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

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.

2001

Asiatic Herpetological Research

Vol. 9, p. 3

>

o

•B 7

9?

o

e

a>

o

o

<U

o5

3-

J I L

J I I I l_L

10 11 12 1 2 3 4 5 6 7 8 9 month

Figure 1 . The seasonal change of Mature coefficient

of ovary (1991-1993)

Eggs. The mean number of eggs spawned in each

batch is about 218. The egg of R. boulengeri is big

with a mean diameter of 3.98 mm, enveloped by 3

layers of gum membrane (The diameter of egg includ-

ing membrane is 15.85 mm). The outer layer is the

thickest, the inner layer the thinnest. The outer layer is

very sticky, so that many eggs connect together to

form a long cluster, the end of egg cluster is stuck on

the wall or to ratty plants under water.

Development of genital glands

Development of ovary. The development of ovary

can be divided into 6 stages.

Stage 1. Multiplication period of ovogonium: The

body weights (BW) of young frogs are 7.3-12.5 g and

the body lengths (BL) are 37.2-43.6 mm. The ovogo-

nium cells, aggregating densely with the ability of

division, have not been surrounded by follicle cells.

Stage 2. Growth of ovocyte (oocyte): When the BW

of young frog are 10.2-19.5 g and BL are 47.2-56.2

mm, ovogonium cells develop into ovocyte cells. The

ovocyte clusters are divided by connective tissue of

the ovary and surrounded by follicle cells.

Stage 3. Transition growth of ovocyte: When BW of

young frog are 36.5-74.2 g and BL are 71.7-84.3 mm,

the volume of ovocyte cells increases. The follicle

cells increase from 1 to 2 layers and the zona pellu-

cida becomes clear.

Early stage 4. Formation of yolk in ovocyte: When

young frogs weigh 58.7-80.8 g and their BL are 81.5-

89.7 mm, oocytes begin rapid growth and particles of

yolk begin to form.

Late stage 4. Ovocyte cells filled with yolk: When

frogs have BW of 79.7-125.8 g and BL of 87.3-98.0

mm, the ovum cells are fully filled with yolk and the

nuclei are isolated in the middle of the cells. The two

layers of follicle cells and zona pellucida are well

marked.

Stage 5. Slanting of nucleus of oocyte: When BW are

86.3-161.5 g and BL are 89.4-1 12.3 mm, the nucleus

inclines to the side of animal pole. At this stage, the

female can ovulate under the action of gonadotrophic

hormone.

Stage 6. Ovary after ovulation: When frogs body

weights decrease to 70.6-102.5 g and body length

decrease to 82.1-100.2 mm. In this stage, corpora

lutea appear in ovary.

Development of testis. The development of testis can

be divided into 5 stages.

Stage 1. Multiplication period of spermatogonium: At

this stage, spermatogonium aggregates densely, and

ranges irregularly.

Stage 2. Formation of sperm-tube: In this stage, the

spermatogonium is separated into sperm-tube by con-

nective tissue of testis and part of spermatogonium in

sperm-tube becomes spermatocyte of the first order.

Stage 3. Differentiation of spermatocyte: In this stage,

the number of spermatogonium in sperm-tube

decreases. On the wall of sperm-tube, there are pri-

mary sperm mother cells, secondary sperm mother

cells and spermatophore formed by spermoblast

respectively.

Stage 4. Formation of sperm: This is the ripe stage of

the testis. The spermatogonium and the spermatocyte

of the first order are few on the wall of sperm-tube

instead of spermatocyte of the secondary order and

spermatophore formed by spermoblast respectively.

After the forming of spermatozoon, the spermatozoon

aggregates densely first, then disperses.

Stage 5. Post-ejecting: Most spermatozoons have

been ejected and only a few could be found in the

sperm-tube.

Female order of ovary development and age at sex-

ual maturity. After metamorphosis, the ovaries of the

young frogs reach stage 1 in 2-3 months; stage 2 in 6-

8 months; stage 3 in 10-13 months; early stage 4 in

21-24 months; late stage 4 and stage 5 in 33-36

months (if ecological conditions are very suitable, in

only 22-24 months), when the females can spawn nat-

urally or artificially. After spawning, the ovaries

reaches stage 6.

Male order of testis development and age at sexual

Vol. 9, p. 4

Asiatic Herpetological Research

2001

to

&

1.2

1.1

1.0

0.9

0.8

ti 0.7

c

CD

'o

0)

o

o

a)

g

ra

0.6-

0.5

A

J I I I L

I I .

10 11 12 1 2 3 4 5 6 7 8 9 month

Figure 2. The seasonal change of Mature coefficient

of testis (1991-1993)

maturity. After metamorphosis, the testes of males

develop into stage 1 in 2-4 months; stage 2 in 8-10

months; stage 3 in 14-18 months; stage 4 in 24-26

months. At this stage the males can mate with the

females.

Type of spawning. In the sexually mature females, 3-

4 grades of oocyte can be seen. This is the cytological

proof of multi-oviposition of Rana boulengeri.

Seasonal change of ovaries. The period from May to

August is the reproductive time of R. boulengeri.

Then the ovaries develop slowly and the mature coef-

ficient of ovary (MCO) decreases to 3.10-3.35%.

From February to April of the second year, the ovaries

develop fast and the MCO reaches its highest value

(5.73-14.5%). The mean value of MCO in May is

8.47% (Fig.l). After May, the ovocytes mature in

batches.

Seasonal change of testes. The testes change little

during the different seasons. The range of change of

mature coefficient of testis (MCT) in a year is 0.55-

1.14%. From April to July, testes develop faster than

in other months. In July the MCT is 1.14%. In Octo-

ber the MCT decreases to 0.55% (Fig.2).

Embryonic development

Li et al. (1993) reported in detail on the embryonic

development of R. boulengeri. Under temperature

conditions of 23±0.5°C, 261.05 ± 0.54 hours were

required for the embryo to complete its development.

The whole development procedure is divided into 25

stags on the basis of both the morphological and the

physiological changes during this period (Pollister

and Moore, 1937; Shumway, 1940). The temperature

summation above 0°C for the embryo to complete the

whole development is 6004.15 degree-hours.

Experiments have shown that temperature can

affect the developmental rate of the embryo of R. bou-

lengeri. Under conditions of 13°C, 16°C, 22°C, 24°C,

26°C and 28°C, the time for the embryo to complete

development is 937.97h, 687.78h, 321.61h, 304.76h,

300. 18h and 295. 17h. In 2°C and 4°C water the

embryo can not develop. In 7°C water the embryo

develops very slowly. In 30°C water, the embryo can

not complete development. The optimum tempera-

tures for embryonic development are 22°-24°C (Li et

al., 1994).

Acknowledgments

My research has been supported by the Asiatic Herpe-

tological Research Society. I would like to thank Pro-

fessor Er-mi Zhao (Chengdu Institute of Biology) for

guiding my research and revising my paper. The

author wishes to thank Changde Teachers' College for

its financial support.

Literature Cited

Li, Hu-Ming, Wen-Jian Li, Xing-Guo Gong, and

Ming-You Ma. 1993. Studies on the nutritional com-

positions and energy content of the flesh of Rana bou-

lengeri. Zoological Research, Kunming, 14(1 ):96.

Li, Wen-Jian, Hu-Ming Li, and Ming Li. 1993. Pre-

liminary studies on the artificial breeding of the young

frog of Rana boulengeri Giinther. Pp. 255-259. In

Zhao, Chen and Papenfuss (eds.), Proceedings of the

First Asian Herpetological Meeting. China Forestry

Press, Beijing.

Li, Wen-Jian and Ming Li. 1993. The early embryonic

development and stages of Rana boulengeri. Pp. 124-

133. In Zhao, Chen, and Papenfuss (eds.). Proceeding

of the First Asian Herpetological Meeting. China For-

estry Press, Beijing.

Li, Wen-Jian and Hu-Ming Li. 1994. Effect of temper-

ature on the early embryonic development of Rana

boulengeri. Pp. 508-512. In Proceedings of the sixti-

eth anniversary of the founding of China Zoological

Society. China Science and Technology Press,

Beijing.

Pollister, A. W and J. A. Moore. 1937. Tables for the

normal development of Rana sylvatica.. The Anatomi-

cal Record 68:489-496.

2001 Asiatic Herpetological Research Vol. 9, p. 5

Shumway W. 1940. Stages in the normal development

of Rana pipiens. The Anatomieal Reeord 78: 1 39- 1 47.

Yuan. Feng-Xia and Xiao-Bo Wen. 1990. A prelimi-

nary study on living and feeding habits of Rana bou-

lengeri in western Hubei Province. Chinese Journal of

Zoology. Beijing 25(2)17-21.

2001

Asiatic Herpetological Research

Vol. 9. pp. 6-8

A New Species of Batrachuperus from Northwestern China

Mingtao Song2, Xiaomao Zeng1, Guanfu Wu1, Zhijun Liu 'and Jinzhong Fu3

Chengdu Institute of Biology, Chinese Academy of Sciences. Chengdu. China 610041. Northern west Institute

of Endangered Animals. Xi'an. China 710032. ~ Department of Zoology. University of Guelph. Guelph. Ontario.

Canada NIG 2W1

Abstract.- We describe a new species of salamander in the genus Batrachuperus from Tsinling Mts. in western

China. The new species is morphologically most similar to B. longdongensis. but differentiated by the absence of

horny covers on palms and tarsa. It represents the most northeastern distribution and the lowest elevation of the

genus.

Key words.- Caudata. Batrachuperus. new species. China. Tsinling

Introduction

The salamander genus Batrachuperus Boulenger.

1878 contains seven species (Frost. 1985). Three spe-

cies occur on the western side of the Tibet Plateau

(Iran and Afghanistan). The remaining four species

are found on the eastern side of the Tibet Plateau in

China. During the tieldwork in 1986 and 1999. we

collected specimens of Batrachuperus from Zhouzhi.

China. This collection marks the most northeastern

distribution of the genus. The specimens are different

from all other described species (Fei et al., 1990). and

we describe them as a new species.

Batrachuperus taibaiensis new spe-

cies (Fig. 1). Taibai Stream Sala-

mander

Holotype: NIEA 860122. An adult male from the

upper stream of Heihe River, near Hua Er Ping Vil-

lage. Zhouzhi County. Shaanxi Province, China

(33.85°N, 107.82°E). collected by M. Song on August

8. 1986. elevation 1260m. The holotype is deposited

Figure 1 . Paratype of Batrachuperus taibaiensis (MVZ 230964).

2001

Asiatic Herpetological Research

Vol. 9, p. 7

at the herpetological collection of Northwest Institute

of Endangered Animals, Xi'an, China.

Paratypes: Allotype. NIEA 860116, an adult

female, collected from the same locality at the same

time as the holotype. Other paratypes include NIEA

860114-5, 860117-9, 860121, 860126-7, 860129-139

and MVZ 230964-65, 230979-86. The NIEA series

are collected from the same locality at the same time

as the holotype and are deposited at the herpetology

collection of Northwest Institute of Endangered Ani-

mals, Xi'an, China. The MVZ series are collected

from the same locality on September 30. 1999 by Z.

Liu, and are deposited at the herpetology collection of

the Museum of Vertebrate Zoology, University of Cal-

ifornia. Berkeley.

Diagnosis: A relatively large, robust stream sala-

mander; distinguished from other members of the

genus by the large size, vomerine teeth arranged in a

"A" shape, and lack of horny cover on palms and

tarsa. Morphologically, the species most closely

resembles B. longdongensis. Both species are distin-

guished from others by large size and vomerine teeth

arranged in a "A" shape. Between the two species, B.

longdongensis differs from B. taibaiensis by the pres-

ence of the horny cover on palms, tarsa, and tail tip, as

well as the presence of gill slits in adults. Batrachupe-

rus taibaiensis is distinguished from geographically

neighboring species, B. tibetanus, by its large size and

the arrangement of vomerine teeth. Most phylogeneti-

cally closely related to B. karlschmidti and B. tibeta-

nus. Its cytochrome b gene sequence differs from B.

longdongensis by 9.2%, from B. karlschmidti by 7%,

and from B. tibetanus by 8.2-8.6% (J. Fu et al.,

unpublished data).

Description: Batrachuperus taibaiensis is a large

(adults males over 217 mm maximum SL), stout spe-

cies. Head moderately depressed, its length from

snout to gular fold longer than its width; snout short

and round. Labial fold well developed, often partially

covers the lower jar. Angle of jaw just behind the pos-

terior corner of eye. Both maxilla and mandible with

tiny teeth. Tongue elliptical, without free end. Ver-

mine teeth four, arranged in "I \" shape.

Body stout. Male body length slightly longer than

tail length and female body and tail length about the

same. Costal grooves 11. Limbs relatively short but

strong; when adpressed, tips of digits do not overlap

and are always separated by 1-3 costal folds in adults,

but contacted in juveniles. Separation is greater in

males. Fingers four, 3-2-4-1 in order of length. Toes

four, 3-2-4-1 in order of length. Most individuals

without palmar and tarsal tubercles. No horny cover

Table 1 . Measurement (range, means) and percentage ratios of each character dimension to SVL. All measure-

ments are in mm.

Measurement

Snout-vent length

Tail length

Head length

Head width

Forelimb length

Hindlimb length

Limb interval

Vomerine teeth

Holotype

Allotype Male (n =11)

Juvenile Female (n=9)

102

115

26

23

26

30

52

105

105

27

25

22

30

51

92.9±12.1

73—110

98.5±13.7

76—115

25.4±2.9

20—29

20.4±1.6

18—23

22.7±2.5

20—27

29.6±3.0

25—36

44.8±7.9

33—59

4

65

60

19

13

18

21

30

102.1 ±7.1

94—111

101.3±7.5

91—113

27.2±1.5

25—28

21.4±1.7

19—25

23.7±1.2

22—26

30.9±2.5

23—34

52.2±8.7

44—68

4

Vol. 9, p. 8

Asiatic Herpetological Research

2001

on palms, tarsa, and ventral side of the fingers, and

toes. Some individuals have horny cover of the very

tips of the fingers and toes. Tail round at the base and

gradually flattened laterally. Tail fin moderately high.

Skin smooth. The measurements of the type speci-

mens are presented in table 1.

Habitat and distribution: This species is only

known from the type locality, which is the most north-

eastern distribution of the genus and the only known

location from the northern slope the Tsinling Mts. It

has also the lowest elevation of the genus (1260m).

This species inhabits in fast moving streams (close

to the headwater of Heihe River). During daytime,

they are found under rocks in the stream, and have

never been observed under rocks on the riverbank.

The stream is well covered by the canopy from both

sides of the river and has steep slopes.

Etymology: The name taibaiensis is derived from

the name of the nearby peak, Taibai Peak, which is the

highest point of Tsinling Mts.

Remarks: Recent phylogenetic study of the genus

supported the species status of B. taibaiensis (Fu et al.

unpublished data). It is the sister group of the clade

including B. karlschimdti and B. tibetanus. Together,

the clade is the sister group of B. yenyuansis and B.

pinchonii.

Acknowledgments

We are grateful to E. Zhao, Z. Kou, Q. Wang and K.

Li for their help. This research was supported by the

Chengdu Diao Science Fund, Southwest Base Fund

and NSFC 30070090 to X. Zeng and National Geo-

graphic Society grant 6591-99 to J. Fu.

Literature Cited

Boulenger, G. A. 1878. Description de deux genres

nouveaux de la famille des salamandrides. Bulletin de

la Societe Zoologique de France 1878:71-72.

Fei, L., C. Ye, and Y. Huang. 1990. Key to Chinese

Amphibia. Chongqing Branch, Science and Technol-

ogy Literature Publishing House, Chongqing,

Sichuan.

Frost, D. R. 1985. Amphibian Species of the World.

The Association of Systematics Collections and Allen

Press, Lawrence, Kansas.

2001

Asiatic Herpetological Research

Vol. 9, pp. 9-22

A New Snake of the Genus Ho/ogerrhum Gur\\her (Reptilia; Squamata;

Colubridae) from Panay Island, Philippines

Rafe M. Brown1- 2, Alan E. Leviton3, John W. Ferner2- 4, and Rogelio V. Sison5

Section of Integrative Biology (C0930) and Texas Memorial Museum, University of Texas, Austin, Texas, 78712,

USA. e-mail: rafe@mail.utexas.edu; Geier Collections and Research Center, Museum of Natural History and

Science, 1301 Western Avenue, Cincinnati, Ohio 45203; - Department of Herpetology, California Academy of

Sciences, San Francisco, California 94118, USA. e-mail: aleviton@CalAcademy.org; Department of Biology,

Thomas More College, Crestview Hills, Kentucky, 41017, USA. e-mail: femerj@thomasmore.edu; Herpetology

Section, Zoology Division, National Museum of the Philippines, Executive House, P. Burgos Street, Manila,

Philippines, e-mail: nmuseum@webquest.com.

Abstract.- We describe a new species of snake in the genus Hologerrhum from two forested localities in Antique

Province, Panay Island, Philippines. To clarify species boundaries, we also redescribe its only known congener,

H. philippinum, on the basis of historical collections and newly-acquired material from the Islands of Luzon,

Marinduque, Polillo, and Catanduanes. The new species is the first Hologerrhum from the Visayan Aggregate

Island Complex and differs from Hologerrhum philippinum in color pattern and scalation of head and body. The

new species is one of several recently described vertebrates from Panay Island. Together, they indicate that

forested regions of the individual islands of the Visayan Aggregate Island Complex (Negros, Panay, Cebu.

Masbate, and other associated smaller islands) contain low levels of taxonomic endemicity that warrant further

study.

Key words.- Colubridae, Hologerrhum, Philippines, Panay Island, Visayas.

Introduction

Giinther (1858) erected the monotypic genus Holo-

gerrhum to accommodate a single specimen from

Hugh Cuming's Philippine collections that had been

deposited in the Natural History Museum, London

(Giinther, 1873; Fig. 1A). Giinther distinguished the

new genus and species from members of the Philip-

pine genus Cyclocorus by the presence of slight

grooves in the enlarged fang-like teeth at the posterior

end of the maxilla. Other slight differences between

Hologerrhum and some species of Cyclocorus, not

emphasized by Giinther but mentioned by other work-

ers (Taylor, 1922a, 1922b; Leviton, 1965). include

color pattern, slight scale pattern differences, and less

strongly enlarged middle series of maxillary teeth in

Hologerrhum.

At the time of the description of Hologerrhum, no

specific (island) locality data were available, but

Giinther later (1879) referred a specimen from Placer,

northeast Mindanao Island to this species. That speci-

men (not seen by us) apparently is a representative of

the genus Cyclocorus Taylor 1922c (vide Boulenger,

1896; see also comment by Taylor, 1 922a: 1 16).

Jan and Sordelli (1870) described Cyclochorus

maculatus (generic name misspelled), on the basis of

a specimen reportedly from Java Island, Indonesia

(Fig. IB). Fischer (1885) followed by recognizing

Cyclochorus lineatus var. maculatus reportedly from

S. Mindanao Island, Philippines, but without refer-

ence to new material. The type specimen of Cyclocho-

rus maculatus later was shown to be a representative

of H. philippinum (Giinther, 1873. 1879; Boettger,

1886; Taylor, 1922a), suggesting locality errors by

both Jan and Sordelli and Fisher. Later, Leviton

(1965) inadvertently included Cyclochorus maculatus

in the synonymy of Cyclochorus lineatus.

Castro de Elera (1895) reported Hologerrhum

philippinum from Baco, Mindoro Island. This impor-

tant specimen could not be examined as it was

destroyed during dissections by a biology class at the

University of Santo Thomas, Manila (R. I. Crombie,

personal communication) but the "Mindoro" locality

information suggests a misidentification of a speci-

men of C. lineatus (Taylor, 1922a; Leviton. 1963,

1965). Griffin (1910) did not include Hologerrhum in

his list of snakes from Polillo but did include the spe-

cies in his key to the Philippine snakes (Griffin, 191 1),

although he erred in attributing the type description to

Boulenger and supplied no precise locality data.

Vol. 9, p. 10

Asiatic Herpetological Research

2001

It was not until E. H. Taylor's extensive work in

the Philippines that specimens of H. philippimtm with

reliable locality data became available. Taylor (1922a,

1922b, 1922c, 1922d) consistently reported that this

species was collected in montane habitats in primary

forest and usually was associated with rocky stream

beds on Luzon and its land-bridge satellite island of

Polillo (Fig. 2). Still, Taylor (1922b:200) considered

the species rare and only obtained eight specimens in

two years of nearly continuous field work.

During the nearly 80 years that have elapsed since

Taylor's work, several additional specimens of H.

philippinum have been collected on Luzon and its

associated land-bridge islands (Marinduque, Polillo,

and Catanduanes; see Specimens Examined; Fig. 2).

During the same period, none have been found on

Mindanao, Mindoro, or any of the other Philippine

islands, thus bolstering the notion that H. philippinum

is endemic to the Luzon Pleistocene Aggregate Island

Complex (Fig. 2; sensu Heaney, 1986; Alcala, 1986;

see also comments by Leviton, 1963).

In 1992, while participating in the National

Museum of the Philippines/Cincinnati Natural His-

tory Museum Philippine Biodiversity Inventory

(PNM/CMNH PBI), one of us (RMB) collected speci-

mens of what appeared to be a distinctive new species

of snake, similar to but obviously specifically distinct

from H. philippinum, at 1025 m elevation on the west

face of Mt. Madja-as, Panay Island. In addition to rep-

resenting a previously unrecognized species, this

specimen appears to be the first vouchered record for

the genus on the Visayan Aggregate Island Complex

(Fig. 2; Heaney, 1986; Hall, 1996, 1998). During the

course of this study we examined all available US and

Philippine museum collections of//, philippinum and

■ ;;*•,

Figure I.The first illustrations of Hologerrhum. (A) Gunther's (1879) drawings of H. philippinum 'and (B) Sordelli's

plate of Cyclochorus maculatus (= H. philippinum) from Jan and Sordelli (1870).

2001

Asiatic Herpetological Research

Vol. 9, p.

Less than 1 20 m

submarine contour

200

km

Luzon -

Mindoro-

Busuanga jj

* Balanes islands

Babuyan islands

Polillo

Mannduque

..■■'

Catanduanes

Palawan

.10"

Leyle

Mindanao

• Hologerrhum philippinum if Hologerrhum dermali

Figure 2. Map of the Philippine islands with the major

Pleistocene aggregate island platforms indicated by

the position of the 120 m underwater bathymetric con-

tour (following Heaney, 1986); known collection locali-

ties of the two species of Hologerrhum are indicated.

Darkened circles represent collection localities for

Hologerrhum philippinum 'Gunther, 1858, stars are

localities for Hologerrhum derma/mew species.

Hologerrhum n. sp. In this paper, we redescrihe H.

philippinum and describe the new species from Panay.

Material and Methods

We surveyed the forested slopes of Mt. Madja-as from

6 April- 1 May 1992, utilizing elevational transects as

described by Ruedas et al. ( 1994) and as modified by

Brown et al. (1995, 1996, in press; Ferner et al., this

issue). Sampling techniques employed a variety of

search and trapping methods, including multiple-per-

son time-constrained searches, sticky traps, and pit-

fall traps (Simmons. 1987). Specimens were fixed in

I07t buffered formalin and transferred to 70% ethanol

approximately 2 months following preservation.

Museum specimens (see Specimens Examined,

below) were examined for color pattern and scale

character differentiation and only data scored by

RMB and AEL were used in an effort to minimize

inter-observer sources of variation (Lee. 1990). We

used Dowling's (1951a. 1951b) methods for counting

scales and expressing scale row reduction formulae,

and applied the Evolutionary Species Concept (Simp-

son. 1961: Wiley. 1978; Frost and Hillis. 1990) in

making taxonomic decisions.

Species accounts

Hologerrhum philippinum Gunther 1858

Figures 3-6

Hologerrhum philippinum. Gunther ( 1 858: 1 86).

Cyclochorus maculatus, Jan and Sordelli

(1870:36; generic name misspelled; the specimen

illustrated is H. philippinum with doubtful locality

data).

Hologerrhum philippinum. Gunther (1873: 171:

specimen is a member of the genus Cyclocorus. vide

Boulenger. 1896).

Cyclochorus lineatus var. maculatus Fischer.

1885:81.

Hologerrhum philippinum, Boettger (1886:115)

Castro De Elera (1895:438: specimen probably

Cyclocorus lineatus): Boulenger (1896:33); Taylor

(1922a:116; I922b:200, 1922c:138); Ross and

Gonzales (1991:67); Brown et al. (1996:13).

Hologerrum philippinum Griffin. 1911:263

(generic name misspelled).

Diagnosis: H. philippinum differs from its conge-

ner, H. dermali, by (1) the presence of 12-30 (vs. 7-

10) pairs of alternating black spots on nuchal region

and anterior dorsum. (2) a pale orange to salmon (vs.

bright yellow) venter, (3) absence (vs. presence) of a

black midventral stripe. (4) labials cream or yellow

(vs. labials bright white with thin midlabial black

stripe). (5) dorsum tan to orangish brown or dark

brick red (vs. dark purplish brown), (6) chin and

throat of adults pale tan to orange, immaculate or with

faint white spots in some specimens (vs. darker pur-

plish brown with black and white ocelli), (7) invari-

able presence of moderate to enlarged pretemporal

(length more than half that of secondary temporal; vs.

pretemporal reduced or absent), and (8) posterior tips

of parietals extend caudally, posterior parietal suture

forming a medially inverted V-shaped cleft (vs. poste-

rior portions of parietals squared off. with no medial

cleft).

Description: Body cylindrical, ventrals convex,

head slightly distinct from neck, not flattened; eyes

small, pupil round: vertebral ridge absent.

Rostral scale much broader than deep, scarcely

visible from above, subtriangular with ventromedial

groove in dorsal aspect: nasal divided, naris piercing

Vol. 9, p. 12

Asiatic Herpetological Research

2001

Figure 3. Live photograph of Hologerrhum ph///pp/numUom the Zambales Mountains of western Luzon Island

(female, PNM 2490; photograph copyright D. Wechlser).

A

B

-« . •A-*"*'

I

£5

¥

D

- ,>_

*

$&■*

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

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.

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

14.

Antique

2001

Asiatic Herpetological Research

Vol. 9, p. 37

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

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).

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.

Alcala, A. C. 1962. Breeding behavior and early

development of frogs of Negros, Philippines Islands.

Copeia 1962: 679-726.

Alcala, A. C. 1986. Guide to Philippine flora and

fauna. Vol X, amphibians and reptiles. Natural

Resource Management Center, Ministry of Natural

Resources and the University of the Philippines,

Manila, Philippines. 195 pp.

Alcala, A. C. and W. C. Brown. 1955. Observations on

amphibians of the Mount Halcon and Canlaon areas,

Philippine Islands. Silliman Journal 2:93-102.

Alcala, A. C. and W. C. Brown. 1955. Discovery of

the frog Conuifer guentheri on Negros Island, Philip-

pines, with observations on its life history. Herpeto-

logical 3: 182-184.

Alcala, A. C. and W. C. Brown. 1956. Early life his-

tory of two Philippine frogs with notes on egg deposi-

tion. Herpetologica 12:241-246.

Alcala, A. C. and W. C. Brown. 1967. Population

ecology of the tropical scincoid lizard, Emoia atro-

costata, in the Philippines. Copeia 1967:596-604.

Alcala, A. C. and W. C. Brown. 1998. Philippine

amphibians, an illustrated field guide. Bookmark Inc.,

Makati City, Philippines. 1 16 pp.

Boulenger, G. A. 1882. Catalogue of the Batrachia

and Salientia s. Ecaudata in the collection of the Brit-

ish Museum. Taylor and Francis, London, xvi + 503

pp.

Brown, James, H., and M. V Lomolino. 1998. Bioge-

ography, 2 Ed. Sinauer Associates, Massachusetts.

Brown. R. M., J. W. Ferner, R. V. Sison. 1995a. Redis-

covery and redescription of Sphenomorphus beyeri

(Reptilia: Lacertilia: Scincidae) from the Zambales

Mountains, Luzon Island, Philippines. Proceedings of

the Biological Society of Washington 108:6-17.

Brown, R. M., J. W. Ferner, and L. A. Ruedas. 1995b.

A new species of lygosomine lizard (Reptilia; Lac-

ertilia; Scincidae; Sphenomorphus) from Mt. Isarog,

Luzon Island, Philippines. Proceedings of the Biolog-

ical Society of Washington 108:18-28.

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, and A. C. Diesmos.

2000. Status of some Philippines frogs referred to

Rana everetti (Anura: Ranidae), description of a new

species, and resurrection of R. igorota Taylor 1922.

Herpetologica 56:81-104.

Brown, R. M., and D. T. Iskandar. 2000. Nest site

selection, larval hatching, and advertisement calls of

Rana arathoonifrom southwestern Sulawesi (Celebes)

Island, Indonesia. Journal of Herpetology 34:404-413.

Brown, R. M., A. E. Leviton, J. W. Ferner, and R. V.

Sison. 2000. A new species of Hologerrhum (Ser-

pentes: Colubridae) from Panay Island, Philippines.

Asiatic Herpetological Research 9:9-22.

Brown, R. M., J. A. McGuire, J. W. Ferner, N. Icaran-

gal, and R. S. Kennedy. 2000. Amphibians and rep-

tiles of Luzon Island, II: preliminary report on the

herpetofauna of Aurora Memorial National Park, Phil-

ippines. Hamadryad 25:175-195.

Brown, W. C. and A. C. Alcala. 1961. Populations of

amphibians and reptiles in submontane and montane

forests of Cuernos de Negros, Philippine Islands.

Ecology 42:628-636.

Brown, W. C. and A. C. Alcala. 1964. Relationships of

the herpetofauna of the non-dipterocarp communities

to that of the dipterocarp forest of southern Negros

Island, Philippines. Senckenbergiana Biologica

45:591-611.

Brown, W. C. and A. C. Alcala. 1970a. The zoogeog-

raphy of the Philippine Islands, a fringing archipel-

ago. Proceedings of the California Academy of

Sciences 38:105-130.

Brown, W. C. and A. C. Alcala. 1970b. A new species

of the genus Platymantis (Ranidae) with a list of the

amphibians know from South Gigante Island, Philip-

pines. Occasional Papers of the California Academy

of Sciences 84:1-7.

Brown, W. C. and A. C. Alcala. 1978. Philippine liz-

ards of the family Gekkonidae. Silliman University

Press, Dumaguete City, Philippines. 146 pp.

Vol. 9, p. 68

Asiatic Herpetological Research

2001

Brown, W. C. and A. C. Alcala. 1980. Philippine liz-

ards of the family scincidae. Silliman University

Press, Dumaguete City, Philippines. 264 pp.

Brown, W. C. and A. C. Alcala. 1982a. A new cave

Platymantis (Amphibia: Ranidae) from the Philippine

Islands. Proceedings of the Biological Society of

Washington 95:386-391.

Brown, W. C, and A. C. Alcala. 1982b. Modes of

reproduction of Philippine anurans. Pp. 416-428. In

Rodin, A. G. J. and K. Miyata (eds.), Advances in her-

petology and evolutionary biology. Museum of Com-

parative Biology, Cambridge, Massachusetts.

Brown, W. C. and A. C. Alcala. 1986. Comparison of

the herpetofaunal species richness on Negros and

Cebu Islands, Philippines. Silliman Journal 33:74-86.

Brown, W. C. and A. C. Alcala. 1994. Philippine frogs

of the family Rhacophoridae. Proceedings of the Cali-

fornia Academy of Sciences 48: 1 85-220.

Brown, W. C, R. M. Brown, and A. C. Alcala. 1997a.

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: 405-42 1 .

Brown, W. C, A. C. Alcala, A. C. Diesmos, and E.

Alcala. 1997b. Species of the guentheri group of

Platymantis with descriptions of four new species.

Proceedings of the California Academy of Sciences

50:1-20.

Brown, W. C, A. C. Alcala, and A. C. Diesmos.

1997c. A new species of the genus Platymantis

(Amphibia: Ranidae) from Luzon Island, Philippines.

Proceedings of the Biological Society of Washington

110:18-23.

Brown, W. C, A. C. Alcala, and A. C. Diesmos. 1999.

Four new species of the genus Plarymantis( Amphibia:

Ranidae) from Luzon Island, Philippines. Proceedings

of the California Academy of Sciences 5 1 :449-460.

Brown, W. C, and D. S. Rabor. 1967. Review of the

genus Brachymeles (Scincidae), with descriptions of

new species and subspecies. Proceedings of the Cali-

fornia Academy of Sciences 34:525-548.

Collins, N. M., J. A. Sayer and T. C. Whitmore. 1991.

The conservation atlas of tropical forests: Asia and the

pacific. Simon and Schuster, New York, U.S.A. 311

pp.

Dickinson, E. C. 1991. Biogeography of Philippine

birds. Pp. 24-54. In Dickinson, E. C, R. S. Kennedy,

and K. C. Parks (eds.), The birds of the Philippines.

British Ornithologists' Union, Dorset, Great Britain.

Diesmos, A. C. 1998. The amphibian faunas of Mt.

Banahao, Mt. San Cristobal, and Mt. Maquiling,

Luzon Island, Philippines. MS thesis, University of

the Philippines at Los Banos, College, Laguna. Philip-

pines. 115 pp.

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 reductions in snakes. Copeia

1951:131-134.

Dubois, A. 1992. Notes sur la classification des

Ranidae(Amphibiens, Anoures). Bulletin of the Men-

suel Society Linneenne de Lyon 61:305-352.

Duellman, W. E., T. Fritts, and A. E. Leviton. 1978.

Museum acronyms. Herpetological Review 9:5-9.

Duellman. W. E. 1993. Amphibian species of the

world: additions and corrections. The University of

Kansas Museum of Natural History, Lawrence, Kan-

sas. 372 pp.

Emerson, S. B., R. F. Inger, and D. Iskandar. 2000.

Molecular systematics and biogeography of the

fanged frogs of southeast Asia. Molecular Phyloge-

netics and Evolution 16:131-142.

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.

Frost, D. R. 1985. Amphibian species of the world.

Allen Press and the Association of Systematic Collec-

tions, Lawrence, Kansas. 732 pp.

Gaulke, M. 1991a. On the diet of the water monitor,

Varanus salvator, in the Philippines. Mertensiella

15:143-153.

Gaulke, M. 1991b. Systematic relationships of the

Philippine water monitors as compared with Varanus

salvator salvator, with a discussion of possible dis-

persal routes. Mertensiella 15:154-167.

Gaulke, M. 1992. Taxonomy and biology of the Phil-

ippine monitors (Varanus salvator). Phillippine Jour-

nal of Science 121:345-381.

Gaulke, M. and U. Fritz. 1998. Distribution patterns

of batagurid turtles in the Philippines. Herpetozoa

11:3-12.

Gaulke, M. In press. Die Herpetofauna von Sibaliw,

einem der letzten Tieflandregenwaldgebiete der West-

Visayas, Philippinen - Teil I: Amphibien und Echsen.

Herpetofauna.

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.

2001

Asiatic Herpetological Research

Vol. 9, p. 69

Gonzales, P. C, and R. S. Kennedy. 1996. A new spe-

cies of Crateromys (Rodentia: Muridae) from Panay,

Philippines. Journal of Mammalogy 77:25-40.

Gyi, K. 1970. A revision of the colubrid snakes of the

subfamily Homolopsinae. Miscellaneous Publications

of the University of Kansas Museum of Natural His-

tory 29:47-223.

Hall, R. 1996. Reconstructing Cenozoic southeast

Asia. Pp. 153-184. In R. Hall and D. Blundell (eds.),

Tectonic Evolution of Southeast Asia. Geological

Society of London, London England.

Hall, R. 1998. The plate tectonics of Cenozoic SE

Asia and the distribution of land and sea. Pp. 99-132.

In R. Hall and J. D. Holloway (eds.), Biogeography

and geological evolution of Southeast Asia. Brack-

huys, Leiden, The Netherlands.

Heaney, L. R. 1985. Zoogeographic evidence for mid-

dle and late Pleistocene land bridges to the Philip-

pines. Modern Quaternary Research of SE Asia

9:127-143.

Heaney, L. R. 1986. Biogeography of small mammals

in Southeast Asia: estimates of rates of colonization,

extinction and speciation. Biological Journal of the

Linnean Society 28: 127-165.

Heaney, L. R. and J. C. Regalado. 1998. Vanishing

treasures of the Philippine rain forest. The Field

Museum, Chicago, Illinois. 88 pp.

Heaney, L. R., D. S. Balete, M. L. Dolar, A. C. Alcala,

A. T. L. Dans, P. C. Gonzales, N. R. Ingle, M. V. Lep-

iten, W. L. R. Oliver, P. S. Ong, E. A. Rickart, B. R.

Tabaranza, Jr. and R. C. B. Utzurrum. 1998. A synop-

sis of the mammalian fauna of the Philippines. Fieldi-

ana Zoology 88:1-61.

Heyer, W. R., M. A. Donnelly, R. W. McDiarmid, L.-

A. C. Hayek, & M. S. Foster. 1994. Measuring and

monitoring biological diversity: standard methods for

amphibians. Smithsonian Institution Press, Washing-

ton DC. 364 pp.

Inger, R. F 1954. Systematics and zoogeography of

Philippine Amphibia. Fieldiana Zoology 33:182-531.

Inger, R. F, and H. Marx. 1965. The systematics and

evolution of the Oriental snake genus Calamaria.

Fieldiana Zoology 49:1-304.

Inger, R. F, H. K. Voris, and P. Walker. 1986. Larval

transport in a Bornean ranid frog. Copeia 1986:523-

525.

Inger, R. I., and F L. Tan. 1996. Checklist of the frogs

of Borneo. Raffles Bulletin of Zoology 44:551-574.

Kennedy, R. S., P.C. Gonzales, E. C. Dickinson, H. C.

Miranda, Jr. & T. H. Fisher. 2000. A guide to the birds

of the Philippines. Oxford University Press, Oxford,

U.K. 369 pp.

Leviton, A. E. 1958. A review of the Philippine

snakes of the genus Oxyrhabdion (Serpentes: Colu-

bridae). Wasmann Journal of Biology 15:285-301.

Leviton, A. E. 1961. Description of a new subspecies

of the Philippine snake Dendrelaphis caudolineatas.

Occasional Papers of the Natural History Museum,

Stanford University 9: 1-6.

Leviton, A. E. 1963a. Contributions to a review of

Philippine snakes, I. The snakes of the genus Oli-

godon. Philippine Journal of Science 9 1 :459-484.

Leviton, A. E. 1963b. Contributions to a review of

Philippine Snakes, III. The genera Maticora and Cal-

liophis. Philippine Journal of Science 92:523-550.

Leviton, A. E. 1963c. Remarks on the zoogeography

of Philippine terrestrial snakes. Proceedings of the

California Academy of Sciences 4: 31:369-416.

Leviton, A. E. 1964a. Contributions to a review of

Philippine snakes, IV The genera Chrysopelea and

Dryophiops. Philippine Journal of Science 93: BO-

Leviton, A. E. 1964b. Contributions to a review of

Philippine snakes, V The snakes of the genus Trim-

eresurus. Philippine Journal of Science 93:251-276.

Leviton, A. E. 1964c. Contributions to a review of

Philippine snakes, VI. The snakes of the genus

Oxyrhabdium. Philippine Journal of Science 93:407-

422.

Leviton, A. E. 1964d. Contributions to a review of

Philippine snakes, VII. The snakes of the genera Naja

and Ophiophagus. Philippine Journal of Science

93:531-550.

Leviton, A. E. 1965. Contributions to a review of Phil-

ippine Snakes, VIII. The snakes of the genus Lycodon

H. Boie. Philippine Journal of Science 94:1 19-140.

Leviton, A. E. 1967. 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. 1968. Contributions to a review of Phil-

ippine Snakes, X. The snakes of the genus Aliaetulla.

Philippine Journal of Science 96:73-90.

Leviton, A. E. 1970a. Contributions to a review of

Philippine Snakes, XI. The snakes of the genus Boiga.

Philippine Journal of Science 97:291-314.

Leviton, A. E. 1970b. Contributions to a review of

Philippine snakes, XII. The Philippine snakes of the

genus Dendrelaphis (Serpentes: Colubridae). Philip-

pine Journal of Science. 97:37 1 -396.

Vol. 9, p. 70

Asiatic Herpetological Research

2001

Leviton, A. E. 1979. Contributions to a review of Phil-

ippine snakes, XIII. The Snakes of the Genus Elaphe.

Philippine Journal of Science 106:99-128.

Leviton, A. E. 1983. Contributions to a review of Phil-

ippine snakes, XIV. The Snakes of the Genus

Xenopeltis, Zaocys, Psammodynastes, and Myerso-

phis. Philippine Journal of Science 1 12:195-223.

Leviton, A. E., and W. C. Brown. 1959. A review of

the snakes of the genus Pseudorabdion with remarks

on the status of the genera Agrophis and Typhlogeo-

phis (Serpentes: Colubridae). Proceedings of the Cali-

fornia Academy of Sciences 29:475-508.

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.

Malnate, E. V, and G. Underwood. 1988. Australasian

natricine snakes of the genus Tropidonophis. Proceed-

ings of the Academy of Natural Science of Philadel-

phia 140:59-201.

McDiarmid. R. W., J. A. Cambell, amd T. A. Toure.

1999. Snake Species of the World: A Taxonomic and

Geographic Reference. The Herpetologists' League,

Washington DC. 511 pp.

McDowell, S. B. 1974. A catalog of the snakes of

New Guinea and the Solomons, with special reference

to those in the Bernice P. Bishop Museum, Part I.

Scolecophidia. Journal of Herpetology 8:1-57.

McGuire, J. A., & A. C. Alcala. 2000. A taxonomic

revision of the flying lizards of the Philippine Islands

(Iguania: Agamidae: Draco), with a description of a

new species. Herpetological Monographs 14:92-145.

Musters, C. J. M. 1983. Taxonomy of the genus Draco

L. (Agamidae. Lecertilia, Reptilia). Zool. Verhandel.,

Leiden. 199:1-20.

Ota, H., and T. Hikida. 1989. A record of a triploid

gecko, Hemidactylus stejnegeri, from the northern

Philippines. Japanese Journal of Herpetology 13:35-

39.

Ota, H., I. S. Darevsky, L. A. Kupriyanova, T. Hikida,

K.-Y. Lue, S.-H. Chen, and T. Hayashi. 1993. Geo-

graphic variation in the parthenogenic lizard Hemi-

dactylus stejnegeri Ota and Hikida 1989

(Gekkonidae: Reptilia), with comments on recently

discovered male phenotypes. Tropical Zoology 6:125-

142.

Regalado, F. B., and Q. B. Franco. 1973. History of

Panay. Central Philippine University, Iloilo City. 594

pp.

Ruedas, L. A., J. R. Demboski & 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.

Ross, C. A., A. C. Alcala, and R. V Sison. 1987. Dis-

tribution of Zaocys luzonensis (Serpentes: Colu-

bridae) in the Visayan Islands, Philippines. Silliman

Journal 34:29-31.

Ross, C. A., and P. C. Gonzales. 1991. Amphibians

and reptiles of Catanduanes Island, Philippine.

National Museum Papers 2:50-76.

Simmons, J. E. 1987. Herpetological collecting and

collections management. Society for the Study of

Amphibians and Reptiles Herpetological Circular

16:1-70.

Sison, R., P. C. Gonzales and J. W. Ferner. 1995. New

records from Panay, Philippines. Herpetological

Review 26: 48-49.

Taylor, E. H. 1920. Philippine Amphibia. Philippine

Journal of Science 16: 213-359.

Taylor, E. H. 1922a. Additions of the herpetological

fauna of the Philippine Islands, I. Philippine Journal

of Science 21: 161-206.

Taylor, E. H. 1922b. Additions to the herpetological

fauna of the Philippine Islands, II Philippine Journal

of Science 21: 257-303.

Taylor, E. H. 1922c. The lizards of the Philippine

islands. Philippine Bureau of Science, Manila, Philip-

pines. 269 pp.

Taylor, E. H. 1922d. The snakes of the Philippine

islands. Philippine Bureau of Science, Manila, Philip-

pines. 312 pp.

Taylor, E. H. 1923. Additions to the herpetological

fauna of the Philippine Islands, III. Philippine Journal

of Science 22:515-557.

Taylor, E. H. 1925. Additions to the herpetological

fauna of the Philippine Islands, IV Philippine Journal

of Science 26:97-1 11.

Taylor, E. H. 1928. Amphibians, lizards, and snakes

of the Philippines, Pp 214-241. In Dickerson, R. E. ,

Distribution of Life in the Philippines. Manila Bureau

of Printing, Manila, Philippines

Whitmore, T. C. 1984. Tropical rain forests of the far

east. Clarendon Press, Oxford, U.K. 221 pp.

Wynn, A. H. and A. E. 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.

Iverson, J. B. 1992. A Revised Checklist with Distri-

bution Maps of the Turtles of the World. Privately

Printed, Richmond. 363 pp.

Iverson, J. B. and W. P. McCord. 1992. A new Chi-

nese turtle of the genus Sacalia (Batagurinae:

Testudines). Proceedings of the Biological Society of

Washington 105(3):426-432.

Iverson, J. B. and W. P. McCord. 1994. Variation in

East Asian turtles of the Genus Mauremys (Batagu-

ridae; Testudines). Journal of Herpetology 28(2): 178-

187.

Lau, M. and Shi, H. 2000. Conservation and trade of

terrestrial and freshwater turtles and tortoises in the

People's Republic of China. Chelonian Research

Monographs 2:30-38.

Leviton, A. E., R. H. Gibbs Jr.. and C. E. Dawson.

1985. Standards in herpetology and ichthyology. Part

I. Standard symbolic codes for institutional resource

Vol. 9, p. 76

Asiatic Herpetological Research

2001

collections in ichthyology and herpetology. Copeia

1985(4):802-821.

McCord, W. P. 1997. 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

27.5

I 25

»

E 22.5

o

1 20

B

S 17.5

« 15

c

9 12.5

co

«> 10

3

<D

o 7.5

z 100 150 200 250 300 350 400

Integrated optical density (Feulgen dye intensity,

DNA content)

Figure 2. Diagram showing the nucleus projection area

in relationship to the integrated optical density (the

DNA content, respectively) of 100 Feulgen stained

erythrocyte nuclei of a diploid standard specimen from

50 km E Gonbad-e-Kavus (3 in Fig. 1), a tetraploid

standard specimen from Kashgar (43) and the diploid

specimen from Kapkan (12a).

Material and Methods

New data from Iran

We used three sources: Martens (unpubl.) recorded

mating calls of green toads from Fasham (4 July 1978,

23.00) and Polur (30 June 1978, 22.30) which were

analyzed according to Stock (1998a). Frynta collected

in 1997 and 1998 (comp. Frynta et al, 1997) green

toads, among them five living specimens from Ghola-

man (MTKD D 41350), Kapkan (MTKD D 41351),

Baghestan (CUP AMPH/IRA/130) and Choqua Zan-

bil (MTKD D 41352, 41353) whose ploidy levels

were analyzed with erythrocyte measurements,

microdensitometrical DNA-measurements and/or

karyological techniques (Schmid, 1978; Stock and

Grosse, 1997a). Stock visited in June 1998 Kerman,

the type locality of Bufo viridis kermanensis Eiselt &

Schmidtler, 1971, and Birjand, the type locality of

Bufo oblongus Nikolsky, 1896. A mating call of a sin-

gle male from Birjand recorded by Stock in the late

phase of the breeding period (8 June 1998, 21.30) was

analyzed as described by Stock (1998a). Seven speci-

mens form Kerman (among them ZFMK 69909 to

69911, MTKD D 40730, 40731, 41348, 41349) and

five adult and six juvenile specimens from Birjand

(among them ZFMK 69901 to 69908, MTKD D

40729, 41346, 41347) were examined either by flow

cytometry as described by Stock et al. (1999) or kary-

ologically according to Schmid (1978). For morpho-

logical comparisons we included type material - Bufo

kavirensis: GNM Ba. ex. 1278 (holotype), GNM Ba.

ex. 1280 (paratype); B. luristanicus: ZMUC R 13221

(holotype); B. oblongus: ZISP 1952.1 (now lecto-

type), ZISP 1952.2 (now paralectotype). Institutional

abbreviations are as listed in Leviton et al. (1985); for

localities see Fig. 1.

New data from Pakistan

In June and July 2000. Stock and Dressel traveled in

the Northern Areas and North West Frontier Province

of Pakistan. In the Hunza valley of the Karakoram

Range near Karimabad (n = 8), and Pasu (n = 50), at

the tributaries of the Gilgit river near Gupis (n = 2), at

the Shandur pass (n = 8), and in the Chitral valley,

near Buni (n = 2) and in Chitral City (n = 8), blood

samples for ploidy determination of anesthetized

adult green toads were taken, stored in 70% ethanol

and refrigerated until flow cytometry according to

Stock etal. (1999).

In Skardu, at the western margin of the type region

("Ladak") of Bufo latastii Boulenger, 1882, blood

samples of 15 adult toads of this species were taken,

among them ZMB 62721 to 62726. A mating call of a

single male (24 June 2000, 21.30) was recorded and

analysed as described (Stock 1998a).

For morphological comparisons we examined the

lectotype of Bufo latastii BMNH 1947.2.21.28 (for-

merly 72.4.17.223; Stock et al., 1999: Fig. 1).

Map (Fig. 1), list of records with ploidy detec-

tion and type localities of nominal Asian

green toad taxa (Appendix)

The map contains most of the localities or regions

where diploid, tetraploid and/or triploid toads have

been recorded in Middle and Central Asia and the

eastern parts of the Middle East after 1976, the year of

the first detection of polyploids. Our map covers all

regions where polyploid green toads have been found.

These records (numbers 1 to 74) from the literature

have been made using various methods for identifica-

tion or determination of the taxa and/or ploidy levels.

Some methods are not unambiguous and might there-

fore cause errors. Consequently, it was necessary to

evaluate the methods which were utilized for the

determination of the ploidy level because not all data

are indisputable. In the case of the numbers 1 to 74,

the present article only includes papers if they either

contain statements on the ploidy of the toads or data

appearing suitable to draw a conclusion to the ploidy

(e.g. call data, see below). However, even if authors

distinguished between diploid and tetraploid toads, in

rare cases, especially in abstracts, the localities of the

records were not precisely published (e.g., Fikhtman,

1989) or it was impossible to deduce from the publi-

Vol. 9, p. 80

Asiatic Herpetological Research

2001

20

175

IS

12.5

10

50 100 150 200 250 300 350

Integrated optical density (Feulgen dye intensrtsy; DNA-content)

Figure 3. Diagram showing the nucleus projection area

in relationship to the integrated optical density (the

DNA content, respectively) of 100 Feulgen stained

erythrocyte nuclei of a diploid standard specimen from

S of Gorgan (2), 50 nuclei of a tetraploid standard

specimen from Kashgar (43), 100 nuclei of the tetrap-

loid specimen from Baghestan (13) and 100 nuclei of

the diploid specimen from Gholaman (1).

cation to the exact ploidy of toads from a concrete

locality (e.g., Castellano and Giacoma, 1998) and/or

the method of ploidy determination was not men-

tioned (e.g., Pisanets & Vasilenko, 1995). Since no

polyploid green toads have been found in the Cauca-

sus (e.g., Kuzmin 1995: 187), we disregarded this

area. If the information about a region but not a spe-

cial position was available, the record is not shown in

the map but listed in the appendix, and marked with a

"?" (instead a number) in the line next to the nearest

concrete record. Furthermore, we present some hith-

erto unpublished single records ("Stock, unpubl.")

from Kazakhstan and Kyrgyzstan.

We used different maps (Anonymous, 1993;

DMAAC) and/or the descriptions and sketch maps in

the literature (e.g., Borkin et al., 1986a) for the local-

ization of the records. If they were available, the

appendix contains the condensed descriptions of the

localities, the geographic position and the method of

ploidy determination.

The second part of the appendix (letters A to V)

contains the 21 Asian type localities shown in Fig. 1

and represents a preliminary precondition for a sys-

tematic discussion. The bibliography in that part only

comprises a choice of the literature which either dis-

cussed the systematic rank of a taxon or shows details

suitable to draw a conclusion to the ploidy. Some

descriptions of taxa either mentioned regions only but

no special type locality (e.g., Boulenger, 1882:

"Ladak" for B. latastii), but were drawn in the map;

others presented a confusing diversity of type locali-

01

E

3

o

iiriiiiliifftinl!

20 40 60

80 100 120 140 160 180 200

DAPI-fluorescence

Figure 4. Histogram obtained by DNA flow cytometry

from a mixture of DAPI stained blood samples with

chicken as the standard (a), CV = 2.63%, a diploid

green toad from Kerman, type locality of Bufo viridis

kermanensis, (b), CV = 3.62%, and a tetraploid green

toad from Birjand, type locality of Bufo ob/ongus, (c),

CV = 3.31%. Total cell number 10 327.

ties covering large parts of Central Asia (e.g., Bed-

riaga, 1898: B. viridis var. pewzowi and var. strauchi

with a type series from 4 or 14 localities, respectively,

distributed from Mongolia to the Pamirs). In the latter

cases, only some localities are shown, and this is men-

tioned in the appendix. In future, such taxonomic

problems should be resolved by careful lectotype des-

ignations leading to type locality restrictions as a

basis of a revision. Finally, the status of some old

names, at least for diploid green toads, still remains

unclear (see ref. in Kuzmin, 1999: 251, 264). They

were not shown in the map or originated from regions

outside of it.

Results and Discussion

New data from Iran

Cytometric and karyological data. A large male

from Kapkan (Fig. 1: 12a) in the East-Iranian part of

the Kopet Dagh which we considered to represent

Bufo viridis turanensis was identified to be diploid

(Fig. 2). A male Bufo viridis ssp. from Gholaman

(Fig. 1:1; Fig. 8C) in the Zagros mountains, was dip-

loid (Fig. 3); two additional males from Choqa Zanbil

(Fig. 1: la) below the south-western foot of the

Zagros mountains were also diploid.

All toads examined from Kerman, the type local-

ity of Bufo viridis kermanensis, were found to be dip-

loid as well (Fig. 4). This agrees with the

contemporaneously published results of Borkin et al.

(2000). We detected (Fig. 3) the first tetraploid toad in

2001

Asiatic Herpetological Research

Vol. 9, p. 81

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.

Mm 3.25 94.75 7 61 3.56 99.04 8.34 7 04 96.83 16.47 3.77 91.21 8.83

Max 3.69 107.58 8.63 3.63 100.95 8.50 7.57 104.13 17.71 4.59 1 10.95 10 74

SD 0.17 - 0 40 0.05 - 0.11 0.19 - 0.45 0.25 - 0.27

Table 2. Erythrocyte size (projection areas of 30 red blood cells per toad were measured) in five tetraploid speci-

mens from Birjand (B. ob/ongus), a tetraploid specimen from Baghestan, six diploid specimens from Kerman (B.

viridis kermanensis), a diploid specimen from Kapkan, a diploid specimen from Gholaman, and two diploid speci-

mens from Choqua Zanbil.

Parameter (urn2)

Locality

(ploidy)

ii

CO M

iS

-5, "

01 r-

S 5

CO

ii

!o N

1"

HI C flj c

C CM

CO <N

E M

II

C Z

(0 —

CO

C\l

Mean of means in the population

Maximal mean in the population

Minimal mean in the population

Largest cell measured

Smallest cell measured

Mean standard deviation in the pop.

353.87

374.30

333.03

463.29

274.07

33.62

335.98

335.98

335.98

413.50

280.11

30.80

256.38

274.93

224.40

358.91

183.95

23.22

272.24

272.24

272.24

318.11

216.37

21.99

248.39

248.39

248.39

322.77

216.21

22.17

CN

II

z

O CL o 21

236 26

181.47

286 32

181.47

286.32

22.94

localities has been controversially discussed (for over-

view see Balletto et al., 1999: Stock et al.. 1999). As

compared with conventionally stained tetraploid karyo-

types (e.g.. Borkin et al.. 1986b, c: Borkin and Kuzmin.

1988; Orlova & Uteshev, 1986; Pisanets. 1978; Roth &

Rab, 1986, 1987; Stock. 1998b; Toktosunov, 1984;

Whu & Zhao. 1987). that of B. oblongus (Fig. 6a. b)

does not exhibit visible differences.

Mating calls (Fig. 7, Table 3). The mating call data

from Lar valley (Andren and Nilson, 1979), Fasham

and Polur suggest that these toads are probably diploid

(see below for methodical questions). The same predic-

tion (Stock. 1998a) is possible for the toads from

Cheshmeh-ye-Sefid-Ab, the type locality of Bufo kavi-

rensis. The mating call data from Polur (2a) and

Fasham (2b) and the data from Lar valley (2b) and

Gorgan (2) refer to the occurrence of (only) diploid

2001

Asiatic Herpetological Research

Vol. 9, p. 83

35

30

n 25

<D

*■»

n

a> 20

v>

3

a.

15

10

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

<S Q.

« 2

£ o3

- 3

CD -Q

« 9

> .§

«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

o

co

-a

c

<

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

§.

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.

(one of) the oldest available name(s) for tetraploids in

Middle and Central Asia.

New data from Pakistan

Cytometric data. Among the triploid populations

reported by Stock et al. (1999), DNA-measurements

in additional specimens confirmed bisexual triploidy

in altogether 82 adult specimens of Bufo pseudorad-

dei baturae from Pasu (47). Exclusively triploids were

also detected by our sample survey along the upper

tributaries of the Hunza (47a) and Gilgit river systems

(48b, c). The record of triploids at the Shandur pass

(48c, 3720 m a.s.l ) and the occurrence of triploids in

the Chitral valley (48d, e) demonstrated their continu-

ous distribution, i.e. a more or less connected range.

In Chitral, sympatric occurrence of triploids with B.

stomaticus was observed.

The flow cytometric measurements (Tab. 1, Fig.

9E) revealed all 15 specimens of Bufo latastii from

Skardu (49) to be diploid. Although the absolute val-

ues cover a relatively large range resulting from the

fixation of samples in ethanol in the field, apparently

the DNA content of B. latastii is about 15 to 20%

higher than that of diploid B. viridis ssp. and B. v. ker-

manensis from Iran. The karyotype of toads from

Skardu comprises 2n = 22 chromosomes; a detailed

cytogenetic study will be published elsewhere. It

remains to be clarified if the karyological study on a

"Bufo spec." from Kashmir (Duda & Opendar, 1971)

really represented B. latastii as Dubois and Martens

(1977) and Roth and Rab (1986) presumed when they

considered this species diploid. In any case, long last-

ing speculations (see appendix: M) on the ploidy of B.

latastii which previously were connected with signifi-

cant uncertainties in the systematics of Central Asian

green toads (Borkin et al. 2000; Stock et al., 1999)

have been finished now.

Mating calls (Table 4). Mating call parameters of a

single diploid male B. latastii from Skardu were

found in the range reported for this species by Dubois

and Martens (1977) from Kashmir and Ladakh. As

discussed by Stock et al. (1999), the B. latastii calls

are rather similar to those of triploid B. pseudoraddei

baturae from Western Karakoram and B. p. pseudo-

raddei from Western Himalayas. This can be either

interpreted as convergences in isolated species or may

result from the participation of B. latastii genes in the

probably allotriploid B. p. baturae, but up to now,

exact explanations are not available.

Morphology and taxonomy. The morphology of

triploids from Shandur Pass (48c) and Chitral (48d, e)

was similar to that of B. pseudoraddei baturae from

Gilgit (Stock et al., 1999); therefore, we consider

them to represent the same taxon.

The 15 toads examined from Skardu (49) were

clearly identified as Bufo latastii Boulenger, 1882 (=

B. siachinensis Khan, 1997). The external morphol-

ogy and the coloration are very similar to the lecto-

type of B. latastii (Fig. 9A-D). Furthermore, the

characteristic ventral dark pattern comprising black-

ish, up to eye-sized, sometimes connected spots and

an abrupt darkening of the caudal third of the belly

including the base of the hind legs was found to occur

in the B. latastii lectotype, the B. siachinensis holo-

type, and always in the toads from Skardu also (Fig.

9C, D).

Vol. 9, p. 86

Asiatic Herpetological Research

2001

Table 5. Evaluation (overview) of methods previously used by different authors for the determination of diploid and

polyploid green toads. For detailed information see text.

Method of determination

Unequivocalness

Applicability in the field Abbr. in the appendix

Karyotype, karyogram, counting of

chromosomes

Flow cytometry

Microdensitometry (Feulgen stain-

ing), cytometry

Electrophoresis

Measurement of erythrocytes

yes

yes, if applied professionally

yes, if applied professionally

no, only prognosis possible

no, only prognosis possible

probably yes, for distinguish-

ing between diploid and poly-

ploid (triploid. tetraploid)

forms, but see B. latastii

External morphology, morphometry no, only prognosis possible

Form and size of the clutch no, often misleading

Mating call analysis

no

no, but blood samples can

shortly be stored in etha-

nol

yes, if blood smears are

stored in 5% formaldehyde

solution

collection of air dried blood

samples easily possible

yes

yes

chrom. count,

flow cytom.

microdens., cytom.

electrophor.

erythr. size

calls

ext. morph.

clutch

Evaluation of methods for the identification

and determination of the ploidy in Asian

green toads (Table 5)

Some forms/species of Middle and Central Asian

green toads have been partly identified using mislead-

ing or ambiguous methods (comp. also Stock and

Grosse, 1997a).

The karyological analysis still remains an essen-

tial tool. The treatment with colchicine and the prepa-

ration of bone marrow, spleen or intestine tissue for

the counting of stained chromosomes (e.g., Schmid,

1978; Macgregor and Varley, 1983) requires the kill-

ing of the animals. The obtaining of blood by heart

puncture with micro-syringes (e.g., Schroer, 1996)

and the determination of the ploidy by chromosome

counts from blood cultures (Castellano and Giacoma,

1998) causes little mortality but requires the transport

of the toads to the laboratory.

Flow cytometry is another unequivocal technique

(Murphy et al., 1997 for review). The preferable stor-

age of blood by freezing in liquid nitrogen is usually

limited in the field. Ethanol fixation of blood obtained

from anaesthized toads allows a storage for some

weeks and the ploidy determination by flow cytome-

try if refrigeration is possible (e.g.. Stock et al., 1999),

but does not allow the exact determination of absolute

DNA-contents.

Feulgen staining of erythrocytes and the

microdensitometrical determination of their DNA-

content provides also unequivocal results (Stock and

Grosse, 1997a). Although better results can be obtained

if applied in the laboratory since the storage of air dried

blood causes a loss of stainability, we recently got good

results with blood smears which were stored in 5%

formaldehyde solution until staining (our unpublished

data). The obtaining of blood smears from the tip of the

finger seems to be of very little damage for the animals,

but requires to prevent the pollution of blood samples

with skin mucus.

Significant differences in the average erythrocyte

size allow a prognosis of the ploidy level (Stock and

Grosse 1997a). Discrimination problems occur

because some populations exhibit values in the over-

lapping range between diploid and tetraploid toads. In

addition, the classification of triploid individuals

remains unsolved. In probably all-triploid populations

from Karakoram, the mean erythrocyte areas exhib-

ited significant differences to diploid and tetraploid

populations (Stock et al., 1999), but the variability of

the individual erythrocyte size shows a large overlap-

ping with diploid and tetraploid specimens. Unfortu-

nately, these phenomena also prevent an exact

analysis of the ploidy level of museum and type spec-

imens with the technique of Mercadal (1981).

Results of protein electrophoresis (e.g., Borkin

and Sokolova 1989, Mezhzherin and Pisanets, 1995a,

1995b) can usually not substitute the preceding ploidy

determination since "electromorphs" may cause mis-

interpretations. Some authors of electrophoretic stud-

2001

Asiatic Herpetological Research

Vol. 9. p. 87

Figure 8. A - Lectotype (present designation) of Bufo ob/ongusN\ko\sky, 1896 (Z ISP 1952.1) from Birjand, E-lran.

B - Topotypic tetraploid female Bufo oblongus\xom Birjand. C - Diploid male from Gholaman. D - Topotypic tetrap-

loid male Bufo ob/ongusUom Birjand. E - Topotypic diploid male of Bufo viridis kermanensis Eiselt & Schmidtler,

1971. F - Tetraploid male from Baghestan. G - Holotype of Bufo kavirensis, Andren & Nilson. 1979. H - Holotype of

Bufo luristanicus Schmidt, 1952.

Vol. 9, p. 88

Asiatic Herpetological Research

2001

ies did not describe whether or how they determined

the ploidy level (e.g.. Mezhzherin and Pisanets.

1991).

At any given temperature mating calls of tetrap-

loid toads exhibited longer pulses and interpulse inter-

vals resulting in lower pulse rates than diploid toads

(Stock 1997a. 1998a). a finding which exhibits paral-

lels in other diploid/tetraploid anuran species (Hyl-

idae, Leptodactylidae) and artificial tetraploids

suggesting that differences are caused by polyploidy

(Gerhardt. 1994: 317).

For triploid toads from northern Kyrgyzstan (Cas-

tellano et al.. 1998) and triploid populations from

northern Pakistan (Stock et al.. 1998. 1999) call

parameters were found to resemble those of tetraploid

toads. Therefore, the mating call analysis seems suit-

able for distinguishing between diploid and polyploid

(triploid. tetraploid) green toads. However, indepen-

dently from polyploidisation. speciation in the B. viri-

dis complex appears to take place also within each

ploidy level and as in other anurans. evolutionary

divergences in sympatry as well as convergences of

call parameters in allopatry can be expected. The lat-

ter seems to be the case in diploid B. latastii. from

which pulse rate data (Dubois and Martens. 1977)

suggested a polyploid species (Stock, 1998a).

Many morphometric traits exhibited differences of

the means but values showed large intersections

between both diploid and tetraploid toads and mor-

phometric parameters were not suitable for ploidy

determination (Stock. 1997b). The study announced

by Roth (1986) on "about 2000 specimens from the

whole range" has never been published (Roth. pers.

comm.). In advance. Roth (1986) considered morpho-

logical characters "useless" for taxonomy but without

determination of the ploidy level of the collection

material. Multivariate analyses confirmed relation-

ships between the ploidy level and the external char-

acter syndrome in diploid and tetraploid green toads

(Stock, 1997b; Castellano et al.. 1998). Triploid toads

from Kyrgyzstan exhibited more distinct differences

to diploid than to tetraploid specimens (Castellano et

al., 1998). Triploid populations in the Karakoram

showed significant differences to both diploid and tet-

Figure 9. A, B - Lectotype of Bufo /<?/<3s///Boulenger,

1882 (BMNH 1947.2.21.28, formerly 72.4.17.223),

scale for B: 1 centimetre. C, D- Male B. /atast//(ZMB

62721 ) examined from Skardu, Baltistan. E - Histogram

obtained by DNA flow cytometry from a mixture of

DAPI stained blood samples with chicken as the stan-

dard (a), CV = 2.94%, and diploid B. /atasti/Uom

Skardu (b), CV =-2.74%. Total cell number 22 744.

raploid toads from Central Asia in many characters

with univariate methods (Stock et al., 1999). A prog-

nosis for the purely morphometric classification of

diploid, triploid and tetraploid toads is possible with

multivariate methods and this appears to be one tool

for the prognostic assortment of preserved specimens

in collections. The prognosis can be improved if mor-

phometric data are combined with erythrocyte size

data (Stock, 1997b; Stock and Grosse, 1997a; Stock et

al., 1999). In any case, the use of exact methods for

ploidy determination is highly recommended (e.g.,

Borkin et al.. 2000).

Pisanets (1987) indicated differences in form and

size of the clutch between diploid and tetraploid toads

\ r— -1 r-

0 20 40 60 80 100120140160180200

DAPI-tluorescence

2001

Asiatic Herpetological Research

Vol. 9, p. 89

from Turkmenistan (Kurukhaudan village. 12). The

character is occasionally misleading (Kuzmin, 1995:

94; Stock and Grosse, 1997a) although it was infre-

quently used for the identification of toads for further

investigations (e.g., Mezhzherin and Pisanets, 1990).

Distribution

In general, the present map (Fig. I) does not com-

pletely reflect the occurrence of green toads whose

ploidy is known but also illustrates the recent degree

of investigation.

Diploid toads (black arrows without starlet). The

records of green toads found to be diploid in Middle

Asia are distributed in the Turan Basin and are also

concentrated at the foothills of the high mountains.

Diploids have been found along the bottom part of the

Kopet Dagh mountains (7, 8, 10, 1 1); a second group of

records is to be seen in the Kafirnigan and Vakhsh river

valleys (23, 25, 28, 29). The last records seem to exem-

plify a dispersion of diploid toads along the rivers into

the mountain valleys. The western and particularly the

northern Tien Shan form a third region with several

records of diploid toads (31a, 36, 38, 39, 59, 59a).

More distant from the mountains researches combined

with ploidy determination have only sporadically been

carried out. Merely some localities with doubtless dip-

loid toads near the lower Amu-Darya (9, 14) and Zer-

avshan river (17, 18) are known. The record of diploid

toads from the Chingzhal river (61) in Kazakhstan by

Golubev (1990) appears to be doubtful (Dujsebayeva et

al., 1997). The main range (maps in Dubois and Mar-

tens, 1977, Stock et al., 1999) of the diploid B. latastii

(N, 49) is the rather humid Kashmir valley and the

surrounding rather arid mountain ranges.

The small and highly generalized map by Kuzmin

(1995: 182) and the more detailed one (Kuzmin,

1999: 479) show either a continuous range or many

separate symbols of (diploid) B. viridis in Middle Asia

and Kazakhstan. Both maps cannot reflect the detailed

records and/or were not based on karyological studies.

Therefore, Kuzmin (1995, 1999) also announced that

the distribution requires additional investigations.

However, Kuzmin's maps refer to the occurrence of

diploid green toads in the Turan Basin along the rivers

whereas the symbols for (diploid) B. viridis in the

Inner Tien Shan and around the Issyk-Kul appear to

be very doubtful (e.g. see Borkin, 1989).

In spite of only few references which contained

data on the ploidy level and on the altitudinal distribu-

tion, the occurrence of diploid toads in Middle Asia

apparently exceeds 1600 m a.s.l. only in rare cases

(appendix; Castellano et al., 1998: "At least in Kaza-

khstan and Kyrgyzstan no diploid populations have

been found above 2000 m a.s.l."). Many papers (sum-

marized e.g. by Meinig, 1995) reporting the occur-

rence of "B. viridis" until 4000 m a.s.l. in Central

Asia, very probably represent data on polyploid toads.

This also may account for Kuzmin's (1999) material

who reported on B. viridis up to 3200 m a.s.l. in Mid-

dle Asia.

Tetraploid toads (white arrows). The tetraploid

toads are largely distributed in the Middle and Central

Asian mountain systems. They have been described

from the Kopet Dagh (5, 6, 8, 11, 12), the Gissar range

including its southern spurs (21 to 27, 27a, 30), the

western Tien Shan (31a, 32, 33, 40), the northern Tien

Shan and the Issyk-Kul gorge (39, 51 to 59, 59b, 60),

the Central Tien Shan (42, 51), the margins of the Tak-

lamakan desert (43, 50?), the region between Zailiyskiy

Alatau and lake Balkhash (60, 37, 37a, 37b), the Dzun-

garian Alatau (60a, 60b), the eastern Tien Shan (63),

and Dzungaria (62, 64, 66 to 72). In the Pamirs, tetrap-

loid toads have been found in the western (34, 35) and

in the central parts (45). Furthermore, Stock (1998b)

found toads from eastern Pamirs (44) to be tetraploid.

They were recently described by Fei et al. ( 1999) as a

new subspecies (T). The present map provides argu-

ments for a rather continuous distribution of tetraploid

toads from Eastern Tien Shan to the Dzungarian Gobi

in Mongolia. Interestingly, eastern of about 80° E only

records of tetraploids and obviously no record of dip-

loid toads have been made. Various records underline

the high ecological plasticity especially of the tetrap-

loid green toads (e.g. Dujsebayeva et al., 1997). The

dry centers of the large continental deserts Kyzylkum,

Karakum, Muyunkum and Betpak-Dala seem to be

not suitable as habitats of (tetraploid) green toads but

this might be also caused by the recent low degree of

investigation. Considering some records of tetraploids

in the eastern plane regions, some additional records

of tetraploid toads in the Turan lowlands can be

expected in future. The tetraploids in the West of the

known range (Kopet Dagh) may have a rather contin-

uous distribution via the mountains of Afghanistan

until those of the Pamiro-Alai-System, but data from

Afghanistan are recently too scarce for a detailed

analysis.

Triploid toads (black arrows with starlet). For the

first time, probable triploidy of three specimens was

detected by Bachmann et al. (1978: "36% more DNA

than diploid Bufo viridis" ) in toads (presumably) orig-

inating from Kabul (48f in Fig.l; Hemmer et al.,

1978: 352, 370). Triploid individuals have been

detected in south-western Turkmenistan (5), in north-

ern Kyrgyzstan (39) and south-eastern Kazakhstan;

the last record was reported without particular locality

Vol. 9, p. 90

Asiatic Herpetological Research

2001

(Borkin et al., 1997, 2000) and we did not draw it in

the map. The rare occurrence of triploid individuals in

the Central Pamirs (45) has been concluded from

isozyme data (Mezhzherin and Pisanets 1990). Most

of the authors supposed a hybrid origin of triploids as

a result of mismatings between diploid and tetraploid

toads or only noted the occurrence in contact zones of

diploid and tetraploid toads (Borkin et al., 1997,

2000). Preliminarily, Lattes (1997) and Cervella et al.

(1997) indicated that triploids from Kyrgyzstan (Kok-

jar, near 39; Castellano et al., 1998) do not originate

from hybridizations between diploid and tetraploids

but seem to be closely related forms of the tetraploids.

Presumably all-triploid populations of green toads

from the Karakoram range and Western Himalayas

(46, 47, 48a) have been discovered by Stock et al.

(1998) and were described as a new subspecies (Stock

et al., 1999). The new data from Pakistan (47a, 48b-e)

suggest that triploids, which form at least at some

localities (e.g., 47) all-triploid gonochoristic popula-

tions, are distributed along the upper tributaries of the

Gilgit river (48b), they live at the Shandur pass (48c,

3720 m a.s.l), and also in the Central Hindukush, i.e.

the Chitral valley (48d, e). Interestingly, in the Kara-

koram exists obviously an eastern limit of the distri-

bution of triploids situated between their most eastern

record (48a) and the most western one (49) of diploid

B. latastii in the gorge of the Indus river. Based only

on morphological characters, Baig (1998) reported on

the sympatric occurrence of "Bufo latastii" and "B.

pseudoraddei" in the "Neelam valley" (= Jhelum val-

ley) of Azad Kashmir in eastern Pakistan.

Generally, the correlation of distributional data on

green toads with global climatic factors appears to be

problematic because the local or microclimatic condi-

tions to which the animals are adapted may differ

strongly from the climatic zone in which any locality

is situated.

Zoogeographic implications. Borkin (1999: 350)

considered "the B. viridis group as an indicator of

southern limits of the Palearctic in arid regions".

Mazik et. al. (1976) as well as Pisanets (1978) already

reported an allopatric occurrence of diploid and tetra-

ploid green toads because they have different ecologi-

cal preferences. This hypothesis is principally

supported by the present map but it seems more

appropriate to call the ranges parapatric (Borkin et al.,

1997). Many authors (Pisanets and Shcherbak, 1979;

Toktosunov, 1984; Borkin et al., 1986b, c; Borkin and

Kuzmin, 1988) arranged the tetraploids of the arid

foothill regions in contrast to those of the high moun-

tains. In our opinion, it appears possible that the limit-

ing climatic factors in the high mountains as well as in

the more continental high plains are similar and are

obviously suitable only for polyploid green toads.

At the moment, the causes for the specific distri-

bution pattern remain speculation. Kuzmin (1995:

189, 1999) refers to the high degree of temperature

tolerance of tetraploid green toads (-30°C to 45°C). In

the high mountain habitats as well as in the continen-

tal deserts strong daily and annual variations of tem-

peratures as well as a high mean annual solar radiance

(e.g. in the northern Asian deserts 2700-2800 h, in the

interior Tien Shan > 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). Experientia 34:331-332.

Baig, K. J. 1998. The Amphibian Fauna of Azad

Jammu and Kashmir with new record of Paa liebigii.

Proceedings Pakistan Academy of Sciences

35(2): 1 17-121.

Balletto, E., M. A. Cherchi and J. Gasperetti. 1985.

Amphibians of the Arabian peninsula. Fauna of Saudi

Arabia 7:318-392.

Balletto, E.L, L. Borkin, S. Castellano, T Dujse-

bayeva, V. Eremchenko, C. Giacoma, A. Lattes, G.

Odierna. 1999. Sistematica e filogenesi nel complesso

di Bufo viridis Laurenti, 1768. Rev. Idrobol. 38 (1/2/

3): 199-220.

Baloutch, M. and H. G. Kami. 1995. Amphibians of

Iran. Tehran University Publ. 2250. (in Farsi).

Bedriaga, J. V. 1898. Wissenschaftliche Resultate der

von N. M. Przewalski unternommenen Reisen. Zoolo-

gischer Theil. vol. Ill, part 1. Amphibien und Rep-

tilien. Kaiserliche Akademie der Wissenschaften, 1st

ed. St. Petersburg.

Birstein, V. Ya. 1981. Osobennosti differencialnikh

okrasok khromosom Anura na primere khromosom

nekototrikh vidov Rana i Bufo. Zoologicheskii Zhur-

nal 60 (2):246-256. (In Russian).

Blair, W. F. 1972: Evolution in the genus Bufo. Uni-

versity of Texas Press, Austin.

Bogart, J. P. 1972. Karyotypes. Pp. 171-232 In W. F.

Blair (ed.), Evolution in the genus Bufo. Univ. Texas

Press. Austin, London.

Boulenger, G. A. 1882. Catalogue of the Batrachia

Salientia in the collection of the British Museum. 2nd

ed. London.

Boulenger, G. A. 1891. Descriptions of new Oriental

reptiles and batrachians. Annals and Magazine of Nat-

ural history (6) 7:279-283.

Borkin, L. Ya. 1989. Uber den Fund einer albinotis-

chen Kaulquappe der tetraploiden Krotenart Bufo

danatensis Pisanetz. 1978 aus dem Issyk-Kul'-See,

Kirgisien. Salamandra 25:122-124.

Borkin, L. Ya. 1999. Distribution of amphibians in

North Africa, Europe, and the former Soviet Union,

pp. 329-420. In Duellman, W. E. (ed.) Patterns of dis-

tribution of Amphibians. John Hopkins University

press. Baltimore, London.

Borkin, L. Ya., I. A. Caune, E. M. Pisanets and Y. M.

Rozhanov. 1986a. Karyotype and genome size in the

Bufo viridis-gwup. pp. 137-141. In Rocek, Z. (ed.),

Studies in Herpetology. Proceedings of the 6th Ordi-

nary General Meeting of the Societas Europea Herpe-

tologica. Prague.

Borkin, L. Ya., Ch. Terbish and I. A. Caune. 1986b.

Tetraploidnaya i diploidnye populyacii zhab gruppy

Bufo viridis iz Mongolii. Doklady Akademii Nauk

SSSR 287 (3):760-764. (In Russian).

Borkin, L. Ya., J. M. Rozanov, Ch. Terbish and I. A.

Caune. 1986c. Raspostranenie, kariologia, tak-

sonomitcheskoe polozhenie i izmenchivostch zhab

gruppy Bufo viridis v Mongolii. pp. 120-143. In Aka-

demiya Nauk SSSR, Institut evolyutsionnoi mor-

Vol. 9. p. 92

Asiatic Herpetological Research

2001

fologii i ekologii zhivotnych imeni A. N. Severtsova

(ed.): Gerpetologicheskie issledovaniya v Mongolskoi

narodnoi respublike. (In Russian).

Borkin L. Ya., and T. M. Sokolova 1989. Electro-

phoretic comparison of diploid and tetraploid toads

of the Bufo viridis group (preliminary data), pp. 36-

38. In Szczerbak, N. N. (ed.), Problems of Herpetol-

ogy. The 7th USSR Herpetol. Conf. Abstracts. Nauk-

ova Dumka Press, Kiev (In Russian).

Borkin, L. Ya., and S. L. Kuzmin. 1988. [Amphibians

of Mongolia, species account], pp. 30-197. In E. I.

Vorob'eva and I. S. Darevski (eds.), Amphibians and

Reptiles of Mongolian Peoples Republic. General

problems. Moskva, Nauka. (In Russian).

Borkin, L. Ya., A. E. Vinogradov, Z. K. Brushko and

S. L. Kuzmin. 1995. New records of tetraploid toads

(Bufo viridis group) from Alma-Ata and Taldy-Kur-

gan region, Kazakhstan. Russian Journal Herpetology

2 (2): 174- 175.

Borkin, L. Ya., V K. Eremchenko and J. M. Rozanov.

1997. Distribution of and contact zones in the diploid-

tetraploid Bufo viridis complex in Central Asia. p. 26,

In Z. Rocek and S. Hart (eds.), Herpetology '97,

Abstracts of the Third World Congress of Herpetol-

ogy 2-10 August. 1997, Prague, Czech Republic. Pra-

gue.

Borkin, L. Ya., J. M. Rozanov, and S. N. Litvinchuk.

2000. Nuclear DNA content in some green toads

(Bufo viridis complex) of Turkey and Iran. Russian

Journal of Herpetology 7 (in press).

Bretagnolle, E, E Felber, E G. Calame and P. Kupfer.

1998. La polyloidie chez les plantes. Article de

synthese. Botanica Helvetica 108 (l):5-37.

Carevsky, S. ,,1925". 1926. Notes on some batra-

chians from the Palearctic region. Ezhegodnik zoo-

logiceskogo muzeya. Leningrad 26 (l/2):74-78.

Castellano, S., C. Giacoma, T. Dujsebayeva, G. Odi-

erna and E. Balletto. 1998. Morphometrical and

acoustical comparison between diploid and tetraploid

green toads. Biological Journal of the Linnean Society

63:257-281

Castellano, S. and C. Giacoma. 1998. Stabilising and

directional female choice for male calls in the Euro-

pean green toad. Animal Behaviour 56 (2):275-287.

DMAAC - Defense Mapping Agency Aerospace cen-

ter. 1986-1993. Tactical pilot charts (TPC). Scale

1 :500.000. Sheets F5 A to D, F6 A to D, F7 A to D, G-

5 A to D, G-6 A to D, G-6 A to D. Different issues

after. 1986. St. Louis, Missouri.

Dubois, A. and J. Martens. 1977. Sur les crapauds du

groupe de Bufo viridis (Amphibiens, Anoures) de

1'Himalaya occidental (Cachmire et Ladakh). Bulletin

Societe Zoologique de France 102:459-465.

Duda, P. L., and K. Opendar. 1971. The karyotype of

Bufo sp. from Kashmir. Chromosome Information

Service 12: 18-20.

Dujsebayeva, T, S. Castellano, C. Giacoma, E. Bal-

letto, and G. Odierna. 1997. On the distribution of the

Bufo viridis complex (Anura. Bufonidae) in Southern

Kazakhstan. Asiatic Herpetological Research 7:27-

31.

Eiselt, J. and J. F. Schmidtler. 1971. Vorlaufige Mittei-

lung uber zwei neue Subspecies von Amphibia salien-

tia aus dem Iran. Annalen des Naturhistorischen

Museums Wien 75:383-385.

Eiselt, J. and J. F. Schmidtler. 1973. Froschlurche aus

dem Iran unter Beriicksichtigung ausseriranischer

Populationsgruppen. Annalen des Naturhistorischen

Museums Wien 77:181-243.

Fei Liang, Ye Chang yuan, Huang Yong zhao, Chen

Xiao nuan. 1999. Taxonomic studies on Bufo viridis

from west China (Amphibia: Anura). Zoological

Research. 20 (4): 294-300.

Fikhtman, B. S. 1989. K diagnostike zelenykh zhab

kompleksa Bufo viridis Kirgisii. Pp. 263-264. In

Shcherbak, N. N. (ed.), Voprocy gerpetologii. Nauk-

ova Dumka. (In Russian).

Frynta, D., J. Moravec, J. Cihakova, J. Sadlo, Z. Hod-

kova, M. Kaftan, P. Kodym, D. Krai, V Pitule, and L.

Shejna. 1997. Results of the Czech biological expedi-

tion to Iran. Part 1. Notes on the distribution of

amphibians and reptiles. Acta Societas Zoologica

Bohemica 61:3-17.

Futuyama, D. J. 1990. Evolutionsbiologie.

Birkhauser. Basel, Bosten, Berlin.

Gerhardt, H. C. 1994. The evolution of vocalization in

frogs and toads. Annual Review Ecology and System-

atics 25:293-324.

Golubev, M. L. 1990. New records of amphibians and

reptiles in Kazakhstan (USSR). Vestnik Zoologii

9:76-78. Kiev. (In Russian).

Gruber, U. 1981. Notes on the Herpeto fauna of Kash-

mir and Ladakh. British Journal Herpetology 6: MS-

ISO.

Hemmer, H., J. F Schmidtler and W. Bohme. 1978.

Zur Systematik zentralasiatischer Griinkroten (Bufo

viridis - Komplex) (Amphibia, Salientia. Bufonidae).

Abhandlungen Museum Tierkunde Dresden 34:349-

384.

Kashchenko, N. E 1909. Gady, sobrannye sredneazi-

atskimi ekspediciami prof. V V Sapozhnikova v.

2001

Asiatic Herpetological Research

Vol. 9, p. 93

1902-1906 i. 1908 gg. Ezhegognik Zoologicheskogo

Muzeya imp. Bd. 14:119-130.

Khan, M. S. 1997. A new toad of the genus Bufo from

the foot of Siachin Glacier, Baltistan, Northeastern

Pakistan. Pakistan Journal Zoology 29:43-48.

Kryukov, V. I., V. T. Rzhepakovskii and V. P. Astapov.

1985. Kariotipicheskaya charakteristika zhab semi

vyborok iz Tadzhikistana. Doklady Akademii Nauk

Tadzhikskoi SSR XXVIII (10): 593-595. (In Russian).

Kudryavcev. B. N.. V. V Ni, L. Ya. Borkin and L.V.

Solov'eva. 1988. Soderzhanie RNK i belka v kletkakh

diploidnykh i tetraploidnykh zhab gruppy Bufo viri-

dis. Citologia 1 1:1324-1331. (In Russian).

Kuzmin S. L. 1995. Die Amphibien RuBlands und

angrenzender Gebiete. Neue Brehm-Biicherei.

Westarp Wissenschaften, Magdeburg.

Kuzmin S. L. 1999. The amphibians of the former

Soviet Union. Sofia. Pensoft. 538 pp.

Leviton A. E., Gibbs jr. R. H., Heal E., Dawson C. E.

1985. Standards in herpetology and ichthyology: Part

I. Standard symbolic codes for institutional resource

collections in herpetology and ichthyology. Copeia

1985: 802-832.

Lumaret, R., J. L.Guillerm, J. Maillet, R. Verlaque.

1997. Plant species diversity and polyploidy in islands

of natural vegetation isolated in extensive cultivated

lands. Biodiversity and Conservation 6 (4): 591-613.

Macgregor, H. C. and J. M. Varley. 1983. Working

with animal chromosomes. A. Wiley-Interscience

Publication. Chicester, New York, Brisbane, Toronto,

Singapore.

Mazik, E. Yu., B. K. Kadyrova and A. T. Tokotosunov.

1976. Osobennosti kariotipa zelenoi zhaby (Bufo viri-

dis) v Kirgizii. Zoologiceskii Zhurnal 55:1740-1742.

(In Russian).

Meinig, H. 1995. Beitrag zur Hohenverbreitung der

Wechselkrote (Bufo viridis). Zeitschrift fur Feldherpe-

tologie (Bochum) 2:91-95.

Mercadal, I. T. 1981. Determinacion del nivel de

ploidia en ejemplares preservados del genero Cer-

atophrys. Amphibia-Reptilia 3/4:205-212.

Mertens, R. 1971a. Die Amphibien und Reptilien

West-Pakistans. 2. Nachtrag. Senckenbergiana biolog-

ica, 52:7-15.

Mertens, R. 1971b. Zur Kenntnis von Bufo luristani-

cus (Salientia, Bufonidae). Salamandra. 7:83-84.

Meyer, F. 1991. Beobachtungen an Bufo danatensis

(Pisanetz, 1978) in der Mongolei (Amphibia, Anura:

Bufonidae). Zoologische Abh. Mus. Tierkd. Dresden

46: 157-164.

Mezhzherin, S. V and E. M. Pisanets. 1990. Genet-

icheskaya struktura i proischozhdenie aziatskikh poli-

ploidnikh zhab Bufo danatensis Pisanets. 1978.

Doklady Akademii Nauk Ukrainskoi SSR, Serya B,

Geologiceskie, chimiceskie i biologiceske. Nauki 0

(8): 71-73. (In Russian).

Mezhzherin, S. V and E. M. Pisanets. 1991. Genet-

icheskaya divergenciya predstavitelei kompleksa

zelenykh zhab Bufo viridis complex fauny SSSR.

Doklady Akademii Nauk 317 (10): 222-226. (In Rus-

sian).

Mezhzherin, S. V and E. M. Pisanets. 1995a. Genetic

structure and origin of the tetraploid toad Bufo danat-

ensis Pisanets, 1978 (Amphibia, Bufonidae) from Cen-

tral Asia: Description of biochemical polymorphism

and comparision of heterozygosity levels in diploid and

tetraploid species. Genetica 31 (1): 43-53. (In Russian).

Mezhzherin, S. V and E. M. Pisanets. 1995b. Genetic

structure and origin of the tetraploid toad Bufo danat-

ensis Pisanets, 1978 (Amphibia, Bufonidae) from Cen-

tral Asia: Differentiation of geographic forms and

genetic relationship between diploid and tetraploid spe-

cies. Genetica 31 (3): 342-352. (In Russian).

Mocquard, F. 1910. Voyage de M. le Dr. Louis Villant

dans l'Asie centrale (Mission Pelliot). Bull. Mus.

Hist. Paris 16:145-153.

Morescalchi, A. (1990): Cytogenetics and the prob-

lem of Lissamphibian relationships. Pp. 1-19 In E.

Olmo (ed.). Cytogenetics of Amphibians and Reptiles.

Advances in Life Sciences. Birkhauser, Basel.

Miiller, M. J. 1996. Handbuch ausgewahlter Klimasta-

tionen der Erde. Forschungsstelle Bodenerosion der

Universitat Trier, Mertesdorf (Ruwertal) 5 (5th ed.).

Trier.

Murphy, R. W., L. A. Lowcock, S. Cheryl, I. S.

Darevsky, N. Orlov, R. D. MacCulloch, and D. E.

Upton. 1997. Flow cytometry in biodiversity surveys:

methods, utility and constraints. Amphibia-Reptilia

18:1-13.

Nikolsky, A. M. 1 896. 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. Napravlen-

nost' zhab v Tyan' -Shane, pp. 203-204. In Darevski, I.

S. (ed.) Voprocy gerpetologii. Nauka. (In Russian).

Ullerich, F.-H. 1966. Karyotyp und DNS-Gehalt von

Bufo bufo, B. viridis, B. bufo x B. viridis und B.

calamita (Amphibia, Anura). Chromosoma 18:316-

342.

Walter, H. and S.-W. Breckle. 1986. Okologie der

Erde. Vol. 4. Stuttgart. Gustav Fischer.

Walter, H. and H. Lieth. 1967. Klimadiagramm-

Weltatlas. 3rd. ed., Gustav Fischer, Jena.

Wu Min and Zhao Yajiang. 1987. A preliminary study

of the karyotype of Bufo viridis Laurenti in Xinjiang.

Zoological Research, Kunming 8 (4):339-342. (In

Chinese with English summary).

Zhao, E.-M. and K. Adler. 1993. Herpetology of

China. Oxford, Ohio, USA, SSAR:l-522pp.

Zlotin, R. I. 1997. Geography and organization of

high mountain ecosystems in the former USSR. pp.

133-159. In Wiegolaski, F E. (ed.) Ecosystems of the

world, vol. 3: Polar and alpine tundra. Elsevier,

Amsterdam.

Vol. 9, p. 96

Asiatic Herpetologieal Research

2001

Apppendix 1

2001

Asiatic Herpetological Research

Vol. 9, p. 97

Vol. 9, p. 98

Asiatic Herpetological Research

2001

2001

Asiatic Herpetological Research

Vol. 9, p. 99

Vol. 9, p. 100

Asiatic Herpetological Research

2001

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.

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,

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