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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.
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$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
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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

8.1

11.0

10.0

12.2

10.8

12.8

13.0

14.4

14.6

14.9

16.3

16.1

23.9

16.5

22.1

16.3

18.1

16.1

12.9

13.1

11.5

8.2

10.0

Chemical properties of the water.

The chemical characteristics of the water inhabited by
R. boulengen were measured and analyzed. The
results are listed in Table 2. On the whole, the water is
clear, thin, neutral and acidulous, with some contents
of Ca, Mg and other minerals.

Vertical distribution. Rana boulengeri are distrib-
uted vertically from 200 to 700 meters above sea
level, but most of them are found from 450 to 650
meters. Older, large adult frogs are mostly found
above 600 meters, while young frogs and tadpoles are
found lower.

Growth and age

When bred in artificial pools, the mean body length
and the mean body weight of the frogs just complet-
ing metamorphosis are 19.8 mm and 0.95 g respec-

tively. When feeding lasts to the end of the year, their
body weight increases to 3.07 g and their body length
to 31.2 mm. After feeding for 2 years, their body
weight grows to 20.50 g, and the body length to 58.1
mm. The growth curves of body weight and body
length are logistic (i.e. "S" shaped) (Li, et al., 1993).
In the field, it was found that the largest male was 261
g with a body length of 134 mm. The largest female
was 202 g with body length of 120 mm.

Reproductive characteristics

Environmental conditions of breeding areas. In the

wild, the adult frogs often spawn under waterfalls or
in shallow pools. The mean water area and mean

water depth of natural spawn areas are 2.16 m" and
0.35 m respectively. The water flows slowly with high
dissolved oxygen. The mean value of pH is 6.3. Gen-
erally, there are small stones, sand, ratty plant and
leaves, or humus on the bottom. There are bryophytes
and algae on the bottom as well as on the sides of the
pools.

Reproductive period. In west Hunan at 563 m above
sea level, R boulengeri begin reproduction in May and
end in August. The reproductive peak is in June and
July. Female frogs may have three clutches, the egg
number of the second and third clutches depend on
the availability of food and environmental conditions.

Temperature. When water temperature reaches
15.5°C R. boulengeri begins to spawn. The optimum
temperatures for large numbers of females to spawn
are 17.7-22.5°C. From July to August, the mean air
temperature of natural spawning sites is 21.4°C, and
the optimum humidity is above 95%.

Reproductive behavior. In middle or late March, R.
boulengeri end their hibernation and feed for a month.
Then the male and female frogs aggregate at spawn-
ing fields. Before mating, male frogs call for about 15
days as a courting period. The ovulation time of
females and fertilization times are generally from
5:00 a.m. to 8:00 a.m. When the female spawns, she
creeps slowly and her cloaca is near the walls of pool
or brooks.

Table 2. Analysis of the water inhabited by R. bou/engeri(mglL). Date of analysis: 5, January, 1990; elevation where
water was sampled: 204m above sea level.

total hardness

6.71

12.92»C

58.36

13.66

0.013

0.183

0.793

0.001

0.036

0.003

Sulfate

N(N02)

N(HN03)

N(NH4)

Water type

0.0148

0.002

0.017

0.0199

4.45

9.89

0.001

0.026

0.075

Thin acidulous

2001

Asiatic Herpetological Research

Vol. 9, p. 3

>
o

•B 7

9?
o

a>
o
o

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

1.2

1.1

1.0

0.9

0.8

ti 0.7

0)
o
o

0.6-

0.5

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

105

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

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

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

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-*"*'

£5

- ,>_

$&■*

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

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Vol. 9, p. 13

Table 1 . Summary of diagnostic characters distinguishing Hologerrhum dermal! {new species; from Panay Island)
from Hologerrhum philippinum (Gunther, 1858; from the Luzon Aggregate Island Complex).

Characters

H. philippinum

H. dermali

Pretemporals

Ventral nuchal blotches

enlarged
-, +

reduced or absent

Lateral black ventral spots

-, +

+, fused into line

Dorsal nuchal spots

12-30

7-10

Midlabial stripe

Dorsal live color

tan to reddish brown

purplish brown

Ventral live color

orange to salmon

bright yellow

Adult throat color

tan to orange (some spotted white)

purplish brown w
ocelli

Midventral dark stripe

Parietal suture

notched

unnotched

Ventral scale at 2nd dorsal
reduction

97-112*

84-97

Subcaudals (females)

42-53

57-61

* Excluding apparently aberrant counts of CAS 61554 (Table 2).

suture between pre- and postnasal; together, nasal
scales square to triangular; dorsal border of nares
formed by thick shelf of prenasal, ventral border
formed by extension of postnasal; internasals as long
as broad, slightly shorter than prefrontals, laterally
contacting both pre-and postnasals, forming a vague
right triangle with 45° face oriented anterolaterally;
loreal single, as large as or only slightly smaller than
ventral preocular, half as high as postnasal, pentago-
nal, surrounded by postnasal, lateral edge of prefron-
tal, dorsal and ventral preoculars, and second
supralabial; prefrontals longer than internasals, with
irregular lateral extensions caused by presence of con-
cave curved suture with preoculars (concave surface
oriented posterolaterally); frontal twice as long as
broad, longer than to equal to its distance to the end of
the snout, shorter than parietals; anterolateral corner
of frontal barely contacting medial point of preocular
(e.g., CAS 31553, 60951, MCZ R-25693-94, PNM
6505), or with substantive contact between frontal and
preocular squeezed off by contact between supraocu-
lar and prefrontal (e.g., CAS 60950, 61554, 62430,
134075; PNM 2120, 2490; USNM 498718, MCZ R-
25695); posteromedial point of frontal extends past
posterior margin of supraoculars for distance shorter
than or equal to length of internasals; supraoculars
very large, nearly as long as and slightly narrower

than frontal; parietals very large, laterally contacting
dorsal postocular, pretemporal, and highly enlarged
secondary temporal, together bordered posteriorly by
three (CAS 61554, 134075), four (CAS 60950,
61553; USNM 498718, MCZ R-25695) or five (CAS
60951, 62430; PNM 2490, MCZ R-25693-94) undif-
ferentiated nuchals; posterior tips of parietals extend
caudally, parietal suture forming a distinct medially
inverted V-shaped cleft, in which a single slightly
enlarged to undifferentiated nuchal (Fig. 6) lies; tem-
porals arranged in three to four irregular vertical rows
with enlarged posttemporals extending caudally
beyond posterior ends of parietals; temporal formu-
lae: (L) 2/1 + 1/1+2, (R) 1 + 1 + 1/1+2 (PNM 2490), (L,
R) 1+2+1/1+2 (CAS 60950), (R, L) 1 + 1+2+3 (MCZ
R-25695), (L) 1 + 1+2+3, (R) 1 + 1+2+2 (CAS 60951),
(L) 1+1+2+3, (R) 1 + 1+2+4 (MCZ R-25694) (L)
1 + 1 + 1/1+2, (R) 1 + 1+2+3 (CAS 61554, PNM 6505),
(L, R) 1+1+1/1+2 (CAS 62430, 61558, 134075;
USNM 318363, 498718; TNHC 60114, MCZ R-
25693, PNM 2120), (L) 1 + 1+2+3, (R) 2+1/1+2
(USNM 319037), (L); pretemporal relatively
enlarged, its length much more than half that of sec-
ondary temporal.

Orbit surrounded by supraocular, two preoculars
(dorsal larger than ventral), two postoculars, and
supralabials 3-5; supralabials eight, fifth largest;

Vol. 9, p. 14

Asiatic Herpetological Research

2001

Table 2. Scale row reduction formulae (Dowling, 1951b) variation in H. philippinum philippinum (Gunther, 1858;
from the Luzon Aggregate Island Complex) and the type series of H. derma// (new species; from Panay Island).

Specimen (sex)

Reduction 1

Reduction 2

Hologerrhum phillippinum

3+4=3(4)

CAS 6095 l(juv.) 19 17

4+5=4(5)

3+4=3(107)

17 15

3+4=3(106)

4+5=4(5)

CAS 60950 (juv.) 19 17

3+4=3(4)

3+4=3(103)

17 15

3+4=3(105)

3+4=3(7)

CAS 61553 (0 19 ' 17

4+5=4(5)

3+4=3(109)

17 15

3+4=3(112)

3+4=3(7)
CAS 61554(0 19 17

4+5=4(6)

3+4=3(93)

17 15

3+4=3(95)

+4(96)

3+4=3(100)

3+4=3(7)

CAS 62430 (juv.) 19 17

3+4=3(6)

3+4=3(101)

17 15

3+4=3(103)

3+4=3(16)

PNM 2490(0 19 17

5+6=5(5)

3+4=3(110)

17 15

4+5=4(109)

5+6=5(5)

PNM 2120(0 19 17

5+6=5(6)

3+4=3(95)

17 15

-4(99)

3+4=3(4)
PNM 2120 (juv) 19 n

3+4=3(4)

3+4=3(6)

USNM 3 19037 (0 19 17

3+4=3(5)

3+4=3(97)

17 15

-4(96)

3+4=3(105)

17-

3+4=3(109)

-15

3+4=3(5)

USNM 318363(0 19 17

3+4=3(7)

3+4=3(100)

17 15

3+4=3(99)

3+4=3(5)

USNM 498718 (m) 19 17

3+4=3(5)

3+4=3(5)
TNHC 60114(0 19 17

3+4=3(8)

-4(100)

17 15

^4(105)

3+4=3(102)

17-

-4(100)

-15

4+5=4(4)
MCZ R-25693 (0 19 17

3+4=3(4)

3+4=3(100)

17 15

3+4=3(100)

2001

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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

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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.

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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.

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Ferner, J. W., R. M. Brown, R. V. Sison, and Robert S.
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2001

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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

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

3
SQ
-

6
z

-2

O RETREAT BURROWS -Q FORAGING BURROWS

JAN

SEP

MAR MAY JUL

MONTH

Figure 3. Monthly number of foraging and retreat burrows of V. panoptes at Fog Bay.

NOV

However, the number of foraging burrows is not
correlated to burrow depths (Table 1 ). The number of
retreat burrows are negatively correlated with the min-
imum sand-surface temperatures (r = 0.463; P =
0.024) and the maximum sand temperature at 50 cm (r
= -0.616; P = 0.033; Table 2). Burrow depths were
also negatively correlated to the minimum sand sur-
face temperature (r = -0.789; P = 0.002) and the maxi-
mum sand temperature at 50 cm (r = -0.64; P = 0.033;
Table 2). The number of foraging burrows was unaf-
fected by sand surface temperature and sand tempera-
ture at 50 cm (Table 2). This equates to more and
deeper retreat burrows being dug, the cooler the sand
temperature at the surface and at 50 cm.

Discussion

The field experiments herein were the first attempt
with this modified design of burrow viewing equip-
ment and an assessment of its effectiveness is war-
ranted to justify the results. The only problem
encountered was when the original silicon glue (a
metal sealant), used to hold the camera to the jar cas-
ing, was suspected of having a corrosive effect on the
camera. The camera stopped working within a short
time and, when inspected, small white spots were
seen on the solder of the circuit board. The camera
worked properly when the spots were cleaned off and

a non-corrosive sealant (glass/window sealant) was
used. From then on, the system provided clear images,
with effective viewing of monitor lizard presence in
burrows and the depth, width and curvature of all bur-
rows. The narrower, rounder burrows of other animals
were easily identifiable at the bottom of foraging bur-
rows.

The number of foraging burrows was independent
of depth or temperature influences. However, the
greater the number of retreat burrows dug, the greater
their depth. The number and depth of retreat burrows
was also negatively correlated to minimum sand-sur-
face temperature each month. This is indicative of an
increased need for the lizards to burrow to an increas-
ing depth as the overnight temperatures cooled.

Cooler sand temperatures at 50 cm also caused an
increase in the number and depth of retreat burrows.
Since thermal diffusion in sand is slow, sand tempera-
ture at 50 cm heats and cools slowly (Packard and
Packard, 1988). The sand temperature at 50 cm at Fog
Bay was the highest overnight when the sand surface
is lowest (Guinea, 1994). Thus, monitors may be bur-
rowing more in the dry season to exploit the warmer
overnight internal sand temperatures. Maximum inter-
nal sand temperatures fall during the dry season and
the monitors therefore dig further to find even warmer
temperatures. Foraging burrows were also responsible
for increasing the number of burrows seen in the dry

Vol. 9, p. 28

Asiatic Herpetological Research

2001

Table 2. Correlation coefficients between the number of retreat and foraging burrows and burrow depth and sand
surface (Tso) minimum and maximum temperatures and sand at 50 cm depth (TS50) minimum and maximum tem-
peratures. * denotes a significant correlation.

Retreat burrows
r P

Foraging
r

burrows
P

Burrow

depth
P

Min. Tso
Max. Tso

-0.643
0.316

0.024*
0.317

0.18

0.227

0.576
0.478

-0.789
0.722

0.002*
0.008*

Min. TS50

-0.296

0.351

-0.145

0.653

-0.086

0.79

Max. TS50

-0.616

0.033*

0.026

0.937

-0.646

0.033*

season, but their depths were constant, and indepen-
dent of temperature.

Of other possible reasons for an increase in moni-
tor lizard burrowing in the dry season, egg ovipositing
is unlikely, as no eggs were ever observed within bur-
rows and V. panoptes. Burrow use to conserve water
loss has been proven important for some arid zone
monitor lizards (Green, 1972, Vernet et al., 1988).
This has not been demonstrated for tropical monitor
lizards, although there is a considerable difference in
humidity and rainfall between the wet and dry seasons
in Australia's wet-dry tropical region (Bureau of
Meteorology, 1989). Water loss in lizards increases
with decreasing ambient humidity (Hillman and Gor-
man 1977) and without physiological adaptations to
prevent water loss, burrowing to moist sand may be
utilised to prevent desiccation by V. panoptes when
ambient humidity decreases. More needs to be investi-
gated on the water economies, and uses of other water
sources, in this monitor to determine if water loss is
influential in increasing burrowing during the dry sea-
son at Fog Bay.

Acknowledgments

I thank Bill Milne, Faculty of Technology, Northern
Territory University for constructing the burrowscope.
Greg Hill and Pam Dyer provided feedback on its
design. Equipment was purchased from Oatley Elec-
tronics. Project funding was by Australian Geo-
graphic, The Queen's Trust for Young Achievers and
The Centre for Tropical Wetlands Management,
Northern Territory University.

Literature Cited

Auffenberg, W. 1983. The burrows of Varanus benga-
lensis: characteristics and use. Records of the Zoolog-
ical Survey of India. 80:375-385.

Bennett, A. F. and K. A. Nagy. 1977. Energy expendi-
ture in free ranging lizards. Ecology 58:697-700.
Blamires, S. J. 1999. Quantifying predation on sea
turtle nests by varanids at Fog Bay. MSc Thesis.
Northern Territory University.

Bureau of Meteorology. 1989. Climate of Australia.
Morphea Press, Canberra.

Capel, V. 1993. Security Systems and Intruder
Alarms. Newnes, Oxford. 267 pp.
Cowles, R. B., and Bogert, C. M. 1944. A preliminary
study of the thermal requirements of desert reptiles.
Bulletin of the American Natural History Museum
83:261-296.

Dyer, P. K., and G.J.E. Hill. 1991. A solution to the
problem of determining the occupancy status of
wedge-tailed shearwater Puffinus pacificus burrows.
Emu 91:20-25.

Dyer, P. K., and K. Aldworth.. 1998. The "burrow-
scope": modifications to burrow viewing equipment.
Emu 98:143-146.

Green, B. 1972. Water loss of the sand monitor lizard
(Varanus gouldii) in its natural environment. Ecology
53:452-457.

Guinea, M. L. 1994. A possible model to explain win-
ter nesting by the flatback turtle, Natator depressus at
Fog Bay, Northern Territory. Pp. 154- 155. In R. James
(ed.), Australian Marine Turtle Conservation Work-
shop. Queensland Department of Environment and

2001 Asiatic Herpetological Research Vol. 9, p. 29

Heritage, Australian Nature Conservation Agency.
Gold Coast.

Hansell, M. H. 1993. The ecological impact of animal
nests and burrows. Functional Ecology 7:5-12.

Hillman, S. S., and G. C. Gorman. 1977. Water loss,
desiccation tolerance and survival under desiccating
conditions in two species of Carribean Anolis. Oeco-
logia 29:105-1 16.

Packard, G. C, and M. J. Packard. 1988. The physio-
logical ecology of reptilian eggs and embryos. Pp.
523-605. In C. Gans and R. B. Huey (eds.). Biology
of the Replilia, Vol. 16. Alan Liss, New York.

Pianka, E. R. 1969. Habitat specificity, speciation and
species density in Australian desert lizards. Ecology
50:498-502.

Vernet, R.. M. Lemire. and C. Grenot 1988. Field
studies on activity and water balance of a desert mon-
itor Varanus griseus (Reptilia: Varanidae). Journal of
Arid Environments. 15:81-90.

2001

Asiatic Herpetological Research

Vol. 9, pp. 30-33

Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx aegyptius
microlepis from the United Arab Emirates

Peter L. Cunningham

P.O. Box 17258, AlAin. United Arab Emirates, email: plc@ emirates. net.ae

Abstract.- Uromastyx aegyptius microlepis are almost exclusively herbivorous and feed on a variety of plant
species with Pennisetum divisum being extensively utilized in the United Arab Emirates. The survival rate, as
determined over a one year period without any rainfall is <50%. Burrow openings are mainly aligned in a
southerly and easterly direction with the average burrow entrance size being 60x155 cm. Temperatures measured
at 30 cm down the burrow entrance are on average 6°C lower than the ambient temperature.

Key words.- Reptilia. Squamata. Agamidae, Uromastyx aegyptius microlepis. United Arab Emirates, ecology

Figure I.The Spiny-tailed lizard, Uromastyx aegyptius
microlepis.

Introduction

Uromastyx aegyptius microlepis (Spiny-tail Lizards;
Fig. 1) belong to the Family Agamidae and occur
throughout the Arabian Peninsula, Iran, Iraq and Jor-
dan (Arnold 1986. Leviton et al. 1992). They are
diurnal ground dwelling lizards that may reach sizes
of up to 60 cm and are generally yellowish-grey in
colour with an impressive spiny club-like tail. Adult
U. a. microlepis are documented as being mainly her-
bivorous, although insects form part of their diet,
while juvenile lizards are thought to be mainly insec-
tivorous (Arnold 1984, Baha El Din 1996, Brown

Figure 2. The study site and habitat of U. a. microlepis
in Abu Dhabi.

982, Highveld and Slimani 1998, Jongbloed 1997,
Manthey and Shuster 1996).

Material and Methods

Twenty mature Uromastyx aegyptius microlepis indi-
viduals in a scattered population, ±35 km northwest of
Al Ain (24°25'07"N and 55°35'01"E; Fig. 2) in the
Abu Dhabi Emirate of the United Arab Emirates
(UAE), were observed for one full day each (sunrise
to sunset). These observations took place during sum-
mer (May and June 1999) and winter (December 1999
and January 2000), respectively. I spent 480 observa-
tion hours studying these lizards. Direct observations
of plant species utilized during their feeding bouts
was conducted using an 8x40 binocular over a dis-
tance of 100m so as not to influence the feeding
behavior. Tracks were also followed once the lizards
had retreated underground to confirm sightings. Fecal
pellets were collected for later analysis. Burrow

2001

Asiatic Herpetological Research

Vol. 9, p. 31

information (orientation, burrow entrance height and
width, temperature above ground and 30 cm down
burrow) was collected from 25 active burrows once
the lizards had retreated.

Survival rate was determined for the 20 study individ-
uals by investigating their burrows for any signs of
activity in June 2000, one year after the first observa-
tions were made.

Results

Diet. Uwmastyx aegyptius microlepis utilized the fol-
lowing 10 plant species:

Class Monocotyledonae

Gramineae (Grass family): Pennisetum divisum,
Stipagrostis plumosa

Class Dicotyledonae

Amaranthaceae (Cockscomb family): Aerva jav-
anica

Asclepiadaceae (Milkweed family): Leptadenia
pyrotechnica

Boraginaceae (Borage family): Moltkiopsis cili-
ata, Heliotropium kotschyi

Chenopodiaceae (Goosefoot family): Haloxylon
salicomicum

Cucurbitaceae (Gourd family): Citrullus colocyn-
this

Leguminosae (Pea family): Tavemiera cuneifolia

Polygalaceae (Milkwort family): Polygala eri-
optera

Survival Rate. Of the 20 study individuals first
observed in May 1999, only 9 were still present in
June 2000.

Table 1 . Vegetation selected by Uromastyx aegyptius microlepis as documented by different authors from the
Arabian Peninsula. * Pulicaria glutinosa observed being utilized in other areas although not in the present study
area.

This Study
UAE- Abu Dhabi

Aerva javanica

Citrullus colocynthis

Haloxylon salicomicum
Heliotropium kotschyi

Leptadenia pyrotechnica
Moltkiopsis ciliata

Pennisetum divisum capitata

Polygala erioptera
*Pulicaria glutinosa
Stipagrostis plumosa
Tavemiera cuneifolia

Jongbloed(1997)
UAE - Sharjah

Fagonia sp.

Pennisetum divisum

Mandevile(1965)
Saudi Arabia

Aristida plumosa
Astragalus gyzensis
Citrullus colocxnthis

Horwoodia dicks on eae
Launaea capitata

Moltkiopsis ciliata
Neurada procumbens

Plantago boissieri

Zygophyllum sp.

Vol. 9, p. 32

Asiatic Herpetological Research

2001

Burrow specifics (n=25). Burrow openings were ori-
entated as follows:

North 2, West 3, South 12 (S/West: 4, South: 5, S/
East: 3) and East 8.

Mean average burrow opening height and width
was 61 cm and 155 cm (n=25), respectively.

Mean average temperature at a depth of 30 cm
down the burrow was 38°C (n=25). The ambient tem-
perature measured at 1.5 m above ground at 13h00
was 44°C.

Discussion

Diet. Ten perennial plant species are selected by Uro-
mastyx aegyptius microlepis as observed during the
study period, with the coarse desert grasses Pennise-
tum division and Stipagrostis plumosa being favored.
Jongbloed (1997) identified 3 plant species selected
by the lizards from the Sharjah area in the UAE while
Mandaville (1965) identified 8 plant species from the
stomach contents of six Uromastyx aegyptius speci-
mens from Saudi Arabia (See Table 1). Foley et al.
(1992) identified 23 plant species, mainly annuals,
selected by the same species in Israel.

As no rain had fallen during the study period no
annuals were observed and therefor the exclusion
from the diet. It would therefor be expected that more
plant species, especially flowering annuals, would be
utilized after rains. Rainfall is highly variable
(± 100mm p. a.) and unpredictable in the UAE (Bot-
tomley 1996) forcing the lizards to rely on the avail-
able perennials. According to Zari (1996 and 1998)
Uromastyx philbyi feed on a variety of annual grasses
and perennial shrubs in western Saudi Arabia. High-
field and Slimani (1998) indicate Uromastyx acan-
thinurus's preference for the family Chenopodia (fat
hen/spinach) as well as the salt tolerant Artiplex genus
and a wide variety of ephemeral plants after rains in
Morocco.

An analysis of 170 fresh adult fecal pellets (85
summer and winter, respectively) confirmed the her-
bivorous nature of U. a.microlepis as only one pellet
(<1%) contained insect remains, that of a Tennebri-
onid beetle. According to Highfield and Slimani
(1998) adult fecal pellets of U. acanthinurus in
Morocco contained 6% insect matter. Brown (1982)
also mentions U. a.microlepis feeding on locusts from
the UAE. It still has to be investigated if the lizards

would show a preference for insects during autumn
and spring although it is expected that the lack of
insects in the diet could be contributed to the fact that
most insects, Tenebrionidae at least, are crepuscular
or nocturnal (Tiger 1996).

Other items consumed by Uromastyx as observed
during the fecal analysis of the pellets include: sheep
droppings (17 pellets), date kernels (4 pellets) and
feathers (3 pellets). It is expected that above men-
tioned items are consumed erroneously by the lizards
as insect material although Brown (1982) suggests
that sheep droppings be consumed to extract undi-
gested plant material. If the sheep droppings, date
kernels and feathers were consumed as insect mate-
rial, it would suggest that the lizards would prey on
insects more often if they were more readily available.

Survival rate. Nine (45%) of the 20 study individuals
survived a period of one year from May 1999 to June
2000. Three of the 1 1 missing individuals were found
as severely malnourished carcasses. The other 8 indi-
viduals not accounted for are presumed to have suc-
cumbed below ground in their burrows, some of
which were covered by sand. No signs of emigration
were determined. No rainfall occurred during this
period in the study area. The area is also frequented
by large numbers of domestic camels and sheep,
which compete directly with the lizards for the avail-
able food. It is suggested that the lack of rainfall and
lack of annual plants as a result of this, together with
the extra pressure of domesticated stock, resulted in
the high mortality observed.

Burrow specfics. Burrow openings were mainly
aligned to the south and east with 80% of the burrow
entranced facing this way. During summer, tempera-
tures consistently reach between 40°C and 47°C
between lOhOO and 16h00. North and west facing
burrow entrances would be hotter than burrows facing
in other directions, especially during midday. During
winter the lizards would also benefit from the south-
erly orientation warming the burrows quicker and
more effectively. Wind direction during summer is
predominantly from the northwest and can cause
severe sandstorms locally known as "shamal" (El-
Ghonemy 1985). This causes lizard whose burrows
face that direction to spend more time on den clearing,
a time consuming an energetically costly affair espe-

2001

Asiatic Herpetological Research

Vol. 9, p. 33

cially during summer when plant availability and pal-
atability are low.

Burrow size is often related to the size of the resi-
dent lizard. The mean average size of 61 x 155 cm
(n=25) is indicative of an adult Uromastyx colony.
Bigger burrow openings have higher temperatures at a
depth of 30 cm, especially when facing to the north or
west.

The mean average temperature of 38°C, during the
heat of the day, at a depth of 30 cm down the burrow
is on average 6°C lower than the ambient temperature.
This gradient is imperative to the survival of the liz-
ards. During the fieldwork it was often found that the
lizards were resting in the burrows at a depth of just
over 30 cm, scurrying deeper once the thermometer
was inserted down the burrow. According to Brown
(1982), the burrows can be more than 6 feet long and
3 to 4 feet deep in the form of a spiral. Jongbloed
(1997) confirms this depth even indicating reports of
up to 12 feet deep burrows. The same author men-
tions that an excavated burrow of 5 feet in depth was 2
feet across and 1 foot high and still not at its end.
Burrows do not only serve as thermoregulatory
havens for Uromastyx aegyptius microlepis, but also
for the Desert Monitor (Varanus griseus) (Pers.obs.)
and certain Larks (Cunningham 2000, Williams et al.
1999). Williams et al. (1999), states that Uromastyx
burrows can potentially reduce Hoopoe Lark (Alae-
mon alaudipes) water loss by as much as 81% during
the hottest periods during summer. Temperatures at a
depth of 30 cm are relatively stable therefor being
exploited by many desert dwelling animals (Love-
grove and Knight-Eloff 1988). Burrows are thus an
effective way of escaping predators, the harsh desert
environment as well as controlling water loss.

Literature Cited

Arnold, E. N. 1984. Ecology of lowland lizards in the
eastern United Arab Emirates. Journal of Zoology,
London 204:329-354.

Arnold, E. N. 1986. A key and annotated checklist to
the lizards and amphisbaenians of Arabia. Fauna of
Saudi Arabia 8:385-435.

Baha El Din, S. 1996. Terrestrial reptiles of Abu
Dhabi. Pp. 124-147. In P.E. Osborne (ed.), Desert
Ecology of Abu Dhabi. Pisces Publications, Newbury,
UK.

Bottomley, N. 1996. Recent climate of Abu Dhabi.
Pp. 36-49. In: P.E. Osborne (Editor), Desert Ecology
of Abu Dhabi. Pisces Publications, Newbury, UK.

Brown, J. N. B. 1982. Spiny-tailed Agamid - Uro-
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Foley. W. J., A. Bouskila, A. Schkolnik and I. Chosh-
niak. 1992. Microbial digestion in the herbivorous liz-
ard Uromastyx aegyptius (Agamidae). Journal of
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Highveld, A. C. and T. Slimani. 1998. The Spiny-
Tailed Lizard at home: Uromastyx acanthinurus in
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tiles and amphibians, July 1998: 76-87.

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Tribulus 7.2:23-24.

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Minton. 1992. Handbook to Middle East Amphibians
and Reptiles. Scociety for the Study of Amphibians
and Reptiles, Oxford, Ohio. 252 pp.

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Arid Environment 38:371-377.

2001

Asiatic Herpetological Research

Vol. 9, pp. 34-70

The Amphibians and Reptiles of Panay Island, Philippines

John W. Ferner1, 2, Rafe M. Brown2, 3, Rogelio V. Sison4 and Robert S. Kennedy2

Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, USA. email:

fernerj@thomasmore.edu; Geier Collections and Research Center, Museum of Natural History and Science,

1301 Western Avenue, Cincinnati, Ohio 45203, USA; Section of Integrative Biology and Texas Memorial

Museum, University of Texas, Austin, Texas 78712, USA. email: rafe@mail.utexas.edu;4 Zoology Division,

National Museum of the Philippines, Executive House, P. Burgos Street, Manila, Philippines, email:

nmzoo@fastmail.i-next.net.

Abstract.- We provide species accounts for 20 amphibians (frogs and toads) and 72 reptiles (one turtle, 36 lizards,
and 35 snakes) from the central Philippine island of Panay and its associated land-bridge islets. Panay is a
moderately-sized island (123,000 km-) that currently is separated from the nearby islands of Negros, Masbate,
and Cebu by shallow seas, indicating that dry land connections once allowed free exchange of flora and fauna
between these land masses at several periods during the Pleistocene. This fact, coupled with the wealth of
knowledge on herpetological communities of Negros and Cebu, has led biologists to assume that the amphibians
and reptiles of Panay are reasonably well known. Our data suggest that this is far from true. Our recent survey
work (1989 until present) has resulted in the discovery of at least six (and probably 12-15) new species of
vertebrates, most of which appear to be high elevation rainforest Panay endemics. In this paper we note
numerous taxonomic problems that await the attention of biologists; these surely will result in an increase of the
known species from Panay. We expect that many widespread species complexes currently of uncertain taxonomic
status will eventually be recognized as Panay endemics, further bolstering the total diversity and degree of
endemism on Panay. The status of Panay herpetofauna warrants particular attention because many of the
suspected new and endemic species appear to be forest obligates. With the near complete removal of Panay 's low
elevation forests and increased pressure on its mid- and upper montane forests, we suspect that many of Panay's
endemics are threatened by extinction. Panay should be regarded as a tropical island with a unique geological
history, a spectacular compliment of habitat types, and a diverse set of species communities - all of which are
beset with a conservation crisis of deforestation and probable extinction. Panay may be a particularly appropriate
model island for large scale conservation efforts aimed at sustainable management of forest resources. However,
before informed conservation and management plans can be enacted, additional biodiversity survey work must
be conducted on Panay.

Key words.- Reptiles, amphibians, SE Asia, Philippines, Panay, Visayan Islands, conservation crisis,
herpetofauna.

Introduction

The central (or "Visayan") Philippine island of Panay
(Figs. 1 , 2), is part of a Pleistocene aggregate island
complex that includes Negros, Cebu, Guimaras, Mas-
bate, Ticao, and several small land-bridge islands
associated with each of these larger land masses.
Although amphibian and reptile species similarity
among Visayan Islands was noted much earlier (Tay-
lor, 1920, 1922a-d, 1928; Inger, 1954; Leviton,
1963c; Brown and Alcala, 1970a), it was not until
Heaney (1985, 1986) explicitly illustrated the position
of the 120 m underwater bathy metric contour in the
Philippines that the land bridge implications of chan-
nel depth became fully appreciated by biologists. It is

now understood that five major Philippine island
groups, (complexes of islands separated by shallow
channels) intermittently formed much larger land
mass amalgamations at various times during the mid-
to late-Pleistocene (Fig. 1). It is presumed that these
events allowed free exchange of fauna and flora via
land-positive connections between the otherwise iso-
lated islands of today. Each of the Philippine Pleis-
tocene aggregate island platforms (Fig. 1) are now
recognized by biogeographers as subprovinces (Tay-
lor, 1928) due to the fact that each supports highly-
celebrated suites of endemic taxa (Taylor, 1928; Dick-
inson, 1991; Heaney and Regalado, 1998; Heaney et
al., 1998; Alcala and Brown, 1998; Kennedy et al.,
2000).

2001

Asiatic Herpetological Research

Vol. 9, p. 35

Figure I.The Philippines (darkly shaded islands), with
the major Pleistocene aggregate island platforms indi-
cated by tracing of the 120 m submarine bathymetric
contour (following Heaney, 1985, 1986). Star = Manilla.

One unfortunate result of the prevailing perspec-
tive since Heaney 's (1985) landmark paper, has been
herpetologists' lack of attention to islands that are
land-bridge (once connected by dry land) to well sur-
veyed members of the same Pleistocene island com-
plex. A case in point is Panay, a central Philippine

island of 123,000 km" with several mountain peaks of
more than 2000 m in elevation (Figs. 1, 2). Perhaps
not unreasonably, numerous biologists over the past
century have assumed or suggested that a species'
presence on the well-surveyed Negros implied its
undocumented presence on Panay as well (see Inger,
1954; Leviton, 1963; Brown and Alcala, 1970, 1978,
1980, 1986; Brown and Rabor, 1967; Alcala, 1986).
Nevertheless, cases of low-level island endemism
have been documented on individual islands within
Pleistocene island complexes (Alcala, 1958, 1962;
Taylor, 1920; 1922a-d; 1923; 1925; Inger, 1954;

0-150

150-400

400-800

800-1200

1200-1600

> 1600

Figure 2. Panay Island, its position in the Philippines indicated by darkened arrow (inset). Discrete shading indi-
cates increasing elevational increments (see key), enclosed numbers indicate collection localities (see Table 1 ),
underlined bold text indicates provinces (boundaries indicated with darkened dashed lines), and darkened circles
indicate major cities.

Vol. 9, p. 36

Asiatic Herpetological Research

2001

Table 1 . Study sites and collection localities for museum specimens of amphibians and reptiles from Panay Island.
"No further data available" indicates specimens for which municipalities were the only available collection data in
museum catalogs. The term "municipality" is synonymous with "town" but municipalities are also the political
entity surrounding towns by these names. Thus, municipalities contain small surrounding villages (Barrios
[="Barangays "] which may contain smaller "Sitios"). It is reasonable to assume that most collections localized to
the municipality level were also collected in the town of the same name. Entries including "and vicinity" indicates
specimens that may have been collected in the municipal town, or they may be from smaller surrounding Baran-
gays. * Combined into a single locality due to the close proximity of collection sites at the common borders of
these municipalities.

Site

Province

Municipality

Notes

Ibajay

Barangay Bugtong-bato

Libacao

Libacao town "and vicinity"

Makato

Makato town "and vicinity"

Malinao

no specific locality data

Malay

Nogas Point

Nabas

Nabas town "and vicinity"

Tangalan

Tangalan town "and vicinity"

Malay

Borocay Isl., no specific locality data

Barbaza

Barbaza town "and vicinity"

Bugasong

Bugasong town "and vicinity"

Culasi

Barangay Alojipan (Mt. Madja-as;
PNM/CMNH site)

Tobias Fornier

Barangay Tobias, Villaflor

Valderrama

Barangay San Agustin, (Mt. Baloy and
vicinity; PNM/CMNH site)

San Remegio

Barangay Aningalan, Aningalan
mountain range

Tibiao

no specific locality data

Caluya

Caluya Isl., no specific locality data

Sibay

Sibay Isl., no specific locality data

Caluya

Semirara Isl., no specific locality data

Hamtik

Barangay Gunisang-an

Sibalom

Barangay Egaiia

San Jose

Barangay Bagumbayan

Capiz

no specific locality data

Marubusao

Burias

lloilo

no specific locality data

lloilo City

lloilo City "and vicinity"

Ajuy

no specific locality data

Calinog

Mt. Baloy and vicinity

Aklan

Antique

10.

Antique

11.

Antique

12.

Antique

13.

Antique

14.

Antique

15.

Antique

16.

Antique

17.

Antique

18.

Antique

19.*

Antique

20.

Capiz

21.

Capiz

22.

lloilo

23.

lloilo

24.

lloilo

25.

lloilo

2001

Asiatic Herpetological Research

Vol. 9, p. 37

Site

Province

Municipality

Notes

26.

lloilo

Barotac Viejo

Barangay San Francisco

27.

lloilo

Carles

no specific locality data

28.

lloilo

Carles

Sicogon Island; including Buaya area

29.

lloilo

Dingle

Bulabog-Putian National Park; PNM/
CMNH Site

30.

lloilo

Estancia

no specific locality data

31.

lloilo

Lambunao

no specific locality data

32.

lloilo

Mandurriao

no specific locality data

33.

lloilo

Pototan

Pototan town "and vicinity"

34.

lloilo

Cabatuan

no specific locality data

35.

lloilo

Estancia

Gigante North Isl., no specific locality
data

36.

lloilo

Estancia

Gigante South Isl., Gabi area

37.

lloilo

Estancia

Gigante South Isl., no specific locality
data

38.

lloilo

Calinog

E. Catalbac ("Calinog town")

39.

lloilo

Estancia

Calagna-an Isl.; Barangkalan and
vicinity

40.

lloilo

Concepcion

ca 1 km NW SBS lyang Beach Resort

41.

lloilo

Concepcion

Pan de Azucar Isl., no specific locality
data

42.

Negros Occidental

Pulupandan

Inampulagan Isl., 8 km W Munic. W.
Pulupandan, Negros Occ. Prov.

43.

lloilo,

Subprov.

Guimaras

Unknown

Guimaras Isl., no specific locality data

44.

lloilo,

Subprov.

Guimaras

Jordan

Guimaras Isl.; Jordan town "and
vicinity"

45.

lloilo,

Subprov.

Guimaras

Unknown

Guimaras Isl.; 2.1 km SW Daan
Banwa

46.

lloilo,

Subprov.

Guimaras

Nueva Valencia

Guimaras Isl.; Nueva Valencia town
"and vicinity"

47.

lloilo,

Subprov.

Guimaras

Buenavista

Guimaras Isl.; Buenavista town

48.

lloilo,

Subprov.

Guimaras

Nueva Valencia

Panobolon Isl.; Nueva Valencia town.

49.

Unknown

"Masaya" (not on available maps)

50.

Unknown

Panay Island: no Further Data

Vol. 9, p. 38

Asiatic Herpetological Research

2001

Brown and Alcala, 1961, 1970a-b, 1978, 1980,
1982a, 1986, 1994; Brown et al, 1997a-b; Brown et
al., 1999), suggesting that the two large islands war-
rant individual attention by biogeographers. Despite
the assumption that many widespread Philippine spe-
cies were present on Panay (e.g., see Alcala, 1986),
biogeographical summaries and taxonomic reviews
historically have listed only nine vouchered (based on
museum specimens) species of snakes, six geckos, six
skinks, and seven frogs (Inger, 1954; Leviton, 1963c;
Brown and Alcala, 1970, 1978, 1980) - 30% of the
herpetofauna considered in the present report.

Until very recently the higher elevation forested
regions of Panay have not been explored by biolo-
gists. At present, the results of only a few preliminary
surveys in a few high elevations localities are avail-
able (Gonzales and Kennedy, 1990, 1996; Sison et al.,
1995; Ferner et al., 1997; Brown et al., 1997a; 1999).

The purpose of this paper is to synthesize museum
records that document the diversity, endemism, status,
and distribution of the amphibians and reptiles from
Panay and to report on several recent collections
resulting from the National Museum of the Philip-
pines/Cincinnati Museum of Natural History Philip-
pine Biodiversity Inventory (1989-1994). Another
primary goal is to draw attention to recently-discov-
ered endemic species of amphibians and reptiles from
Panay and to stress the need for immediate faunal sur-
veys on this tropical island beset by a conservation
crisis of deforestation.

Material and Methods

Study sites and collection locations are presented in
Table 1 and shown in Figure 2. Additional site and
habitat information for many TNHC, PNM, CAS,
FMNH, and CMNH specimens may be found in the
museum catalogs. The PNM/CMNH Philippine
Biodiversity Inventory team conducted field studies at
sites 11 and 13 (Figs. 3-9) and various members of
the team (particularly RVS and RMB), visited other
locations (Figs. 10-14) to do more limited surveying.
We established elevational transects in a variety of
habitat types (Ruedas et al., 1994, as modified by
Brown et al. 1995a, 1996, 2000) and utilized standard
collection and specimen preservation techniques
(Simmons, 1987; Heyer et al., 1994). Detailed exami-
nation of all material was conducted by RMB and
JWF.

We follow the taxonomy of Taylor (1922c), Brown
and Alcala (1978, 1980) and Brown et al. (1995a;
1995b) for gekkonid and scincid lizards. The taxon-
omy of Inger (1954, 1996), Frost (1985), Duellman

(1993), Brown and Alcala (1994), Inger and Tan
(1996), Alcala and Brown (1998) and Emerson et al.
(2000), was consulted for amphibians and we include
(in parentheses) the unsubstantiated taxonomic
hypotheses of Dubois (1992) for reference. While no
suitable taxonomy currently is available for Philippine
agamids (see Taylor, 1922c), we consulted Musters
(1983) and McGuire and Alcala (2000) for identifica-
tion of our Draco specimens. Snake taxonomy was
based on Taylor (1922a), Leviton's "Contribution to a
Review of Philippine Snakes" series (Leviton 1961,
1962, 1963a-c, 1964a-d, 1965, 1967, 1968, 1970a-b,
1979, 1983, 1985), Leviton and Brown (1959), Inger
and Marx (1965), Malnate and Underwood (1988),
Wynn and Leviton (1993), McDiarmid et al. (1999),
and Brown et al. (1999). Scale counts on snakes were
taken using the methods of Dowling (1951a, 1951b).
Field techniques in our surveys have been described
in Ruedas et al. (1994) and Brown et al. (1995a; 1996;
2000). Museum abbreviations follow Duellman et al.
(1978) and Leviton et al. (1985).

In order to examine large scale faunal similarities
between Panay and other large island of the Philip-
pines, we assessed overall herpetofaunal simmilarities
by calculating coefficients of similarity using a simple
index (C - 2W/a+b; see Brown and Lomolino, 1998,
for review) for the amphibians and reptiles of Panay
and other islands. Our data for these calculations were
based on all available literature (above) and updated
conservative estimates of the taxonomy of Philippine
amphibians and reptiles (Brown, Crombie, Diesmos,
unpublished data).

Results

We present records for 20 amphibians (anurans; Figs.
15-25), and 72 reptiles (one turtle, 36 lizards, and 35
snakes; Figs 26-52) from Panay and its associated
land-bridge islets. The results of faunal similarity cal-
culations are presented in Figure 53. Individual spe-
cies accounts, with comments on their status, are
presented below.

2001

Asiatic Herpetological Research

Vol. 9, p. 39

Species Accounts

Amphibia

Anura

Bufonidae

Bufo marinus (Linnaeus) (Fig. 15)

Introduced on most inhabited islands of the Philip-
pines (Inger, 1954; Diesmos, 1998; Alcala and Brown,
1998), this species is particularly common in agricul-
tural areas and near human habitation. It is wide-
spread and abundant in disturbed and agricultural
areas on Panay.

Localities and specimens: (Site 1) PNM 1 144 — 47,

1228-33; (Site 11) CMNH 4958-59, PNM 1289-96,
2552-54; (Site 12) PNM 1232-33; (Site 19) CMNH
5197; (Site 23) USNM 339985-86; (Site 24) FMNH
61482-84; (Site 31) PNM 1 144-47; (Site 33) USNM
339987-88, 340062.

Microhylidae

Kaloula conjuncta negrosensis Taylor (Fig.
16)

Taylor (1920, 1922a) first collected this form on
Negros and Guimaras islands. He (Taylor, 1922a) rec-
ognized it as a full species but Inger (1954) placed K.
negrosensis in the synonymy of Luzon's Kaloula con-
juncta; no further taxonomic studies have been forth-
coming. Kaloula conjuncta negrosensis is both a
burrower and a climber (Taylor, 1920; Alcala, 1962,
1986); it has been found in the detritus of the forest
floor (750-1075 m on Negros; Alcala, 1962) and in
the leaf axils and holes of trees (Alcala 1962). They
also emerge and congregate around water that collects
in pools and ditches in the rainy season (Taylor, 1920,
1922a). Our Mt. Madja-as specimen was found in a
tree hole 10 cm from the ground (tree < 0.5 m dbh).
Specimens from near the base of Mt. Baloy were
found in breeding aggregations around water buffalo
wallows near a large river. We suspect that the
Visayan populations represent a distinct evolutionary
lineage and that they will eventually once again be
recognized as a full species once new data become
available (Brown and Diesmos, unpublished data).

Localities and specimens: (Site 8) CAS 127890;
(Site 1 1) PNM 2555; (Site 13)TNHC 56340-46; (Site
16) CAS 127510-11, 127591; (Site 18) CAS 127815;
(Site 47) CAS 124446.

Kaloula picta (Dumeril and Bibron) (Fig. 17)

This frog is found at low elevations (100-200 m) in
open areas near human habitations (Boulenger, 1882;
Alcala, 1956, 1958). It is a burrowing species that
conceals itself under leaf litter and topsoil until the
beginning of the rainy season. Choruses may contain
hundreds of individuals (Alcala, 1962; Alcala and
Brown, 1998). This appears to be the first published
account of this species on Panay.

Localities and specimens: . (Site 16) CAS 127617-
37; (Site 18) CAS 127816, 127827; (Site 23) CAS-
SU 14219-20; USNM 78079-80, 78842.

Kaloula sp.

In 1992, while conducting survey work at Barangay
Alojipan (Site 11), we heard the distinctive honking
calls of a forest species of Kaloula. Although we were
unable to locate and collect specimens, we noted that
the calls were superficially similar to Kaloula kalin-
gensis (Taylor, 1922a; single honk, delivered approxi-
mately every 15 to 20 minutes) from Luzon. Neither
Kaloula kalingensis nor the related K. kokacii (Ross
and Gonzales, 1991) have been reported from the
nearby island of Visayas, although recent field work in
the last remaining low elevation forests of Negros
(Municipality of Ayungon, Negros Oriental Prov.; C.
N. Dolino and A. C. Diesmos, personal communica-
tion) reveals the presence of a forest species (also pre-
viously unrecorded) with single honking calls there as
well. Gaulke (in press) recently has discovered a pop-
ulation of Kaloula in NW Panay that may be the same
as that heard (but not collected) previously on Panay
and Negros.

Localities and specimens: none.

Ranidae

Limnonectesc\. leytensis (Boettger) (Fig. 18)

This species is widely distributed in patches and con-
sidered common in some localities on the nearby
Negros (Alcala, 1986; Alcala and Brown, 1998).
However, it has been collected from only one locality
on Panay (Sison et al., 1995). On Negros, this species
inhabits coolor high elevation mountain streams
between 150 and 900 m (Alcala, 1962) and probably
breeds terrestrially but deposits eggs in close proxim-
ity of water (Alcala and Brown, 1956; Alcala, 1962;
Brown and Alcala, 1982b). When hatching, terrestrial
embryos of these and related species fall, are carried
by males, or are washed into water (Alcala, 1962; see
also Inger et al., 1986; Brown and Iskandar, 2000).
We find it doubtful that Visayan specimens identified

Vol. 9, p. 40

Asiatic Herpetological Research

2001

Table 2. List of known species from Panay and smaller, nearby, land-bridge islands and other islands within the
political boundaries of major Panay Island provinces. Entries include Panay species with vouchered specimens in
major museum collections (see text and species accounts for discussion of taxonomy used) and the first published
accounts by authorities utilizing museum specimens. IR = island record or first published record from within major
Panay island provinces. (1 Previously reported from Panay, nearby islets, or Visayan sea by Alcala (1986) but with-
out specific reference to specimens. 2 See also Gaulke (in press). 3 Recorded from land-bridge islets but currently
not recorded from the mainland of Panay. 4 A record from Semirara Isl., within the political boundary of Panay's
Antique Province, but land-bridge to Mindoro Isl.; not likely to be discovered on Panay in the future. 5 Apparent
major range extension, in need of verification or based on specimens with locality data probably in error.)

Bufo marinus

Kaloula conjuncta negrosensis
Kaloula picta
Kaloula sp.

Limnonectes d leytensis
Limnonectes visayanus
Occidozyga laevis
Platymantis corrugatus
Platymantis dorsalis
Platymantis negrosensis
Platymantis insula tus
Platymantis panay en sis
Platymantis sp. 1
Platymantis sp. 2
Platymantis sp. 3
Rana cancrivora cancrivora
Rana erythraea
Ranad. everetti
Rana vittigera
Po/ypedates leucomystax
Cuora amboinensis amboinensis
Bronchocela sp.
Draco spilopterus
Hydrosaurus pus tula tus
Gonocephalus sp.
Cosymbotus platyurus
Cyrtodactylus annulatus

Inger, 1954

IR (see Taylor, 1920; 1922a)1

IR2

Inger, 1954

IR2

Sisonet al., 1995

Brown and Alcala, 1970b3

Taylor, 1920; Inger, 1954; Brown et al., 1997a

Inger, 1954

Sison etal., 1995

Inger, 1954

Gaulke and Fritz, 1998

Taylor, 1922c

Musters, 1983

IR2

Sison etal., 1995

Brown and Alcala, 1978

IR2

2001

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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

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

Brown and Alcala, 1980

Brown and Alcala, 19804

IR5

IR3

Brown and Alcala, 1980

Sison etal., 1995

IR2

Ferneretal., 1997

Brown and Alcala, 1980

IR5

Brown and Alcala, 1980

Sison etal., 1995

Gaulke, 1991a, 1991b, 1992

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

Leviton, 1963c; Ingerand Marx, 1965

Gyi, 1970

Sisonetal., 1995

Leviton, 1970b

Leviton, 1963c, 1970b

Leviton, 1979

IR2

Leviton, 1965

Sisonietal., 1995

IR2

Sisonietal., 1995

Brown etal., 1999

Leviton, 1963c; Malnate and Underwood, 1988

Ross etal., 1987

Leviton 1963b, 1963c

IR1

Wynn and Leviton, 1 993

IR? (see McDiarmid et al., 1999)

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!

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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.

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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

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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

Cfl

(T,

■;

■a

ID
_Q

Ifl

■?

I/I

IT,

i—

, —

.£_

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).

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

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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).

A. ^^k

■ w

:5^£>*rr.

.^dMk

.>\

Fig. 35.

gigante (Site 37: Photo RMB)

Fig. 36. Gekko gigante eggs in cave crevice
(Site 37; Photo: RMB).

~» - «*

- > V~M^^;

Fig. 37. Gekko mindorensis (Photo: RMB)

Fig. 38. Brachymeles boulengehtaylon (Photo:
RMB).

Vol. 9, p. 62

Asiatic Herpetological Research

2001

-■• " -«p

Fig. 39. Brachymeles boulengeri taylori (Photo: Fig. 40. Brachymeles tridactylus (Photo: RMB).
RMB).

Fig. 41 . Mabuya multicarinata borealis (Photo: Fig. 42. Sphenomorphus jagori grandis (Photo:
J. McGuire). ' RMB).

Fig. 43. Tropidophorus grayi (Photo: RMB).

Fig. 44. Varanus salvator nuchalis, dorsal view
(Photo: J. McGuire).

Fig. 45. Varanus salvator nuchalis, ventral view Fig. 46. Python reticulatus (Photo: RMB).
(Photo: J. McGuire).

2001

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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).

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.

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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).

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2001

Asiatic Herpetological Research

Vol.9, pp. 77-100

A Review of the Distribution of Diploid, Triploid andTetraploid Green Toads
{Bufo viridis Complex) in Asia Including New Data from Iran and Pakistan1

Matthias Stock1, Daniel Frynta2, Wolf-Rudiger Grosse1, Claus Steinlein3 and

Michael Schmid3

Martin-Luther-Universitdt Halle-Wittenberg, Institutfiir Zoologie, Domplatz 4, D - 06099 Halle/ Saale,

Germany, e-mail: stoeck@zoologie.uni-halle.de. ' Charles University, Department of Zoology, ViniCnd 7,

128 44 Praha, Czech Republic. * Universitat Wiirzburg, Institutfiir Humangenetik, Biozentrum, Am Hubland,

D - 97074 Wiirzburg, Germany.

Abstract.- A summary of the distribution of green toads containing most of the data published after the discovery
of polyploid forms (1976) including a map, an index and a bibliography are presented and discussed. 21 Asian
type localities of hitherto described nominal green toad taxa are shown. The tetraploids are distributed in high
mountains and extremely continental regions with strong climatic shifts. Records of triploid specimens are
situated in supposed contact zones between the parapatric diploid and tetraploid toads at foothills of Middle
Asian high mountains, and triploid bisexual populations occur in the Karakoram and West-Himalayas. Habitats
of diploids appear to be restricted to lowlands and valley grounds. Polyploids seem to be more resistant. The
methods hitherto used for the determination of the ploidy level and their applicability are evaluated. We show
new data on this species complex from Iran including cytometric, karyological, bioacoustic and morphological
data and we draw taxonomic conclusions for tetraploid Bufo oblongus, diploid Bufo viridis kermanensis, and
probably diploid Bufo kavirensis. New information on the distribution of triploids in northwestern Pakistan based
on flow cytometric measurements is presented. The ploidy level of Bufo latastii is revealed to be diploid.

Key words.- Amphibia, Bufonidae, Bufo viridis complex, Bufo oblongus lectotype, Bufo viridis kermanensis,
Bufo kavirensis, Bufo latastii, Bufo pseudoraddei pseudoraddei, Bufo pseudoraddei baturae, distribution, type
localities, Asia, Iran, Pakistan, chromosomes, ploidy determination, calls, systematics

This paper is a chapter of the doctoral dissertation of M. Stock.

Introduction

Since the discovery of tetraploid forms of the Bufo
viridis complex in the northern Tien Shan (Bachmann
et al„ 1978; Mazik et al., 1976) many records of dip-
loid, triploid and tetraploid green toads have been
made, especially on the territory of the former Soviet
Union and some few in other countries. New findings
as the detection of possibly all-triploid, gonochoristic
populations of this complex in the Karakorum range of
Pakistan (Stock et al., 1998, 1999) provide arguments
that the number of investigations in this species com-
plex will increase during the next years because not
only zoogeographic and phylogenetic but also cytoge-
netic, bioacoustic and biochemical questions will be
studied. Since a fundamental summary of the data on
the territory of the former Soviet Union was done by
Borkin et al. (1986a), the development of knowledge in
this species complex has been dramatically increased.

Hence, the present paper was aimed to summarize the
recent information about the distribution which is one
of the preconditions for the understanding of the natu-
ral history of these toads. The data on chorology will
also be an important prerequisite for a revision of sys-
tematics and nomenclature in this species complex.

We discuss the hitherto applied techniques of
ploidy determination, the size of the (known) range of
polyploid green toads, any detectable correlation of
their occurrence with climatic/ecological factors, and
we try to enlarge the knowledge about the ploidy level
of taxa hitherto described from various type localities.

Because the knowledge on the ploidy of green
toads from the territory of Iran and Pakistan is very
scarce, we also present and discuss some new, sporad-
ically collected data from different sources, and first
results of a field excursion to Pakistan.

Vol. 9, p. 78

Asiatic Herpetological Research

2001

Figure 1 . Map of Middle and Central Asia and parts of the Middle East with records of diploid, triploid and tetraploid
green toads after 1976. For code numbers see appendix.

2001

Asiatic Herpetological Research

Vol. 9, p. 79

A *loi

a&£.3^ A

'^^AA

□

A ^jS A 1

■Is

D bonoaa-e-r\avus (aipioia;

A Kashgar (tetraploid)

■ Kapkan (diploid)

27.5

I 25

E 22.5

1 20

S 17.5

« 15

9 12.5

«> 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

* A

■

*.*4t£\4

■
■

* *4d

SSftf*^*4

rA-i"

a J»*~i

i^Hrv i

D™

A Kashgar (tetraploid)

H Gholaman (diploid)
1 i

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-

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

- —

•ill*-

ins ii b

12 3 4

•1 W XX AA

5 6 7 8

IX III M

9 10 11

Ii it ii a

12 3 4

II 11 |« M

5 6 7 8

* * «■ ->.

9 10 11

Figure 5. Chromosomes of diploid male Bufo viridis
kermanensis Uom Kerman. a - Giemsa-stained
metaphase, b - Giemsa-stained karyotype, c - Q-
banded metaphase, d - Q-banded karyotype.

Iran near Baghestan (Fig. 1: 13; Fig. 8F) in the moun-
tainous region north-eastern of the Central Iranian
Plateau (abstract by Stock et al.. 1998). A sample of
five toads from Birjand (Fig. 1: 13a), the type locality
(Fig. 1: E) of Bufo oblongus, was found to be tetrap-
loid (Fig. 4). Table I outlines the results of the flow
cytometric measurements in specimens from Kerman
and Birjand. Table 2 summarizes the results of eryth-
rocyte size measurements. The erythrocyte sizes were
typical of diploid and tetraploid green toads, respec-
tively, according to Stock and Grosse (1997a). The
mean DNA content of B. oblongus (17.02 pg) rather
corresponds to that of tetraploids from Kashgar (17.5
pg; Stock, 1998b) measured with the same technique,
but during another session.

The Giemsa-staining (Fig. 5a, b) revealed a diploid
karyotype of B. viridis kermanensis. These convention-
ally stained chromosomes posses the characters of
those from some other Eurasian diploid Bufo viridis
which have been examined until now (Birstein. 1981;
Bogart. 1972; Roth and Rab. 1987; Schmid. 1978:
Ullerich. 1966). The pair 6 terminally exhibits in its
long arms a secondary constriction which is caused by
the telomeric position of the nucleolus organizer
regions (NORs) in B. viridis (see also Roth and Rab,
1987; Schmid 1978). The Quinacrine-banding in
chromosomes of B. viridis kermanensis (Fig. 5c. d)
shows differently intensive fluorescing chromosomal
regions in pairs 6 to II, but distinct bright fluorescence

»» .

1 h m

1 2 3

Mi mi in*

4 5 6

XXXX MAI MM

7 8 9

1 *
UJVMM

4 5 6

■vkX «MI xxk»

Figure 6. Chromosomes of a tetraploid female Bufo
oblongus Uom Birjand. a - Giemsa-stained metaphase,
b - Giemsa-stained karyotype, c - Q-banded
metaphase, d - Q-banded karyotype.

was only found in short and long arms of pairs 6 and 8.
Birstein (1981) detected Q-bands in a B. viridis from
the Crimea in pairs 6 to II (the most distinct ones in
both arms of pairs 7, 10, and II). In a triploid taxon
(Bufo pseudoraddei balurae) of the B. viridis complex
from the Karakoram. we found Q-bands in the long
arms close to the centromere of triplet 1, in the short
arms of triplets 6 and 7 and in both arms of triplets 8 to
11 (Stock etal., 1999).

Similarly, the karyotype of the tetraploid Bufo
oblongus exhibits Q-bands in chromosomes of the
quartets 1, 6 to 11 (Fig. 6c. d). The most interesting
linding is the occurrence of Q-bands in the short arms
of only two out of the chromosomes of quartet 6. These
two chromosomes have also larger long arms than the
two remaining Q-negative chromosomes, and there-
fore, we conclude that the Q-positive pair also repre-
sents the only one which carries the telomeric NORs as
Roth and Rab (1987) found in tetraploid toads from
Kyrgyzstan. We are preparing a detailed cytogenetic
study. Like in the triplet 1 of triploids from Karakoram.
one or two chromosomes of quartet I in B. oblongus
posses Q-bands in their long arms. Furthermore, addi-
tional differences are visible in the occurrence and
position of Q-bands among the chromosomes in each
of the quartets 7. 8 and 10 (Fig. 6c. d). These observa-
tions in B. oblongus provide arguments for considering
this tetraploid form allopolyploid. The question of
autoploidy or alloploidy of polyploids from various

Vol. 9. p. 82

Asiatic Herpetological Research

2001

Table 1 . DNA content in diploid toads from Kerman (Bufo viridis kermanensis), diploid toads from Choqua Zanbil
(B. wridisssp.), tetraploid toads from Birjand [Bufo ob/ongus), and diploid B. iatastiiUom Skardu measured by flow
cytometry of DAPI stained erythrocytes with chicken nuclei (= 2.34 pg DNA/nucleus) as a standard. SD = standard
deviation.

Kerman, diploid

Zanbil,

diploid

rjand, tetraploid

Skardu,

diploid

(N = 7)

(N =

15)

. .

. ,

*,~

f .

, ,

en
Q.

■o

CO
Q.

■o

—

■d

C
0)

C
CD

■o

Q
CD

Mean 3.43

100

8.02

3.59

100

8.42

7.27

100

17.02

4.13

100

9.68

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)

CO M

-5, "
01 r-
S 5

!o N

HI C flj c

C CM

CO <N
E M

C Z
(0 —

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

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

n 25

<D
*■»

a> 20
v>

<►
<>

<>
<>

, *

A -

■■
■■

1 «.^i

A M

A
▲
D

Tetraploid. Middle and Central Asia
(Stock, 1998a, b)

Birjand

Diploid, Middle Asia (Stock, 1998a)

Lar Valley (Andren and Nilson,
1979)

Cheshmeh-ye-Sefied-Ab (Andren
and Nilson, 1979)

Fasham

Polur

Linear (Tetraploid, Middle and
Central Asia (Stock, 1998a. b))

• Linear (Diploid, Middle Asia (Stock,
1998a))

10 15 20 25

Water temperature (°C)

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

<S Q.

« 2
£ o3

- 3

CD -Q

« 9

> .§
«a
of "5

> I
CO E"

* S

? -D

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E 2

^ Q.
O ,
If) oi

!<§

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5 S

E «
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E 5.

o >-

^= x:
e/> £
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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)

o
to

•w v-/ ^

-a-

C\J

o
co

t~-

-a
c
<

ffl

O.
CO

Q.
CO

■o

•CD

■o

C
CO

•D
C
<

CO
CD

Cvl_

„ — , t

CD CO

2,
o

CD
C\i

CM
CO

C\i

o o in
o o C\J

N ^ CO

CM
CD

en
cvi

■w- CM t-

T- Y- f- ^

in in

Cvi C\i CO CD TJ-

o
co

f- cm co ■* in

§.

CM-

■o

— '

■2

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.

*■

*-*
a

■o

■
1

c
o

'to

o t

O CO
— cb

ra.E

o ^_

CD o

c
o

t
2
3P

a
a

3
£ c

■
W

22
to

a
«

3
a.

| o
E c

ro ffi
■a 3
c o-

3 S>

Li- —

IS CO
CO =
^ CO

= o

CO-,

o£

3
O
</)

B. lalastii

Kargil and Tang-

30-

8-13

1200-1300

2.5-6.1

no data

Dubois and

marg,

Pahalgam

data

[data from
spectrograms]

Martens
(1977)

B. lalaslii

Skard

u(49)

54.3

39.8

10.62

1470

4.3-6.9 [3]

4.9-69

present

(diploid)

[3]

paper

B. pseudorad-

Gilgit (48a) and

7-27

27.3-

19-

7.38-

1378-1636

1.2-4.2

2.0-6.8

Stock et al.

dei baturae

Pasu (47, P)

86.2

56.7

21.28

[25]

(1999).

(triploid)

unpubl.

(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).

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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

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, 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

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).

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Vol. 9, p. 96

Asiatic Herpetologieal Research

2001

Apppendix 1

L\£' Ploidy Topographic description Source Method

Iran. Zagros Mountains. Gholaman. 30 km W Khorram „„..„, _,„„
Abad. 33=25' N. 48° 1 2' E preSent paper

erythr. size, microdens.

Iran. SW-slope of Zagors mountains. Khuzestan
Province. Choqa Zanbil. 32°31'N. 48°32' E. 560 m
a.s.l.

present paper

low cytom.

iran. N-slope of Elburz mountains, valley 15 km S
Gorgan, approx. 1 100 m a.s.l.

Stock (1995)

chrom count, of larvae and
uveniles

Iran. S-slope of Elburz mountains. N of Theran. near
Polur. approx. 2350 m a.s.l.

Martens unpublished, present paper

calls

- Iran, S-slope of Elburz mountains, N of Theran, valley
NE of Fasham. aporox. 2540 m a.s.l.

Martens unpublished, present paper

calls

Iran, S-slope of Elburz mountains. Lar valley NE

Andren and Nilson (1979). present paper

calls

Iran. Tehran Province, N environs of Tehran, garden
of Plant Pests and Diseases Research Institute

Borkin et al. (2000)

flow cytom.

Iran. Tehran Province, approx. 70 km W of Karaj.
Karpuz-Abad village

Borkin et al. (2000)

flow cytom.

Iran, NE. frontier zone near Turkmenistan, approx. 50
km NE Gonbad-e-Kavus, 250 m a.s.l.. 37°38' N, 55°29'
E

Stock (1997a), Stock and Grosse (1997a),
Stock (1998a)

chrom. count, of larvae, adults,
erythr. size

Turkmenistan Nebil-Dagskii Rayon N-slope of Stflck ( , Qg7a) StSck wd Grosse ( 1997a), chrom. count, of larvae, adults,
Bolshoi Balkhan. approx. 15 km S of Oglanly village, stock (1998a) erythr size, calls
500 m a.s.l., 39°43' N, 54°29' E

Turkmenistan. Ashgabadskava oblast near Danata

Pisanets (1978), Borkinetal. (1986a) chrom. count

-"-

Mezhzherin and Pisanets (1990), Pisanets
(1992a)

ext morph.

4n, 3n

Pisanets (1978)

chrom. count

Borkinetal (1986a)

chrom. count.

Mezhzhenn and Pisanets (19901. Mezhzhenn
and Pisanets ( 1991 ). Mezhzherin and Pisanets
(1995a. b)

chrom. count and/or ext morph.

Ataev (1987), Pisanets (1992a)

ext morph.

1 Pisanets (1992b)

chrom count, ext morph.

Turkmenistan, Ashgabadskaya oblast, stream 2-4 km
SE of Danata village and warm spring approx. 4 km SE
of Danata. 200 m a s 1

Stock ( 1997a), Stock and Grosse (1997a),
Stock (1998a)

chrom. count, of larvae, adults,
erythr. size, calls

"South-west Kopet-Dagh", not exactly localized

Pisanets and Vasilenko (1995)

method not described

Turkmenistan, Kuruchsudon-Reserve, Kopet-Dag (not
exactly localized)

Borkinetal (1986a)

chrom. count, and/or flow cytom.

Turkmenistan, Kyzyl-Arvatskii Rayon. Kopet-Dag-
Range, valley approx. 25 km SW of the station Bami. S
of the pass. 750 m a.s.l., 38°37' N. 56°38' E

Stock (1997a). Stock and Grosse ( 1 997a),
Stock (1998a)

chrom. count of larvae, erythr. size,
microdens.

Turkmenistan. Aydere oasis, 1 Kara-Kala

Borkinetal. (1986a)

chrom count and/oi flow cytom

Turkmenistan. Adere-oasis. E Kara-Kala

Borkinetal. (1986a)

chrom. count and/or flow cytom.

4n i Turkmenistan, Kara- Kala Mezhzhenn and Pisanets (1995a. b)

chrom. count and/or ext. morph.

Turkmenistan, lake Sarykamysh Borkinetal (1986a)

chrom. count

„ ., ._ — ~ o ut. Schneider and Egiasarvan (1995),
Kazahkstan. Guryev Town. Chornaya Rechka Dujsebaveva et al. ( 1997)

calls

Turkmenistan. Ashgabadskava oblast. Bacharden Pisanets ( 1992a)

ext morph.

.". Mezhzhenn and Pisanets ( 1991 1 method not mentioned

Turkmenistan, Ashgabadskaya oblast, Bacharden Mezhzhenn and Pisanets (1995a. b) chrom. count.and/or ext. morph.

Turkmenistan. Ashgabadskaya oblast. S Bacharden,
approx. 10 km W Kelyata, 500 m a.s.l., 38°14' N,
57°31'E

Stock (1997a). Stock and Grosse (1997a),
Stock (1998a)

chrom. count, of larvae, erythr. size,
microdens.

Turkmenistan. Ashgabadskaya oblast near Ashgabad

Borkinetal. (1986a)

chrom. count

Mezhzherin and Pisanets ( 1 990), Mezhzherin
and Pisanets (1991 1

ext. morph.

Turkmenistan, Ashgabad, village Kodzh Mezhzhenn and Pisanets (1991)

method not mentioned

Mezhzhenn and Pisanets (1995a. b)

chrom count and/or ext morph.

Turkmenistan, Ashgabadskaya oblast near Ashgabad ! Pisanets and Vasilenko (1995)

method not described

Pisanets (1991)

chrom. count

2n Turkmenistan, Ashgabadskava oblast, Ashgabad Stock (1995)

chrom. count, of larvae

4n Turkmenistan, Ashgabad Roth and Rab (1 9861

chrom. count.

Turkmenistan, village Kuruchaudan Pisanets (1987)

clutch

12a

Iran, 20 km WNW Kapkan. Khorasan, 37°22' N,
58°32'E, 1670 m a.s.l.

present paper

erythr. size, microdens.

Iran. 10 km NE Baghestan. Khorasan. N of Kuh-e-
Kalat 34°09' N. 58°25' E. 1900 m a.s.l.

Stock et al. (1998a). present paper

chrom. count, erythr. size,
microdens. and flow cytom

13a

Iran, Khorasan, Birjand, 32°33' N, 59° 10' N, about
1500 m a.s.l.

present paper

chrom. count, flow cytom.

13b

Iran. Kerman Province, Kerman, 30° 18' N, 57°05' E,
1860 m a.s.l.

present paper

chrom. count, flow cytom.

13c

Iran. Kerman Province, Bahr-e-Aseman Mountains,
Sarduiyeh village in Sarduiyeh District 2500 m a.s.I.

Borkin et al. (2000)

flow cytom.

, Turkmenistan, lake Shach-Senem (not exactly
zn [localized, SE of 9)

Borkinetal. (1986a)

chrom. count.and/or flow cytom

Turkmenistan, Ashgabadskaya oblast near Iolotan

Mezhzherin and Pisanets (1990), Mezhzherin
and Pisanets (1991)

ext. morph.

Badchyz-Reserve, AkarChechme Pisanets (1978)

chrom. count

Borkin et al. (1986a)

chrom. count

_ " .

Mezhzhenn and Pisanets (1995a, b)

chrom. count.and/or ext. morph.

Uzbekistan. Buchara

Roth and Rab (1986)

chrom. count

2001

Asiatic Herpetological Research

Vol. 9, p. 97

Uzbekistan. Samarkand Borkrn et al (1986a) chrom cuunl

Uzbekistan. Dzhisakskaya oblast Ravon Farislt Stock (1997a), Stock and Grossed 997a), chrom. count, of larvae, adults,

Nurauu-Reserve N-slope r of Nuratau Range, 900 - | „ k , 9Qg ^ ■ ca,K
1600 ma.s.l.. 40 35 N. 66 30 E | '

Uzbekistan, Dzhisakskaya oblast. Rayon Farish, S Stock (1997a), Stock and Grosse (1997a), chrom. count, of larvae, erythr. size,
bank of Aidar-Kul near of Nuratau- Range, 300 m as. 1. Stock (1998a) microdens., calls

Tadzhikistan. 60 km E of mouth of Janob into
Zeravshan river, right bank of Zeravshan river, near
Pasucax

Kryukovetal. (1985)

chrom. count.

Tadzhikistan, Gissar- Range. Ziddi. 3000 m a.sl.

Borkinetal. (1986a)

chrom count

-In

Tadzhikistan. Ramit-gorge, Chuhgaram

Borkinetal. (1986a)

chrom count

4n Tadzhikistan. Ramit-gorge. Chuligaram Mezhzhenn and Pisanets ( 1995a, b)

chrom. count and/or ext morph

4n ■"- Pisanets (1992b)

chrom. count., ext. morph

, Tadzhikistan, 1 5 km SW of the mouth of Sardan-
n Miena and Sorbo. gorge of Kafinugan, near Javroz

Kryukovetal. (1985)

chrom count.

2n and 4n

Tadzhikistan, Dushanbe

Roth and Rab (1986)

chrom. count.

4n Tadzhikistan, N Dushanbe

Kryukovetal. (1985)

chrom. count

Uzbekistan, Gissar- Range, 20 km of mouth of
Tupalang-Darya. 800 m a si

Borkinetal. (1986a)

chrom. count.

Tadzhikistan, 90 km SW Dushanbe. Kafirnigan-valley

Borkinetal. (1986a)

chrom. count.

1 adzhikistan, 90 km S Dushanbe. Kafirnigan-valley.
near lsambai

Pisanets (1992b)

chrom. count., ext. morph.

4n Tadzhikistan. S Pamir, near Lyangar

Mezhzhenn and Pisanets (1990)

ext. morph.

4n Tadzhikistan. S Pamir, near Lyangar

Mezhzhenn and Pisanets (1991)

method not mentioned

4n 1 -"-1800m a.sl Pisanets 1 1992b)

chrom. count . ext. morph

27a

4n Tadzhikistan, near Faizabad Mezhzhenn and Pisanets (1991)

method not mentioned

4n I ad/hikistan. near Faizabad Pisanets (1991)

chrom count

Tadzhikistan. near Faizabad

Mezhzhenn and Pisanets ( 1995a. b)

chrom count. and/or ext. morph.

- " - .localization using the information:„I20 km N of
Dushanbe" in:

Pisanets (1992a)

ext. morph.

. " _

Pisanets 1 1992b)

chrom. count., ext. morph.

ladzhikistan. Vakh valley, environs of Obi-Garm Mezhzhenn and Pisanets 1 1991 1

method not mentioned

Tadzhikistan. Obi-Gann. 2300 m a.s.l. Pisanets (1992a)

ext. morph.

ladzhikistan. Obi-Garm Mezhzhenn and Pisanets (1995a, b)

chrom. count. and/or ext. morph

. ".

Pisanets (1992b)

chrom count . ext. morph

27b

fadzhikistan. mouth of Komarou-river. 2000 ma.s.l.
(not exactly localized)

Borkinetal. (1986a)

chrom. count.

Tadzhikistan. 10-12 km SW of the mouth of Vakhsh
and Javansu. near Kurgan-Tiube

Kryukovetal. (1985)

chrom count.

Tadzhikistan. 1 0- 1 2 km W of the mouth of Vakhsh and
Javansu. left of Vachsh, Kuibyshevskii Rayon

Kryukovetal. (1985)

chrom. count.

Tadzhikistan, 50 km NE of mouth of Vakhsh into
Pvandzh. near Dusti

Kryukovetal. (1985)

chrom. count.

2n Tadzhikistan. Chirik

Borkinetal. (1986a)

chrom. count.

environs of Chaartuz. Chirik

Pisanets (1991 i

chrom. count.

Tadzhikistan, Beshkenlskre peski, valley of Kafinugan
nver. environs of Chaartuz

Mezhzhenn and Pisanets ( 1991)

method not mentioned

-"-

Pisanets (1992a)

ext morph.

Tadzhikistan, S of Aktau-Range, 80 km S Dushanbe

Kryukovetal. (1985)

chrom. count

Uzbekistan, Tashkent

Borkinetal. (1986a)

chrom count and/or flow cy torn.

. " .

Kudryavcev et al. (1988)

cytophotometry

. " .

Roth and Rab (1986)

chrom. count

Uzbekistan, Tashkent, 450 m as.l., 41°16' N. 69°13' E

Stock [1997a), Stock and Grosse (1997a)
Stock (1998a]

chrom. count of larvae, adults,
erythr. size, calls

31a

2n Kazakhstan, environs of Chimkent

Mezhzherin and Pisanets (1995a, b)

chrom count and/or ext. morph.

4n Kazakhstan, environs of Chimkent Mezhzhenn and Pisanets ( 1991)

method not mentioned

31b

Kazakhstan. Kyzyl-kum desert, 50 km SW of Bairkurn DuJ,ehayeva e, al. ( 1997), Castellanoet al.
village, Chimkent region. 250 ma.s.l. 67°25 N, 42 02 ,„-Ug '

chrom count

31c

Kazakhstan, Chimkent region. Jagbagly village, 1 100
mas.l..70i32,N,42°25'E

Dujsebayeva et al. (1997), Castellano et al.
(1998)

chrom. count.

Uzbekistan. 80 km E Tashkent, entry of Chatkal-
Reserve. 5 km SE Burchmulla, 900 m a.s.l., 41°35' N,
70°07' E

Stock ( 1 997a), Stock and Grosse (1997a),
Stock (1998a)

chrom. count. and/or erythr. size,
calls

4n Uzbekistan. Kuraminsky-Range, 3000 ma.s.l.

Borkin et al. (1986a)

chrom. count

Tadzhikistan. S Pamir, near Khorog

Mezhzherin and Pisanets (1990)

ext morph.

Tadzhikistan. Gorno-Badakhshan. S Pamir, near
Khorog

Mezhzherin and Pisanets (1991)

method not mentioned

- " - Mezhzhenn and Pisanets (1995a. b)

chrom count and/or ext morph

-"- Pisanets (1992a)

ext morph

ladzhikistan, S Pamir, near Ishkashim Mezhzhenn and Pisanets ( 1990i

ext morph.

, Tadzhikistan, Gorno-Badakhshan, S Pamir, near

Ishkashim

Mezhzherin and Pisanets (1991)

method not mentioned

4n Tadzhikistan. S Pamir, near Ishkashim Me/hzhenn and Pisanets ( 1995a, b)

chrom. countanoVor ext. morpjt
chrom count

2n | Kazakhstan, Dzhambul Roth and Rab ( 1 986)

36a

(outside the map ) Kazakhstan. Tengis Lake. 1 2 km W Dieterich leg., examined by Stock,
ol Abaya village, about 50 40 N 69 4(1 1 unpublished

flow cytometry (one specimen)

Kazakhstan, desert near Burubaital, S Balchash lake
region

Egembergdieva ( 1983) cited by Borkin et al.
(1986a)

unknown, original paper not
available

4n ! Kazakhstan, desert near Burubaital,SBalchash lake Borkinetal (1995) flow cytometry

region

Vol. 9, p. 98

Asiatic Herpetological Research

2001

37a

Kazakhstan. Karaoi village. 350 m as 1 , 74 47' N. Dujsebayeva et al ( 1997). Castellano et al.
45°54'E (1998)

:hrom. count., calls

37b

Kazakhstan. Zhidely Channel. Hi River Delta, 370 m Dujsebayeva et al. (1997), Castellano et al.
a.s.i.,75°12'N.45°18'E (1998)

:hrom. count., calls

_,. „ ,„„ , Mazik. Kadyrova and Toktosunov (1976) and
Kyrgyzstan, Chu-valley. 650 m a.s.l. C|ted bv Borkin et al . ( 1 9Sh., 1

:hrom. count.

Kyrgyzstan, Chu-valley Toktosunov and Mazik (1977) method not mentioned

Kvfvzstan Tulek Castellano et al (1998) chrom count, calls

Kyrgyzstan, Tyulyok, not exactly localized | Borkin et al. (1986a) chrom. count.and/or flow cytom.

Kyrgyzstan, Frunze (Bishkek) Bachmann et al. (1978) cytophotometry

Borkin et al 1 1986a)

chrom count, and/or flow cytom.

3n
4n

Kyrgyzstan. S Bishkek Kuzmin(1995 187)
Kyrgyzstan. Frunze (Bishkek) Borkin et al. (1986a)

chrom. count.

Kvrgvzstan. environs of Frunze (Bishkek) Mezhzhenn and Pisanets ( 1991 ) jmefhod not mentioned

Bachmann et al. (1978) cytophotometrv

Kyrgyzstan. Bishkek (Frunze)

Stock ( 1 997a ), Stock and Grosse ( 1 997a)

chrom. count.and/or erythr. size

Kyrgyzstan. Kyrgyzskn Khrebet, 1200 m a.s.l. and
2500 m a.s.l.. not exactly localized

Fikhtman(1989)

method not mentioned

2n, 3n, 4n

Kyrgyzstan. Kok-jar, 25 km S of Bishkek, approx.
1300 m a.s.l., artificial bassin about 6 km from the
villare

Castellano etal. (1998)

chrom. count., calls

Kyrgyzstan, Toktogul-valley. 900 m a.s.l.

Toktosunov (1984) and cited by Borkin et al.

(1986a)

chrom. count.

2n?

Kyrgyzstan, SW slope of Fergana- Range, near
Arslanbob, 1 800 m a.s.1.

Pisanets and Shcherbak ( 1979)

method not described

.".

Stock (unpublished)

chrom. count, (one juvenile animal)

_ " .

Borkin et al. (2000)

method nut mentioned

Kyrgyzstan, Arpa-valley. 3500 m a.s.l.

Toktosunov (1984) and cited by Borkin et al.
(1986a)

chrom. count.

China, Kashgar, 39°29'N, 76°02'E. 1350 m a.s.l.

Stock (1998b)

chrom. counl . flow cytom . and/or
ervlhr size, calls

China, E-Pamir. Taxkurgan, 37°47'N, 75° WE, 3350 m
a.s.l.

Stock et al. (1998a), Stock (1998b)

chrom. count., calls

Tadzhikistan, Central Pamir, lake Jashilkul. 3734 m

Toimastov (1989)

ext. morph.

- " - (also 3n, 2n ?) Mezhzhenn and Pisanets ( 1 990)

ext. morph.

_ " - Mezhzhenn and Pisanets (1991)

method not mentioned

- " - (also 3n, 2n ?) Pisanets (1992a)

est morph

- " - (also 3n, 2n ?) Pisanets (1992b) chrom. count., ext. morph.

- " - (also 3n, 2n ?)

Mezhzhenn and Pisanets ( 1 995a, b) chrom. count.and/or ext morph.

Pakistan. Karakoram, Sust, Hunza-Valley, 36°46iN,
74°50'E, 2950 m a.s.l.

Stock etal. (1998. 1999)

chrom. count., flow cytom., and/or
erythr. size, calls

, Pakistan. Karakoram. Pasu, Hunza- Valley. 36°30'N,
ia 74°52'E. 2600-2800 m a.s.l.

Stock et al. ( 1 998, 1 999 ). present paper

chrom. count., flow cytom., and/or
erythr. size, calls

47a

Pakistan. Northern Areas, Hunza-Valley, river bank.
Karimabad near Ganesh, 36°I8'N, 74°4FE, 2060 m
a.s.l.

present paper

flow cytom.

Pakistan, W-Himalayas, Upper Swat valley. Kulalai,
35° 1 9'N, 72°36'E, 1 750 ma.s.1.

Stock etal. (1998, 1999)

chrom. count, only one specimen),
and/or erythr. size, calls

48a

Pakistan, Northern Areas, Gilgit, 35°54'N, 74°24'E,
1550 m a.s.l.

Stock etal. (1998, 1999)

chrom. count., flow cytom.. and/or
erythr. size, calls

48b

Pakistan, Northern Areas, Gupis, 36°14'N, 73°27'E,
opposite Yasin valley, 2160 m a.s.l.

present paper

flow cytom.

48c

Pakistan, Shandur Pass, Lake, border of NWFrontier
Prov. and Northern Arears, 3720 m a.s.l.

present paper

flow cytom.

48d

Pakistan, NWFrontier Prov.. Hindu-Kush, Buni,
approx. 36°20'N, 72°20'E, approx. 1900 m a.s.l.

present paper

flow cytom.

48e

Pakistan, NWFrontier Prov., Hindu-Kush, Chitral City,
35°53'N. 71°47'E, 1480 m a.s.1.

present paper

flow cytom.

48f

3n?

Afghanistan. Kabul ?. 34°31iN, 69°12'E, sample not
surely localized for Kabul

Bachmann et al. (1978): "36% more DNA
than diploid B. viridis ", Hemmer et al. (1978)

microdens.

Pakistan, Northern Areas (Baltistan). Himalaya,
Satpara river SW of Skardu, 35"17- N, 75°37'E. 2300
m a.s.1., see also M and N

present paper

flow cytom., chrom. count

50(?)

China, Xinjang; Hotan; localization in the present map
not sure

Whu Mm and Zhao Yajiang (1987)

chrom. count.

Kyrgyzstan. Naryn

Stock (unpublished)

chrom. count.

Kyrgyzstan. Kara-Kudzhur-valley

Toktosunov (1984) and cited by Borkin et al.
(1986a)

chrom. count.

52-57

Kyrgyzstan. whole environs of Issyk-Kul. 1670 m a.s.l

Mazik etal. (1976)

chrom. count

.-.

Toktosunov (1984) and cited by Borkin et al.

(1986a)

chrom. count.

Roth and Rab ( 1 987b) , chrom. count.

Borkin ( 1 989) partly chrom count.

.".

Mezhzhenn and Pisanets I !99Sa. b)
Fikhtman(1989)

chrom. count.and/or ext. morph.

.".

method not mentioned

Kyrgyzstan, Chu-valley. approx. 20 km W Rybache,

Stock (1997a), Stock and Grosse (1997a)

erythr. size

4n Kyrgyzstan. SW bank of Issyk-kul

Me/hzhenn and Pisanets ( 1991 )

method not mentioned

2001

Asiatic Herpetological Research

Vol. 9, p. 99

,, . Kyrgyzstan. Issyk-kul. N-bank near Sary-Kamysh. Stock ( 1997a), Stock and Grosse (1997a). —al- .:„ —n.
34 n 1670 m a.s.l.. 42°29'N,76°20'E ISlock (1998a) eryuir. size, cans

Kvrgvzstan. Issyk-kul [Castellano et al (1998)

chrom. count., calls

Kyrgyzstan, Issyk-kul. S-bank nearTamga, 1670 m
a.s.l

Stock (1997a). Stock and Grosse (1997a).
Stock (1998a)

chrom. count, of adults, erythr. size,
calls

Kyrgyzstan. Rayon Tyub. village I-runze

Borkin (1989)

partly chrom count

Kyrgyzstan. NO-bank of Issyk-Kul. Rayon Tyub. near
Kuturga

Borkin (1989)

partly chrom. count.

Kyrgyzstan. Issyk-kul. N-bank near Chon-Oryuktu.
1670 m a.s.l.

Stock (1997a). Stock and Grosse (1997a)

chrom. count of adults, erythr. size

Kyrgyzstan. Kemin-valley. 2500 m a.s.l.

Toktosunov (1984) and cited by Borkin et al.
(1986a)

chrom count.

Kazakhstan. Almaty

Birstein (1981). Pisanets ( 1 99 1 )

chrom. count

Kazakhstan. Almaty ] Mezhzhenn and Pisanets ( 1 99 1 )

method not mentioned

Kazakhstan, Almaty Mezhzherin and Pisanets ( 1995a. b)

chrom. count. and/or ext morph.

Kazakhstan. Almaty Borkin et al. (1995)

flow cytometry

Kazakhstan. Almaty. 900 m a.s.l., 76°55' N, 43° 15' E

Dujsebayeva et al. (1997). Castellano et al.
(1998)

chrom. count. , calls

59a

Kazakhstan, Kopa, 20 km S of Kopa station. 75"47' N.
4< 25' E

Dujsebayeva et al. (1997), Castellano et al
(1998)

chrom. count, calls

59b

Big Almaty Lake. Zailiskii Alatau Range. 2300 m
as 1 , 77°N. 43°04' E

Dujsebayeva et al (1997), Castellano et al
(1998)

chrom. count., calls

4n Kazakhstan. Kapchagav. Ili-river

Borkin etal. (1986a). Borkin et al (1995)

chrom. count and/or flow cytom.

60a

. Kazakhstan. Bashn, 1 km S Altyn-Emel Mountain
Range 44' 10' N. 78 "45' E

Borkin etal. (1995)

flow cytometry

60b

4n S foothills of the Koyandytau Mountain Range

Borkin etal. (1995)

How cytometry

, Ayan-Saz Point. Borokhudzir river valley, between
Kovandvtau and Dsunganan Alatau Mountains

Borkin etal. (1995)

flow cytometry

-, ^ 4n Kazakhstan, valley of river Chingzhal, basin of lake
' ' Alakol. 6- 1 0 km near Andreevka

Golubev(1990)

method not mentioned

4n Kazakhstan. Andreevka

Mezhzhenn and Pisanets (1995a. b)

chrom count and/or ext. morph.

4m Kazakhstan. Taldy-Kurganskaya oblast

Pisanets 1 1992b)

chrom. count., ext. morph.

4n Kazakhstan. Ucharal Mezhzhenn and Pisanets (1995a, b)

chrom. count and/or ext. morph.

4n China. Xinjang; Wusu (Usu ?)

Whu Mm and Zhao Yajiang 1 1987)

chrom. count.

China. E-Tian Shan (E-Narat Shan), Kunas 43°14'N,
84 40'E, :i45M,

Stock (1998b)

chrom. count., calls

Kazakhstan. Aksnr-Farm. Zaysan-lake

Borkin etal. (1986a)

chrom count

SW-part of Zaysan-gorge, 1 2 km NW of settlement
Maikapchagai near Aksiir-Farm

Shcherbak and Golubev ( 1981)

method not mentioned

China, Xinjang, Wulumuqui (Urumqi)

Whu Min and Zhao Yajiang (1987)

chrom. count

Mongolia, river Ded-Nariin-Gol, S-slope of Mongolian
Altai. 2000 m a.s.l., 1 2 of the detailed map in:

Borkin and Kuzmin (1988)

mostly chrom count.

Mongolia, oasis Chug-Bulag, Bulgan-Somon, Chovd-
Aimak

Borkin etal. (1986b)

chrom. count.

corresponds to 1. 2 and 3 of the detailed map in

Borkin and Kuzmin (1988)

chrom. count.

Mongolia, Ml kin W of settlement Bulgan, ( :hovd
Aimak. nver Bulgan-Gol

Meyer (1991)

ext. morph.

Mongolia, spring Chujten-Bulak. S-slope of Mongolian
Altai, 2000 m a.s.l., spring Bayan-Mod, 15 km N of
spring Icher-Tol, 1600 m spring Icher-Tol, 83 km NW
of settlement Bulgan, 1 600 m a.s.l., spring Chavchig-
Us or Chavchig-Bulag-spring Shara-Bulag

Borkin and Kuzmin (1988)

partly chrom. count.

Mongolia, river Uench-Gol, 46°n.B., 92°w. L., 1350 m
a.s.l. ( 1 1 of the detailed map in:

Borkin and Kuzmin (1988)

partly chrom. count.

Mongolia, spring Jaman-Usny-Bulag or Jaman-Us. 30
km E of settlement Uench, 1 800 m a.s.l., 1 2 of the
detailed map in:

Borkin and Kuzmin (1988)

partly chrom. count.

Mongolia, oasis Ushigiin-Us, Dsungarian Gobi, 90 km
SW of settlement Bulgan. 9 of the detailed map in:

Borkin and Kuzmin ( 1988), Orlova and
Uteshev(l986)

partly chrom. count

4n'

Mongolia, spring Domdzhigiin-Us. Dsunganan Gobi.
85 km S of settlement Bulgan. 10 of the detailed map
in:

Borkin and Kuzmin ( 1 988), Orlova and
Uteshev(1986)

partly chrom. count

China, Xinjang; Hami prefecture, 1.' km M
Koumenzi. 2090 m a.s.l.

Zhao and Adler( 1993)

ext morph.

China. Xinjang. Hami Whu Min and Zhao Yajiang (1987)

chrom. count

Iran, l.unstan. Shah Bazan. km 324 of the Transiranian ^h™*' ' n o^m'' ^^q,,^- „
•, .„ , , , e D , , , ,. ■ Schmidt er (1969 , Mertens 1971b), Eiselt
railway, type locality of Bufo {surdus ) lunsmmcus a[)d Schmidtler ( , 973 . see als0 0 ^ s

no ploidy determination

2n?

Iran. NW pan of Central Iranian Plateau. Cheshmeh-ye
Sefied-Ab, type locality of Bufo ku\ir?n.sis

AndnSn and Nilson (1979), Stock (1998a):
diploid'?, present pajper

calls

Iran, Kerman, type locality of Bufo viridis
kermunensi*

Eiselt and Schmidtler ( 1 97 1 ), Hemmer et al.
(1978), present paper

chrom. count, flow cytometry

Turkmenistan, Achgabadskaya oblast, Danata village,
type locality of Bufo danaiensis

Pisanets (1978). Borkin and Kuzmin (1988),
see also number 5

chrom count

E-Iran, Ssaman Shakhi mountains near Birjand, type
locality of Bufo oblongus

Nikolsky (1896 1897), Eiselt and Schmidtler
(1973). Roth (1986). Borkin and Kuzmin
( 1 988). present paper

chrom. count, flow cytometry

Hemmeret al. (1978). Pisanets and Shcherbak
Tadzhikistan. Dushanbe, type locality of Bufo i in dis (1979), Roth (1986), Borkin etal. (1986b), no ploidy determination of the type
turanensis Kuzmin (1995, 1999), Mezhzherin and series

Pisanets (1995a. b). see also number 23

Vol. 9, p. 100

Asiatic Herpetological Research

2001

c -, Tadzhikistan, SW-part Beshkent desen, near Shaartus. PisanetM 1991 1, Pisanets el al (1996), •aasA
type localitv of Bufo shaarmsiensis Kuzmin (1999). see also number 29

„ .. . ~ .. , , ... rn_r / • -j- \ Eiselt and Schmidller ( 1 973 ). Borkin and
Pakistan, Pishin, type locality of Bufo (vmdts ) Kuzmn ,98g S(6ck ■ a, 999 Borkl[) e(

zugmayen 'al.(2000)

no ploidy determination

3n and/or
4n(?)

Pisanets and Shcherbak (1979: diploid
Kyrgyzstan. Arslanbob. type locality of Bufo viridis without determination), Borkin and Kuzmin
asiomontanus (1988), Borkin et al. (1997), Kuzmin (1999),

Borkin et al. (2000)

chrom. count, (only one triploid
specimen tested by Stock, unpubl.l.
calls

„ „ _. . . , „ . ,„, Kashchenko(1909). Stock (1997a), Stock and
Kyrgyzstan, Kokmoinok. terra typica of Bufo Qws%e , 99? R mjn , 999 see ^ chrQm

(vind«)iim<:otor number 52 to 57

China. Kashgar. one of three type localities of Bnjo Mocquard (1910). Borkin and Kuzmin (1988),
nouenei sensu Mocquard; type locality of the B Stock ( 1 998b), additional papers are cited in
nouettei lectotvDe the last one

chrom. count., calls, flow cytometry

Pakistan, Shinu village near Siachen glacier, type
locality of Bufo siachinensis

Khan ( 1 997 ). Baig ( 1 998 ), Stock et al. ( 1 998,
1999): junior synonym of B. latastii. see N
and 49

chrom. count., flow cytometry

N-India. Ladakh. a locality was not exactly described,
terra typica of Bufo latastii

Boulenger (1882: 295), Dubois and Martens
(1977), Hemmer et al. (1978), Pisanets and
Shcherbak (1979), Gruber (1981 ), Borkin et
al. (1986a). Roth (1986). Borkin and Kuzmin
(1988), Khan (1997). Baig (1998), Kuzmin
( 1999), Stock et al. (1999). see M and 49

chrom. count, flow cytometry

Pakistan, Swat valley, Mingora province, terra typica
of Bufo (viridis ) pseudoraddei pseudoraddei

Mertens (1971a), Pisanets and Shcherbak
(1979). Roth (1986), Borkin and Kuzmin
(1988), Baig (1998), Kuzmin (1999), Stock et
al. (1999), additional papers are cited in the
last one

chrom. count, (only one specimen).
calls

N-Pakistan, Karakoram, type locality of Bufo
pseudoraddei baturae

Stock etal. (1999)

chrom. count., calls, flow cytometry

SE-Iran, Bazman, Tamin in Sarhad, Duz-Ab. Ziaret in
Sarhad. type locality of Bufo persicus

Nikolsky ( 1 899). Carvsky ("1925". 1 926):
"synonym of B surdus ", Schmidt (1955),
Schmidtler and Schmidtler ( 1969). Eiselt and
Schmidtler (1973), see also R and S

no ploidy determination

Belutchistan (W-Pakistan?), not exactly localized, terra f^X'a'n^Schrnidt'ler ( 1969?. Eisllt^d
typica of Bufo surdus Schmidtler ( 1 973). see also Q and S

no ploidy determination

SW-lran. Mehkuh. 70 km S of Shiraz, type locality of Schmidtler and Schmidtler (1969), Eiselt and
Bufo surdus annulatus Schmidtler ( 1 973)

no ploidy determination

China, type locality of Bufo ssp. taxtorensis

Fei et al. (1999), Stock (1998b)

no ploidy determination in ssp.
description, Stock (1998b): chrom
count, (only one specimen), calls

China, Kok-Far (= Kokyar). type locality of the
depicted specimen of Bufo viridis var. pewzowi , three
other type localities for the series

Bedriaga (1898: 61 and Fig. 2. plate I),
Hemmer et al ( 1 978). Borkin and Kuzmin
(1988), Fei etal. (1999)

no ploidy determination

China. Tschik-Tym (= Qiktim). Turfan, type locality of
Bufo viridis var. grum-grzinuiiloi ; "Turfan": one of
14 type localities of Bufo viridis var. strauchi

Bedriaga (1898: 61). Hemmer etal. (1978).
Borkin and Kuzmin ( 1 988 )

no ploidy determination

2001

Asiatic Herpetological Research

Vol.9, pp. 101— 106|

First Record of the Smooth-Backed Parachute Gecko Ptychozoon lionotum
Annandale 1905 from the Indian Mainland

Samraat Pa war' and Sayantan Biswas2

Wildlife Institute of India, Chandrabani, Dehradun- 248 001, India. Present addresses: '3/21, Mohanwadi,
AlandiRoad, Yerawada, Pune-4 11006, India. 2 30/3 Jheel Road, Calcutta-70003 1 , India.

Abstract.- The smooth-backed parachute gecko, Ptychozoon lionotum is reported from the mainland India for the
first time. The nearest known previous record was from Pegu, Myanmar, about 700 km southeast of the previous
location. The species was collected in Langtlai and seen in the Ngengpui Wildlife Sanctuary, both in south
Mizoram. The collected individual was kept in captivity for four and a half months, during this time,
opportunistic observations on activity pattern, food habits, escape and parachuting behavior were made. Both
individuals showed slow, deliberate pre-escape movement previously unrecorded for Ptychozoon. Information
on morphological characters and morphometric measurements is presented. Explanations for the disjunct
distribution are discussed.

Key words.- Reptilia, Gekkonidae, Ptychozoon, parachute gecko, Northeast India, Myanmar, distribution,
biogeography, behavior

Ptychozoon is a genus of arboreal geckos distributed
over much of Southeast Asia, primarily in moist tropi-
cal evergreen and semi-evergreen forests (Brown,
1999; Brown et al. 1997; Smith, 1935). At present, six
species are recognized under the genus: Ptychozoon
kuhli, P. horsfieldii, P. lionotum, P. intermedium, P.
rhacophorus and P. trinotaterra (Brown et al., 1997;
Brown, 1999). To date, the only species reported for
India is Ptychozoon kuhli, from the Nicobar Islands
(De Rooij, 1915; Smith, 1935; Tiwari, 1961). We
present here the first record of the smooth-backed
parachute gecko Ptychozoon lionotum based on two
records from the state of Mizoram (21°56'N to
24°31'N and 92°16'E to 93°26'E) in Northeastern
India (Fig. 1).

The first individual was sighted on 29th June 1998
in Lawngtlai town of south Mizoram during a short
survey. Subsequently, on 21st April 1999, a second
individual was sighted by SP from the vicinity of
Ngengpui Wildlife Sanctuary (NWLS; 22°21'24" N
to 22°30'06" N and 92°45'12" E to 92o50'20•' E) in
south Mizoram, during a herpetofaunal community
study (Pawar, 1999). We could only collect the first
individual, and although SP could get a superficial
look at the second one before it escaped (see below),
we presume that it was the same species as the
straight-line distance between the two sites is only
about 40 km. Both the localities lie in the low to mid-
elevation region of south Mizoram. The vegetation is
of the tropical (moist) evergreen type, corresponding
to Northern Tropical Evergreen Forest (lb/c2; Cham-

20° N

18°N

BAY OF BENGAL

16° N

0 50 100 150lrilom«OT

96° E

Figure 1. Present record (1) of Ptychozoon lionotum
(BNHM 1445) from south Mizoram (India), along with
nearest previous record (2) from Pegu (Myanmar).

Vol. 9, p. 102 Asiatic Herpetological Research 2001

Table 1 . Mensural (in mm) and meristic measurements of two specimens of Ptychozoon lionotum . The
vouchered record reported here (BMNH 1445) is compared with one of the syntypes (ZSI 2601). The latter spec-
imen was fully discolored and severely mutilated so many characters were not discernable (NA) and so measure-
ments beyond mm were not possible.

Character

Head length

BNHM 1445
16.8

ZSI 2601

Head width

16.8

Head height

12.7

Eye diameter

4.8

Eye to nostril distance

8.2

Eye to snout distance

11.3

Eye to ear distance

7.7

Inter orbital distance

10.3

Inter narial distance

3.7

Tympanum diameter

2.3

Neck length

11.9

Snout to forelimb length

36.0

Axilla to groin length

47.0

Body flap width (Greatest width from base of flap to tip)

8.1

Body flap length (From axilla to groin)

39.6

Fore arm length

18.7

Fore limb length

27.8

Femur length

13.6

Tibia length

12.7

Hind leg length

22.9

Hind foot length

39.6

Length of I st Toe

6.3

Length of IV th Toe

9.4

Snout to vent length

94.6

Tail length

93.0

Tail width

7.6

Tail depth

6.3

Terminal tail flap length

20.1

Terminal tail flap width

10.9

Supralabials

10/11

Infralabials

9/9

Transverse dorsal bands in the axilla-groin region

Number of lobes fused before straight flap

2001

Asiatic Herpetological Research

Vol. 9, p. 103

Character

Supranasals in contact

BNHM 1445

ZSI 2601

No. of tail lappets

Subdigital lamellae (L/R)

Finger 1

11/10

Finger II

11/13

Finger III

12/16

Finger IV

15/15

Finger V

14/14

Toel

11/11

Toe II

12/12

Toe 111

16/15

Toe IV

14/14

ToeV

14/14

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,

46 .47

Iraq

\_ Iran

KERMANSHAH ■
J PROVINCE Ty

locality

CT Hamedan

□
Kermanshah^

Lorestan"^_^

50 km

Figure 2. The type locality of Eremias (Eremias) mon-
tanus, vicinity of the Village of Siah-Darreh, about
60km northeast of the city of Kermanshah, Kerman-
shah Province, western Iran.

revised Eremias and divided it into two distinguished
genera based on morphological characters: Mesalina
Gray and Eremias Fitzinger (see under systematic dis-
cussion).

As the first record of Eremias, in this paper we
describe and introduce a new species of this genus
from the upland and mountainous regions of Kerman-
shah province, western Iran at about 1800 m eleva-
tion. This province is located on the western periphery
of the Iranian Plateau (Fig. 1 ) and a major part of it is
covered by the Zagros Mountains. The type locality of
Eremias (Eremias) montanus (sp. nov.) is located in
an upland area surrounded by the Zagros Mountains

Vol. 9, p. 108

Asiatic Herpetological Research

2001

with steppe vegetation (e.g., Astragalus, Euphorbia,
Zygophyllurn), about 60 km northeast of city of Ker-
manshah, Kermanshah province, western Iran (Fig.
2).

Eremias montanus, new species
(Figs. 3-4)

Holotype and type locality: An adult female, Field
number P198, collected by the senior author on 14
August 1995 from the upland regions of the Zagros
Mountains, 60 km northeast of city of Kermanshah
(47° 5'E, 34° 52'N), Kermanshah Province, western
Iran , at about 1800 m elevation
Paratypes: Two adult females. Field number PI 99-
P200, other information as for the holotype.
Diagnosis and comparison: A small-sized lacertid,
maximum snout-vent length (SVL)= 57.2mm, tail
length = 95mm, with 13-14 longitudinal and 27-28
transverse rows of ventral plates, slightly converging
posteriorly; with 63-67 small, granular scales across

middle of dorsum. A species belonging to the subge-
nus Eremias: subocular reaches mouth edge; one fron-
tonasal; two supraoculars which are not completely
separated from frontal and frontoparietals; distance
between the femoral pores being narrow; color pattern
"striped"; inhabitant of steppe and mountain-steppe
landscapes (Szczerbak, 1974: 83).

On the other hand, it differs from all other species
of its relevant subgenus based on having several dis-
tinguishing characters; the color pattern is distinctive
and it is distinguishable from all other species in this
character i. e., dorsum uniformly dark-brown without
spots and ocelli, interrupted by five light longitudinal
stripes; the vertebral stripe bifurcating on the nape,
two paravertebrals on each side; a broad dorso-lateral
stripe containing one or two regularly arranged light
spots (different from the other Eremias species in this
character); Furthermore, it differs from each species
of the typical subgenus in the following character
combinations (Bischoff and Bohme, 1980; Bohme
and Szczerbak, 1991; Rastegar-Pouyani and Nilson,
1997; Szczerbak, 1974) :

From Eremias nigrolateralis Rastegar-Pouyani
and Nilson, 1997 in having a much smaller size, lack
of separation of the third pair of submaxillary shields
by granular scales (0% versus 100%), lower count of
gulars (23-24 versus 41-42), variable number of sub-
maxillary shields (33% versus 0%), reach of femoral

Figure 3. Eremias (Eremias) montanus holotype.

Figure 4. Eremias (Eremias) montanus paratypes.

2001

Asiatic Herpetological Research

Vol. 9, p. 109

pores to the knee (100% versus 0%), and distinct dif-
ferences in color pattern.

From Eremias persica Blanford, 1874 in having a
smaller size, lower count of gulars (23-24 versus 28-
38), the absence of distinctly keeled upper caudal
scales (100% versus 75%), variable number of sub-
maxillary shields (33% versus 4%) and distinct differ-
ences in color pattern.

From E. velox (Pallas, 1771) in having a smaller
size, in the absence of distinctly keeled upper caudal
scales (100% versus 0%), variable number of sub-
maxillary shields (33% versus 5%) and in color pat-
tern.

From E. strauchi Kessler, 1878 in having a
smaller size, the absence of distinctly keeled upper
caudal scales (100% versus almost 0%), variable
number of submaxillary shields (33% versus 9%) and
in color pattern.

From E. lalezharica Moravec, 1994 in having
variable number of submaxillary shields (33% versus
0%), higher count of dorsals (63-68 versus 54-59), no
contact of gulars with the second pair of submaxillary
shields, lack of a small scale between prefrontals,
and distinct differences in color pattern.

From E. afghanistanica Bohme and Szczerbak,
1991 in a much higher count of dorsal scales (63-68
versus 44-46), lower count of gulars (23-24 versus 25-
28), variable number of submaxillary shields (33%
versus 0%), and in color pattern.

From E. nikolskii Bedriaga, 1905 in a higher
count of dorsals (63-68 versus 45-59), variable num-
ber of submaxillary shields (33% versus 8%), and in
color pattern.

From E, regeli Bedriaga, 1905 in a higher count
of dorsals (63-68 versus 43-61), higher number of
scales in the 9th- 10th caudal annulus (27-28 versus
17-25), the absence of distinctly keeled upper caudal
scales (100% versus 0%), variable number of sub-
maxillary shields (33% versus 3%), and in color pat-
tern.

Description of holotype: An adult female, preserved
in 75% ethyl alcohol in good condition; body small
and moderately depressed; a species of the subgenus
Eremias (Szczerbak, 1974:83); five pairs of submaxil-
lary shields, first three pairs in contact, the fourth and
fifth pairs widely separated; first pair of submaxillary
shileds as large as the fifth and in contact with mental
anteriorly, with first and second infralabials laterally;
the fifth sabmaxillary pair each in contact with the
fourth pair anteriorly, being surrounded by 8 granular
scales laterally and posteriorly; 7-8 supralabials, 4-5
of which anterior to subocular which borders the

mouth; two large nasals in contact with rostral anteri-
orly, with first and second supralabials laterally, and
with frontonasal and first loreal posteriorly, the former
being single, broader than long and laterally in contact
with first loreal and posteriorly with prefrontals; two
prefrontals each smaller than frontonasal and almost
as long as broad and laterally in contact with second
loreal and posteriorly with frontal and granules of
supraocular region; only frontonasal with distinct
concavity; frontal almost as long as prefrontal and
frontonasal together, broadened and posteriorly and
laterally partly in contact with supraoculars (and
partly separated from the latter by 2-3 large scales, not
by granules) and posteriorly with frontoparietals; two
frontopariatals almost as large as a single supraocular,
laterally being in contact with the second supraocular,
and posteriorly with interparietal and parietals, the
former being small and relatively lozenge-shaped,
surrounded by frontoparietals and parietals; two vey
large and plate-like parietals, almost as long as broad,
being in contact behind interparietal; a vestigial
occipital; two loreals, first one small, surrounded by
first two supralabials, nasal, frontonasal, and the sec-
ond loreal which is distinctly large with an evident
keel; 6-6 supraciliaries, separated from supraocular by
a series of 42-44 granules; postocular elongate, sur-
rounded by granules anteriorly; temporal region-
mostly covered by granular scales becoming large
towards the orbit, more than 100 on each side; tym-
panic scale distinct and elongated obliquely, almost
the same size on both sides; tympanum vertically
elongated, slightly larger than orbit; no distinct
supratemporal; subocular huge, broader than long
with a distinct ridge being extensively in contact with
the lower edge of orbit; lower eyelids with a semi-
translucent membrane made up of about 22 enlarged
scales; collar well pronounced , not serratted, made up
of 10 scales, the two medial ones the largest; gular
fold weakly developed, 23-24 gulars from symphysis
of chin shields to median gular, becoming enlarged
posteriorly; 13-14 longitudinal and 27-28 transverse
rows of almost squarish ventral plates from collar to
hindlimbs; anterior series of ventrals to some extent
irregular, median ventral longer than broad; dorsal
scales juxtaposed, smooth, granular, becoming slighly
larger posteriorly, 63-68 scales across the middle of
dorsum, and about 160-164 scales in a single row
from occiput to a point just above the vent; proximal
caudals larger than posterior dorsals but the change
being gradual; caudals becoming large, elongate, and
slightly keeled distally, arranged in distinct whorls,
27-28 scales in the 10th whorl behind the vent; upper
forearm covered dorsally by enlarged, juxtaposed, and
almost lozenge-shaped scales; lower forearm covered

Vol. 9, p. 110

Asiatic Herpetological Research

2001

with granules; upper hindlimbs covered externally by
granules, externally by large shields; tibia covered
dorsally by slightly pointed granules, ventrally by
large plates which are slightly keeled, two plates in a
transverse row; no fringes on the toes, 18-20 uni-and
bi-carinate lamellae under fourth toe, proximal part of
lower fourth toe containing two complete rows of
lamellae, distal part with a single row (in this charac-
ter it is quite different from all other species of its rel-
evant subgenus); 18-19 femoral pores in each side, the
two series separated anteriorly by a narrow space con-
sisting of three scales; preanal region encompassing
24 large shields, the four median ones being the larg-
est; 6 plates in longitudinal row from the space
between femoral pores to anterior edge of the vent.

Coloration and color pattern: Dorsum uniformly
dark-brown without spots and ocelli, interrupted by
five light, narrow longitudinal stripes: one vertebral
bifurcating on the nape, two paravertebrals on each
side; a broad dorso-lateral stripe containing one or
two regularly arranged light spots; a ventro-lateral
series of dark-brown spots, to some extent forming a
stripe; upper surface of head uniformly olivish-brown;
temporal and labial regions suffused by dark-brown;
submaxillary region light-gray-cream; ventral region
dirty white, suffused by bluish-brown; upper surface
of limbs dark-brown containing numerous light spots;
proximal upper surface of tail brown, distal part light
brownish-gray; lower surfaces of limbs and tail yel-
lowish-white.

Description of paratypes: The paratypes, two adult
specimens, one male (PI 99) and one female (P200)
approximate the holotype in almost all pertinent
details. However, there are some minor differences
between paratypes and the holotype as follows:

Male paratype: five submaxillary shields on the right
side but only four on the left, the fifth vestigial; 13-14
longitudinal and 30-31 transverse ventral plates; 10-
1 1 collars, 3-4 median ones the largest; 23-24 gulars
in a longitudinal row from chin shields to collar; 20-
20 femoral pores, separated by three small scales; 23-
24 lamellae under fourth toe; 23-27 scales around
10th tail annulus; 8-9 labials, 5 of them anterior to
subocular; 6-6 lower labials; lower nasal resting on
the first two supralabials; temporal scale vestigial; 62-
63 scales around widest part of dorsum; 162-167
scales in a single longitudinal row from occiput to
vent.

Measurements (mm): SVL = 58.5; TL = incomplete;
Forelimb = 24; Hindlimb = 39; HL = 15.5.
Female paratype: only four pairs of submaxillary
shield, the fifth vestigial; 15-17 longitudinal and 31-

33 transverse rows of ventral plates; 9 collars, 4-5
median ones the largest; 24-25 gulars in a single lon-
gitudinal row from chin shields to collar; 20-20 femo-
ral pores, separated by a space corresponding to two
scales; 25-26 scales on the 10th tail annulus, 24-25
subdigital lamellae under fourth toe; 8-9 supra- and 4-
6 infralabials; temporal scales relatively small; frontal
separated from the first supraocular by two large
scales (not by granules); 65-67 scales around widest
part of dorsum; 163-167 scales in a single longitudi-
nal row from occiput to vent.

Measurements (mm): SVL - 52.9; TL = incomplete;
Forelimb = 23; Hindlimb = 35.5; HL = 13.6.

In color pattern they are quite similar to the holotype.

Habitat: During field work on the western regions of
the Iranian Plateau in 1995, we surveyed the Zagros
Mountains and the nearby mountain steppes in the
northern parts of Kermanshah province. 60 km north-
east of Kermanshah city, in the highland steppes, in
vicinity of the Village of Siah-Darreh in an area
named Sarpal, we came across to three specimens of a
new taxon of the genus Eremias, described here as a
new species. The habitat, which is surrounded by the
Zagros Mountains, is characterized by a mountain-
steppe; the vegetation is luxuriant steppe association:
mainly Astragalus, Euphorbia, Artemisia, and
Amigdalus as well as various species of the families
Graminaceae and Compositeae (Fig. 5).

Since it is a mountainous region with relatively
high elevation, the winter being harsh and cold, the
summer being mild and rather short. All the speci-
mens were foraging around the shrubs probably look-
ing for prey. They were quite shy and wary and very
difficult to capture. When alarmed, they took refuge
under the shrubs or inside the underground holes. One
of the most effective anti-predatory adaptations
evolved in these lizards is the ability to lose the tail
(autotomy) when being touched by the predators (or
collectors). Hence, we could only collect one speci-
men with a complete tail and the other specimens lost
their tails during capturing.

In September 1998, we re-surveyed the type local-
ity in order to find more specimens of this species but
without success. Whether it being a relictual and rare
species, confined only to the type locality, or being
distributed over a wider area in the western margin of
the Iranian Plateau is yet to be established.

With regard to the occurrence of Eremias monta-
nus in the highlands of Kermanshah province, the
Procter record of E. velox, as the westernmost record,
from Kuretu (Iran-Iraq border) (Procter, 1921:252)
should seriously be reconsidered.

Asiatic Herpetological Research

Vol. 9. p. I 1 1

js»*h»-

"ym

-*>

•" E

ilfc

•r

■w

■-**

Figure 5. Habitat and type locality of Eremias (Eremias) montanus. 60 km northeast of Kermanshah, vicinity of
Siah-Darreh village, Kermanshah Province, Western Iran.

Etymology: Eremias montanus is so named as it is
apparently restricted in distribution to the upland and
mountainous steppes of northeastern regions of Ker-
manshah province, western Iran.

Taxonomic account: As mentioned before, so far, the
most complete work done on the complicated genus
of Eremias (sensu lato) is of Szczerbak (1974) who
studied almost all species and species complexes of
this genus throughout the range. Based on morpholog-
ical characters and geographic distribution, S/.czerbak
(1974) subdivided the inclusive genus Eremias (s. 1.)
into two distinct genera; the genus Mesalina as a
north African and lowland southwest Asian clade, and
the genus Eremias (sensu stricto) which is mainly
occurring in Central and northeast Asia. (Szczerbak.
1974).

Furthermore. Szczerbak (1974) subdivided Ere-
mias (s.s) into five distinct subgenera: Eremias (Szcz-
erbak, 1974: 83), Rhabderemias (Szczerbak, 1974:
201 ), Ommateremias (Szczerbak, 1974: 146). Parere-
mias (Szczerbak, 1974: 22-23), and Scapteria ( Szcz-
erbak, 1974:247).

Except for the subgenus Pareremias, which is a
Central and east Asian clade, all of the major species
groups of the genus are presented on the Iranian Pla-

teau (Anderson, 1999). Arnold ( 1986) who studied the
hemipenes of lacertids supported the Szczerbak's sub-
generic names. In a more recent study Arnold placed
Eremias as the sister (axon of a clade including Acan-
thodactylus, Mesalina, and Ophisops-Cabrita
(Arnold. 1989:238, 240 ). But Mayer and Benir
( 1994) have proposed a different scenario. According
to these authors, Eremias is the sister taxon of
Mesalina and both of them belong to a larger clade
also containing Omanosaura and Ophisops. They
believe that Eremias is not closely related to Acantho-
dactylus.

The Czech Biological Expedition to Iran in 1996
collected 8 specimens of an undetermined species of
Eremias related to E. persica from the Zagros Moun-
tains in Esfahan province at about 2000-2200 m ele-
vation (Frynta et al., 1997: 9-10). Whether it is a new
taxon or just a variety off. persica is yet to be known.

Material examined: Eremias montanus (n = 3): P
198-200 (Field number), from around the Siah-Darreh
Village (about 1800 m elevation). 60 km northeast
city of Kermanshah. Kermanshah province, western
Iran.

Vol. 9. p. l :

Asiatic Herpetological Research

2001

Eremias nigrolateralis (n = 2): GNHM. Re. ex.
5147-5148, from 150 km northeast of Shiraz. Fars
province, south-central Iran.

Eremias persica (n = 4): GNHM. Re. ex. 5159-
5162. from 150 km northeast of Shiraz. Fars province,
south-central Iran.

Eremias persica (n = 28) : GNHM. Re. ex. 5163-
5190, from 45 km east of Arak on the road to Qum,
Markazi province, north-central Iran.

Eremias persica (n = 4) : GNHM. Re. ex. 5191 -
5194, from 65 km west of Tehran, between Eshtehard-
Saveh, Tehran province, northern Iran.

Eremias persica (n = 2) : GNHM. Re. ex. 5195-
5196. from 45 km east of Golpaygan, Esfahan prov-
ince, central Iran.

Eremias persica (n = 4) : GNHM. Re. ex. 5197-
5200. from 50 km north of Delijan on the road to
Qum. Markazi province, north-central Iran.

Eremias persica (n = 1) : GNHM. Re. ex. 5201,
from 50 km north of Abadeh. Fars province, south-
central Iran.

Eremias persica (n = 1) : GNHM. Re. ex. 5202.
from 50 km east of Hamedan on the road to Qazvin,
Hamedan province, western Iran.

Eremias persica (n = 3) : GNHM. Re. ex. 5203-
5205. from 5 km west of Takestan on the road to Zan-
jan, Zanjan province, northwestern Iran.

Eremias velox (n = 4) : GNHM. Re. ex. 5122(1-
4). from around the Carin River. 250 km E-SE Almaty
(Alma Ata). Kazakhstan.

Eremias velox (n = 2) : GNHM. Re. ex. 5120(1-
2), from Mulali Kurozek. eastern Kazakhstan.

Eremias velox ( n = 2 ) : GNHM. Re. ex. 5121(1-
2), from the Taldi Korgau District, northeast Kazakh-
stan.

Eremias velox (n = 2) : GNHM. GK. 18881 (1-2).
from Archenjan Village ( 1 ), and 30 km north of Mary
(2), Turkmenistan.

Eremias strauchi (n = 3) : GNHM. Re. ex. 441 1
(1-3), from Golestan National Park, Mazandaran
province, northeastern Iran.

Abbreviations : GNHM. Re. ex. = Gothenburg Natu-
ral History Museum. Reptilia exotica; GNHM. GK. =
Gothenburg Natural History Museum. General Kata-
logue.

Acknowledgements

We wish to thank the Razi University authorities (Ker-
manshah-Iran) for their generous help and support
during field work in western Iran.

We thank the Gothenburg Natural History Museum
(Gothenburg-Sweden) for loan of Eremias specimens.
Also we thank Dr. Michael Golubev for translation of
the relevant Russian literature.

Literature Cited

Anderson, S. C. 1999. The Lizards of Iran. Society for
the study of Amphibians and Reptiles 442 pp.

Arnold, E. N. 1986. The hemipenis of lacertid lizards
(Sauria : Lacertidae): structure, variation and system-
atic implications. Journal of Natural History 20:1221-
1257.

Arnold, E. N. 1989. Towards a phylogeny and bioge-
ography of the Lacertidae : relationships within an
Old- World family of lizards derived from morphol-
ogy. Bulletin of the British Museum (Natural His-
tory). Zoology 55(2):209-257.

Bischoff, W. and W. Bohme. 1980. Der systematische
Status der turkischen Wustenrenner des Subgenus
Eremias (Sauria: Lacertidae). Bonner Zoolgische
Beitrage, N. F. 26:297-306.

Bohme. W. and N. N. Szczerbak. 1991. Ein neuer
Wustenrenner aus dem Hochland Afghanistans. Ere-
mias (Eremias ) afghanistanica sp. n. (Reptilia : Sau-
ria : Lacertidae). Bonn. Zool. Beitr. 42:137-141.

Frynta. D.. J. Moravec, J. Cihakova, J. Sadlo, Z. Hod-
kova, M. Kaften. P. Kodym. D. Krai. V. Pitule. and L.
Sejna. 1997. Results of the Czech Biological Expedi-
tion to Iran. Part 1. Notes on the distribution of
amphibians and reptiles Acta Societatis Zoologicae
Bohemicae 61 :3-17.

Mayer. W. and G. Benyr. 1994. Albumin-Evolution
und Phylogenese in der Familie Lacertidae (Reptilia:
Sauria). Annalen Naturhistorischen Museums in Wien
96B:62 1-648.

Moravec. J. 1994. A new lizard from Iran. Eremias
(Eremias ) lalezharica sp. n. (Reptilia: Lacertilia:
Lacertidae). Bonner Zoolgische Beitrage 45(1 ):6 1-66.

Procter. J. B. 1921. Further lizards and snakes from
Persia and Mesopotamia. Journal of the Bombay Nat-
ural History Society 28(1 ):25 1-253

Rastegar-Pouyani. N. and G. Nilson. 1997. A New
Species of Eremias (Sauria: Lacertidae) from Fars
Province, South-Central Iran. Russian Journal of Her-
petology4(2):94-101.

Szczerbak, N. N. 1974. Yashchurki Palearktiki (Pale-
arctic species of Eremias ). Kiev. 295 pp.

2001

Asiatic Herpetological Research

Vol.9, pp. 113-121

Lizards of the Northern Mongolian Deserts: Densities and Community

Structure

KONSTANTIN A. ROGOVIN1, DMITRY V. SEMENOV1, AND GEORGY I. SHENBROT2

'A.N.Severtzov Institute of Ecology and Evolution, Russian Academy of Science, Leninsky pr. 33 Moscow

1 17071, Russia, ~ Ramon Science Center, Ben-Gurion University of the Negev, P.O. Box 194, Mizpe Ramon,

80600, Israel

Abstract.- Spatial organization and population densities of three-species lizard community was studied in the
Gobi Desert, Mongolia. To evaluate the effect of habitat variables on the distribution and abundance of each
species we used the stepwise procedure of factor selection with ANOVA on each step. To describe the
distribution of species' spatial niches in the space of environmental variables, we used stepwise discriminant
function analysis (DFA). The number of species in 1-ha grid areas varied from 0 to 4. Phrynocephalus versicolor
was the only species distributed over the 91% of grids occupied. There was a positive relationship between
distribution and local species abundance. A set of two to three habitat variables determined the abundance of each
species. The result of DFA signify to the well pronounced segregation, but not even distribution of species spatial
niches in the space of resources.

Key words.- Lizards, Mongolia, community, ecology, density

91* 96* 101* 106°

Fig 1. Map of Mongolia and location of sites where data were collected.
Eastern Gobi, 4-Barun-Churay Basin.

111* 116'

Southern Gobi, 2-Western Gobi, 3-

Introduction

The reptile communities of Mongolian deserts are
characterized by several specific features discussed
elsewhere (Ananjeva and Semenov, 1986; Borkin and
Semenov, 1984; Munkhbajar, 1976; Semenov and
Borkin, 1986; Semenov and Shenbrot, 1988). There
are few species in the fauna with low species richness

at sites, a low level of species turnover between habi-
tats, low abundance of most species and high domi-
nance of only one species, Phrynocephalus versicolor.
A few common species have rather broad spatial
niches, diverse behavioral and physiological charac-
teristics (diverse range of thermobiological patterns,
wide active search for food items, etc.).

Vol. 9, p. 114

Asiatic Herpetological Research

2001

The degree of interest in comparative studies of
structure and function of reptile communities
increased dramatically after seminal papers written by
E. Pianka (Pianka, 1973, 1975). Most of the ensuing
studies were devoted to the species rich and diverse
communities of desert lizards in Australia, south-
western North America and southern Africa (Case,
1983; Fuentes, 1976; Henle, 1989; Inger, Colwell,
1977; Millado et al., 1975; Pianka,1986; Scheibe,
1987; Shenbrot et al., 1992; Simbotwe, 1984). At the
same time study of species poor communities in the
Central Asian desert can provide a significant infor-
mation not only in comparison with other continents,
but also can help us to understand better which factors
rule in reality structure and dynamic of lizard commu-
nities of many species. Up to now there were only two
examples of such studies made in China (Chang et al.,
1993;Luietal.,1992).

The main objective of this paper was to study the
specific features of spatial organization and popula-
tion densities of three-species lizard community in the
Gobi Desert, Mongolia. The study was conducted dur-
ing a long-term research program on the biodiversity
of the Mongolian desert biota, and was sponsored by
the Permanent Soviet-Mongolian Biological Expedi-
tion.

Material and Methods

Mongolian desert

A map (Fig. 1) illustrates the location of desert
regions of Mongolia. Three desert regions to the south
of the Altai Mountains are partly separated from one
another by chains of low mountains and hills. These
three are Trans-Altai Gobi (South), Alashan Gobi
(East) and Sungarian Gobi or Barun-Churay Basin
(West) (Yunatov, 1950). Besides these deserts there
are desert areas between the Altai and Hangai moun-
tains, usually called Western cold deserts, and some
arid lands in the Great Lakes and Ubsu-Nur Basins
(northwest).

Three arid subzones of the Mongolian desert are
defined (Sokolov and Gunin, 1986): extra — arid
desert (<50 mm of rainfall per year), real desert (50—
100 mm per year) and steppe — like desert (100—150
mm per year). However, the climatic border that
restricts the distribution of plants (Kazantseva, 1986;
Volkova et al., 1986) and animals (Podtyazhkin and
Orlov, 1986; Semenov and Borkin, 1986) exists only
between the southern part of Trans- Altai Gobi (< 50
mm per year) and the northern waste belt of desert
lands with more predictable precipitation (100-200
mm per year). The narrow real desert subzone appears

transitive with no specific features of vegetation.
Thus, only the southern (extra-arid) and northern sub-
zones are well pronounced.

The southern desert occurs mostly in the Trans-
Altai Gobi and is characterized by a few very dry,
unproductive biotopes inhabited by five lizard species
(two agamids, two gekkonids, and one Eremias spe-
cies). Among this group only one species, Phryno-
cephalus versicolor is common in the northern
subzone (Semenov and Borkin, 1986).

The northern deserts, which extend to the south-
west, west, and south-east of Mongolia and along
both slopes of the Mongolian Altai and Gobi-Altai
mountains, are characterized by pronounced microre-
lief and rich vegetation, although the main landscape
types are the same as in the southern subzone. The
vegetation in rock and gravel valleys consists of
perennial grass (Stipa), forbs, onions and succulents,
and a variety of annual plant species. Shrub vegetation
is often associated with foothills and sand dunes, or is
spread along the dry river beds (Lavrenko, 1978). The
difference in climate between western and eastern
parts of the northern Mongolian desert is not pro-
nounced (Murzaev, 1952); some differences exists in
the composition of the flora (Yunatov, 1950).

Lizard species

There are four lizard species inhabiting northern
Mongolian deserts: Alsophylax pipiens, Phrynoceph-
alus versicolor, Eremias przewalskii and E. multiocel-
lata. Among these, only the three last mentioned
species are abundant and relatively widespread.

Data collection

We collected data during three field trips to the Mon-
golian northern deserts in June-August 1985, 1986
and 1988. Forty five 1-ha grids were established in the
Northern desert subzone (see map, Fig 1 ). Grids were
distributed so that they covered the whole range of
habitats from the middle slopes of the mountains to
the clay basins and sand dunes. Each habitat type was
sampled equally. Two factors determined the number
of grids at a desert region: diversity of habitat types
and abundance of lizards. Each grid was divided into
25 smaller sample plots, 20 x 20 m. The centers of
sample plots were marked with 50 cm aluminum
stakes. Lizards were sampled by repeated, regular
search of established grids during two to four consec-
utive days during periods of their maximal diurnal
activity. Nearly all encountered lizards were captured
by hand. Most of the surveys accounting were con-
ducted before the appearance of hatchlings. In the rest
of the cases hatchlings were not counted. Each cap-

2001

Asiatic Herpetological Research

Vol. 9, p. 115

Table 1. Density (no. ha ) of lizard species.

</>

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o

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c

(A
0)

/? versicolor

E przewalskii

E. multiocellata

1.7±0.9 52.8+8.9 54.4±8.7 36.6±8.2 5.5±4.5 41.2+6.3 0.8±0.4

0-3 24-91 32-83 10-106 1-10 22-64 0-3

0 0 0 2.6±1.3 0 16.3±6.3 0.9±0.5

0-18 0-44 0-4

0 2.1+1.3 0.4±0.4 0.5±0.4 0 2.0±1.8 0
0-12 0-2 0-5 0-11

tured lizard was marked by toe-clipping (the most dis-
tal phalange only) and/or by a number painted with
permanent color marker on its back, and released.
Densities of lizard species were estimated by the total
number of animals caught on a grid. The reliability of
density estimates was verified by recapture of marked
individuals and by the observation of marked lizards
on grids. Two days were sufficient to catch > 80% of
lizards.

A 0.5 kg soil sample was taken from the center of
each plot for laboratory texture analysis. The number
of shrubs (by species) in each plot was counted in a
circle, 5 m in radius. To determine vegetation cover
and volume by height layers within each plot, the
height and diameter of crown of the shrubs (up to 30
shrubs of each species in the grid) were measured.
The abundance of annuals was evaluated by clipping

all annuals on 0.25 m" sample plots (4 sample plots
placed at random in each of 25 plots in 1-ha grid
area). Twenty three parameters were used in the sub-
sequent analysis (Table 2). Data on 1 125 descriptions
of sample plots and 1710 records of 3 lizard species
were used in the analysis. In total, we recorded 1528
individuals of the agamid lizard P. versicolor, 142 of
the lacertid lizard Eremias przewalskii and 40 E. mul-
tiocellata.

Statistical data processing

To evaluate the effect of habitat variables on the distri-
bution and abundance of each species we used the
stepwise procedure of factor selection (Shenbrot,
1988) with ANOVA of each step. Before analysis, the
original values of species' densities and habitat vari-
ables were transformed to an interval scale with five
intervals for habitat variables and three intervals for
species' abundance variables. The influence of each
variable was estimated separately by ANOVA and the
variables with nonsignificant effect were omitted. In
the next step, each of variables that was not omitted
earlier was added separately to the most powerful
variable to select the most powerful pair of variables.
This procedure was repeated until all possible vari-
ables were included in the analysis or until the addi-
tion of the new variable does not increase the
proportion of explained dispersion.

To describe the distribution of species' spatial
niches in the space of environmental variables (eco-
logical space) and to reduce the dimensionality of this
space, we used stepwise discriminant analysis. Dis-
criminant axes were calculated based on the data set
consisting of the values of habitat variables for each
point of lizard's registrations and with lizard species
as a grouping variable.

Vol. 9, p. 116

Asiatic Herpetological Research

2001

Results

Densities and distribution through habitat
types

The results of lizard density and diversity estimations
on the 1-ha grids indicate the low local species diver-
sity in Mongolian deserts (Table 1). The number of
species in our grid areas varied from zero to three.
Among 45 grids there were two grids with no lizards,
21 grids with only one species. 18 grids with two spe-
cies and four grids with three species. Phrynoceph-
alus versicolor was the only species distributed over
the most number of grids (91% of grids occupied).
Eremias przewalskii was found on 33%, E. multioce-
lata on 24% and A. pipiens on 4% of grids. Regarding
distribution through the main habitat types (Table 1),
P. versicolor was also the most widely distributed spe-

cies but with the pronounced habitat preference. The
second was E. multeocellata, and the third was E.
przewalskii. The last species had the most restricted
habitat preference, namely sandy-loess hills in saline
depressions with shrub vegetation of Nitraria sp.

The abundance of P. versicolor was positively cor-
related with its broad distribution. The density of this
species varied from 1 to 106 individuals per 1-ha
(Table 1). The second most abundant E. przewalskii
(44 individuals per lha maximum), and the third was
E. multiocelata (11 individuals per 1-ha maximum).
All three species coexisted at rather high densities in
saline depressions with sandy-loess hills covered with
Nitraria sp. shrubs. There were no pronounced nega-
tive correlation between the densities of two dominant
species on grids situated within this habitat type (P.
versicolor-E. przewalskii: R"=0.04, n=20, ns).

Table 2. Designation and description for the 23 habitat variables included in the analysis.

Mnemonic

Variable

Unit

RCK

GRW

CLY

SCS

SCH

WDS

WDD

NRB

FRB

AGR

ANN

ALL

PGR

MIC

HAL

sue

HLX

NIT

SHC

SV1

SV2

SV3

SV4

Content of rocks in the soil

Content of gravel in the soil

Content of clay in the soil

Sand cover area

Sand cover height

Dry river bed area

Dry river bed depth

Number of rodent burrows

Abundance of annual forbs

Abundance of annual grasses

Overall abundance of annual grasses and forbs

Perennial Allium covet

Perennial grass cover

Cover of microphyllous shrubs

Cover of halophytuos shrubs

Cover of small succulent shrubs

Cover of Haloxylon

Cover of Nitraria

Overall shrub cover

Perennial plant crown volume at the level 0-25 cm

Perennial plant crown volume at the level 25-50 cm

Perennial plant crown volume at the level 0.5-1 m

Perennial plant crown volume at the level 1-2 m

no/sq.m.

/o /o

o/ o/
/o /o

0/ O/

/o /o
o/o.

Vo%

2001

Asiatic Herpetological Research

Vol. 9, p. 117

1
0.5

-0.5

-1

AGR

1
0.5

0
-0.5

-1

NIT

SHV2

0.5

-0.5

-1

GRV

1
0.5

0
-0.5

-1

Fig 2. Habitat use profiles of P. versicolor. For each
habitat variable rank, mean use by species is plotted
as a difference between capture frequency on plots of
this rank and capture frequency of all plots. The size of
histograms therefore denote the selectivity (positive or
negative) of habitat use. For mnemonics see Table 2

Spatial niches

There was a statistically significant influence of habi-
tat variables on the distribution and abundance of all
three studied lizard species (Table 3). For each of
these species we extracted a set of two to three habitat
variables, determining 6.6-28.7% of observed vari-
ance in abundance. Densities of two species (P. versi-
color, E. przewalskii) were moderately affected by
habitat variables, whereas density of E. multiocellata
was weakly affected.

There was a statistically significant influence of
habitat variables on the distribution and abundance of
all three studied lizard species (Table 3). For each of
these species we extracted a set of two to three habitat
variables, determining 6.6-28.7% of observed vari-
ance in abundance. Densities of two species (P. versi-
color, E. przewalskii) were moderately affected by
habitat variables, whereas density of E. multiocellata
was weakly affected.

CLY

0.5

-0.5
-1

Fig. 3. Habitat use profiles of E przewalskii. For expla-
nation see Fig. 2.

FRB

MIC

Fig. 4. Habitat use profiles of E multiocellata. For
explanation see Fig.2.

Patterns of habitat usage based on selected vari-
able sets for each species are presented on Figures
2-4. Phrynocephalus versicolor clearly avoided
microsites with low gravel content, very low and very
high annual grass abundance, moderate and high
shrub crown volume in the level 0.25-0.5 m and pre-
ferred microsites with moderate gravel content, mod-
erate annual grass abundance and very low shrub

Vol. 9, p. 118

Asiatic Herpetological Research

2001

Table 3. Summary of ANOVA analyses of influence of habitat variables on individual species' abundance. Values
are proportions of total dispersion determined by given variable. Total proportion of variance determined by a set
of variables may be greater than sum of influences of individual variables as a result of high-order interactions.

Species V a r

i a b 1 e s

Total

GRV CLY AGR

FRB MIC

NIT

SHV2

P. versicolor 0.0226 0.0387

0.0458

0.1900

E. przewalskii 0.0593

0.1592

0.0102

0.2872

E multeocellata

0.0180 0.0232

0.0662

crown volume in the level 0.25-0.5 m (Fig. 2). Ere-

Discussion

mias przewalskii avoided microsites with high and
very high clay content, very low Nitraria cower, very
low shrub crown volume in the level 0.25-0.5 m, and
preferred microsites with low clay content, moderate
to high Nitraria cower, moderate to high shrub crown
volume in the level 0.25-0.5 m (Fig. 3). Eremias mul-
tiocellata avoided microsites with very high and very
low forb abundance and microphyllous shrub cover,
and preferred microsites with moderate forb abun-
dance and rather high microphyllous shrub cover (Fig.
4).

Structure of ecological space occupied

The results of reducing habitat space dimensionality
using discriminant analysis showed that division of
this space by lizard species occurred along the first
two axes. Both axes accounted in sum for 100 % of
variance and reflected some complex environmental
gradients (Table 4). The first axis represented a gen-
eral gradient of decreasing rock and gravel content in
the soil as well as an increase in the sand cover area,
sand mound height, number of rodent burrows and
general shrub (especially Nitraria) cover. This axis
characterized the between-habitat component of envi-
ronmental variation rather than within-habitat varia-
tion, and described in general spatial segregation of
lizard species according to their preference of physi-
ognomically distinctive habitats. The second axis
reflected an increase of dry river bed area, Allium
abundance and microphyllous shrub cover, and char-
acterized both between- and within-habitat compo-
nents of species segregation. The first axis described
habitat division between E. przewalskii and two other
species, whereas the second axis reflected habitat seg-
regation between E. multiocellata and two other spe-
cies (Fig. 5). All three lizard species had spatially
remote niche centers and less than 30% niche overlap
(Fig.5).

Our data confirm the general opinion of low reptile
species richness in Mongolian deserts. There were
only four 1-ha grids where three lizard species coex-
isted. If we take into account rare encounters with
snake species, the maximum reptile species number is
five. Another opinion about wide spatial niches of
Gobian lizards (Semenov and Borkin, 1986) partly
contradicts our results. At the one hand P. versicolor is
the an abundant species that can be found everywhere
in the Gobi desert, including sandy, clay and rocky
habitats. This feature of its distribution distinguishes
this species greatly from its western congeners that
share habitat types in Middle Asia (Shenbrot et al.,
1991 ). At the other hand, P. versicolor did have well
pronounced spatial preferences as it is seen from
Table 1. The two species of Eremias lizards had dis-
tinctly non-preferable habitats. Eremias przewalskii
was mainly restricted to one habitat type, and E. mul-
tiocellata to two habitat types.

Results of analysis of microhabitat preferences
demonstrate the existence of significantly distinct
environmental variables determining spatial distribu-
tion of each lizard species. The variables elucidated
can be regarded as axes of species spatial niche. Spa-
tial niches for P. versicolor and E. przewalskii
appeared to be determined by three axes of environ-
mental variation and for E. multiocellata by two axes.
The distribution of species along each axes can be
interpreted in accordance with the biological charac-
teristics of each species. The absence of P. versicolor
at microsites with dense vegetation is explained by
typical Phrynocephalus locomotion on straightened
legs and by the group-specific visual orientation when
foraging (in contrast to Eremias, which look for food
items using olfaction). Phrynocephalus versicolor
preferred microsites with moderate gravel content,
moderate annual grass abundance and very low shrub
crown volume, which characterizes the species as an
inhabitant of stony and gravel desert valleys (Fig. 2).
Eremias przewalskii selected microhabitats in accor-

2001

Asiatic Herpetological Research

Vol. 9, p. 119

Fig. 5. Seventy-five percent confidence ellipses for the species observations on two discriminant axes (DF1 and
DF2). Em- Eremias multiocellata, Ep- Eremias przewalskii, Pv - Phrynocephalus vesicolor.

DF1

dance with food abundance and a species-specific tac-
tic of antipredator behavior. It preferred microsites
with moderate to high Nitraria cover and moderate to
high shrub crown volume at 0.25-0.5 m (Fig. 3). In
summer E. przewalskii feed predominantly on berries
and young green twigs of Nitraria, and also find pro-
tection under the dense cover of the crowns of low
spiny shrubs. In contrast to E. przewalskii, E. multio-
cellata avoided microsites with very high annual plant
abundance and shrub cover, but also preferred micro-
sites with moderate and rather high values of these
variables (Fig. 4). This difference can be interpreted
in accordance with thermobiological and size charac-
teristics of two Eremias species. The larger species, E.
przewalskii is not so quick as the smaller E. multiocel-
lata. The first one digs well in soft soil, and ther-
moregulates climbing on or escaping under the shrub
periodically, being active throughout the day. Small E.
multiocellata that occupy habitats with relatively low
vegetation cover (with small sparsely-distributed
shrubs) must cross open sites in search for food items
and escape predator's attacks by quickly rushing into
small shrubs or burrows. This species reduces heat by
escaping into burrow during the day-time.

Another result of the above comparison is the pos-
itive relationship between distribution and local abun-
dance among the species considered. Distribution

here means not the size of the species range area, but
the number of sites where each species was found.
Phrynocephalus versicolor was encountered on 41
grid areas and had the highest population density
(mean: 37.3, median: 26 ind/ha, maximum: 106 ind/
ha). With the edge-effect correction (Semenov, 1991)
maximum density was 70.02 ind/ha (175.5 g/ha biom-
ass). Eremias przewalskii was found on 1 5 grid areas
and was the second abundant species (mean: 9.5,
median: 4 ind/ha, maximum: 44 ind/ha). Eremias mul-
tiocellata was found on 1 1 grid areas and its maxi-
mum density was 12 ind/ha (mean: 3.6, median: 2 ind/
ha). Alsophilax pipiens was met on two grids with
density 3 ind/ha.

Brown (1995) explains this rather common rela-
tionship by the "Hutchinsonian niche model" (see
also for one species, Brown 1984), suggesting that
"the species that is slightly more tolerant of some abi-
otic conditions or biotic interaction or is slightly bet-
ter able to use some resource should not only be able
to occur in more places but also to attain higher abun-
dance in some of those places." In the approach we
use here niche breadth reflects microhabitat require-
ments of each species, namely the range of microcon-
ditions where each lizard species occurred. Diversity
of these microhabitats is not necessarily correlated
with the diversity of macrohabitats as well as with the

Vol. 9, p. 120

Asiatic Herpetological Research

2001

Table 4. Summary of discriminant analysis of the habi-
tats of lizard species. DF1 and DF2 are the first two
components (all are significant, P<0.001). Mnemonics
for habitat variables are from Table 1 .

DF1

DF2

Eigenvalue

0.351

0.102

Chi-square

676.1

165.5

Cumulative % of variance

62.58

37.42

Factor loading

RCK

-0.350

0.295

GRV

-0.291

0.124

SCS

0.544

-0.221

WDD

-0.091

0.241

NRB

0.442

0.253

SHH

0.627

0.190

ALL

-0.141

0.252

MIC

-0.021

0.410

NIT

0.642

0.193

SHC

0.650

0.148

SV1

0.712

0.067

SV2

0.693

-0.069

size of the landmass studied. Phrynocephalus versi-
color that was distributed everywhere in the Gobi, and
in many different macrohabitats used a rather limited
range of microconditions (Fig. 2). In this respect its
spatial niche was not broader than the niche of E. mul-
tiocellata, and was even narrower in comparison with
E. przewalskii. (Fig. 5). Eremias przewalskii which
inhabits a rather limited range of macrohabitats is
characterized by relatively broad requirements for
microconditions along the first discriminant axis
which represents the main direction of spatial segre-
gation of lizard species.

The structure of ecological space occupied by
three lizard species has a complex character, explain-
ing both macrohabitat and microhabitat segregation of
spatial niches. Primarily, each discriminant axis char-
acterizes the range of variation of microconditions in
the study area. This can characterize macrohabitats
only if variation in a set of variables included into
analysis reflects the macrohabitat variation. In our
case DF1 and DF2 possess such a feature: DF1 char-
acterizes better the between-habitat component of
spatial niche segregation and DF2 characterizes more

the within-habitat component. Three species of lizards
share microconditions in two-dimensional ecological
space, so that the niche centers appeared to be almost
equally distant from the geometrical center of the
model (Fig. 5). The two species of Eremias lizards
share ecological space to a greater extent than each
Eremias with P. versicolor. Niche overlap between
species was less than 15%.

It seems difficult to speculate about processes that
led to such spatial relationships. Interspecific compe-
tition could contribute to spatial segregation of Ere-
mias lizards in past, and could determine the low level
of niche overlap. The competitive relationship
between E. przewalskii and P. versicolor is unclear. In
a pair of neighboring grid areas at one location within
one habitat type when one species is in high density,
the other one is usually at low density and vise-versa.
However, there were no correlation between popula-
tion densities of these two dominant species when all
grid areas within one habitat type were put together.
This result is easily explainable because different
localities must have different upper limits of species
densities according to the local environmental capaci-
ties. However, the climatic conditions of Gobi Desert
are so unfavored for reptiles (extremely low winter
temperatures, great interannual, between- and within-
seasonal contrasts in precipitation and temperatures)
that it appears unrealistic to assume the space of
resources is saturated by individuals, and that popula-
tions are at equilibrium.

Acknowledgments

We are very grateful to Dr. David Ward (Ben-Gurion
University of the Negev) for helping with the manu-
script and to Dr. Natalia B. Ananjeva (Zoological
Institute of St. Petersburg) for reading the manuscript.
This is publication no. 12510-5241-9 of IEE and no.
113ofRSC.

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2001

Asiatic Herpetological Research

Vol. 9, pp. 122-129

The Morphology and Size of Blood Cells of Lacerta rudis bithynica
Murat Sevinc1 and Ismail H. UGurta§

Uludagg University, Science and Art Faculty, Department of Biology, 16059 Bursa, Turkey. To whom
correspondence should be addressed e-mail: smurat® uludag.edu.tr

Abstract.- In this study, the morphology of the blood cells of Lacerta rudis bithynica is described using Wright's
technique. The sizes of erythrocytes and their nuclei, leukocytes (monocytes, lymphocytes, basophils,
neutrophils, and eosinophils), and thrombocytes of L. rudis bithynica were measured using an ocular micrometer
at a magnification of 1600X with an oil immersion objective. The results of this study are compared with
previous work on other reptile species.

Key words.-
morphology.

Lacertidae, Lacerta rudis bithynica, erythrocyte, leukocyte, thrombocyte, measurement.

Introduction

The first studies on the blood of reptiles described the
cellular structures, often comparing them with those
of other vertebrates. Literature on the haematology of
reptilian blood is based on few studies and is usually
concerned with European species (Saint Girons,
1970).

Recent studies have concentrated on single species
(Tiliqua sp., Cannon et al., 1988; Cyrtopodion sca-
brum, Canfield and Shea, 1996). Various authors have
described different circulating blood cells of different
reptile species (Taylor and Kaplan, 1961; Heady and
Rogers ,1962; Hartman and Lessler, 1964; Szarski
and Czopek, 1966; Duguy, 1970; Saint Girons, 1970,
Cannon et al., 1988, Canfield and Shea, 1996). Other
authors have studied seasonal (Hutton, 1960; Cline
and Waldman, 1962; Haggag and et. al, 1966) or sex-
ual (Altland and Thompson, 1962) variation in the
number of blood cells of different reptile species. In
addition, authors have studied the number of blood
cells of different reptile species (Charipper and Davis,
1932; Baker and Cline, 1932; Hutton, 1961; Altland
and Thompson, 1962; Hutchinson and Szarski, 1965;
Engbretson and Hutchinson, 1976). Finally, authors
have studied haemoglobin and hematocrit content of
blood and hematopoiesis of different reptile species
(Altland and Thompson, 1958; Hutton, 1961; Goin
and Crawford, 1965; Engbretson and Hutchinson,
1976; Newlin and Ballinger, 1976).

In Turkey, haematological studies have been con-
ducted on humans and economically important ani-
mals. However, there are no haematological studies
on the Turkish reptiles.

In this study, our aim is to describe and measure
blood cells (erythrocyte, leukocyte, thrombocyte) of
Lacerta rudis bithynica (Squamata: Lacertidae). This
study is the first of its kind on a Turkish species.

Material and Methods

In this study, 31 individuals (17 male) of Lacerta
rudis bithynica (Squamata: Lacertidae) were exam-
ined. The study was carried out between June and
August 1998. The specimens were collected from
Uludagg (Bursa) at an altitude of 1745 m. Blood was
obtained by cardiac puncture of the lizards (Canfield
and Shea, 1988). Immediately after the blood was
obtained in heparinized capillary tubes, the blood
smears were prepared. Three to five blood smears
were prepared per individual. The smears were air-
dried and stored until stained with Wright's stain
(Hartman and Lessler, 1964). Twelve drops of
Wright's stain were dropped on the slides and allowed
to remain on the slide one and a half minutes before
rinsing with a phosphate buffer (pH=6.5). The slides
were allowed to stand for ten minutes at room temper-
ature and were then washed with distilled water and
allowed to dry.

On each slide fifty mature erythrocytes and their
nuclei, ten thrombocytes, and ten leukocytes (mono-
cyte, lymphocyte, eosinophil, basophil and neutro-
phil) were measured by means of an ocular
micrometer at a magnification of 1600 x with an oil
immersion objective. Cell sizes were calculated from
the measurements.

2001

Asiatic Herpetological Research

Vol. 9,

123

10/""

Figure 1 . Erythrocytes, an infected erythrocyte and a
mitosis dvision.

Results

I. Erythrocytes

Erythrocytes are nucleated, oval cells. Their nuclei are
also oval, more or less regular and centrally located
(Fig. 1). The cytoplasm of mature erythrocyte
appeared light and dark pink and homogeneous under
Wright's stain. The nuclei of mature erythrocytes are
chromophilic. In some blood smears, immature eryth-
rocytes are seen. They are characterised by a rounded
form, blue cytoplasm and a large nucleus. Mitotic fig-
ures are also present and in some smears, intracorpus-
cular parasites are seen (Fig. 1). Parasites alter the
shape and size of erythrocytes remarkably. When
intracorpuscular parasites are seen, immature erythro-
cytes and mitotic figures are abundant (Fig 1 ). Intrac-
orpuscular parasites alter the shape and size of
infected erythrocytes. The shape and size of other
erythrocytes that are not infected by intracorpuscular
parasites are normal.

20
15

•s io

14 41 15 05 1

5 71

[6.17 [630

M6M

I Erythrocyte
I Nucleus

June

July August

Figure 2. Erythrocyte and nucleus lengths of Lacerta
rudis bithynica qnq\ three months.

to
8
6
4
2
0

7.63 8.08 S 24 ■

BR ™

[3.5s [3 55 lj_74

Erythrocyte
Nucleus

June

July

August

Figure 3. Erythrocyte and nucleus widths of Lacerta
rudis bithynica over three months.

In June mean length of mature erythrocytes was
14.41 urn (±0.77 standard deviations, with a range of
12.20-16.47 urn). In July, the mean length of mature
erythrocytes was 15.05 urn (±0.79, 12.81-17.08 pm).
In August, the mean length of mature erythrocytes
was 15.71 urn (±0.79, 12.81-18.30 um). Other mea-
surements are given in Tables 1, 2 and 3. There are no
significant differences in erythrocyte and nucleus
sizes between females and males. Based on Tables 1 ,
2, and 3 and Figs. 2 and 3, it appears that there were
little monthly variations in erythrocyte and nucleus
sizes among June, July and August.

II. Leukocytes

1. Eosinophils. In blood smears stained by Wright
technique, eosinophils are circular, and the cytoplasm

Table 1. Erythrocyte dimensions of Lacerta rudis bithynica with the standard deviations in June. EL: Erythrocyte
length; EW: Erythrocyte width; ES: Erythrocyte size;NL: Nucleus length; NW: Nucleus width; NS: Nucleus size.

EL(um)

EW (um)

EL/EW

ES (Mm2)

NS/ES

Maximum

16.47 ±0.77

9.15 ±0.48

2.27 ±0.13

110.41 ±8.00

0.30 ± 0.02

Minimum

12.20 ±0.77

6.71 ± 0.48

1.42 ±0.13

64.26 ± 8.00

0.13 ±0.02

Mean

14.41 ±0.77

7.63 ± 0.48

1.89 ±0.13

86.46 ± 8.00

0.20 ± 0.02

NL(um)

NW (um)

NL/NW

NS (Mm2)

Maximum

7.32 ± 0.40

4.88 ± 0.34

2.20 ±0.19

24.53 ±2.19

Minimum

4.88 ± 0.40

3.05 ± 0.34

1.25 ±0.19

11.68 ±2.19

Mean

6.17 ±0.40

3.55 ± 0.34

1.75 ±0.19

17.25 ±2.19

Vol. 9, p. 124

Asiatic Herpetological Research

2001

Table 2. Erythrocyte dimensions of Lacerta rudis bithynica\oqe\!r\er with the standard deviations in July. EL: Eryth-
rocyte length; EW: Erythrocyte width; ES: Erythrocyte size; NL: Nucleus length; NW: Nucleus width; NS: Nucleus

size.

EL (Mm)

EW (pm)

EL/EW

ES (nm2)

NS/ES

Maximum 17.08 ±0.79 9.15 ±0.49 2.16 ±0.12 122.68 ±8.95 0.28 ± 0.02

Minimum 12.81 ±0.79 6.71 ± 0.49 1.53 ±0.12 67.47 ± 8.95 0.13 ±0.02

Mean 15.05 ±0.79 8.08 ± 0.49 1.86 ±0.12 95.62 ± 8.95 0.18 ±0.02

NL(um)

NW (Mm)

NL/NW

NS (Mm2)

Maximum
Minimum

Mean

7.30 ± 0.70 4.27 ± 0.32

5.40 ± 0.70 3.05 ± 0.32

6.30 ± 0.70

3.55 ± 0.32

2.40 ±0.17
1.33 ±0.17

1.79 ±0.17

24.53 ±2.38
13.14 ±2.38
17.67 ±2.38

is stained light red. Eosinophils contain circular to
elongate cytoplasmic granules stained brilliant red
(Fig 4). Eosinophils are different from neutrophils in
that, eosinophils' granules are stained bright red and
neutrophils' granules were stained dim red.

In June, the mean diameter of eosinophils was
12.82 pm, (±1.71 standard deviations, with a range of
9.93-15.25 um). In July, the mean diameter was
13.29 um (±1.25, 10.98-15.25 um). In August, the
mean diameter was 13.80 pm (±1.37, 12.20-15.25
um).

There were no significant differences in eosinophil
diameters between females and males. Based on
Tables 4, 5, and 6 and Fig. 8, it appears that there was
little monthly variation in the diameter of eosinophils
during the three months.

2. Basophils. Basophils are easily recognised. They
are small and circular cells. Nuclei stained blue by
Wright technique are commonly obscured by chro-
mophilic circular granules. These cytoplasmic gran-
ules are large and stained dark purple. In the blood

Erythrocytes j^k

•lO/*"-

smears, they resemble mulberries (Fig. 5). The gran-
ules are so dense that nucleus stained dim blue is
rarely seen.

In June, the mean diameter of basophils was 8.55
um, (±0.61 standard deviations, with a range of 7.32-
9.15 pm). In July, the mean diameter was 9.02 pm
(±0.24, 8.54-9.15 pm). In August, the mean diameter
was 9.00 pm (±0.45, 7.93 -10.37 pm).

There were no significant differences in basophil
diameters between females and males. Based on
Tables 4, 5, and 6 and Fig. 8, it appears that there was
little monthly variation in the diameter of basophils
during the three months.

3. Neutrophils. Neutrophils are circular cells like
eosinophils (Fig. 6). These cells are also called as het-
erophils. They have cytoplasmic granules. The gran-
ules are circular and stained dim red. Cytoplasm is
stained light red.

In June, the mean diameter of neutrophils was
1 0. 1 5 pm, (± 1 . 1 9 standard deviations, with a range of
9.15-13.42 pm). In July, the mean diameter was 10.49

10/u.n^

Figure 4. Erythrocytes and an eosinophil.

Figure 5. Erythrocytes and a basophil.

2001

Asiatic Herpetological Research

Vol. 9, p. 125

Table 3. Erythrocyte dimensions of Lacerta rudis bithynica together with the standard deviations in August. EL:
Erythrocyte length; EW: Erythrocyte width; ES: Erythrocyte size; NL: Nucleus length; NW: Nucleus width; NS;
Nucleus size.

EL (urn)

EW (urn)

EL/EW

ES (urn2)

NS/ES

Max

18.30 ±0.76

9.76 ± 0.48

2.33 ±0.12

140.20 ±8.96

0.30 ± 0.02

Min

12.81 ±0.76

6.71 ± 0.48

1.57 ±0.12

73.60 ± 8.96

0.12 ±0.02

Mean

15.71 ±0.76

8.24 ± 0.48

1.91 ±0.12

101.72 ±8.96

0.18 ±0.02

NL (urn)

NW (pm)

NL/NW

NS (pm2)

Max

7.93 ± 0.51

4.27 ± 0.31

2.40 ± 0.20

24.53 ± 2.20

Min

4.88 ± 0.51

3.05 ±0.31

1.28 ±0.20

11.68 ±2.20

Mean

6.48 ±0.51

3.74 ±0.31

1.74 ±0.20

19.05 ±2.20

Table 4. Leukocyte and thrombocyte measurements of L rudis bi/bynica with the standard deviations in June.

Lym-
phocyte
(Mm)

Monocyte
(Mm)

Neutrophil
(Mm)

Basophil
(Mm)

Eosinophil
(Mm)

Thrombo-
cyte
Length
(Mm)

Throm-
bocyte
Width
(Mm)

Max

8.54 ± 0.81

12.81 ±

13.42 ±

9.15±

15.25 ±

7.32 ± 0.49

4.88 ±

1.23

1.19

0.61

1.71

0.54

Min

4.27 ±0.81

9.32 ± 1.23

9.15 ± 1.19

7.32 ±
0.61

9.93 ± 1.71

4.88 ± 0.49

3.05 ±
0.54

Mean

6.12 ±0.81

11.10±

10,15 ±

8.55 ±

12.82 ±

6.12 ±0.49

3.72 ±

1.23

1.19

0.61

1.71

0.54

Table 5. Leukocyte and thrombocyte measurements of L. rudis biibynica w\tk the standard deviations in July.

Lym-
phocyte
(Mm)

Monocyte
(Mm)

Neutrophil
(Mm)

Basophil
(Mm)

Eosinophil
(Mm)

Thrombo-
cyte
Length
(Mm)

Throm-
bocyte
Width
(Mm)

Max

7.32 ± 0.41

15.25±

12.20 ±

9.15±

15.25±

7.32 ± 0.52

4.88 ±

1.5.0

0.97

0.24

1.25

0.42

Min

6.10 + 0.41

9.15 ± 1.50

8.54 ± 0.97

8.54 ±
0.24

10.98 ±
1.25

6.10 ±0.52

3.66 ±
0.42

Mean

6.62 ±0.41

11.46 ±

10.49 ±

9.02 ±

13.29 ±

6.62 ±0.52

4.08 ±

1.50

0.97

0.24

1.25

0.42

Vol. 9, p. 126

Asiatic Herpetological Research

2001

Table 6. Leukocyte and thrombocyte measurements of L rudis bithynicamVn the standard derivations in August.

Lym-
phocyte
(Mm)

Monocyte
(urn)

Neutrophil
(pm)

Basophil
(pm)

Eosinophil
(pm)

Thrombo-
cyte
Length
(pm)

Throm-
bocyte
Width
(pm)

Max

9.15 ±0.58

15.25±

12.20±

10.37 ±

14.03 ±

7.32 ± 0.35

5.49 ±

1.29

0.97

0.45

1.37

0.49

Min

6.10 ±0.58

9.15± 1.29

8.54 ± 0.97

7.93 ± 0.45

12.20 ±
1.37

5.49 ± 0.35

3.05 ±
0.49

Mean

6.53 ± 0.58

11.21 ±

10.77 ±

9.00 ± 0.45

13.80 ±

6.33 ± 0.35

4.22 ±

1.29

0.97

1.37

0.49

um (±0.97, 8.54-12.20 pm). In August, the mean
diameter was 10.77 um (±0.97, 8.54-12.20 um).

There were no significant differences in neutrophil
diameters between females and males. Based on
Tables 4, 5, 6 and Fig.8, it appears that there was little
monthly variation in diameter of neutrophils during
the three months.

4. Monocytes . Monocytes are round cells with round
nuclei. The cytoplasm is stained blue and the nucleus
is stained purple by Wright's technique. The mono-
cyte's cytoplasm is more abundant than lymphocyte's
cytoplasm. Nuclei vary in shape (Fig. 7). Nuclei may
be nodular, but they are not lobular like granulocytes.
Sometimes nuclei are horseshoe-shaped.

In June, the mean diameter of monocytes was
1 1.10 um, (±1.23 standard deviations, with a range of
9.32-12.81 pm). In July, the mean diameter was 1 1.46
pm (±1.50, 9.15-15.25 pm). In August, the mean
diameter was 1 1 .2 1 um (± 1 .29, 9. 1 5- 1 5.25 pm).

There were no significant differences in monocyte
diameters between females and males. Tables 4, 5, 6
and Fig. 8 show that there was little monthly variation
in the diameter of monocytes during the three months.

5. Lymphocytes . Lymphocytes are round cells like
monocytes, but smaller (Fig. 7). The nuclei contain
many parts of the cell. The nucleus is stained purple,
cytoplasm, rarely seen, is stained blue.

In June, the mean diameter of lymphocytes was
6.12 um, (±0.81 standard deviations, with a range of
4.27-8.54 um). In July, the mean diameter was 6.62
pm (±0.41, 6.10 - 7.32 um). In August, the mean
diameter was 6.53 um (±0.58, 6.10-9.15 pm).

There were no significant differences in lympho-
cyte diameters between females and males. Based on
Tables 4, 5, 6 and Fig. 8, it appears that there was little
monthly variation in diameter of lymphocytes during
the three months.

III. Thrombocytes

Thrombocytes are small cells like lymphocytes, but
they are oval and smaller than lymphocytes (Fig. 9).
Their nuclei are highly chromophilic and stained pur-
ple. Cytoplasm is rarely seen.

In June, the mean length of thrombocytes was 6.12
pm (±0.49 standard deviations, with a range of 4.88-
7.32 pm). In July, the mean length of thrombocytes

~^~

Jfc

¥ ^SfeP^ Neutrophil

Erythrocytes

, * f

\0f*-*

Lymphocyte

lO/*""

Figure 6. Erythrocytes and a neutrophil.

Figure 7. Erythrocytes, a monocyte and a lymphocyte.

2001

Asiatic Herpetological Research

Vol. 9. p. 127

□ June
■ July
E3 August

lymphocyte monocyte neutrohpil basophil eosinophil

Leukocytes

Figure 8. Leukocyte diameters of Lacerta rudis bithynica over three months.

♦

Thrombocytes

Ery throe ytes

\Of*-r

Figure 9. Erythrocytes and thrombocytes.

6 12

ml-

662

1408

6.33

3.72

I4-

■

I Length
I Width

June

August

Figure 10. Thrombocyte sizes of Lacerta rudis
bithynica over three months.

was 6.62 um (±0.52, 6.10-7.32 urn). In August, the
mean length of thrombocytes was 6.33 um (0.35,
5.49-7.32 um).

In June, the mean width of thrombocytes was 3.72
(±0.54 standard deviations, with a range of 3.05-4.88

um). In July, the mean width of thrombocytes was
4.08 um (±0.42, 3.66-4.88 um). In August, the mean
width of thrombocytes was 4.22 um (0.49, 3.05-5.59
Mm).

There were no significant differences in thromb-
ocytes sizes between females and males. Based on
Tables 4, 5, 6 and Fig. 10, it appears that there was lit-
tle monthly variation in thrombocytes sizes during the
three months.

Discussion

Investigations carried out by various authors (Hart-
man and Lessler, 1964; Szarski and Czopek, 1966;
Saint Girons, 1970) reported that the sizes of the
erythrocytes vary in members of the four orders of
reptiles. Within the class Reptilia, the largest erythro-
cytes are seen in Sphenedon punctatus, turtles and
crocodilians. The erythrocytes of lizards vary greatly
in size depending on the family and sometimes even
within one family (Saint Girons, 1970). The smallest
erythrocytes are found in the lizard family Lacertidae
(Saint Girons, 1970).

In the present study, erythrocyte morphology and
the results of erythrocytes sizes (Table 1, 2 and 3) are
agreement with the other results carried out by Hart-
man and Lessler (1964), Szarski and Czopek (1966),
and Saint Girons (1970).

In one of the studies on the leukocytes of the
rough tail Gecko Cyrtopodion scabrum, a bright-field
and phase-contrast study Cannon et al. (1996),

Vol. 9, p. 128

Asiatic Herpetological Research

2001

reported that the neutrophils were not observed, but
the other leukocytes were observed.

Another study on morphological observations on
the erythrocytes, leukocytes and thrombocytes of blue
tongue lizards by Canfield and Shea (1988) reported
that all types of leukocytes were observed.

Saint Girons (1970) and Canfield and Shea (1988)
divided granulocytes into neutrophils, basophils and
eosinophils on the basis of light microscopy. However
Cannon et al. (1996) divided granulocytes into baso-
phils and eosinophils on the basis of bright-field and
phase-contrast microscopy.

Heady and Rogers (1962) divided leukocytes into
neutrophils, small acidophils, eosinophils, lympho-
cytes and monocytes on the basis of light microscopy
in Pseudemys elegans. They, except for monocytes,
gave the sizes of leukocytes and reported that eosino-
phils and neutrophils were numerous than the other
leukocytes. Taylor and Kaplan (1961) also divided
leukocytes into neutrophils, basophils, eosinophils,
lymphocytes and monocytes on the basis of light
microscopy in turtles.

In this study, it appears that on the basis of light
microscopic findings there are three main types of
granulocytes and two types of agranulocytes in L.
rudis bithynica and also the size of all kinds of leuko-
cytes are given in Table 4, 5 and 6.

In the present study, the descriptions and sizes
(Table 4, 5 and 6) of thrombocytes are comparable to
other descriptions. Canfields and Shea (1988)
reported that thrombocyte morphology at the light
microscopic level is influenced by the degree of
aggregation and degranulation. Saint Girons (1970)
reported that thrombocytes are small, oval cells char-
acterised by elongate, centrally located highly chro-
mophilic nuclei. The cytoplasm is almost colourless
(faintly acidophilic) and hence difficult to see in a
blood smears. Taylor and Kaplan (1961) reported the
same findings in turtles.

Literature Cited

Altland, P. D. and E. C. Thompson. 1958. Some fac-
tors affecting blood formation in turtles. Proceedings
for the Society of Experimental Biology and Medi-
cine. 99:456-459.

Baker, E. G. S. and L. E. Kline. 1932. Comparative
erythrocyte count of representative vertebrates. Pro-
ceedings of the Indiana Academy of Sciences 41:417-
418.

Canfield, P. J. and G. M. Shea. 1988. Morphological
observations on the erythrocytes, leukocytes and

thrombocytes of blue tongue lizards (Lacertilia: Scin-
cidae, Tiliqua). Anatomia, Histologia, Embryologia.

17:328-342.

Cannon, M. S., D. A. Freed and P. S. Freed. 1996. The
leukocytes of the roughtail gecko Cyrtopodion sca-
brum: a bright-field and phase-contrast study. Anato-
mia, Histologia, Embryologia. 25:1 1-14.

Charipper, H. A. and D. Davis. 1932. Studies on the
arneth count. A study of the blood cells of Pseudemys
elegans with special reference to the polymorphonu-
clear leukocytes. Q. J. Experimental Physiology.
21:371-382.

Cline, M. J. and T. A. Waldmann. 1962. Effect of tem-
perature on red cells in the alligator. Proceedings for
the Society of Experimental Biology and Medicine.
111:716-718.

Duguy, R. 1970. Numbers of blood cells and their
variations. Pp. 93-104. In Gans (ed.), Biology of the
Reptilia, Vol. 3, Morphology C. Academic Press, New
York.

Engbretson. G. A. and V H. Hutchinson. 1976. Eryth-
rocyte count, hematocrit and haemoglobin content in
the lizard Liolaemus multiformis. Copeia 1976:186.

Goin, C. J. and C. G. Jackson. 1965. Haemoglobin
values of some amphibians and reptiles from Florida.
Herpetologica 21:1 45-146.

Haggag, G., and et al. 1966. Hibernation in reptiles II.
Changes in blood glucose, haemoglobin, red blood
cells count, protein and nonprotein nitrogen. Compar-
ative Biochemistry and Physiology. 17:335-339.

Hartman, F. A., and M. A. Lessler. 1964. Erythrocyte
measurements in fishes, amphibians and reptiles. Bio-
logical Bulletin. 126:83-88.

Heady, J. M. and T. E. Rogers. 1963. Turtle blood cell
morphology. Proceedings of the Iowa Academy of
Sciences. 69:587-590.

Hutchinson, V H. and H. Szarski. 1965. Number of
erythrocytes in some amphibians and reptiles. Copeia
1965:373-375.

Hutton, K. E. 1960. Seasonal physiological changes
in the red eared turtle Pseudemys scripta elegans.
Copeia 1960:360-362.

Hutton, K. E. 1961. Blood volume, corpuscular con-
stants and shell weight in turtles. American Journal of
Physiology 200:1004-1006.

Newlin, M. E. and R. E. Ballinger. 1976. Blood hae-
moglobin concentration in four species of lizards.
Copeia 1976:392-394.

Saint Girons, M. C. 1970. Morphology of the circu-
lating blood cells. Pp. 73-91. In Gans (ed.), Biology of

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the Reptilia, Vol. 3, Morphology C. Academic Press,
New York.

Szarski, H. and G. Czopek. 1966. Erythrocyte diame-
ter in some amphibians and reptiles. Bulletin de
l'Academie Polonaise des Science. Classe 2. Serie des
Sciences Biologiques. 14(6):437-443.

Taylor, K. and H. M. Kaplan. 1961. Light microscopy
of the blood cells of Pseudemyd Turtles. Herpetolog-
ica 17:186-192.

2000

Asiatic Herpetological Research

Vol. 9, pp. 130-138

Buccopharyngeal Morphology and Feeding Ecology of Microhyla ornata

Tadpoles

Muhammad S. Khan

7740 NW 47th St., Lauderhill, FL 33351 USA

Abstract.- The oropharyngeal morphology of Microhyla ornata tadpoles is described. Anatomical peculiarities
are correlated to feeding ecology. Microhylid anatomical features are discussed and compared with ranoid
tadpoles.

Key words.- Amphibia, Salientia, Microhylidae, Microhyla ornata, larval oropharyngeal morphology, feeding
ecology.

Introduction

The common southeast Asian narrow-mouth frog
Microhyla ornata is widely distributed throughout
Punjab, Sindh. N.W.F.P. and Azad Kashmir (Khan,
1974; 1979; 1988; Khan andTasnin, 1987). Normally,
its small size makes it inconspicuous and difficult to
detect in the field. However, after a summer heavy
downpour, the characteristic rasping call of Microhyla
ornata is easily distinguishable from rest of the local
amphibians (Khan and Malik, 1987b). Microhyla
ornata readily takes refuge under vegetation, leaf lit-
ter, logs, stones, in holes and fissures in the ground,
and often is mistaken as a juvenile of some larger spe-
cies.

At mid-monsoon, M. ornata invades large water
bodies, which, by this time, are filled with water and
have developed thick planktonic growth. Solitary
males perch well away from water among marginal
vegetation to call. Eggs are laid in patches of jelly,
which float at water surface as "egg-rafts" (Khan,
1982b).

The present study describes oropharyngeal morphol-
ogy of Microhyla ornata tadpole, at Stage 35 and cor-
relate it to the tadpole's feeding ecology. Moreover, it
is compared with already known morphologies of
sympatxic tadpoles belonging to the genera Bufo and
Rana (Khan and Malik, 1987a; Khan and Mufti,
1994b, 1995).

Material and Methods

Tadpoles for the present study were collected during
the summers of 1986-88 from different localities
along the northwestern border of Rabwah City (Khan
and Malik, 1987b). They were netted at midstream
with a hand net. Microhyla ornata tadpoles from
Ghakkhar, District Gujranwala, Punjab, Pakistan

(Khan, 1974) and Azad Kashmir (Khan, 1979), were
used as comparative material.

Collection, preservation and storage methodology
followed Khan (1982b), while surgical procedures
and descriptive terminology are from Khan and Malik
(1987a) and Khan and Mufti (1994b, 1995), except
that for M. ornata tadpole, the lateral cuts through
buccopharyngeal walls, must pass through mid-eye,
unlike ranoid tadpoles where cuts pass below the eye.
Fine particulate mucilage-trapped material accumu-
lating in the filter cavities is cleared by a jet of water
from an ordinary eyedropper. Drawings of the buc-
copharyngeal surfaces were made with the help of
camera lucida.

Tadpoles at Stage 35 were selected, since at this
stage, they have already attained maximum size and
their characteristic organs are fully developed and
functional. The tadpole at this stage is voraciously
feeding and its digestive system is functioning at its
full capacity. Shortly after this stage, metamorphic
changes start occurring.

For identification of Stage 35, tadpoles were com-
pared with Khan's (1965) table of normal develop-
ment. Data for present study are recorded from 10
specimens.

Description

External morphology

The tadpole's body is perfectly streamlined; the head
is dorsoventrally depressed while its belly is laterally
compressed and oval in dorsal profile. The snout is
countersunk, displacing mouth anterodorsally. The
tail is more than twice the length of the head and
body. Broad caudal fins narrow abruptly in the poste-

2000

Asiatic Herpetological Research

Vol. 9, p. 131

buccal floor arena
(BFA)

preglottal

papilla

laryngeal disc

ventral
vellum

filter cavity

nfralabial papillae
tongue analage

buccal pocket

lottis

BFA papillae

buccal pocket
canal

trachea

esophagus

Morphology of surgically exposed buccopharyngeal floor of Microhyla omata\adpo\e at Stage 35 (scale

Figure 1.
1 mm).

nor half of the tail, passing into a delicate long flagel-
lum.

The body is widest at the level of laterally dis-
posed small eyes. The nostrils are imperforate; their
position is marked by heavily pigmented anterolateral
pits lying just anterior to eyes. The mid- ventral spira-
cle is close to posterior ventral end of the abdomen,
with a distinct prespiracular valve (Khan 1982b).

The horizontal mouth has a median U-shaped cleft in
the middle of the lower lip which remains perma-
nently open (Khan and Mufti, 1994a, Fig. 2). Pres-
ence of iridiocytes in the abdominal wall, give it a
characteristic silvery shine, which is lost within two to
three weeks, on preservation. A median dorsal band of
melanophores covers the brain and extends onto the
base of eyes and the nasal pits.

Measurements (in mm). Body length 5.2-5.8; tail
length (including flagellum) 12.6-13.7; total length
17.8-18.9; greatest breadth of body (at the level of
eyes) 2.7-2.9; greatest depth of body (at level of spira-

cle) 3.2-3.4; interorbital space 2.3-2.4; internarial
space 0.5-1.95; tail muscle height (at base) 1.7-2.0;
tail fin height (at midtail) 4-4.4; length of tail flagel-
lum 2.0-2.3.

Internal Morphology

Buccal region. Khan and Mufti (1994b; 1995) distin-
guished a tadpole's buccal cavity in two functional
units: anterior food gleaning part and posterior food
retrieving part. The dorsoventrally depressed head and
peculiar position of the mouth in M. ornata tadpoles
have affected the form and shape of the tadpole's buc-
cal cavity; reducing the food gleaning part and widen-
ing the food retrieval part which occupies most of the
buccal region.

Ventral buccal (Fig.l). The floor of the food glean-
ing part consists of vertical U-shaped prelingual
chamber which opens out through mouth at snout top
and is lined by a series of three simple infralabial
papillae. Posteriorly, it opens in food retrieval part of

Vol. 9, p. 132

Asiatic Herpetological Research

2000

postnarial ridge

prenarial papilla

nans

buccal roof arena
(BRA)

postero-
lateral
BRA ridge

narial papilla
BRA papilla

pressure
cushions

1 mm

buccal
roof

glandular
area

I — dorsal
vellum

esophagus

Figure 2. Morphology of surgically exposed buccopharyngeal floor o\Microhyla oma/a\adpo\e at Stage 35 (scale
1 mm).

the buccal. A non-papillated tongue analage, a conical
thickening which is broader anteriorly pointed poste-
riorly, guards the opening of the food gleaning part
into the food retrieval part. The spacious food retrieval
part forms the main buccal cavity. The buccal floor
arena (BFA) is rectangular, laterally raised with a
median shallow passage. A group of 12-14 fine
tipped, large, flat BFA papillae lie on lateral sides of
anterior end of trachea. A pair of smaller midpocket
papillae lie at the level of mid-pockets. A buccal
pocket is a long, narrow, club shaped longitudinal slit
running anteroposteriorly on each lateral side of the
BFA with a posterior narrow canal connecting it with
the pressure cushions. Anterior, wider, parts of buccal
pockets have 3-6 pre- and 5-8 post pockets, small,
conical papillae.

The trachea is a long cylindrical pipe that extends
medially to the base of the BFA, carrying the glottis
far anterior to the ventral velum and dividing it into
lateral halves. The thin lipped glottis is 100%
exposed, lies on a bulbous laryngeal disc, and is

guarded by a long preglottal papilla which is tipped
left.

The broad ventral velum has a strong spicular sup-
port. It covers about l/2-l/4th of the underlying bran-
chial baskets and consists of three distinct long and
deep filter cavities. The free margin of the velum is
smooth, with a single broad projection above third fil-
ter plate and is covered by a narrow strip of minute
secretory pits. Rows of melanophores run along the
lateral sides of the buccal arena and are aggregated on
the sides of the tongue analage; a row runs along lat-
eral sides of trachea.

Dorsal buccal (Fig. 2). The food gleaning part of the
buccal is roofed by a broad prenarial arena, which has
a median V-shaped depression with an anterior pre-
narial papilla and several pustules. The position of the
imperforate naris is marked by a shallow depression
from which a flat ribbon like twisted narial papilla
hangs down in the buccal cavity. A thin delicate, nar-
row, papillated, postnarial ridge dorsally delimits nar-
ial region and the food gleaning part of the buccal.

2000

Asiatic Herpetological Research

Vol. 9, p. 133

laryngeal disc

secretory ridg

glottis
trachea

Figure 3. Underside of ventral velum showing details of filter cavities (diagramatic).

The food retrieval part is roofed by a spacious,
broader than long, buccal roof arena (BRA), which is
featured mainly by a distinct posterolateral BRA
ridge, the free margin of which is cut into flat fine
tipped BRA papillae, increasing in length from out
smallest, to inside longest, with blunts on their inner
sides, while smaller are without blunts. The BRA sur-
face is pustular.

The broad glandular area of the buccal roof is
divided into lateral rectangular halves and is covered
with dense minute secretory pits. The dorsal velum
narrows gradually mesoid, at mid-BRA it staggers to
continue with that of other side.

Branchial region. Branchial baskets are more than
twice the length of the buccal. Three filter cavities are
distinct in each branchial basket. The first filter cavity
is largest while the third is smallest. The filter cavities
are always packed with fllocular matter. The third is
tilted outwards, partially blanketing the second. About
half to one third of the filter cavities are covered with
velum. A distinctly ridged oval torus (Fig. 3:1, II, III) is
present in each filter cavity. The subvelar surface is
profusely ridged with fine transverse secretory ridges

(Fig. 3) that run in line with toric ridges, which are
edged with fine secretory pores.

A tight filter ruffle cover the surface of filter
plates. The number of filter rows on filter plates vary
from minimum 9 on the 4th ceratobranchial to 23 on
second (Table 1 ). The filter ruffle is 3° dense with ter-
tiary foldings. Successive filter rows abut across fully
canopied deep filter canals. The filter ruffle covers
both sides of second and third filter plates (Fig. 3: cb2,
cb3).

Three pressure cushions are distinct on posterolat-
eral sides of the dorsal pharynx (Fig. 4). The first and
second are four times longer than broad. The first, and
outer-most, is continuous anteriorly with the buccal
pocket of its side through a buccal pocket canal.
Meanwhile, the third, innermost, is broadest and has a
median hook-like appendage. Deep ciliary groove
runs along posterior border of the pressure cushions
towards esophageal orifice along posterior sides of the
pressure cushions.

The lungs at Stage 35 are well developed, each
running along the dorsolateral sides of the abdominal
cavity, extending to the posterior end of the abdomen.
Anterior half of the lung is broad, with well-devel-

Table 1. Branchial elements of Microhyla ornata tadpoles at Stage 35 (Ant=anterior aspect; B=breadth; cb=cerato-
branchial; L=length; Post=posterior aspect). Data from 10 specimens, all measurements in mm.

Ceratobranchial

Filter plate

Filter rows/side

cb. 1
cb.2
cb.3
cb.4

3.5

1.8

3.7

1.5

2.5

1.4

1.9

1.2

Posterior

15-18

13-16

20-23

16-17

8-9

Vol. 9, p. 134

Asiatic Herpetological Research

2000

Table 2. Comparison of microhylid and ranoid tadpole.

Character

Microhylid

Ranoid

Head

Belly

Color

Tail length

Tail muscle

Tail fins

Tail tip

Spiracle

Belly wall

Mouth

Oral disc

Nostrils

Narial flap/papillae

Infralabial papillae

Lingual papillae

Postnarial papillae

Lateral ridge papilla

Median ridge

Buccal musculature

Branchial basket

Pharyngeal/buccal ratio

Prenarial ridge

BRA/BFA papillae

Glottis

Glottal disc

Glottal position

Glottal papilla

Trachea

Secretory tissue

Subvelar secretory tissue

Filter cavities

Maximum filter rows

Position of tori

Filter cavity depth

depressed

compressed

transparent

2-3 times body

narrow

broad

produced in a flagellum

median ventral

silver-shiny

antero-dorsal

absent

imperforate

flap

simple

absent

small on a membrane

absent

poor

longer than broad

75%

absent

flat fine tipped

form membranes

1 00% exposed

bulbous

anterior to vellum

present

long

single celled

ridged

15-23

all filter cavities

as long as deep

depressed

Drab + pattern

2-2.5 times

broad

moderate

round/pointed

dextral

transparent or drab

anterior/antero-ventral

present

perforate

papillae

palmate

present

long solitary

present

well developed

broader than long

45-50 %

pustules or ridge

short forked blunted

distinct no membranes

not or partiall exposed

concealed

posterior to vellum

absent

multicellular

ridged/pitted

2/3

10-14

absent or first cavity

longer than deep

2000

Asiatic Herpetological Research

Vol. 9, p. 135

buccal pocket canal

pressure cushions

pressure cushion
hook

ciliary
groove

Figure 4. Morphologyofposterolateralpartofdorsal buc-
copharyngeal region, showing details.

oped air sacs, it gradually narrows down caudally and
its terminal part is a dark pigmented cord.

Ecological Correlates

The adult Microhyla ornata is sharply contrasted from
its sympatric ranoids in its choice of breeding site and
mode of egg deposition. Usually its breeding sites are
deep ponds, which develop rich planktonic bloom
during summer. Such sites are quite restricted and rare
in temperate Punjab, unlike sympatric ranoids who
breed in temporal sites that dry in the summer. Inter-
mittent unpredictable rains or irrigation seepage saves
such populations of tadpoles (Khan and Malik,
1987b).

Table 1 and 2 summarize peculiar features of M.
ornata tadpole necessary to suit its lentic habits: per-
fectly streamlined transparent body and long broad
finned tail with independently vibratile terminal fla-
gellum, lateral eyes, median ventral spiracle, abdomi-
nal silver-shine, are adaptations to nektic habits of the
M. ornata tadpole. A school of tadpoles swimming at
midstream is almost invisible from above the water
due to tadpole's transparent bodies. Meanwhile
abdominal coloration makes them invisible in water
since the water surface viewed from inside appears
shiny due to reflection of light from water. A vibrating

tail flagellum and the jet of water from median ventral
spiracle supports the microhylid tadpole at mid-
stream, while its buoyancy is maintained by dorsally
placed air filled lungs. A long broad finned tail helps
the tadpole to react quickly to any stimulus including
enemies and food in the water column above and
below.

Microphagous tadpoles are characterized by an
exaggerated oropharyngeal region (Wassersug, 1980).
All oropharyngeal modifications distinguishing
microhylid tadpoles from bufonids and ranids are
adaptations to microphagy: reduction of infralabial
cartilage, corresponding reduction in prelingual and
prenarial arenas: displacement of mouth to snout top;
simple infralabial papillae; absence of lingual papil-
lae; membranous postnarial and BRA papillary
ridges; divided fine pitted buccal glandular zone; fine
ridged subvelar and toric glandular surfaces; broad
medially divided spicularly supported ventral velum;
compact thick filter ruffle on broad and deep filter
plates; deep filter cavities with distinct tori. These
microhylid structures are part of obligate micropha-
gus filter feeding technology. Similarly long tubular
trachea, bulbous laryngeal cartilage, distinct anteri-
orly displaced glottis with a preglottal papilla, and
dorsal, long, air filled lungs are not only efficient parts
of an efficient pulmonary aeration system, but at the
same time, serve to maintain hydrostatic balance of
the tadpole at mid-stream position. The complex mor-
phology of pressure cushions and their connection
with buccal pockets are a part of a system which
maintains a sustained drainage of excessive buccal
water in pressure cushions, providing necessary tur-
gidity during each stroke of bucco-pharyngeal pump-
ing. Turgid cushions act as pharyngeal valves in the
process and play key role in sustained pumping of
food-laden water current in buccopharyngeal passage.

Morphologically, the Microhyla ornata tadpole is
a perfect model suited for sustained midstream swim-
ming. It has all the necessary oropharyngeal technol-
ogy to filter and feed on the planktonic bloom in
deeper ponds in temperate Punjab.

Microhylid Tadpole Characters

The microhylid tadpole is ranked as Type 2 in Orton's
(1953) classification. It markedly differs in morpho-
logically from ranoid Type 4 macrophagus tadpole
(Table 2). Microhylid is the only tadpole, in Punjab
riparian ecosystem, occupying unique midstream
feeding niche, while rest of the sympatric ranoid tad-
poles are bottom grazers and in no way compete with
microhylid tadpole. Distinguishing morphological
characteristics of Microhyla ornata tadpoles are:

Vol. 9, p. 136

Asiatic Herpetological Research

2000

External morphology

1. Transparent body and tail, silver-shine on abdomen.

2. Head dorso-ventrally depressed, body and tail later-
ally compressed.

3. Antero-dorsal mouth, without keratinized oral disc
and other associated organs.

4. Eyes prominent, laterally disposed on head.

5. Tail broad finned, its tip produced into a terminal
vibratile flagellum.

6. Spiracle median-ventral, mid-abdominal, squarish
opening, with a prespiracular flap.

7. Anal tube straight, median-ventral anal opening.

8. Schools of tadpoles swim at midstream schools,
never rest at bottom. Capable of making spontaneous
movements from midstream to darker parts of the
pond to avoid intruders, shortly reappearing at the
same midstream site.

Internal morphology

9. Opercular chamber extending to vent.

10. Smooth broad ventral vellum, divided into right
and left halves.

1 1 . Long pipe like trachea carries, bulbous glottal car-
tilage,

for forward in buccal cavity. Glottis thin lipped, 100%
exposed.

12. Preglottal papillae guarding glottis.

13. No lingual papillae.

14. No lateral ridge papilla and median ridge.

15. Pharyngeal region exaggerated, about 75 % of
oropharyngeal region, branchial baskets large with
distinct, deep filter cavi ties, which are usually full of
fllocular matter in dissected tadpoles.

16. Tight filter mesh, maximum number of filter rows
13-23 on either sides of the ceratobranchials.

17. Branchial food traps with microscopic openings of
secretory glandular tissue borne on fine parallel ridges
forming distinct crescentric torus in each branchial
cavity.

18. Imperforate nares with a foliaceous broad narial
palp, descending into the buccal cavity.

19. Ceratohyal with a ventrally directed lateral arm,
and an an tero-posterior median arm.

20. Broad based BRA and BFA papillae, which usu-
ally coalesced to form fine membranes.

21. Narrow lateral buccal pockets with distinct con-
nection with pressure cushions, forming an elaborate
system to control function of pressure cushions.

Microhylids are phylogenatically connected with
ranoids through tadpoles with intermediate morpholo-
gies like Psedohemisus granulosa (Wassersug, 1984)
and Otophryne robusta (Wassersug and Pyburn,
1987).

Discussion

Independent vibratile distal caudal flagella are charac-
teristic of mid-stream swimming microphagus tad-
poles (Wassersug, 1980, 1989; Wassersug and Sperry,
1977; Nishikawa and Wassersug, 1988, 1989; Hoff
and Wassersug, 1986). Maintenance of midstream
position is made possible by independent movements
of the caudal flagellum and ventrally directed continu-
ous water from the spiracle giving a sustained upward
thrust (Khan, 1982a, 1991). Apart from morphologi-
cal differences in structure of notochord and arrange-
ment of caudal nerves, microhylids and ranoids differ
in the site of generation of propulsive locomotory
waves. In ranoids, waves are generated at the tail; the
tip acts as a steer. In microhylids, waves are generated
at the end of the caudal flagellum.

The limited tail musculature and bulky form of the
amphibian tadpole restricts its movements so that it
cannot evade its potential enemies, fishes, niads, etc.,
(Khan and Mufti, 1994b; 1995). Amphibian tadpoles
rely on a reduced conspicuousness. Its drab spotted
pattern blends well against natural aquatic back-
ground with moderate to thick vegetation (Caldwell,
et al., 1981; Gatten et al., 1984; Kehr and Basso,
1990; Khan and Mufti, 1994 b. 1995). The microhylid
tadpole, which is exposed at midstream, solves this
problem differently. Transparency of its body reduces
its shadow at pond bottom and its abdominal shine
blends well against water surface reflecting sun rays,
making it invisible from inside pond as well from out
side, to its predators.

Microhyla omata has a larger buccal volume than
ranoids. It constantly pumps large amounts of water to
get food (Seale and Wassersug, 1979; Wassersug,
1980). Due to its specialized feeding habits, several
elements universally present in ranoid tadpoles are
missing in its oropharyngeal morphology: median
ridge, lateral ridge papillae and lingual papillae.
Moreover, papillae in the food retrieval part are a part
of particulate food guiding membrane rather particu-
late food retrieving sieves. Moreover, microhylids
have fine-ridged oral and branchial glandular system
with fine pits, a specialization to entrap finest particu-
late food. Deep filter cavities, long filter plates and
tight filter mesh are more a part of food retrieval sys-
tem rather respiratory in function (Wassersug and
Murphy, 1987). Particulate food filtering capacity of

2000

Asiatic Herpetological Research

Vol. 9, p. 137

filtering system is enhanced by development of a fine-
ridged torus in each filter cavity. A well-developed
pulmonary system with long tubular trachea, a totally
exposed guarded glottism, and inflated dorsal func-
tional lungs are adaptations to the midstream sus-
tained swimming and are efficient respiratory organs
(Khan, 1991).

Tadpoles of the microhylid genera Otophryne and
Psdudohemisus show both ranoid as well as micro-
hylid characteristics. Frogs of Family Microhylidae
and Ranidae have distinctive adult morphology, how-
ever definition of microhylid larva stands only on
imperforate naris, since the ranoids have always per-
forated naris (Wassersug, 1989).

Tadpoles of M. ornata from Pakistan differ in mor-
phological details from those collected from Thailand
by Inger (1985) in having longer infralabial papillae,
more filter rows, presence of postnarial membranous
ridge, BRA and BFA membranous papillae, mesially
divided dorsal buccal glandular zone, and a single
preglottal papilla. Moreover, no silver shine is
reported on the belly of tadpoles from Thailand, as is
reported from India (Rao, 1917; Azad Kashmir
(Khan, 1979) and District Jhang, Punjab Pakistan
(Khan 1982a). Flower (1899) reported morphological
differences among adults and larvae of this species
from Malay peninsula and Siam, while Liu (1950) has
reported dextral anal tube in tadpoles from China,
which is straight in Pakistani tadpoles. These morpho-
logical differences may refer to geographical races
within this widely distributed southeast Asian species
of narrow-mouth frogs.

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2001

Asiatic Herpetological Research

Vol. 9, pp. 139-141

Variation in Pelobates syriacus of Turkey

Ismail H. UGurta§

Science and Art Faculty, Uludag University, Department of Biology, Bursa, Turkey

Abstract.- Morphometric data and color patterns of Pelobates syriacus from different regions of Turkey were
compared. Pelobates syriacus from Edirne appear to be distinct in terms of color pattern. Pelobates syriacus from
Seydi§ehir appear to be distinct in terms of morphometric measurements.

Key words.- Pelobates syriacus, Turkey, morphometry, color pattern

BLACK SEA

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0 26

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MEDITERRANEAN

100

200 km

Figure 1 .The places where Pelobates syr/acushave been collected in Turkey. The key to the letters is found in
Table 2. Dark circles = The localities where we collected specimens. Open circles = The localities where speci-
mens were collected before. 1. Buyukdolluk, 2. Babaeski, 3. Altin§ehir, 4. Terkos, 5.Yassioren, 6. Adapazari, 7.
Sogutlu, 8. Karasu, 9. Poyrazli Lake, 10. £erkesji, H.Terme, 12. Karacabey, 13. Bursa, 14. §akran, 15. Bostanli,
16 Bornova, 17. Ivrindi, 18. Seydi§ehir,19. Karata§, 20. Iskenderun, 21. Van.

Pelobates syriacus Boettger 1889 was first described
from specimens collected from Hayfa-Israel. Later,
Mertens (1923) examined two specimens, found in
Belesuwar near the Azerbaijan and Iran border.
Because of their long and narrow skull and the round
and raised frontoparietals, he recognised them as
another subspecies, Pelobates syriacus boettgeri.

Other specimens from Macedonia were described
by Karaman (1928) as the subspecies Pelobates syria-
cus balcanicus. Miiller (1932) criticized this classifi-
cation based on the shape of the skull. He emphasised
that skull shape can be variable. Another subspecies

living around Tiflis, Georgia was named Pelobates
syriacus transcaucasicus by Delwig (1927). Accord-
ing to Gilsen (1937), this subspecies is intermediate in
morphology between P. s. boettgeri and P. s. syriacus,
and Terentjev and Chernov (1965) are convinced that
P. s. transcaucasicus is a synonym of the nominate
race. Furthermore, Eiselt and Schmidtler (1973) pro-
posed P. s. boettgeri as a synonym for P. s. syriacus.

The taxonomic status of Turkish Pelobates syria-
cus is poorly studied. Mertens (1953) described a
young sample collected in Van (Turkey) and sug-
gested that it was P. s. boettgeri. Zaloglu (1964) stud-

Vol. 9, p. 140

Asiatic Herpetological Research

2001

Table 1. Percentage of each color pattern in populations of Pelobates syriacus.

Population

Edirne

Istanbul

Adapazari

Samsun

Bursa

Balikesir-izmir

Seydi§ehir

Adana-lskenderun

Van

100

56.66

40.00

3.34

11.54

65.38

68.42

21.05

10.53

73.92

21.73

4.35

15.00

75.00

5.00

7.70

92.30

66.00

34.00

14.38

14.28

71.44

ied Pelobates syriacus from the Turkish region of
Izmir. He made an osteological comparison of chra-
cters used to differntiate the subspecies of P. syriacus.
He pointed out that these characters showed a wide
range of variation, and refrained from classifying the
Izmir P. syriacus into subspecies.

In this study, samples (77 male, 57 female, 25
juveniles, 35 larvae) collected from 9 different areas
(Istanbul, Adapazari, Samsun, Bursa, Izmir-Balikesir,
Seydi§ehir, Adana, Van) were investigated to obtain
more information about variation in Pelobates syria-
cus living in Turkey (Fig. 1 ).

Results and Discussion

There are four kinds of dorsal patterns on Pelobates
syriacus (Fig. 2). The dorsal patterns are not sexually
dimorphic. The percentage of dorsal patterns are fol-
lows: A: 36.08 %, B: 25.95 %, C: 13.92 %, D: 24.05
%. When all the samples are compared, it is seen that
the dorsal pattern of Edirne population is different
than the other populations (Table 1 ).

All of the adults from Edirne have big green dots
narrow raised surrounding on the back. The back-
ground color of the back is light yellow-green. On this
background there are connected raised surrounding
with brown-green dots. Those dots also appear on all
extremities. Furthermore, on the background color,
and within the brown-green dots, are yellow and red
dots. These dots are missing on the upper side of the
front fingers. Dots on the body become smaller on the
side. All the Edirne population have C pattern type
(Fig. 2).

In terms of morphological measurements, Pelo-
bates syriacus populations living in Turkey are very
similar. Here, the exception is the Seydi§ehir popula-
tion which have shorter indices of tibia length (Table
2).

The differences in the color patterns of adult ani-
mals from the Edirne population and the morphomet-

Figure 2. Types of color patterns in Pelobates syria-
cus. A: Spots are irregularly distributed and isolated.
B: Two or more spots combine to form irregular islets.
C: The edges of the spots are wavy and connected by
thin bands. D: Spots form lengthwise bands.

2001 Asiatic Herpetological Research Vol. 9. p. 141

ric differences of the Seydi§ehir population should be
explored using biochemical data in order to determine
whether the Pelobates syriacus from these regions
represent different taxa.

Literature Cited

Boettger, O. 1889. Ein neuer Pelobates aus Syrien.
ZoologischerAnzeiger 12:144, 1889.

Delwig, W. 1927. Eine neue Art der Gattung Pelo-
bates Wagler aus dem zentralen Transcaucasus. Zool-
ogischer Anzeiger 75, Heft 1/2:24.

Eiselt, J. and J. F. Schmidtler 1973. Froschlurche aus
dem Iran unter Beriick chtigung ausser-iranischer
Populationgruppen. Annalen Naturhistorischen Muse-
ums in Wien 77:181-243.

Gislen, T. 1937. On the history of evolution and distri-
bution of the European pelobatids. Zoogeographica
3:119.

Karaman, S. 1928. Contribution a 1' Herpetologie de
Jugoslavja. Bulletin de la Societe Scientifique de Sko-
plje 4. 2:129.

Mertens, R. 1923. Beitrage zur Kenntnis der Gattung
Pelobates Wagler Senckenbergiana 5(3/4): 1 1 8- 1 28.

Mertens, R. 1953. Weiters zur Kenntnis der Herpeto-
fauna der Asiatischen Tiirkei. Istanbul Univeritesi
Fakultesi Mecmausi Seri B18: 373-375.

Muller, L. 1932. Beitrage zur Herpetologie der
Sudosteuropaischen Halbinsel. Teil I, Zoologischer
Anzeiger 100, Heft 11/12:301-309.

Teretnjev, P. and S. A. Chernov. 1965. Key to
Amphibians and Reptiles. Israel Program for Scien-
tific Translations, Jerusalem.

Zaloglu, §. 1964. Ege bolgesinde bulunan Pelobates
syriacus'un morfoloji, osteoloji ve biyolojisi ile ilgili
ara§tirmalar. Ege Univeritesi. Fen Fakultesi ilmi Rap.
Ser. No. 16:1-50.

2001

Asiatic Herpetological Research

Vol. 9, pp. 142-144

Translation: A New Species of the Turtle Genus Ctav#(Testudoformes:

Testudinidae)

Ming-tao Song

Shaanxi Institute of Zoology, Xi 'an Province, China
(Current Address: Northern west Institute of Endangered Animals, Xi'an, China 710032)

Original English Abstract.- Cuora pani, sp. nov. (Figs. 1, 2). Holotype, SIZ 80170, an adult male. Allotype, SIZ
80171, an adult female. All of these type specimens were collected from Xujiaba (alt. 420 m) of Pingli County in
Shaanxi Province, on June 17, 1981 by the author, and are preserved in the Shaanxi Institute of Zoology. This
new species is similar to Cuora yurmanensis (Boulenger), but differs from the latter in having the median keel
not conspicuous and no lateral keels; a brown narrow band extending from behind eye to the neck; the suture
between gulars 1.5 times as long as that between humerals and much shorter than those between pectorals and
between anals, but longer than that between femorals; the plastron yellow, with black sutures; and the limbs
brown, without any markings.

Song, M. T. 1984. A New Species Of The Turtle Genus Cuora (Testudoformes: Testudinidae). Acta Zootaxonom-
ica Sinica 9(3):330-332. (In Chinese with English abstract)

There are six species belonging to the genus Cuora
Gray, 1855 (Pritchard, 1967; Sichuan Institute of
Biology, 1977), that are distributed in Thailand, Cam-
bodia, Malaysia, Indonesia, the Philippines, and
southern provinces of China.

In 1981, two Cuora specimens were found during a
survey of Mt. Dabashan, Shaanxi Province. These
specimens are different from the six known species
and are recognized as a new form. Its description is
given below.

Cuora pani, sp. nov. (figs. 1-2)

The holotype, SIZ 80170, is an adult male. The allo-
type, SIZ 80171, is an adult female. Both of the type
specimens were collected from Xujiaba (alt. 420 m)
of Pingli County in Shaanxi Province, on June 17,
1981 by the author, and are preserved in the Shaanxi
Institute of Zoology.

Diagnosis

The new species is similar to Cuora yurmanensis
(Boulenger), but differs from that species by having a
median keel that is not conspicuous and no lateral
keels; a brown narrow band extending from behind
the eye to the neck; the seam between the gulars 1.5
times as long as the seam between the humerals and
much shorter than those between pectorals and
between anals, but longer than that between femorals;
the plastron yellow, with black markings along the
seams; the limbs are brown and without any mark-
ings.

Description of Holotype

The carapace is flattened, with an inconspicuous
median keel. The cervical scute is small. The anterior
margin of the first vertebral is flaring, the anterior
margin wider than the posterior margin. The second
vertebral is square-shaped, with both sides are slightly
protruding. The third and the fourth vertebrals are as
wide as long, with the fourth vertebral being wider
than long. The first pleural is the longest, the second is
wider and as long as the third, and the fourth the
smallest. There are twelve marginals. The first mar-
ginal is the widest, the third to seventh and the ninth
to tenth are slightly flared.

The plastron is rounded anteriorly and notched
posteriorly. The plastron can completely close the
shell and is united to the carapace by ligamentous tis-
sue. The ligamentous tissue is also between the pecto-
rals and abdominals. The length of gular seam is 1.5
times the humeral seam, and much shorter than pecto-
ral, abdominal, or anal seam. The pectoral seam is as
long as abdominal seam, the anal seam a little shorter,
and the abdominal seam is nearly twice the length of
the gular seam; femoral seam much shorter, only a lit-
tle longer than humeral seam. There are a pair of anals
that are notched posteriorly.

The head is moderate in size, smooth on top, and
rather rough in the occipital region. The snout is
pointed, projecting over the upper jaw. The diameter
of the orbit is about the same of the length of snout.
Upper jaw is slightly curved and a little longer than
the lower jaw.

2001

Asiatic Herpetological Research

Vol. 9. p. 143

| = 1 cm

Figurel. The holotype (SIZ 80170), an adult male, in
dorsal view.

Figure 2. The allotype (SIZ 801 70), an adult female, in
ventral view.

The forelimb is covered by imbricate scales poste-
riorly and with a transverse series of large scales on
ventrally. The hindlimb is covered by scales medially
and on the tarsus. There are five claws on the forelimb
and four on the hindlimb. The webbing between the
digits is well developed. The tail is short, conical in
shape. The tail is covered by granular scales on its
back, and covered with paired scales forming a longi-
tudinal groove ventrally.

Color in life

The carapace is light brown. The plastron and ventral
side of marginals are yellow, with broad black bars
along the seams. The top of the head is olive, dark
gray laterally, with two brown stripes behind the orbit
and along tympanum to neck. The tympanum is light
gray. The lower jaw and chin are grayish-yellow. The
back of the neck is brown, but lighter ventrally. The
shoulders are light yellow and the axilla are yellow.
The limbs and tail are light brown above and gray
below. The pelvic region and the area behind the fern-

Vol. 9. p. 144

Asiatic Herpetological Research

2001

ora are light yellow. The claws are brown with their
tips yellow.

Allotype

The seams of the plastron are not as black as in the
holotype, the back of tail is covered with a few large
scales.

Acknowledgments

This translation was provided by Ermi Zhao with per-
mission of Acta Zootaxonomica Sinica. The figures
were provided by Ming-Tao Song.

Table 1 . Measurements of types (in mm).

Literature Cited

Department of Herpetology. Sichuan Institute of Biol-
ogy (Zhao. E.. Y.-m. Jiang, and Y. Shen). 1977. [Sys-
tematic Key to Chinese Reptilia]. Science Press,
Beijing. 1 10 pp. (in Chinese).

Pope. C. H. 1935. The reptiles of China. Natural His-
tory of Central Asia 10:28-35.

Smith. M.A. The Fauna of British India. Including
Ceylon and Burma. Reptilia and Amphibia. Vol. I.-
Loricata, Testudines. Taylor and Francis. London. 185
pp.

„_ C

£L

*3 CD

O) CD

E
o

■o

O
CO

CJ
CO
a.

II he
apac

CD ^

-C CO

(f)

CO CJ

Holotype Male 32 18 16 18 120 86 40

Allotype Female 33 18 18 27 115 81 37

33.3

32.2

2000

Asiatic Herpetological Research

Vol. 9, pp. 145-148

Translation: Testudo graeca anamurensis ssp. nov. from Asia Minor

Heinz Weissinger

Richard-Gebhardtgasse 4. 3423 St. Andra-Wordern, Austria

Weissinger. H. 1987. Testudo graeca anamurensis ssp. nov. aus Kleinasien. OGH-Naehriehten (Wien) 10/1
14-18

The South-western coast of Turkey, from the Bey
mountains near Antalya eastwards up to Mersin is the
area of distribution of a Testudo graeca which is char-
acterized hy an elongated, trapezium-shaped and
strongly flattened carapace. This southcoast-popula-
tion was so conspicuous among the more than 200
specimens which I examined and measured in Turkey,
that I consider them a geographic subspecies and
describe them as follows:

Testudo graeca anamurensis nov. ssp.

Material

Type: One female. Anamurum; NMW 30795:1
Paratypes: One male, Anamurum; NMW 30795:2df
One female. Side; NMW 30795:3
Both alive in the author's collection

Terra typica

Beach of Anamurum. 7 km. west of Anamur. SW
coast of Turkey.

Diagnosis

Differs from Testudo graeca ibera Pallas 1814 by
its considerably Hatter and narrower (elongated trape-
zium-shaped) carapace. The length of the carapace is
two-and-a-half times its height and one-and-a-half
times its width. In most cases the carapace pattern is a
completely broken, obvious blotchy pattern. The plas-
tron has a sharply demarcated pattern of brown to
black blotches, which can cover the plastral area for
20 - 100 7c. In 80 CA of cases, 100% at the terra typica,
the animals have a white-yellow chin and throat. The
head can be uniformly dark to irregularly spotted with
yellow. A triangle on the snout - as in Testudo graeca
terrestris Forskal 1775 - may be present only in early
juvenile stages. In the subspecies most abundant in
Asia Minor. Testudo graeca ibera Pallas 1814. the
carapace is oval, domed regularly and has a smooth
posterior margin. Its ground colour is bright olive with
large yet individual blotches. The head and extremi-
ties are most often uniformly dark. The plastron usu-
ally shows indistinctly bordered blotches.

Figures 1-3. (Top and bottom left) Testudo graeca
anamurensis ssp. nov.; paratype. female NMW
30795:3.

Figure 4. (Bottom right) Testudo gracea. Ssp. anamu-
rensis, male. Anamurum (left); ssp. ibera, female,
Koyegegiz (right).

Table 1. Measurements (straight-line distance) of the
type specimen and a paratype

NMW 30795:1 NMW 30795:3

210mm

Carapace

260 mm

length

Carapace
width

160 mm

Carapace

105 mm

height

135 mm

88 mm

Vol. 9, p. 146

Asiatic Herpetological Research

2000

303 tan

9 Testudo graeca ibera

C) Testudo graeca ibera "lerrestris"

O Testudo graeca "lerrestris"

J^ Testudo graeca anamurensis ssp. nov.

Figure 5. Localities of Testudo graeca in Turkey with dates that tortoises were observed (Weissinger, 1 986). 1 , ca.
70 km NW of Ankara, 5/18; 2, Dogansehir, open oak forest, 5/20; 3, 4 km from Diyabakir, 5/22; 4, 40 km from Mar-
din; 5, Birecik, 5/24; 6, Nizip, in the direction of Ganziantep, dry wadi, about 45g steep loam walls with tortoise bur-
rows, 5/24; 7, ca. 10 km N of Islaniye, 5/24; 8, 20 km S of Iskenderund, 5/25; 9, ca. 20 km N. of Iskenderund, 5/26;
10, ca. 25 km of Fg, cornfield, 5/26; 1 1 , 1 km from road crossing Ceyhan - Gaziantep, lava field, 5/26; 12, Ayas, 5/
27; 13, Korykos-Kizkale, in the direction of Silifke, 5/27; 14, Bozyazi, onion field, 5/28; 15, Anamurum, 7 km W of
Anamur, 5/29; 16, ca. 20 km W of Anamur, 5/29; 17Pine forest, ca. 40 km W of Anamur, towards Gazipasa, 5/29;
18, Side, sand dunes, ruin-meadows, 5/30; 19, 20 km from Antalya, 5/31; 20, 10 km from Antalya, towards Kugla,
5/31; 21, Kemer, 5/31; 22, 10 km from Kemer, 5/31; 23, Mountain pass, ca. 800 m above sea level, towards Kas,
5/31; 24, Vavi, ca. 800 m above sea level, towards Kas, 5/31; 25, Kinis, wheat field, 6/1; 26, Letoon, tilled field, 6/
1 ; 27, Esen, pine forest, 6/1 ; 28, Kargi, 6/1 ; 29, Koygegiz, riverine forest with wet meadows 6/2; 30, Ula, 6/2; 31 ,
Gokovalskele, 6/2; 32, Yatagan, 6/2; 33, Soke, towards Kusadasi, 6/2; 34, Selcuk, 6/3; 35, Pamucak, 6/4; 36, 2 km
from Ayvacik, 6/6; 37, 15 km from Ayvacik, 6/6; 38, 20 km from Kesan, towards Greece (European Turkey).

Etymology

I name this South-west coast race after its terra typica.
Anamurum, Testudo graeca anamurensis.

Distribution

The area of distribution of this race extends along
the Southwest coast of Turkey from the Bey Daglari
in the West to the plain of Mersin in the East. Most
were found in the area between Side and Anamurum.

Acknowledgments

This translation was provided by Peter Paul van Dijk
with some additions by Heinz Grillitsch. The transla-
tion was done with the permission and full agreement
of the Austrian Herpetological Society. Figures 1-4
were provided by Jim Buskirk who, in turn, recieved

Weissinger's original slides from his widow. The edi-
tors of AHR would like to thank Heinz Grillitsch for
his assistance in publishing this translation.

Literature Cited

Bodenheimer, F. S. 1935. Animal life in Palestine. An
introduction to the problems of animal ecology and
zoogeography. Ludwig Mayer, Jerusalem. 506 pp.

Eiselt. T J., and Spitzenberger, F. 1967. Ergebnisse
Zoologischer Sammelreisen in derTiirkei: Testudines.
(Results of zoological collecting expeditions in Tur-
key : Testudines) - Annalen des Naturhistorischen
Museums in Wien. Serie B. fur Botanik und Zoologie
70: 357-378.

2000

Asiatic Herpetologia.il Research

Vol.9, p. 147

Figure 6. The type specimen of Testudo graeca ana-
/ra//-<?/75/5Weissinger 1987, NMW 30795:1.

Figure 7. A male paratype of Testudo graeca anamu-
/ms/sWeissinger 1987, NMW 30795:2.

Obst. F.-J.. and Meusel. W. 1974. Die Landschild-
kroten Europas (The tortoises of Europe) A. Ziemsen
Verlag. Wittenberg Lutherstadt

Wermuth. H. 1958. Status und Nomenklatur der Mau-
rischen Landschildkrote, Testudo graeca, in SW-
Asien und NO-Afrika. (Status & Nomenclature of T.
graeca in SW Asia & NE Africa). Senckenbergiana
Biologica. 39 : 149-153.

Wermuth. H.. and Mertens. R. 1961. Schildkroten -
Krokodile - Briickenechsen. (Turtles. Crocodiles &
Tuatara). Fischer Verlag. Jena 422 pp.

Werner, F 1902. Die Reptilien- und Amphibienfauna
von Kleinasien. (The reptile and amphibian fauna of
Asia Minor). Sit/^ungsberichte Akademie Wissen-
schaften. Mathematisch-Naturwissenschaftliche

Klasse 111: 1057-1121.

Vol. 9, p. 148

Asiatic Herpetological Research

2000

Appendix

This translation was prepared by Peter Paul van Dijk
with some additions by Heinz Grillitsch of the
Naturhistorische Museum Wien (NMW). This trans-
lation is published with the full permission of the
Austrian Herpetological Society. In addition to the
original figures. Heinz Grillistach and Alice Schuma-
cher (NMW) have provided additional figures of the
original type series (Figs. 6-8). Uwe Fritz (pers.
comm.). states that there are three additional
paratypes in the collections of the Staatliches
Museum fur Tierkunde Dresden (MTKD 29200-
29202). Finally, the female paratype that Weisssinger
lists as NMW 30795 is currently catalogued as NMW
31031 (Fritz and Grillitsch, pers. comm).

Figure 8. A female paratype of Testudo graeca anamu-
/•ews/sWeissinger 1987, NMW 31031 (formerly
30795:3).

2001

Asiatic Herpetological Research

Vol. 9, pp. 149-150

Book Review: A Guide to the Fauna of Iran

A Guide to the Fauna of Iran. Eskandar Firouz. Iran University Press; Tehran. 2000. vi + 491 pp

The first attempt to provide a comprehensive verte-
brate zoology of Iran was that of William T. Blanford
in 1876. Since that time, there have been treatises on
various groups of vertebrates (e. g., mammals: Lay.
1967; Harrington, 1977; Ziaie, 1996; birds: Scott, et
al., 1975; reptiles: Latifi, 1984 (snakes); Anderson,
1979 (turtles, crocodiles, and amphisbaenians), 1999
(lizards); amphibians: Baloutch and Kami, 1995;
fishes: Coad, 1987, 1995. A Guide to the Fauna of
Iran by Firouz is the first attempt since Blanford to
bring all of the vertebrates of Iran together into a natu-
ral history guide.

Although the text is in Farsi (Persian), it is acces-
sible to Western readers as a comprehensive list of the
vertebrate species of Iran, since there is a Latin and
English index, color illustrations with Latin species
names, and lists of taxa with both Latin species names
and English colloquial names. Most genera are repre-
sented by color illustrations, fishes by paintings,
amphibians and reptiles by photographs, birds by
paintings, and mammals by both. A list of the princi-
pal works consulted is provided (pp. 423-432). This
list constitutes a good beginning bibliography for any-
one interested in the vertebrate zoology and natural
history of Iran.

For this book, the author has consulted the zoolo-
gists currently working on each of the vertebrate
groups to assure that the species lists are up to date.
The work covers 164 vertebrate families and 1054
species. Emphasis is given to conservation topics,
including threatened and endangered species, prob-
lems of exotic introductions, and the ecological con-
sequences of environmental change. The natural
history sections include consideration of the zoogeog-
raphy of Iran.

A renewed interest in zoology has occurred
recently in Iran, the many universities including
departments of biology and zoology are attempting to
build knowledgeable faculties in these disciplines.
This emphasis is largely due to the efforts of the
author, Eskandar Firouz. Prior to the Islamic Revolu-
tion, Firouz was the principal architect of the conser-
vation, natural history legislation and implementation
in Iran through the Department of Environment, the
establishment of Protected Regions and National
Parks, and the regulation of hunting and fishing.
These efforts were well on the way to setting a world
standard for conservation and the study of natural his-
tory at the time of the revolution. Interests in natural

history have survived the period of resistance to per-
ceived "Westernization" in Iran, and young scholars
and conservationists must now face the daunting chal-
lenge of rebuilding the edifice of conservation initi-
ated by Firouz and his colleagues in the 1960s and
early 70s. The present book is yet another major con-
tribution to that effort, and there should be a well-used
copy in every local Department of Environment
office, university and department library and in the
personal libraries of individual zoologists and ecolo-
gists in Iran. It will also be a useful reference in spe-
cialized libraries, natural history museums, and
conservation institutions in the West and in the coun-
tries of the former Soviet Union.

Steven C. Anderson

Department of Biological Sciences, University of the

Pacific, Stockton, CA 95211 USA and Department of

Herpetology, California Academy of Sciences, Golden

Gate Park, San Francisco, CA, 94118 USA. email:

asaccus@aol. com

Literature Cited

Anderson, S. C. 1979. Synopsis of the turtles, lizards,
and amphisbaenians of Iran. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, 41(22):501-528.

Anderson. S. C. 1999. The Lizards of Iran. Society for
the Study of Amphibians and Reptiles, Ithaca, NY, vii
+ 442 pp.

Baloutch, M. and H. G. Kami. 1995. Amphibians of
Iran. Tehran University Publications, Tehran, 177 pp.
(In Farsi [Persian]).

Blanford, W. T. 1876. Eastern Persia. An Account of
the Journeys of the Persian Boundary Commission,
1870-1872, vol. 2. The Zoology and Geology. Mac-
millan and Co., London, viii + 516 pp.

Coad, B. W. 1987. Zoogeography of the freshwater
fishes of Iran. In : Krupp, et al. (Eds.). Proceedings of
the Symposium of the Fauna and Zoogeography of
the Middle East. Beihefte zum Tubinger Atlas des
Vorderen Orients. Reihe A (Naturwissenschaften)
28:213-228.

Coad, B. W. 1995. Freshwater fishes of Iran. A check-
list. Institute of Landscape Ecology of the Academy
of Science of the Czech Republic, 29( 1 ): 1 -64.

Vol. 9, p. 150

Asiatic Herpetological Research

2001

Harrington, F. A. Jr. 1977. A Guide to the Mammals
of Iran. Department of the Environment, Tehran, 89
pp.

Latifi, M. 1984. The Snakes of Iran. Department of
the Environment, Tehran, 231 pp. (in Farsi [Persian];
English Translation, 1991, Society for the Study of
Amphibians and Reptiles, Oxford, Ohio).

Lay, D. M. 1967. A study of the mammals of Iran.
Fieldiana Zoology 54:282 pp.

Scott, D. A., H. Moravej Hamadani, and A. Adhami
Mirhosseyni. 1975. Parandegan-e Iran (The Birds of
Iran). Department of the Environment, Tehran (In
Farsi [Persian]; Latin, English, and French names).

Ziaie, H. 1996. Field Guide to the Mammals of Iran.
Department of the Environment, Tehran (In Farsi
[Persian]; Latin and English names).

2001

Asiatic Herpetological Research

Vol. 9, pp. 151-152

Book Review: Four Recent Handbooks for Turkey

Tiirkiye Amfibiliri. The Amphibians of Turkey by

Necla Ozeti and I. Yilmaz, 1994. Izmir, 219 pp., 40
col. photos in pis., 92 text-figs. (In Turkish with
English summary).

Tiikiye Omurglilari, Amfibiler edited by Ali Demir-
soy, 1996. Meteksan; Ankara, vi + 69 pp., numerous
maps and figs. (In Turkish).

Tiikiye Omurglilari, Siiriingenler edited by Ali
Demirsoy, 1996. Meteksan; Ankara, viii + 205 pp.,
numerous maps and figs. (In Turkish).

Turkish Herpetofauna (Amphibians and Reptiles)

by Ibrahim Baran and Mehmet Atatiir. 1998. Republic
of Turkey Ministry of Environment: Ankara, x + 214
pp., 122 col. pis., 7 text-figs. (In English).

Until the publication of these books, the standard
herpetological summary publications for Turkey had
been Ba§oglu and Ozeti (1973) and Ba§oglu and
Baran (1977, 1980). The four handbooks reviewed
here accompanied us on a recent herpetological expe-
dition to western and southern Turkey. The combina-
tion of these references enabled us to quickly identify
most species in the field.

Turkish Herpetofauna by Baran and Atatiir has a
color photograph of each species, illustrated keys, one
paragraph each on identification, habitat, biology, and
distribution. The distribution sections include brief
statements of the overall distribution and the range
within Turkey. Where subspecies are recognized, a
short distribution statement is given for each. There is
no descriptive differentiation of subspecies. There are
no distribution maps. The color photographs are of
good quality and usually enabled quick identification
of live specimens. Photos and text are on glossy paper
and this hardback book is sturdily bound. The bibliog-
raphy includes 53 references, and while not exhaus-
tive, serves as a good introduction to the
herpetological literature of Turkey. The nomenclature
is reasonably current, with the exception, of course, of
changes (e.g. the break-up of Eumeces and Coluber)
since its publication. It is the only one of these hand-
books to include Rana dalmatina Schneider, Sinch,
and Nevo, 1992. Of the four, this book got most use
on the trip, particularly as it is in English.

The two paper-bound books edited by Demirsoy,
Tiikiye Omurglilari, Amfibiler and Tiikiye Omurg-
lilari, Siiriingenler are useful to Western herpetolo-
gists, in particular, because they have a distribution
map for each species, usually accompanied by a black
and white illustration, often a photograph, but some-
times one or more line drawings. Although the books

are printed on high quality stock, many of the photo-
graphs have not reproduced well.

The tabular format of species presentation is
somewhat unusual and innovative. These tables are
the same for each species and include Latin and Turk-
ish names, the author, date; one or more illustrations;
distribution map for Turkey, showing both range and
spot localities; remarks on habitats, relationships, tax-
onomic status, etc.; conservation status (rare, endan-
gered, vulnerable, widespread, etc.); overall
distribution and range within Turkey; type locality;
greatest size; morphological characteristics; color pat-
tern; reproduction; time from hatching to maturity;
conservation status of habitats; defense mechanisms;
population densities; potential hazards; zoogeo-
graphic origins; and conservation proposals for the
future. A table listing the fauna summarizes the global
and Turkish distributions, zoogeographic source, pre-
sumed geological time of arrival in Turkey, biogeo-
graphic and conservation status, and special remarks.

The books also contain descriptions and com-
ments on higher taxa for the general reader. The bibli-
ographies are extensive, much more so than that of the
Baran and Atatiir handbook in English. Contributors
to these volumes are prominent Turkish herpetolo-
gists: Varol Tok, Ibrahim Baran, Mehmet Atatiir, Abi-
din Budak, and Mehmet Oz. These volumes are
obviously designed for the serious Turkish zoology
student or wildlife professional and should serve this
purpose well. They appear to be part of a larger series
covering the entire Turkish fauna. Subspecies are cov-
ered individually, although there are some strange
inconsistencies: sometimes there is a separate table
and map for the species as a whole, in addition to the
subspecies accounts, and sometimes not. The usage of
generic names is sometimes confusing, as for exam-
ple: Cyrtodactylus (-Cyrtopodion) (Mediodactylus)
heterocercus and Agama (-Laudakia) stellio so that it
not clear which generic name is recognized by the
authors. As I don't read Turkish, I can't comment on
errors in the text. Some of the range maps may be too
general, especially in those cases where spot distribu-
tions are not shown. For example, Laudakia stellio is
shown to extend to broad contact with the Iranian bor-
der, whereas it has never been recorded for Iran. We
used the maps of these volumes in conjunction with
the photographs and English text of the Baran and
Atatiir book and this greatly facilitated our work.

Tiirkiye Amfibiliri. The Amphibians of Turkey by

Ozeti and Yilmaz is aimed primarily at Turkish zool-
ogy students and serves as an introduction to amphib-

Vol. 9, p. 152

Asiatic Herpetological Research

2001

iology. It is an update, or second edition of Ba§oglu
and Ozeti (1973). The first chapter is an introduction
to nomenclature, and in particular, to the taxonomy of
the Turkish amphibian fauna. One chapter deals with
the biology of amphibians in general, including mor-
phology, reproduction, development, distribution, and
folklore. Two subsequent chapters cover these topics
in more detail, one for salamanders and one for frogs.
These chapters are arranged by family and include
families not found in Turkey. They include keys to the
Turkish genera and species and spot locality maps.
These maps are useful, if somewhat cluttered (several
taxa per map); localities are numbered and named in
the figure legends. Keys to the generic level for egg
masses and larvae have been provided. The descrip-
tions and discussions of morphology are accompanied
by useful illustrations taken from several sources (not
always attributed). There is a short chapter on meth-
ods and techniques, a glossary, and a bibliography of
181 references, including the most important techni-
cal papers for the fauna, an excellent introduction to
the literature of the Turkish amphibian fauna. At the
end of the book are 1 3 plates of black and white pho-
tographs illustrating the covered taxa and some habi-
tats. The photographs (or at least the printing of them)
are of rather poor quality, although they are adequate
to illustrate pattern differences between the species.
An English summary includes a comprehensive key to
the amphibians of Turkey. This key, along with the
English figure legends, and specific collecting locali-
ties, makes this book particularly useful to non-Turk-
ish speaking herpetologists.

Steven C. Anderson

Department of Biological Sciences, University of the

Pacific, Stockton, CA 9521 1 USA and Department of

Herpetology, California Academy of Sciences, Golden

Gate Park, San Francisco, CA, 94118 USA. email:

asaccus @ aol. com

Literature Cited

Ba§oglu, M. and N. Ozeti 1973. Tiirkiye Amfibileri.
The Amphibians of Turkey. Ege Universitesi Fen
Fakultesi Kitaplar Serisi No. 50. Ege Universitesi
Matbaasi; Bornova/Izmir, 155 pp., 9 pis.

Ba§oglu, M. and I. Baran. 1977. Tiirkiye
Suriingenleri. Kisim I. Kaplumbaga ve Kertenkeleler.
The Reptiles of Turkey. Part I. The Turtles and Liz-
ards.Ege Universitesi Fen Fakultesi Kitaplar Serisi
No. 76. Ilker Matbaasi; Bornova/Izmir, 255 pp., 16
pis.

Ba§oglu, M. and I. Baran. 1980. Tiirkiye
Suriingenleri. Kisim II. Yilanlar.The Reptiles of Tur-
key. Part II. Snakes. Ege Universitesi Fen Fakultesi
Kitaplar Serisi No. 81. Ilker Matbaasi; Bornova/Izmir,
218 pp.

2001

Asiatic Herpetological Research

Vol. 9, pp. 153-153

Book Review: Wild About Reptiles.
Field Guide to the Reptiles and Amphibians of the UAE

Wild About Reptiles. Field Guide to the Reptiles and Amphibians of the UAE. by Marycke Jongbloed. 2000.
Barkers Trident Communications; London. I 16 pp.. numerous unnumbered color photographs. Order from Natu-
ral History Book Service: www.nhbs.com

This attractive little spiral-bound book is the first field
guide to the amphibians and reptiles of the United
Arab Emirates. Most of the 68 species covered are
illustrated with good color photographs. There are no
keys, but the color photographs will serve to identify
most live specimens. Descriptions are sketchy and the
emphasis is on natural history. The book should serve
well as an introduction to desert reptiles, sea snakes
and sea turtles, and the few amphibians of the region.
English speakers who work in the Emirates, particu-
larly members of the natural history societies, will
find this a handy reference. Visiting herpetologists.
while they will still want the keys and descriptions of
Arnold (1986). Balletto, et al. (1986). Gasperetti
(1988). Leviton and Anderson (1967), and Leviton, et
al. ( 1992). will want this for quick identification. The
technical papers that emphasize the herpetology of the
Emirates are few in number (see Arnold. 1984: Levi-
ton and Anderson, ) and only Arnold ( 1984) concerns
the ecology and natural history of this region of the
Arabian Peninsula.

Jongbloed takes a highly personal and anecdotal
approach in this book, demonstrating her enthusiasm
for these creatures. She includes field notes taken by
her friend. John Norman Bishop "Bish" Brown.
Because so little professional herpetological work has
been done in the Emirates, and those studies that have
been carried out have dealt mainly with systematics.
using preserved specimens, this book is almost the
only source of natural history information for these
groups in the UAE. The natural history observations
are informal, but still, informative. The bibliography
is sketchy and disappointing and will not lead the
beginning naturalist very deeply into the literature of
the UAE.

I recommend this field guide to any naturalist, pro-
fessional or amateur, planning to travel to the UAE.

Anyone seriously interested in the herpetofauna of
Southwest Asia in general or the Arabian Peninsula in
particular will want this in his library.

Steven C. Anderson

Department of Biological Sciences, University of the

Pacific, Stockton, CA 9521] USA and Department of

Herpetology, California Academy of Sciences, Golden

Gate Park, San Francisco, CA, 94118 USA. email:

asaccus @ aol. com

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 check list to
the lizards and amphisbaenians of Arabia. Fauna of
Saudi Arabia 8:384-435.

Balletto. E.. M. A. Cherchi, and J. Gasperetti. 1986.
Amphibians of the Arabian Peninsula. Fauna of Saudi
Arabia 7:318-392.

Gasperetti. J. 1988. Snakes of Arabia. Fauna of Saudi
Arabia 9:169-450.

Leviton, A. E. and S. C. Anderson. 1967. Survey of
the Sheikhdom of Abu Dhabi. Arabian Peninsula. Part
II: Systematic account of the collection of reptiles
made in the Sheikhdom of Abu Dhabi by John Gas-
peretti. Proceedings of the California Academy of Sci-
ences (4) 35(91:157-192.

Leviton, A. E.. S. C. Anderson. K. Adler, and S. A.
Minton. 1992. Handbook to Middle East Amphibians
and Reptiles. Society for the Study of Amphibians
and Reptiles. Oxford. Ohio, vii + 252 pp.

2001

Asiatic Herpetological Research

Vol. 9, pp. 154-155

Obituary: Sherman Anthony Minton Jr.

Muhammad S. Khan

7740 NW, 47th Street Lauderhill, FL.33351, USA

Dr. Sherman Anthony Minton Jr., died of cancer in
Indianapolis. Indiana, USA. on 15 June 1999, at the
age of 80. He was born on the 24 February 1919, in
New Albany. Indiana, where he also attended school.
He had a fondness for amphibians and reptiles even in
his childhood. He was the eldest child of Senator
Sherman Minton Sr.. on whose suggestion young
Sherman chose medicine instead of law, since medi-
cine was closer to zoology. He obtained a BS in Zool-
ogy in 1939 and an MD in 1942 at Indiana University.
He served during World War II in the U.S. Navy from
1943-1946. After the war, Sherman spent 1947-1948
at the University of Michigan Zoology Department,
taking Herpetology and Microbiology. He later joined
the faculty of Indiana University School of Medicine,
in the Department of Microbiology and Immunology,
where he remained until he retired as Emeritus Pro-
fessor in 1984. Dr. Minton's career presents a beauti-
ful amalgamation of medicine, microbiology, and
herpetology.

Sherman married Madge Alice Shortridge Ruther-
ford on 10 October 1943. while both were in military
service. He was in the Navy and she was flying with
the Women Air Force Service Pilots (WASP). Interest-
ingly. Madge has also been interested in snakes from
her childhood, so their relationship was also profes-
sional. They co-authored eight publications including
two books. Venomous Reptiles (Minton and Minton
1969. 1980) and Giant Reptiles (Minton and Minton.
1973).

Minton took a break from Indiana ( 1958-1962) to
teach at the Basic Medical Sciences Institute. Karachi.
Pakistan (now Postgraduate Medical Center), as part
of the U.S. AID program. He taught at the institute,
reorganized the medical teaching courses, and headed
the institute for a short time.

He pioneered herpetological studies in Pakistan.
Both he and his wife Madge Minton traveled 44,000
miles in different parts of Pakistan, collecting
amphibians and reptiles and information about them
from the local people. One of the results of this is the
well-illustrated paper on the amphibians and reptiles
of Sind and Las Bela (1962). In 1965. he visited Iran
and Pakistan under the sponsorship of American
Museum of Natural History and collected material for
his main book on the herpetology of Pakistan ( 1966).

Sherman, with Madge's constant support and help,
produced over 170 articles, books and monographs.
He was a pioneer in the study of venomous reptiles
and toxicology. He was on the editorial board of the
journals Toxicon and Clinical Toxicology. He was the
chairman and a member of the major professional
societies in his field and received numerous awards
and honors for his work. He also made a significant
contribution to the herpetology of Indiana

From 1972-1980 he joined several expeditions to
different oceans to study biology of sea snakes and
other venomous sea animals. He was visiting profes-
sor in the Department of Zoology, University of New
England in Australia during 1980.

In remembrance of Dr. Minton's services to the
herpetology of Pakistan, the following fossil Ameri-

2001 Asiatic Herpetological Research Vol. 9, p. 155

can frog and Pakistani lizard and snakes have been
named after him:

Proacris mintoni Holman, 1961

Coluber karelini mintonorum Mertens, 1969

Gymnodactylus mintoni Golubev and Szczerbak,
1981

Typhlops madgemintonai shermanai Khan, 1999

Dr. Sherman A. Minton, Jr., is survived by his
widow Madge Rutherford Minton, and three daugh-
ters.

Literature Cited

Golubev. M., and Szczerbak, N. N. 1981. A new spe-
cies of Gymnodactylus Spix, 1923 (Reptilia, Sauria,
Gekkonidae) from Pakistan. Vestnik Zoolii 1981:40-
45, (in Russian).

Holman, A. 1961. A fossil frog from the Lower
Miocene of Florida. Copeia 1961(3):354-355.

Khan, M. S. 1999. Two new species and a subspecies
of blind snakes of genus Typhlops from Azad Kashmir
and Punjab, Pakistan. Russian Journal of Herpetology
6(3): 231-240.

Mertens. R. 1969. Die Amphibien und Reptilien West-
Pakistans. Stuttgarter Beitrage zur Naturkunde 197: 1-
96.

Minton, Jr., S. A. 1962. An annotated key to the
amphibians and reptiles of Sind and Las Bela, West
Pakistan. American Museum Novitates 208 1 : 1-21.

Minton, Jr., S. A. 1966. A contribution to the herpetol-
ogy of West Pakistan. Bulletin of the American
Museum of Natural History 134(2): 31-184.

Minton Jr., S. A., and Minton, M. R. 1969. Venomous
Reptiles. Charles Scribner's Sons, New York. 274 pp.

Minton Jr., S. A., and Minton, M. R. 1973. Giant Rep-
tiles. Charles Scribner's Sons, New York. 345 pp.

Minton Jr., S. A., and Minton, M. R. 1980. Venomous
Reptiles. Charles Scribner's Sons, New York. 308 pp.

Stewart, M. M. 2000. Madge and Sherman Minton.
Copeia, 2000(1): 304-309.

2001 Asiatic Herpetological Research Vol. 9, pp. 156-160

Guidelines for Manuscript Preparation and Submission

NOTE: AHR now requires all tables to be submitted camera-ready. Please send
hard copy, or electronic images of tables in the following format:

Table 1 .AHR Standard table format. Note double line under heading row. Please use a sanserif font
(Helvetica 1 0pt or Ariel 10pt).

Column 1 Column 2 Column 3 Column 4

12.4 43321.5 9.1 0.01

12.1 91020.4 0.6 0.02

Summary

Manuscripts must:

1) be written in English.

2) be of letter quality (laser printed or typewritten on bond paper).

3) include camera ready tables and figures (if any).

4) include complete and accurate literature citations.

5) include complete and accurate localities with latitude and longitude.

6) include a camera ready map illustrating regions discussed (when applicable).

Tips for electronic submission

• Do not type authors names in all capitals in literature cited.

• Do not use two spaces following a period, or for any other purpose.

• Do not attempt to recreate the format of the journal in your manuscript. Please use only sim-
ple formatting limited to italics, boldface, and underline.

Manuscripts failing to meet these criteria will be returned without review for correction.

Purpose and Content

Asiatic Herpetological Research publishes articles concerning but not limited to Asian herpetology. The editors
encourage publications from all countries in an attempt to create an open forum for the discussion of Asian her-
petological research.
Articles should be in standard scientific format and style. The following sections should be included:

Title

The title should reflect the general content of the article in as few words as possible. The editors encourage titles
that summarize the main findings of the article.

Names and Addresses

The names and addresses of all authors must be complete enough to allow postal correspondence. Please include
email and World Wide Web addresses if applicable.

Abstract

The abstract should briefly summarize the nature of the research, its results, and the main conclusions. Abstracts
should be less than 300 words.

2001 Asiatic Herpetological Research Vol.9, p. 157

Key Words

Key words provide an index for the filing of articles. Key words provide the following information (when appli-
cable): 1 ) Taxonomy (e.g. Reptilia, Squamata, Gekkonidae, Gekko gecko). 2) Geography (e.g. China. Thailand).
3) Subject (e.g. taxonomic validity, ecology, biogeography). The order of taxonomy, geography, and subject
should be observed.

Text

Manuscripts must be in English and spelling must be correct and consistent. Use Webster's New International
Dictionary for reference. For clarity, use active voice whenever possible. For example, the following sentences in
active voice are preferable to those in passive voice.

Active voice: "Lizards were extremely common on the site." and "I examined three female snakes."

Passive voice: "Lizards were observed to be extremely common on the site." and "Three female snakes were
examined."

Abbreviation

Do not abbreviate unless the full phrase has already appeared. Scientific names may be abbreviated only if they
have appeared fully in the same paragraph. Never begin a sentence with an abbreviation of a scientific name.

Statistics

Statistics must be accompanied by sample sizes, significance levels, and the names of any tests. Investigators
should pay careful attention to independence and applicability of tests, and randomness of samples. One of the
most frequent examples of nonindependence is the use of multiple, paired t-tests instead of analysis of variance
(anova). In general, multiple tests on the same data set are not valid. Descriptive statistics are in many cases more
appropriate than inferential statistics.

Standard Format

Manuscripts following standard format should include introduction, methods, results, and discussion sections.
While other formats are acceptable, the editors encourage the use of standard format. Please do not type in all
capital letters.

Introduction

The introduction typically states the significance of the topic and reviews prior research.

Material and Methods

This section should clearly state where, when, and how research was carried out. Include sample sizes. Protocols
designed by other investigators must be properly cited. Research materials and their manufacturers should be
listed. The reader must be able to replicate the methods of the author(s).

Results

This section states the results and their significance to the investigation. Figures and tables may be used to clarify,
but not to replace, results statements in the text. Statistics should be used when applicable. Large amounts of data
should be avoided, or included as an appendix at the end of the article.

Discussion

The discussion is a synthesis of the introduction and the results. No new information should be discussed unless
it was presented in the results section. New findings should be discussed in relation to prior research. The
author(s) should feel free to present several possible interpretations of the results. The editors particularly
encourage suggestions of future research in Asian herpetology.

Vol. 9, p. 158 Asiatic Herpetological Research 2001

Manuscript Preparation

Overview

Please do not attempt to replicate the formatting style of AHR in your manuscript. All formatting except italics
will be removed in the production process. Bold and underlined text should be used only to identify section head-
ing levels (see below). Extraneous formatting is counterproductive and increases the production costs of the jour-
nal. There are a few simple guidelines that authors must follow.

Section Headings

Articles will be published using three section heading styles. All heading levels must be on their own line, and
left justified. For the purposes of manuscript submission. Level 1 heading is bold, and generally reserved for
Introduction, Material and Methods, Results, and Discussion; Level 2 is italic, and Level 3 is underlined.

Figures

Figures must be referenced in order in the text. Each figure illustration (line art or photograph) submitted must
be "camera ready" for publication with no modifications necessary other than reduction. AHR does not publish
"plates"; please refer to these as figures numbered sequentially. Do not write on figure; do not mount more than
one figure to a sheet. AHR cannot be responsible for redrawing, touching up, or otherwise modifying figure illus-
trations for authors. In addition, figure illustrations submitted must:

1 ) be of publication quality with typeset text.

2) be mounted on a separate 21.5 x 28 cm (8.5 x 1 1 inch) sheet with figure number on back.

3) be on a separate sheet from figure legend.

4) not have poor type or handwriting on the face of the figure.

5) The TIFF file format is preferable for electronic versions of figures, but Photoshop, JPEG, or PICT formats are
acceptable. Resolution of electronic versions of figures must be at least 600 dpi for line art, or 300 dpi for gray-
scale and color images.

6) Figures will be reduced to either 1 column (3.25") or two columns (6.5").

Times Roman typeface is preferred. In order to avoid wasted effort, please follow the above instructions care-
fully. Please note: AHR will not alter or lay out figures for publication. Any figure requiring modification will be
returned, and may cause significant delay in publication.

Figure Legends. Figure legends should be typed on a separate sheet. Legends should explain the figure without
reference to the text. A figure and legend should make sense if separated from the rest of the article. For example:

Figure 2. Lateral view of live Psammodynastes pulverulentus holding a prey lizard (Anolis car-
olinensis). Note buccal tissue surrounding the enlarged anterior maxillary and dentary teeth of
the snake.

Color Figures. AHR may publish color figures at the discretion of the editors. AHR is now published both on
paper and electronically. Printing costs of color figures may be required for the paper version of AHR. Color fig-
ures will be published free of charge in the electronic version. If you submit color figures, please indicate if you
wish them to be considered for publication in color in the paper version. Otherwise, they will be converted to
black and white in the paper version.

Tables

Tables must be referenced in order in the text. AHR now requires that all tables be submitted camera-ready in
AHR's standard format. See Table 1 (above) and prior copies of AHR for examples. Please use a sanserif font
(Helvetica 1 Opt or Ariel lOpt).

Typeface

Twelve point type is preferred. Supply a detailed list of special characters (greek letters, male or female symbols,
etc.) that are not part of a standard font.

2001 Asiatic Herpetological Research Vol. 9, p. 159

Literature Cited

Accurate and standard references are a crucial part of any article. This is especially important when dealing with
publications from many different countries. The reader must be able to precisely identify any literature cited.
References in the text must be checked for consistency with references in the literature cited section. All refer-
ences cited in the text must be in the literature cited section. The literature cited section may not contain any ref-
erences not mentioned in the text. Articles containing inaccurate or inconsistent literature citations will be
returned for correction.

References in Text. 1 ) References to articles by one or two authors must include both surnames in the order they
appear in the original publication. References to articles by more than two authors must include the first author's
surname, followed by "et al." 2) The year of article follows the authors, separated only by a space. 3) References
with the same author and year are distinguished by the lower case characters "a, b, c, . . ." 4) References cited in
text are listed in alphabetical order by first author.

For example, "My results also incorporate literature records (Marx et al., 1982; Marx and Rabb, 1972;
Mertens, 1930; Pope, 1929; Wall, 1909, 1910a, 1910b, 1910c)."

References in Literature Cited. 1 ) References must include all authors, in the order that they appear in the orig-
inal publication; "et al." is never used in a literature cited section. 2) The first author is listed surname first, ini-
tial(s) last. All other authors are listed initial(s) first, surname last. 3) References with the same author and year
are distinguished by the lower case characters, "a, b, c, . . ." 4) References cited are listed in alphabetical order by
first author. 5) Names of journals are not abbreviated. See below for examples:

Journal article

Dial, B. E. 1987. Energetics and performance during nest emergence and the hatchling frenzy in loggerhead sea
turtles (Caretta caretta). Herpetologica 43(3);307-315.

Journal article from a journal that uses year instead of volume

Gatten, R. E. Jr. 1974. Effect of nutritional state on the preferred body temperatures of turtles. Copeia
1974(4):912-917.

Journal article, title translated, article not in English

Ananjeva, N. B. 1986. [On the validity of Megalochilus mystaceus (Pallas, 1776)]. Proceedings of the Zoological
Institute, Leningrad 1 57:4- 1 3. (In Russian).

Note that for Acta Herpetologica Sinica, the year must precede the volume number. This is to distinguish
between the old and new series, and between 1982-1987, Vols. 1-6 (new series) and 1988 with no volume number,
numbers 1 and 2 (new series).

Cai, M., J. Zhang, and D. Lin. 1985. [Preliminary observation on the embryonic development of Hyiwbius chin-
ensis Guenther]. Acta Herpetologica Sinica 1985, 4(2): 177- 180. (In Chinese).

Book

Pratt, A. E. 1892. To the snows of Tibet through China. Longmans, Green, and Co., London. 268 pp.

Article in book

Huey, R. B. 1982. Temperature, physiology, and the ecology of reptiles. Pp. 25-91. In C. Gans and F. H. Pough
(eds.). Biology of the Reptilia, Vol. 12, Physiological Ecology. Academic Press, New York.

Government publication

United States Environmental Data Service. 1968. Climatic Atlas of the United States. Environmental Data Ser-
vice, Washington, D. C.

Abstract of oral presentation

Arnold. S. J. 1982. Are scale counts used in snake systematics heritable? SSAR/HL Annual Meeting. Raleigh,
North Carolina. [Abstr].

Vol. 9, p. 160 Asiatic Herpetological Research 2001

Thesis or dissertation

Moody, S. 1980. Phylogenetic and historical biogeographical relationships of the genera in the Agamidae (Rep-
tilia: Lacertilia). Ph.D. Thesis. University of Michigan. 373 pp.

Anonymous, undated

Anonymous. Undated. Turpan brochure. Promotion Department of the National Tourism Administration of the
People's Republic of China, China Travel and Tourism Press, Turpan, Xinjiang Uygur Autonomous Region,
China.

Asiatic Herpetological Research reserves the copyrights to all material published therein, except that excluded
by permission of the editors. Any material under a prior copyright submitted to Asiatic Herpetological Research
must be accompanied by the written consent of the copyright holder.

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ures will be reduced to either 1 column (3.25") or two columns (6.5").

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NOTE: AHR now requires all tables to be submitted camera-ready. Please send
hard copy, or electronic images of tables. Please use a sanserif font (Helvetica
10pt or Ariel 10pt).

Book Review: Wild About Reptiles. Field Guide to the Reptiles and Amphibians

of the UAE 153

Obituary: Sherman Anthony Minton Jr 154

Guidelines for Manuscript Preparation and Submission 156

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ISSN 1051-382^

Wen-Jian Li. The Reproductive Bioiogy of Rana boulengeri

Mingtao Song. Xiaomao Zeng, Guanfu Wu. Zhijun Liu, and Jinzhong Fu. A New Species of
Batrachuperus from Northwestern China t>

Rafe M. Brown, Alan E. Leviton, John W. Ferner. and Rogelio V. Sison. A New Snake of the

Genus Hologerrhum Gunther (Reptilia; Squamata; Coluhridae) from Panay Island, Philippines

M. Farid Ahsan and Shayla Parvin. The First Record of Ptyas korros (Coluhridae) from
Bangladesh 23

Sean J. Blamires. Influence of Temperature on Burrow Use hy the Monitor Lizard Varanus
panoptes of the Coastal Dunes at Fog Bay, Northern Australia 25

Peter L. Cunningham. Notes on the Diet, Survival Rate, and Burrow Specifics of Uromastyx
aegyptius microlepis from the United Arab Emirates 30

John W. Ferner. Rafe M. Brown, Rogelio V. Sison, and Robert S. Kennedy. The Amphibians
and Reptiles of Panay Island. Philippines 34

James Ford Parham and Haitao Shi. The Discovery of Mauremys iversoni-like Turtles at a
Turtle Farm in Hainan Province, China: The Counterfeit Golden Coin 71

Matthias Stock, Daniel Frynta. Wolf-Riidiger Grosse, Claus Steinlein, and Michael Schmid. A
Review of the Distribution of Diploid, Triploid and Tetraploid Green Toads (Bujo viridis
complex) in Asia Including New Data from Iran and Pakistan 77

Samraat Pawar and Sayantan Biswas. First Record of the Smooth-Backed Parachute Gecko
Ptychozoon lionotum Annandale 1905 from the Indian Mainland 101

Nasrullah Rastegar-Pouyam and Eskandar Rastegar-Pouyani. A New Species of Eremias
(Sauria: Lacertidae) from Highlands of Kermanshah Province, Western Iran 107

Konstantin A. Rogovin, Dmitry V. Semenov, and Georgy I. Shenbrot. Lizards of the Northern
Mongolian Deserts: Densities and Community Structure 113

Murat Sevinc and Ismail Hakki Ugurta§. The Morphology and Size of Blood Cells of Lacerta
rudis bithynica 122

Muhammad S. Khan. Buccopharyngeal Morphology and Feeding Ecology of Microhyla
ornata tadpoles 1 30

Ismail H. Ugurtas. Variation in Pelobates syriacus of Turkey 139

Translation: Mingtao Song. A New Species of the Turtle Genus Cuora (Testudoformes:
Testudinidae j 142

Translation: Heinz Weissingcr. Testudo graeca anamurensis ssp. nov. from Asia Minor 145

Book Review A Guide to th~ Fauna Of Iran . . 149

Book Review: Four Receni Handbooks for Turkey .'...'. 1^1

(Continued on inside of back cover)

Harvard MCZ Llbran

llll

3 2044 066 300 427

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