 Table 3. Summary of ANOVA analyses of influence of habitat variables on individual species' abundance. Values are proportions of total dispersion determined by given variable. Total proportion of variance determined by a set of variables may be greater than sum of influences of individual variables as a result of high-order interactions. Species V a r i a b 1 e s Total GRV CLY AGR FRB MIC NIT SHV2 P. versicolor 0.0226 0.0387 0.0458 0.1900 E. przewalskii 0.0593 0.1592 0.0102 0.2872 E multeocellata 0.0180 0.0232 0.0662 crown volume in the level 0.25-0.5 m (Fig. 2). Ere- Discussion mias przewalskii avoided microsites with high and very high clay content, very low Nitraria cower, very low shrub crown volume in the level 0.25-0.5 m, and preferred microsites with low clay content, moderate to high Nitraria cower, moderate to high shrub crown volume in the level 0.25-0.5 m (Fig. 3). Eremias mul- tiocellata avoided microsites with very high and very low forb abundance and microphyllous shrub cover, and preferred microsites with moderate forb abun- dance and rather high microphyllous shrub cover (Fig. 4). Structure of ecological space occupied The results of reducing habitat space dimensionality using discriminant analysis showed that division of this space by lizard species occurred along the first two axes. Both axes accounted in sum for 100 % of variance and reflected some complex environmental gradients (Table 4). The first axis represented a gen- eral gradient of decreasing rock and gravel content in the soil as well as an increase in the sand cover area, sand mound height, number of rodent burrows and general shrub (especially Nitraria) cover. This axis characterized the between-habitat component of envi- ronmental variation rather than within-habitat varia- tion, and described in general spatial segregation of lizard species according to their preference of physi- ognomically distinctive habitats. The second axis reflected an increase of dry river bed area, Allium abundance and microphyllous shrub cover, and char- acterized both between- and within-habitat compo- nents of species segregation. The first axis described habitat division between E. przewalskii and two other species, whereas the second axis reflected habitat seg- regation between E. multiocellata and two other spe- cies (Fig. 5). All three lizard species had spatially remote niche centers and less than 30% niche overlap (Fig.5). Our data confirm the general opinion of low reptile species richness in Mongolian deserts. There were only four 1-ha grids where three lizard species coex- isted. If we take into account rare encounters with snake species, the maximum reptile species number is five. Another opinion about wide spatial niches of Gobian lizards (Semenov and Borkin, 1986) partly contradicts our results. At the one hand P. versicolor is the an abundant species that can be found everywhere in the Gobi desert, including sandy, clay and rocky habitats. This feature of its distribution distinguishes this species greatly from its western congeners that share habitat types in Middle Asia (Shenbrot et al., 1991 ). At the other hand, P. versicolor did have well pronounced spatial preferences as it is seen from Table 1. The two species of Eremias lizards had dis- tinctly non-preferable habitats. Eremias przewalskii was mainly restricted to one habitat type, and E. mul- tiocellata to two habitat types. Results of analysis of microhabitat preferences demonstrate the existence of significantly distinct environmental variables determining spatial distribu- tion of each lizard species. The variables elucidated can be regarded as axes of species spatial niche. Spa- tial niches for P. versicolor and E. przewalskii appeared to be determined by three axes of environ- mental variation and for E. multiocellata by two axes. The distribution of species along each axes can be interpreted in accordance with the biological charac- teristics of each species. The absence of P. versicolor at microsites with dense vegetation is explained by typical Phrynocephalus locomotion on straightened legs and by the group-specific visual orientation when foraging (in contrast to Eremias, which look for food items using olfaction). Phrynocephalus versicolor preferred microsites with moderate gravel content, moderate annual grass abundance and very low shrub crown volume, which characterizes the species as an inhabitant of stony and gravel desert valleys (Fig. 2). Eremias przewalskii selected microhabitats in accor- 2001 Asiatic Herpetological Research Vol. 9, p. 119 Fig. 5. Seventy-five percent confidence ellipses for the species observations on two discriminant axes (DF1 and DF2). Em- Eremias multiocellata, Ep- Eremias przewalskii, Pv - Phrynocephalus vesicolor. fa Q DF1 dance with food abundance and a species-specific tac- tic of antipredator behavior. It preferred microsites with moderate to high Nitraria cover and moderate to high shrub crown volume at 0.25-0.5 m (Fig. 3). In summer E. przewalskii feed predominantly on berries and young green twigs of Nitraria, and also find pro- tection under the dense cover of the crowns of low spiny shrubs. In contrast to E. przewalskii, E. multio- cellata avoided microsites with very high annual plant abundance and shrub cover, but also preferred micro- sites with moderate and rather high values of these variables (Fig. 4). This difference can be interpreted in accordance with thermobiological and size charac- teristics of two Eremias species. The larger species, E. przewalskii is not so quick as the smaller E. multiocel- lata. The first one digs well in soft soil, and ther- moregulates climbing on or escaping under the shrub periodically, being active throughout the day. Small E. multiocellata that occupy habitats with relatively low vegetation cover (with small sparsely-distributed shrubs) must cross open sites in search for food items and escape predator's attacks by quickly rushing into small shrubs or burrows. This species reduces heat by escaping into burrow during the day-time. Another result of the above comparison is the pos- itive relationship between distribution and local abun- dance among the species considered. Distribution here means not the size of the species range area, but the number of sites where each species was found. Phrynocephalus versicolor was encountered on 41 grid areas and had the highest population density (mean: 37.3, median: 26 ind/ha, maximum: 106 ind/ ha). With the edge-effect correction (Semenov, 1991) maximum density was 70.02 ind/ha (175.5 g/ha biom- ass). Eremias przewalskii was found on 1 5 grid areas and was the second abundant species (mean: 9.5, median: 4 ind/ha, maximum: 44 ind/ha). Eremias mul- tiocellata was found on 1 1 grid areas and its maxi- mum density was 12 ind/ha (mean: 3.6, median: 2 ind/ ha). Alsophilax pipiens was met on two grids with density 3 ind/ha. Brown (1995) explains this rather common rela- tionship by the "Hutchinsonian niche model" (see also for one species, Brown 1984), suggesting that "the species that is slightly more tolerant of some abi- otic conditions or biotic interaction or is slightly bet- ter able to use some resource should not only be able to occur in more places but also to attain higher abun- dance in some of those places." In the approach we use here niche breadth reflects microhabitat require- ments of each species, namely the range of microcon- ditions where each lizard species occurred. Diversity of these microhabitats is not necessarily correlated with the diversity of macrohabitats as well as with the Vol. 9, p. 120 Asiatic Herpetological Research 2001 Table 4. Summary of discriminant analysis of the habi- tats of lizard species. DF1 and DF2 are the first two components (all are significant, P<0.001). Mnemonics for habitat variables are from Table 1 . DF1 DF2 Eigenvalue 0.351 0.102 Chi-square 676.1 165.5 Cumulative % of variance 62.58 37.42 Factor loading RCK -0.350 0.295 GRV -0.291 0.124 SCS 0.544 -0.221 WDD -0.091 0.241 NRB 0.442 0.253 SHH 0.627 0.190 ALL -0.141 0.252 MIC -0.021 0.410 NIT 0.642 0.193 SHC 0.650 0.148 SV1 0.712 0.067 SV2 0.693 -0.069 size of the landmass studied. Phrynocephalus versi- color that was distributed everywhere in the Gobi, and in many different macrohabitats used a rather limited range of microconditions (Fig. 2). In this respect its spatial niche was not broader than the niche of E. mul- tiocellata, and was even narrower in comparison with E. przewalskii. (Fig. 5). Eremias przewalskii which inhabits a rather limited range of macrohabitats is characterized by relatively broad requirements for microconditions along the first discriminant axis which represents the main direction of spatial segre- gation of lizard species. The structure of ecological space occupied by three lizard species has a complex character, explain- ing both macrohabitat and microhabitat segregation of spatial niches. Primarily, each discriminant axis char- acterizes the range of variation of microconditions in the study area. This can characterize macrohabitats only if variation in a set of variables included into analysis reflects the macrohabitat variation. In our case DF1 and DF2 possess such a feature: DF1 char- acterizes better the between-habitat component of spatial niche segregation and DF2 characterizes more the within-habitat component. Three species of lizards share microconditions in two-dimensional ecological space, so that the niche centers appeared to be almost equally distant from the geometrical center of the model (Fig. 5). The two species of Eremias lizards share ecological space to a greater extent than each Eremias with P. versicolor. Niche overlap between species was less than 15%. It seems difficult to speculate about processes that led to such spatial relationships. Interspecific compe- tition could contribute to spatial segregation of Ere- mias lizards in past, and could determine the low level of niche overlap. The competitive relationship between E. przewalskii and P. versicolor is unclear. In a pair of neighboring grid areas at one location within one habitat type when one species is in high density, the other one is usually at low density and vise-versa. However, there were no correlation between popula- tion densities of these two dominant species when all grid areas within one habitat type were put together. This result is easily explainable because different localities must have different upper limits of species densities according to the local environmental capaci- ties. However, the climatic conditions of Gobi Desert are so unfavored for reptiles (extremely low winter temperatures, great interannual, between- and within- seasonal contrasts in precipitation and temperatures) that it appears unrealistic to assume the space of resources is saturated by individuals, and that popula- tions are at equilibrium. Acknowledgments We are very grateful to Dr. David Ward (Ben-Gurion University of the Negev) for helping with the manu- script and to Dr. Natalia B. Ananjeva (Zoological Institute of St. Petersburg) for reading the manuscript. This is publication no. 12510-5241-9 of IEE and no. 113ofRSC. Literature Cited Ananjeva, N. B., and D. V. Semenov. 1986. [Some aspects of ecology of Mongolian desert lizards]. Nature and biological resources of Mongolian Peo- ple's Republic. Moscow. [Abstr.] (In Russian). Borkin, L. J., and D. V.Semenov. 1984. [Distribution of Phrynocephalus versicolor in South Mongolia]. VIII All-Union Zoogeographical Conference. Mos- cow. [Abstr.] (In Russian). Brown J. H. 1984. On the relationship between abun- dance and distribution of species. American Naturalist 124:255-279. 2001 Asiatic Herpetological Research Vol. 9, p. 121 Brown J. H. 1995. Macroecology. The University of Chicago Press. Chicago and London. 269 pp. Case, T. J. 1983. Niche overlap and the assembly of island lizard communities. Oikos 41(3):427-433. Chang Cheng, Quang Chen, and Nai-Fa Liu. 1993. [The coexisting mechanism of two species of lizards in the same community]. Journal of Lanzhou Univer- sity, Natural Scienes 29(3): 180- 183. (In Chinese). Fuentes, E. R. 1976. Ecological convergence of lizard communitiesin Chile and California. Ecology 57:1- 17. Henle, K. 1989. Ecological segregation in an assem- blage of diurnal lizards in arid Australia. Acta Oeco- logica 10(1): 19-35. Inger, R. F, and R. K. Colwell. 1977. Organization of contiguous communities of amphibians and reptiles in Thailand. Ecological Monographs 47:229-253. Kazantseva, T I. 1986. [Distribution and dynamics of the overground phytomass]. Pp. 106-114 In V.E. Sokolov and PD. Gunin (eds.). [Deserts of Trans- Altai Gobi]. Nauka, Moskva. (In Russian) Lavrenko, E. M. 1978. [On vegetation of steppes and deserts of Mongolian People's Republic]. Problemy osvoeniya pustyn. Ashkhabad 1:3-18. (In Russian). Liu Nai-Fa, Ren-De Li, and Xiao-Cheng Liang. 1992. [Community structure of desert lizard in Gansu Province]. Acta Zoologica Sinica 38(4):377-384. (In China). Millado, J., F Amores, F F. Parreno, and F Hiraldo. 1975. The structure of a Mediterranean lizard com- munity. Donana Acta Vertebrata 2 (2): 145- 160. Munkhbayar, Kh. 1976. [Amphibians and reptiles of Mongolian People's Republic]. Ulaanbaatar. 167 pp (In Mongolian). Murzaev, E. M. 1952. [Mongolian People Republic]. 472pp. Geografgiz. Moskva. (In Russian) Pianka, E. R. 1973. The structure of lizard communi- ties. Annual Review of Ecology and Systematics 4:53-74. Pianka, E. R. 1975. Niche relations of desert lizards. Pp. 292-314. In M.L. Cody and J.M. Diamond (eds). Ecology and Evolution of Communities. Harvard University Press. Pianka, E.R. 1986. Ecology and natural history of desert lizards. Princeton University Press. Princeton, New Jersey. 208 pp. Podtyazhkin, O. I. and V. N. Orlov. 1986. [Faunoge- netic grouping of rodents in Mongolian arid zone]. Pp. 124-136. In V.E. Sokolov (ed.). [Zoogeographical regionalization of Mongolian People's Republic]. Institute of Animal Evolutionary Morphology and Ecology. Moscow. (In Russian). Scheibe, J. S. 1987. Climate, competition, and the structure of temperate zone lizard communities. Ecol- ogy 68(5): 1424-1436. Semenov, D. V. 1991. [Variations of characteristics of animal spatial distribution in Phrynocephalus ver- sicolor populations (Reptilia; Agamidae]. Ekologiya, Sverdlovsk 1991(3):77-81 (In Russian). Semenov, D. V, and L. J. Borkin. 1986. [Amphibians and reptiles]. Pp. 1 14-119. In V E. Sokolov and P. D. Gunin (eds.). [The deserts of the Transaltai Gobi. The natural features, ecosystems, and regionalization]. Nauka, Moscow. (In Russian). Semenov, D. V, and Shenbrot G. I. 1988. [Lizard den- sities in some Mongolian regions]. Natural features and resources of some regions of Mongolian People's Republic. [Abstr.] (In Russian). Shenbrot, G. I. 1988. [Evaluation of factors that affect lizard distribution and abundance in deserts of the South of Bukhara Oblast (Uzbek SSR, USSR)]. Ekologiya, Sverdlovsk 1988 (5):51-56. (In Russian). Shenbrot, G.I. 1991. On the environmental factors affecting desert lizard distribution and abundance. 6th Ordinary General Meeting of Societas Herpetologica Europaea. Budapest. [Abst.]. Shenbrot, G. I., K. A. Rogovin, and A. V Surov. 1992. Comparative analysis of the spatial organization of the desert lizard communities in Middle Asia and in Mexico. Pp. 147-188. In V Sokolov, G. Halffter and A. Ortega (eds). Vertebrate ecology in arid zones of Mexico and Asia. Inst, de Ecologia A.C., Centro de Investigaciones Biologicas de Baja California Sur A.C., and MAB-UNESCO. Simbotwe, M. P. 1984. Resource partitioning in a woodland reptile community of the Kafue flats, Zam- bia. African Journal of Ecology 22:281-287. Sokolov, V E. and P. D. Gunin (eds.). 1986. [Deserts of Trans- Altai Gobi]. Nauka. Moskva. 207 pp. (In Russian). Volkova, G.A., E. I. Rachkovskaya, and I. T. Feodor- ova. 1986. [General features of plant distribution]. Pp. 84-95. In V. E. Sokolov and P. D. Gunin (eds.). [Deserts of Trans-Altai Gobi]. Nauka. Moskva. (In Russian). Yunatov, A. A. 1950. [The main features of vegetation of Mongolian People's Republic]. Proceedings of Mongolian Comission 39. AS USSR Publishers. Mos- cow, Leningrad. 223 pp. (In Russian). 2001 Asiatic Herpetological Research Vol. 9, pp. 122-129 The Morphology and Size of Blood Cells of Lacerta rudis bithynica Murat Sevinc1 and Ismail H. UGurta§ Uludagg University, Science and Art Faculty, Department of Biology, 16059 Bursa, Turkey. To whom correspondence should be addressed e-mail: smurat® uludag.edu.tr Abstract.- In this study, the morphology of the blood cells of Lacerta rudis bithynica is described using Wright's technique. The sizes of erythrocytes and their nuclei, leukocytes (monocytes, lymphocytes, basophils, neutrophils, and eosinophils), and thrombocytes of L. rudis bithynica were measured using an ocular micrometer at a magnification of 1600X with an oil immersion objective. The results of this study are compared with previous work on other reptile species. Key words.- morphology. Lacertidae, Lacerta rudis bithynica, erythrocyte, leukocyte, thrombocyte, measurement. Introduction The first studies on the blood of reptiles described the cellular structures, often comparing them with those of other vertebrates. Literature on the haematology of reptilian blood is based on few studies and is usually concerned with European species (Saint Girons, 1970). Recent studies have concentrated on single species (Tiliqua sp., Cannon et al., 1988; Cyrtopodion sca- brum, Canfield and Shea, 1996). Various authors have described different circulating blood cells of different reptile species (Taylor and Kaplan, 1961; Heady and Rogers ,1962; Hartman and Lessler, 1964; Szarski and Czopek, 1966; Duguy, 1970; Saint Girons, 1970, Cannon et al., 1988, Canfield and Shea, 1996). Other authors have studied seasonal (Hutton, 1960; Cline and Waldman, 1962; Haggag and et. al, 1966) or sex- ual (Altland and Thompson, 1962) variation in the number of blood cells of different reptile species. In addition, authors have studied the number of blood cells of different reptile species (Charipper and Davis, 1932; Baker and Cline, 1932; Hutton, 1961; Altland and Thompson, 1962; Hutchinson and Szarski, 1965; Engbretson and Hutchinson, 1976). Finally, authors have studied haemoglobin and hematocrit content of blood and hematopoiesis of different reptile species (Altland and Thompson, 1958; Hutton, 1961; Goin and Crawford, 1965; Engbretson and Hutchinson, 1976; Newlin and Ballinger, 1976). In Turkey, haematological studies have been con- ducted on humans and economically important ani- mals. However, there are no haematological studies on the Turkish reptiles. In this study, our aim is to describe and measure blood cells (erythrocyte, leukocyte, thrombocyte) of Lacerta rudis bithynica (Squamata: Lacertidae). This study is the first of its kind on a Turkish species. Material and Methods In this study, 31 individuals (17 male) of Lacerta rudis bithynica (Squamata: Lacertidae) were exam- ined. The study was carried out between June and August 1998. The specimens were collected from Uludagg (Bursa) at an altitude of 1745 m. Blood was obtained by cardiac puncture of the lizards (Canfield and Shea, 1988). Immediately after the blood was obtained in heparinized capillary tubes, the blood smears were prepared. Three to five blood smears were prepared per individual. The smears were air- dried and stored until stained with Wright's stain (Hartman and Lessler, 1964). Twelve drops of Wright's stain were dropped on the slides and allowed to remain on the slide one and a half minutes before rinsing with a phosphate buffer (pH=6.5). The slides were allowed to stand for ten minutes at room temper- ature and were then washed with distilled water and allowed to dry. On each slide fifty mature erythrocytes and their nuclei, ten thrombocytes, and ten leukocytes (mono- cyte, lymphocyte, eosinophil, basophil and neutro- phil) were measured by means of an ocular micrometer at a magnification of 1600 x with an oil immersion objective. Cell sizes were calculated from the measurements. 2001 Asiatic Herpetological Research Vol. 9, 123 10/"" Figure 1 . Erythrocytes, an infected erythrocyte and a mitosis dvision. Results I. Erythrocytes Erythrocytes are nucleated, oval cells. Their nuclei are also oval, more or less regular and centrally located (Fig. 1). The cytoplasm of mature erythrocyte appeared light and dark pink and homogeneous under Wright's stain. The nuclei of mature erythrocytes are chromophilic. In some blood smears, immature eryth- rocytes are seen. They are characterised by a rounded form, blue cytoplasm and a large nucleus. Mitotic fig- ures are also present and in some smears, intracorpus- cular parasites are seen (Fig. 1). Parasites alter the shape and size of erythrocytes remarkably. When intracorpuscular parasites are seen, immature erythro- cytes and mitotic figures are abundant (Fig 1 ). Intrac- orpuscular parasites alter the shape and size of infected erythrocytes. The shape and size of other erythrocytes that are not infected by intracorpuscular parasites are normal. 20 15 •s io s 0 14 41 15 05 1 5 71 [6.17 [630 M6M I Erythrocyte I Nucleus June July August Figure 2. Erythrocyte and nucleus lengths of Lacerta rudis bithynica qnq\ three months. to 8 6 4 2 0 7.63 8.08 S 24 ■ BR ™ [3.5s [3 55 lj_74 Erythrocyte Nucleus June July August Figure 3. Erythrocyte and nucleus widths of Lacerta rudis bithynica over three months. In June mean length of mature erythrocytes was 14.41 urn (±0.77 standard deviations, with a range of 12.20-16.47 urn). In July, the mean length of mature erythrocytes was 15.05 urn (±0.79, 12.81-17.08 pm). In August, the mean length of mature erythrocytes was 15.71 urn (±0.79, 12.81-18.30 um). Other mea- surements are given in Tables 1, 2 and 3. There are no significant differences in erythrocyte and nucleus sizes between females and males. Based on Tables 1 , 2, and 3 and Figs. 2 and 3, it appears that there were little monthly variations in erythrocyte and nucleus sizes among June, July and August. II. Leukocytes 1. Eosinophils. In blood smears stained by Wright technique, eosinophils are circular, and the cytoplasm Table 1. Erythrocyte dimensions of Lacerta rudis bithynica with the standard deviations in June. EL: Erythrocyte length; EW: Erythrocyte width; ES: Erythrocyte size;NL: Nucleus length; NW: Nucleus width; NS: Nucleus size. EL(um) EW (um) EL/EW ES (Mm2) NS/ES Maximum 16.47 ±0.77 9.15 ±0.48 2.27 ±0.13 110.41 ±8.00 0.30 ± 0.02 Minimum 12.20 ±0.77 6.71 ± 0.48 1.42 ±0.13 64.26 ± 8.00 0.13 ±0.02 Mean 14.41 ±0.77 7.63 ± 0.48 1.89 ±0.13 86.46 ± 8.00 0.20 ± 0.02 NL(um) NW (um) NL/NW NS (Mm2) Maximum 7.32 ± 0.40 4.88 ± 0.34 2.20 ±0.19 24.53 ±2.19 Minimum 4.88 ± 0.40 3.05 ± 0.34 1.25 ±0.19 11.68 ±2.19 Mean 6.17 ±0.40 3.55 ± 0.34 1.75 ±0.19 17.25 ±2.19 Vol. 9, p. 124 Asiatic Herpetological Research 2001 Table 2. Erythrocyte dimensions of Lacerta rudis bithynica\oqe\!r\er with the standard deviations in July. EL: Eryth- rocyte length; EW: Erythrocyte width; ES: Erythrocyte size; NL: Nucleus length; NW: Nucleus width; NS: Nucleus size. EL (Mm) EW (pm) EL/EW ES (nm2) NS/ES Maximum 17.08 ±0.79 9.15 ±0.49 2.16 ±0.12 122.68 ±8.95 0.28 ± 0.02 Minimum 12.81 ±0.79 6.71 ± 0.49 1.53 ±0.12 67.47 ± 8.95 0.13 ±0.02 Mean 15.05 ±0.79 8.08 ± 0.49 1.86 ±0.12 95.62 ± 8.95 0.18 ±0.02 NL(um) NW (Mm) NL/NW NS (Mm2) Maximum Minimum Mean 7.30 ± 0.70 4.27 ± 0.32 5.40 ± 0.70 3.05 ± 0.32 6.30 ± 0.70 3.55 ± 0.32 2.40 ±0.17 1.33 ±0.17 1.79 ±0.17 24.53 ±2.38 13.14 ±2.38 17.67 ±2.38 is stained light red. Eosinophils contain circular to elongate cytoplasmic granules stained brilliant red (Fig 4). Eosinophils are different from neutrophils in that, eosinophils' granules are stained bright red and neutrophils' granules were stained dim red. In June, the mean diameter of eosinophils was 12.82 pm, (±1.71 standard deviations, with a range of 9.93-15.25 um). In July, the mean diameter was 13.29 um (±1.25, 10.98-15.25 um). In August, the mean diameter was 13.80 pm (±1.37, 12.20-15.25 um). There were no significant differences in eosinophil diameters between females and males. Based on Tables 4, 5, and 6 and Fig. 8, it appears that there was little monthly variation in the diameter of eosinophils during the three months. 2. Basophils. Basophils are easily recognised. They are small and circular cells. Nuclei stained blue by Wright technique are commonly obscured by chro- mophilic circular granules. These cytoplasmic gran- ules are large and stained dark purple. In the blood

— £\ »*10 / 1 ^ ~ f»12 \ V f ^*13E X H>l7-s O 022 1 fvij ' °24 0 27 (5 S>21^ \ °s/ & 9^~£l 0 26 \M8 ' / ■IS/' » 023 20 | ^s- I » — * MEDITERRANEAN O 0 100 200 km Figure 1 .The places where Pelobates syr/acushave been collected in Turkey. The key to the letters is found in Table 2. Dark circles = The localities where we collected specimens. Open circles = The localities where speci- mens were collected before. 1. Buyukdolluk, 2. Babaeski, 3. Altin§ehir, 4. Terkos, 5.Yassioren, 6. Adapazari, 7. Sogutlu, 8. Karasu, 9. Poyrazli Lake, 10. £erkesji, H.Terme, 12. Karacabey, 13. Bursa, 14. §akran, 15. Bostanli, 16 Bornova, 17. Ivrindi, 18. Seydi§ehir,19. Karata§, 20. Iskenderun, 21. Van. Pelobates syriacus Boettger 1889 was first described from specimens collected from Hayfa-Israel. Later, Mertens (1923) examined two specimens, found in Belesuwar near the Azerbaijan and Iran border. Because of their long and narrow skull and the round and raised frontoparietals, he recognised them as another subspecies, Pelobates syriacus boettgeri. Other specimens from Macedonia were described by Karaman (1928) as the subspecies Pelobates syria- cus balcanicus. Miiller (1932) criticized this classifi- cation based on the shape of the skull. He emphasised that skull shape can be variable. Another subspecies living around Tiflis, Georgia was named Pelobates syriacus transcaucasicus by Delwig (1927). Accord- ing to Gilsen (1937), this subspecies is intermediate in morphology between P. s. boettgeri and P. s. syriacus, and Terentjev and Chernov (1965) are convinced that P. s. transcaucasicus is a synonym of the nominate race. Furthermore, Eiselt and Schmidtler (1973) pro- posed P. s. boettgeri as a synonym for P. s. syriacus. The taxonomic status of Turkish Pelobates syria- cus is poorly studied. Mertens (1953) described a young sample collected in Van (Turkey) and sug- gested that it was P. s. boettgeri. Zaloglu (1964) stud- Vol. 9, p. 140 Asiatic Herpetological Research 2001 Table 1. Percentage of each color pattern in populations of Pelobates syriacus. Population B D Edirne Istanbul Adapazari Samsun Bursa Balikesir-izmir Seydi§ehir Adana-lskenderun Van - - 100 - 56.66 40.00 3.34 - 11.54 11.54 11.54 65.38 68.42 21.05 - 10.53 73.92 21.73 - 4.35 15.00 75.00 5.00 5.00 7.70 - - 92.30 66.00 - - 34.00 14.38 14.28 . 71.44 ied Pelobates syriacus from the Turkish region of Izmir. He made an osteological comparison of chra- cters used to differntiate the subspecies of P. syriacus. He pointed out that these characters showed a wide range of variation, and refrained from classifying the Izmir P. syriacus into subspecies. In this study, samples (77 male, 57 female, 25 juveniles, 35 larvae) collected from 9 different areas (Istanbul, Adapazari, Samsun, Bursa, Izmir-Balikesir, Seydi§ehir, Adana, Van) were investigated to obtain more information about variation in Pelobates syria- cus living in Turkey (Fig. 1 ). Results and Discussion There are four kinds of dorsal patterns on Pelobates syriacus (Fig. 2). The dorsal patterns are not sexually dimorphic. The percentage of dorsal patterns are fol- lows: A: 36.08 %, B: 25.95 %, C: 13.92 %, D: 24.05 %. When all the samples are compared, it is seen that the dorsal pattern of Edirne population is different than the other populations (Table 1 ). All of the adults from Edirne have big green dots narrow raised surrounding on the back. The back- ground color of the back is light yellow-green. On this background there are connected raised surrounding with brown-green dots. Those dots also appear on all extremities. Furthermore, on the background color, and within the brown-green dots, are yellow and red dots. These dots are missing on the upper side of the front fingers. Dots on the body become smaller on the side. All the Edirne population have C pattern type (Fig. 2). In terms of morphological measurements, Pelo- bates syriacus populations living in Turkey are very similar. Here, the exception is the Seydi§ehir popula- tion which have shorter indices of tibia length (Table 2). The differences in the color patterns of adult ani- mals from the Edirne population and the morphomet- Figure 2. Types of color patterns in Pelobates syria- cus. A: Spots are irregularly distributed and isolated. B: Two or more spots combine to form irregular islets. C: The edges of the spots are wavy and connected by thin bands. D: Spots form lengthwise bands. 2001 Asiatic Herpetological Research Vol. 9. p. 141 ric differences of the Seydi§ehir population should be explored using biochemical data in order to determine whether the Pelobates syriacus from these regions represent different taxa. Literature Cited Boettger, O. 1889. Ein neuer Pelobates aus Syrien. ZoologischerAnzeiger 12:144, 1889. Delwig, W. 1927. Eine neue Art der Gattung Pelo- bates Wagler aus dem zentralen Transcaucasus. Zool- ogischer Anzeiger 75, Heft 1/2:24. Eiselt, J. and J. F. Schmidtler 1973. Froschlurche aus dem Iran unter Beriick chtigung ausser-iranischer Populationgruppen. Annalen Naturhistorischen Muse- ums in Wien 77:181-243. Gislen, T. 1937. On the history of evolution and distri- bution of the European pelobatids. Zoogeographica 3:119. Karaman, S. 1928. Contribution a 1' Herpetologie de Jugoslavja. Bulletin de la Societe Scientifique de Sko- plje 4. 2:129. Mertens, R. 1923. Beitrage zur Kenntnis der Gattung Pelobates Wagler Senckenbergiana 5(3/4): 1 1 8- 1 28. Mertens, R. 1953. Weiters zur Kenntnis der Herpeto- fauna der Asiatischen Tiirkei. Istanbul Univeritesi Fakultesi Mecmausi Seri B18: 373-375. Muller, L. 1932. Beitrage zur Herpetologie der Sudosteuropaischen Halbinsel. Teil I, Zoologischer Anzeiger 100, Heft 11/12:301-309. Teretnjev, P. and S. A. Chernov. 1965. Key to Amphibians and Reptiles. Israel Program for Scien- tific Translations, Jerusalem. Zaloglu, §. 1964. Ege bolgesinde bulunan Pelobates syriacus'un morfoloji, osteoloji ve biyolojisi ile ilgili ara§tirmalar. Ege Univeritesi. Fen Fakultesi ilmi Rap. Ser. No. 16:1-50. 2001 Asiatic Herpetological Research Vol. 9, pp. 142-144 Translation: A New Species of the Turtle Genus Ctav#(Testudoformes: Testudinidae) Ming-tao Song Shaanxi Institute of Zoology, Xi 'an Province, China (Current Address: Northern west Institute of Endangered Animals, Xi'an, China 710032) Original English Abstract.- Cuora pani, sp. nov. (Figs. 1, 2). Holotype, SIZ 80170, an adult male. Allotype, SIZ 80171, an adult female. All of these type specimens were collected from Xujiaba (alt. 420 m) of Pingli County in Shaanxi Province, on June 17, 1981 by the author, and are preserved in the Shaanxi Institute of Zoology. This new species is similar to Cuora yurmanensis (Boulenger), but differs from the latter in having the median keel not conspicuous and no lateral keels; a brown narrow band extending from behind eye to the neck; the suture between gulars 1.5 times as long as that between humerals and much shorter than those between pectorals and between anals, but longer than that between femorals; the plastron yellow, with black sutures; and the limbs brown, without any markings. Song, M. T. 1984. A New Species Of The Turtle Genus Cuora (Testudoformes: Testudinidae). Acta Zootaxonom- ica Sinica 9(3):330-332. (In Chinese with English abstract) There are six species belonging to the genus Cuora Gray, 1855 (Pritchard, 1967; Sichuan Institute of Biology, 1977), that are distributed in Thailand, Cam- bodia, Malaysia, Indonesia, the Philippines, and southern provinces of China. In 1981, two Cuora specimens were found during a survey of Mt. Dabashan, Shaanxi Province. These specimens are different from the six known species and are recognized as a new form. Its description is given below. Cuora pani, sp. nov. (figs. 1-2) The holotype, SIZ 80170, is an adult male. The allo- type, SIZ 80171, is an adult female. Both of the type specimens were collected from Xujiaba (alt. 420 m) of Pingli County in Shaanxi Province, on June 17, 1981 by the author, and are preserved in the Shaanxi Institute of Zoology. Diagnosis The new species is similar to Cuora yurmanensis (Boulenger), but differs from that species by having a median keel that is not conspicuous and no lateral keels; a brown narrow band extending from behind the eye to the neck; the seam between the gulars 1.5 times as long as the seam between the humerals and much shorter than those between pectorals and between anals, but longer than that between femorals; the plastron yellow, with black markings along the seams; the limbs are brown and without any mark- ings. Description of Holotype The carapace is flattened, with an inconspicuous median keel. The cervical scute is small. The anterior margin of the first vertebral is flaring, the anterior margin wider than the posterior margin. The second vertebral is square-shaped, with both sides are slightly protruding. The third and the fourth vertebrals are as wide as long, with the fourth vertebral being wider than long. The first pleural is the longest, the second is wider and as long as the third, and the fourth the smallest. There are twelve marginals. The first mar- ginal is the widest, the third to seventh and the ninth to tenth are slightly flared. The plastron is rounded anteriorly and notched posteriorly. The plastron can completely close the shell and is united to the carapace by ligamentous tis- sue. The ligamentous tissue is also between the pecto- rals and abdominals. The length of gular seam is 1.5 times the humeral seam, and much shorter than pecto- ral, abdominal, or anal seam. The pectoral seam is as long as abdominal seam, the anal seam a little shorter, and the abdominal seam is nearly twice the length of the gular seam; femoral seam much shorter, only a lit- tle longer than humeral seam. There are a pair of anals that are notched posteriorly. The head is moderate in size, smooth on top, and rather rough in the occipital region. The snout is pointed, projecting over the upper jaw. The diameter of the orbit is about the same of the length of snout. Upper jaw is slightly curved and a little longer than the lower jaw. 2001 Asiatic Herpetological Research Vol. 9. p. 143 | = 1 cm I cm Figurel. The holotype (SIZ 80170), an adult male, in dorsal view. Figure 2. The allotype (SIZ 801 70), an adult female, in ventral view. The forelimb is covered by imbricate scales poste- riorly and with a transverse series of large scales on ventrally. The hindlimb is covered by scales medially and on the tarsus. There are five claws on the forelimb and four on the hindlimb. The webbing between the digits is well developed. The tail is short, conical in shape. The tail is covered by granular scales on its back, and covered with paired scales forming a longi- tudinal groove ventrally. Color in life The carapace is light brown. The plastron and ventral side of marginals are yellow, with broad black bars along the seams. The top of the head is olive, dark gray laterally, with two brown stripes behind the orbit and along tympanum to neck. The tympanum is light gray. The lower jaw and chin are grayish-yellow. The back of the neck is brown, but lighter ventrally. The shoulders are light yellow and the axilla are yellow. The limbs and tail are light brown above and gray below. The pelvic region and the area behind the fern- Vol. 9. p. 144 Asiatic Herpetological Research 2001 ora are light yellow. The claws are brown with their tips yellow. Allotype The seams of the plastron are not as black as in the holotype, the back of tail is covered with a few large scales. Acknowledgments This translation was provided by Ermi Zhao with per- mission of Acta Zootaxonomica Sinica. The figures were provided by Ming-Tao Song. Table 1 . Measurements of types (in mm). Literature Cited Department of Herpetology. Sichuan Institute of Biol- ogy (Zhao. E.. Y.-m. Jiang, and Y. Shen). 1977. [Sys- tematic Key to Chinese Reptilia]. Science Press, Beijing. 1 10 pp. (in Chinese). Pope. C. H. 1935. The reptiles of China. Natural His- tory of Central Asia 10:28-35. Smith. M.A. The Fauna of British India. Including Ceylon and Burma. Reptilia and Amphibia. Vol. I.- Loricata, Testudines. Taylor and Francis. London. 185 pp. .c .c O) D) c „_ C £L .c sz a> 5 2 *3 CD c cn "D en .c CD CD O) O) CD CD E o ■o CD .c T3 c CD O CO a. CJ CO a. CD II he apac CD CO CO CO CD CD ^ Q. CD CD CD CD £ CO CO .C -C CO in (f) I I I O O CO CO CJ Holotype Male 32 18 16 18 120 86 40 Allotype Female 33 18 18 27 115 81 37 33.3 32.2 2000 Asiatic Herpetological Research Vol. 9, pp. 145-148 Translation: Testudo graeca anamurensis ssp. nov. from Asia Minor Heinz Weissinger Richard-Gebhardtgasse 4. 3423 St. Andra-Wordern, Austria Weissinger. H. 1987. Testudo graeca anamurensis ssp. nov. aus Kleinasien. OGH-Naehriehten (Wien) 10/1 14-18 The South-western coast of Turkey, from the Bey mountains near Antalya eastwards up to Mersin is the area of distribution of a Testudo graeca which is char- acterized hy an elongated, trapezium-shaped and strongly flattened carapace. This southcoast-popula- tion was so conspicuous among the more than 200 specimens which I examined and measured in Turkey, that I consider them a geographic subspecies and describe them as follows: Testudo graeca anamurensis nov. ssp. Material Type: One female. Anamurum; NMW 30795:1 Paratypes: One male, Anamurum; NMW 30795:2df One female. Side; NMW 30795:3 Both alive in the author's collection Terra typica Beach of Anamurum. 7 km. west of Anamur. SW coast of Turkey. Diagnosis Differs from Testudo graeca ibera Pallas 1814 by its considerably Hatter and narrower (elongated trape- zium-shaped) carapace. The length of the carapace is two-and-a-half times its height and one-and-a-half times its width. In most cases the carapace pattern is a completely broken, obvious blotchy pattern. The plas- tron has a sharply demarcated pattern of brown to black blotches, which can cover the plastral area for 20 - 100 7c. In 80 CA of cases, 100% at the terra typica, the animals have a white-yellow chin and throat. The head can be uniformly dark to irregularly spotted with yellow. A triangle on the snout - as in Testudo graeca terrestris Forskal 1775 - may be present only in early juvenile stages. In the subspecies most abundant in Asia Minor. Testudo graeca ibera Pallas 1814. the carapace is oval, domed regularly and has a smooth posterior margin. Its ground colour is bright olive with large yet individual blotches. The head and extremi- ties are most often uniformly dark. The plastron usu- ally shows indistinctly bordered blotches. Figures 1-3. (Top and bottom left) Testudo graeca anamurensis ssp. nov.; paratype. female NMW 30795:3. Figure 4. (Bottom right) Testudo gracea. Ssp. anamu- rensis, male. Anamurum (left); ssp. ibera, female, Koyegegiz (right). Table 1. Measurements (straight-line distance) of the type specimen and a paratype NMW 30795:1 NMW 30795:3 210mm Carapace 260 mm length Carapace width 160 mm Carapace 105 mm height 135 mm 88 mm Vol. 9, p. 146 Asiatic Herpetological Research 2000 303 tan 9 Testudo graeca ibera C) Testudo graeca ibera "lerrestris" O Testudo graeca "lerrestris" J^ Testudo graeca anamurensis ssp. nov. Figure 5. Localities of Testudo graeca in Turkey with dates that tortoises were observed (Weissinger, 1 986). 1 , ca. 70 km NW of Ankara, 5/18; 2, Dogansehir, open oak forest, 5/20; 3, 4 km from Diyabakir, 5/22; 4, 40 km from Mar- din; 5, Birecik, 5/24; 6, Nizip, in the direction of Ganziantep, dry wadi, about 45g steep loam walls with tortoise bur- rows, 5/24; 7, ca. 10 km N of Islaniye, 5/24; 8, 20 km S of Iskenderund, 5/25; 9, ca. 20 km N. of Iskenderund, 5/26; 10, ca. 25 km of Fg, cornfield, 5/26; 1 1 , 1 km from road crossing Ceyhan - Gaziantep, lava field, 5/26; 12, Ayas, 5/ 27; 13, Korykos-Kizkale, in the direction of Silifke, 5/27; 14, Bozyazi, onion field, 5/28; 15, Anamurum, 7 km W of Anamur, 5/29; 16, ca. 20 km W of Anamur, 5/29; 17Pine forest, ca. 40 km W of Anamur, towards Gazipasa, 5/29; 18, Side, sand dunes, ruin-meadows, 5/30; 19, 20 km from Antalya, 5/31; 20, 10 km from Antalya, towards Kugla, 5/31; 21, Kemer, 5/31; 22, 10 km from Kemer, 5/31; 23, Mountain pass, ca. 800 m above sea level, towards Kas, 5/31; 24, Vavi, ca. 800 m above sea level, towards Kas, 5/31; 25, Kinis, wheat field, 6/1; 26, Letoon, tilled field, 6/ 1 ; 27, Esen, pine forest, 6/1 ; 28, Kargi, 6/1 ; 29, Koygegiz, riverine forest with wet meadows 6/2; 30, Ula, 6/2; 31 , Gokovalskele, 6/2; 32, Yatagan, 6/2; 33, Soke, towards Kusadasi, 6/2; 34, Selcuk, 6/3; 35, Pamucak, 6/4; 36, 2 km from Ayvacik, 6/6; 37, 15 km from Ayvacik, 6/6; 38, 20 km from Kesan, towards Greece (European Turkey). Etymology I name this South-west coast race after its terra typica. Anamurum, Testudo graeca anamurensis. Distribution The area of distribution of this race extends along the Southwest coast of Turkey from the Bey Daglari in the West to the plain of Mersin in the East. Most were found in the area between Side and Anamurum. Acknowledgments This translation was provided by Peter Paul van Dijk with some additions by Heinz Grillitsch. The transla- tion was done with the permission and full agreement of the Austrian Herpetological Society. Figures 1-4 were provided by Jim Buskirk who, in turn, recieved Weissinger's original slides from his widow. The edi- tors of AHR would like to thank Heinz Grillitsch for his assistance in publishing this translation. Literature Cited Bodenheimer, F. S. 1935. Animal life in Palestine. An introduction to the problems of animal ecology and zoogeography. Ludwig Mayer, Jerusalem. 506 pp. Eiselt. T J., and Spitzenberger, F. 1967. Ergebnisse Zoologischer Sammelreisen in derTiirkei: Testudines. (Results of zoological collecting expeditions in Tur- key : Testudines) - Annalen des Naturhistorischen Museums in Wien. Serie B. fur Botanik und Zoologie 70: 357-378. 2000 Asiatic Herpetologia.il Research Vol.9, p. 147 Figure 6. The type specimen of Testudo graeca ana- /ra//- Rafe M. Brown, Alan E. Leviton, John W. Ferner. and Rogelio V. Sison. A New Snake of the Genus Hologerrhum Gunther (Reptilia; Squamata; Coluhridae) from Panay Island, Philippines 9 M. Farid Ahsan and Shayla Parvin. The First Record of Ptyas korros (Coluhridae) from Bangladesh 23 Sean J. Blamires. Influence of Temperature on Burrow Use hy the Monitor Lizard Varanus panoptes of the Coastal Dunes at Fog Bay, Northern Australia 25 Peter L. Cunningham. Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx aegyptius microlepis from the United Arab Emirates 30 John W. Ferner. Rafe M. Brown, Rogelio V. Sison, and Robert S. Kennedy. The Amphibians and Reptiles of Panay Island. Philippines 34 James Ford Parham and Haitao Shi. The Discovery of Mauremys iversoni-like Turtles at a Turtle Farm in Hainan Province, China: The Counterfeit Golden Coin 71 Matthias Stock, Daniel Frynta. Wolf-Riidiger Grosse, Claus Steinlein, and Michael Schmid. A Review of the Distribution of Diploid, Triploid and Tetraploid Green Toads (Bujo viridis complex) in Asia Including New Data from Iran and Pakistan 77 Samraat Pawar and Sayantan Biswas. First Record of the Smooth-Backed Parachute Gecko Ptychozoon lionotum Annandale 1905 from the Indian Mainland 101 Nasrullah Rastegar-Pouyam and Eskandar Rastegar-Pouyani. A New Species of Eremias (Sauria: Lacertidae) from Highlands of Kermanshah Province, Western Iran 107 Konstantin A. Rogovin, Dmitry V. Semenov, and Georgy I. Shenbrot. Lizards of the Northern Mongolian Deserts: Densities and Community Structure 113 Murat Sevinc and Ismail Hakki Ugurta§. The Morphology and Size of Blood Cells of Lacerta rudis bithynica 122 Muhammad S. Khan. Buccopharyngeal Morphology and Feeding Ecology of Microhyla ornata tadpoles 1 30 Ismail H. Ugurtas. Variation in Pelobates syriacus of Turkey 139 Translation: Mingtao Song. A New Species of the Turtle Genus Cuora (Testudoformes: Testudinidae j 142 Translation: Heinz Weissingcr. Testudo graeca anamurensis ssp. nov. from Asia Minor 145 Book Review A Guide to th~ Fauna Of Iran . . 149 Book Review: Four Receni Handbooks for Turkey .'...'. 1^1 (Continued on inside of back cover) Harvard MCZ Llbran llll 3 2044 066 300 427