“4 CHINESE

| (ALERPETOLOGICAL

\

| IRBSEARCHI

Volume 2 1988-1989 ©

Chinese Herpetological Research :

Editor

ERMI ZHAO

Academia Sinica, Chengdu Institute of Biology, Chengdu, Sichuan, China

Associate Editors

KELLAR AUTUMN

Dept. of Environmental Studies, University of

California, Santa Cruz, California, USA

J. ROBERT MACEY

Museum of Vertebrate Zoology, University of

California, Berkeley, California, USA

THEODORE J. PAPENFUSS

Museum of Vertebrate Zoology, University of

California, Berkeley, California, USA

Editorial Board

KRAIG ADLER

Cornell University, Ithaca, New York, USA

NATALIA B. ANANJEVA

Zoological Institute, Leningrad, USSR

LEO BORKIN

Zoological Institute, Leningrad, USSR

BIHUI CHEN

Anhui Normal University, Wuhu, Anhui, China

YUANCHONG CHEN

Shanghai Institute of Biochemistry, Shanghai,

China

ILLYA DAREVSKY

Zoological Institute, Leningrad, USSR

WILLIAM E, DUELLMAN

University of Kansas, Lawrance, Kansas, USA

HAJIME FUKADA

Sennyuji Sannaicho, Higashiyamaku, Kyota, Japan

MEIHUA HUANG

Zheijiang Medical University, Hangzhou, Zhejiang,

China

ROBERT F. INGER

Field Museum, Chicago, Illinois, USA

KUANGYANG LUE

National Taiwan Normal University, Taipei,

Taiwan

HIDETOSHI OTA

Department of Biology, University of the

Ryukyus, Nishihara, Okinawa, Japan

ANMING TAN

University of California, Berkeley, California,

USA

DATONG YANG

Kunming Institute of Biology, Kunming, Yunnan,

China

Chinese Herpetological Research is published by the Chinese Society for the Study of

Amphibians and Reptiles (CSSAR) as a cooperative venture between the Chengdu Institute of Biology,

Academia Sinica, and the Museum of Verrtebrate Zoology, University of California. We encourage authors

from all countries to submit articles concerning, but not limited to, Asian herpetology.

Authors should submit letter quality manuscripts in English and the original language. All tables

and figures must be of publication quality and limited to 21.5 x 28 cm (8.5 x 11 inch) sheets. A map of

study areas must accompany each manuscript, when applicable.

All correspondence within the People's Republic of China should be directed to Ermi Zhao, Editor,

Chengdu Institute of Biology, P.O. Box 416, Academia Sinica, Chengdu, Sichuan Province, People's

Republic of China.

All correspondence outside of the People's Republic of China and requests for subscription should be

directed to CSSAR, Foreign Memberships, Museum of Vertebrate Zoology, University of California,

Berkeley, California, USA 94720.

Subscriptions are $20 for unbound issues, as they become available, The yearly bound volume is

available for $20. Both can be purchased for $40. Please specify which materials you wish to recieve.

Make checks or money orders payable in US currency to CSSAR. If you do not have access to US

currency, please notify us, and we will make other arrangements.

Cover: Phrynocephalus axillaris, from Dunhuang, Jiuquan Prefecture, Gansu Province, China.

CHINESE

[EIERPETOLOGICAL

IRESEARCEH

Volume 2, No. I May, 1988

CONTENTS

ZHAO, ERMI, J. ROBERT MACEY, AND THEODORE J. PAPENFUSS!. A New Species of

Rana from Ningxia Hui Autonomous Re Stones cere sets sense niasaasanuaeas 1

MACEY, J. ROBERT, THEODORE J. PAPENFUSS, AND ERMI ZHAO2. The Snakes of

Ningxia Hui Autonomous Region as an Indication of a Herpetofaunal Corridor........ 4

AUTUMN, KELLAR, AND YAO-ZHAO WANG3. Preliminary Observations on the Ecology

of Phrynocephalus axillaris and Eremias velox in the Turpan Depression, Xinjiang

Uygur Autonomous Region, China................:sscsscssssceeseessscesseeereesseetscenss 6

1 Sino-Soviet-American Arid Asian Desert Regions Research Paper no. 1.

2 Sino-Soviet-American Arid Asian Desert Regions Research Paper no. 2.

3 Sino-Soviet-American Arid Asian Desert Regions Research Paper no. 3.

A New Species of Rana from Ningxia Hui Autonomous Region

ERMI ZHAO!, J. ROBERT MACEY2 AND THEODORE J. PAPENFUSS2

1Chengdu Institute of Biology, Academia Sinica, Chengdu, China

2Museum of Vertebrate Zoology, Univ. of California, Berkeley, USA

FIG. 1. Holotype CIB 80001 - CAS 166936 of Rana tenggerensis

Two specimens of Rana were

collected during the joint 1987 Chengdu

Institute of Biology and University of

California Expedition to the deserts of

northern China. They were found along the

banks of the Yellow River (Huang He)

where it flows along the edge of the

Tengger Desert in Ningxia Hui

Autonomous Region. Both specimens

were collected after dark in pools along the

banks of the river. Upon examination of

these specimens we determined that they

represent an undescribed species (Fig.1).

This species, named after the Tengger

Desert, is closely related to Rana

nigromaculata Hallowell. Figure 2 shows

the distribution of both forms. The

distribution shown for Rana nigromaculata

was compiled from Dixon (1956), Lui

(1950), Lui & Hu (1961), Moriya (1955),

Okada (1927), Kawamura & Nishioka

(1979), Shannon (1956), and Bannikov

(1977). The nomenclature used follows

that of Kawamura & Nishioka (1979).

Rana tenggerensis sp. nov.

Holotype: CIB 80001 - CAS 166936, an

adult male from along the north shore of the

Yellow River (Huang He), at Shapotou

Desert Research Station, Shapotou (37°

30'N 104° 58'E), Yinnan Prefecture,

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 2

(Do Rana nigromaculata

A Rana tenggerensis

area of uncertainty and

? questionable record for Qinghai Province

FIG. 2. The distribution of Rana nigromaculata and Rana tenggerensis. The question mark between

Rana nigromaculata and Rana tenggerensis represents an area of uncertainty. The other question mark to

the southwest of Rana tenggerensis represents a questionable record from Qinghai Province.

Ningxia Hui Autonomous Region, China.

The specimen was collected August 14,

1987 by J. Robert Macey and Theodore J.

Papenfuss. The Holotype is deposited at

the Chengdu Institute of Biology.

Paratype: CAS 166808, an adult male

from along the south shore of the Yellow

River (Huang He), at Shenjiatan (37° 28'N

105° 18' E), Yinnan Prefecture, Ningxia

Hui Autonomous Region, China. The

specimen was collected August 13, 1987

by J. Robert Macey and Theodore J.

Papenfuss. The specimen is deposited at

the California Academy of Sciences.

Diagnosis: This new form, Rana

tenggerensis, is a close relative of Rana

nigromaculata Hallowell, but differs from

the latter in having 1) a wider head, 2) the

nostril closer to the eye than to the tip of the

snout, 3) the tibia to snout-vent length

proportionally shorter, 4) full webbing

with almost no notch, 5) a more rugose

back, 6) no dark spots on the lips, 7) no

dark stripe in front of the shoulder.

Literature Cited

BANNIKOV, A. G., DAREVSKY, I. S., ISHCHENKO,

E. G., RUSTAMOV, A. K., AND SHCHERBAK,

N. N. 1977. Guide to amphibians and reptiles

of the USSR fauna. Prosveshchenie, Moskva. 1-

415. (in Russian.)

DIXON, J. R. 1956. A collection of amphibians and

reptiles from west central Korea. Herpetologica

12:50-56.

LIU, C. C. 1950. Amphibians of western China.

Fieldiana, Zool. Mem., v.2, 400 pp., 100 text

figs.

LIU, C.C. AND C. HU. 1961. Tailless Salientia

in China. Kolsuo-Choppan [Scientific Press],

Peiking, 364 pp, 151 fig., 28pls. (In Chinese.)

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 3

KAWAMURA, T. AND M. NISHIOKA, 1979.

Isolating mechnanisms among the water frog

species distributed in the Palearctic Region.

Mitt. Zool. Mus. Berlin, Bd. 55, Heft I, 171-

185.

MORIYA, K. 1955. Local races in the pond frog,

Rana nigromaculata, with special reference to

their distribution. Bull. Biogeograph. Soc. Japan

(recent conceptions of Japanese Fauna) 16-

19:354-359. (In Japanese.)

OKADA, Y. 1927. A study on the tailless

batrachians of Japan. Ann. Zool. Japon. 11:97-

103.

SHANNON, F. A. 1956. The reptiles and

amphibians of Korea. Herpetologica 12:22-49.

hinese Her,

vetological Research

The Snakes of Ningxia Hui Autonomous Region as an Indication

of a Herpetofaunal Corridor

J. ROBERT Macey!, THEODORE J. PAPENFUSS! AND ERMI ZHAO2

IMuseum of Vertebrate Zoology, Univ. of California, Berkeley, USA

2Chengdu Institute of Biology, Academia Sinica, Chengdu, China

Few herpetological expeditions

have made collections in Ningxia Hui

Autonomous Region and the snake fauna is

poorly known. Only two species,

Psammophis lineolatus and Agkistrodon

intermedius have been reported (Tian &

Jiang 1986). A live specimen of

Agkistrodon strauchii was examined by the

third author at Zhejiang Medical University

in Hangzhou. It was said to have come

from the Liupan Mountains. During the

joint 1987 Chengdu Institute of Biology

and University of California Expedition to

the deserts of northern China, three new

snake records were obtained from Ningxia

Hui Autonomous Region. The specimens

were all collected along the north shore of

the Yellow River (Huang He), at Shapotou

Desert Research Station, Shapotou (37° 30'

N 104° 58' E), Yinnan Prefecture, Ningxia

Hui Autonomous Region, China.

The three species we found at

Shapotou are Coluber spinalis, Elaphe

dione and Rabdophis tigrina lateralis. The

Coluber spinalis was found in a bush in an

area of low, rolling sand dunes adjacent to

the Yellow River. It was active during the

late morning in an area where

Phrynocephalus przewalskii were very

abundant. The specimens of Elaphe dione

and Rabdophis tigrina lateralis were found

in an area of cultivated fields near the river.

This site is located at the edge of the

sandy Tengger Desert. In this area the

Yellow River begins to broaden into an

interior delta after passing through a rocky

stretch with little vegetation. Here the river

flows north-northeast looping through the

Gobi Desert and finally flowing south and

east to northeastern China (Fig. 1).

As the Yellow River descends in

altitude from the Tibetan Plateu, the annual

rain fall decreases from 500 - 750 mm to

100 - 250 mm in the Gobi Desert loop of

the river. Following the Yellow River

upstream from the Pacific Coast, the same

effect occurs. Annual precipitation drops

from 500 - 750 mm to 100 - 250 mm.

Subsequently the Gobi Desert loop of the

Yellow River provides a mesic connection

from the Tibetan Plateu across the arid

Gobi Desert to the coastal region of

northeastern China. This connection

provides a route for dispersal of mesophilic

species.

Two of the snake species reported

here, Elaphe dione and Rabdophis tigrina

lateralis, follow this corridor. The direction

of dispersal for Elaphe dione cannot be

inferred to be from northeastern China,

because of its occurrence from the Black

Sea in Europe across the USSR (Bannikov

et al. 1977), Mongolia, and much of

northern China. Bufo raddei uses the

Yellow River as a mesic refuge in the same

manner as Elaphe dione and also occurs

widely to the northeast and west of the

Gobi Desert loop of the Yellow River. The

distribution of these two species in northern

China is similar in that both species have

their coastal southern distributional limits in

Jiangsu Province (Tien and Jiang 1986).

Rabdophis tigrina laterals enters the Gobi

Desert by way of the Yellow River. Its

overall distribution is throughout eastern

China except the tropical south. Another

example of the use of the Yellow River as a

corridor is found in frogs of the Rana

nigromaculata species group which also are

distributed mainly in eastern China except

the tropical south. The outlying species

occurring in the Ningxia Hui Autonomous

Region is the recently described Rana

tenggerensis (Zhou et al. 1988). It is

interesting to note that both Rabdophis

tigrina and Rana nigromaculata share

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 5

FIG. 1. The location of the Yellow River of China with approximate rainfall contours to illustrate the

aridity of the Gobi Desert loop. The location of Shapatou is represented by a dot. The arrows show

possible uni- and bi-directional dispersal routes along the corridor.

further similarity in distribution and occur

in Korea, Japan, and the southern Pacific

coastal region of the USSR. The other

species of snake reported here, Coluber

spinalis, does not follow the Yellow River

into the Gobi Desert. It is a wide-ranging

species in northern China that can be found

in both mesic and xeric conditions (Pope

1935).

Literature Cited

BANNIKOV, A. G., DAREVSKY, I. S., ISHCHENKO,

E. G., RUSTAMOV, A. K., AND N. N.

SHCHERBAK. 1977. Guide to amphibians and

reptiles of the USSR fauna. Prosveshchenie,

Moskva, 1-415. (In Russian.)

POPE, C. H. 1935. The reptiles of China. Natural

history of Central Asia, Vol.10. New York.

604 pp., figs., pls.1-27, map.

TIEN, W. S. AND Y. M. JIANG (eds). 1986.

Handbook of Chinese Amphibia and Reptilia.

Kolsuo-Choppan [Scientific Press], Beijing. 164

pp., 20pls.

ZHAO, E., J. R. MACEY AND T. J. PAPENFUSS.

1988. A new species of Rana from Ningxia Hui

Autonomous Region. Chinese Herp. Res.

2(1):1-3.

hinese Herpetological Research

Preliminary Observations on the Ecology of Phrynocephalus

axillaris and Eremias velox in the Turpan Depression, Xinjiang

Uygur Autonomous Region, China

KELLAR AUTUMN! AND YAO-ZHAO WANG2

1Cowell College, University of California, Santa Cruz, CA 95064

2Chengdu Institute of Biology, Academia Sinica, Chengdu, China

Abstract.-During the joint Chengdu Institute of Biology and the University of California expedition to the

deserts of western China in Summer 1987 we gathered data on thermoregulatory behavior and microhabitat

utilization in Phrynocephalus axillaris and Eremias velox. These data demonstrate a clear niche segregation

based on the use of bushes. The floral species diversity is very low, with only two species sampled on our

site. P. axillaris seems to make no use of the bushes in its daily activities, while E. velox spends most of

its time within 0.04 m of a bush. Asa result, P. axillaris is restricted in its activity period to mornings

and afternoons.

Introduction

The Turpan Depression of Xinjiang

Uygur Autonomous Region, China (Fig. 1)

is the second lowest point on earth (-150

m) and has been geographically isolated

since before the Pleistocene (Xi 1985).

The simplicity of both the floral and

herpetofaunal communities in the Turpan

Depression (as well as most of the Chinese

deserts) makes it an ideal region in which to

study lizard ecology. While participating in

a cooperative expedition between the

Chengdu Institute of Biology, Academia

Sinica, China, and the University of

California, USA, in the summer of 1987,

we recorded some preliminary ecological

data on Phrynocephalus_ axillaris

(Agamidae) (Fig. 2) and Eremias velox

(Lacertidae) (Fig. 3) at a site near the town

of Turpan. Teratoscincus scincus

(Gekkonidae) and Psammophis lineolatus

(Colubridae) were also present on the site.

The logistics of the expedition prevented

us from sampling over a longer period of

time, so our data are not by any means

representative and should be interpreted

only as a single sample in time and space.

To the best of our knowledge, this is the

first study in desert lizard ecology in the

People's Republic of China.

Methods

The study site, 4.4 km west of the

main mosque in Turpan (42° 56'N 89° 10'

E) on the Turpan-Jiaohe Road, Turpan

Prefecture, Xinjiang Uygur Autonomous

Region, People's Republic of China, was

representative of much of the undisturbed

vegetated habitat near Turpan. Sunrise at

Turpan was at 9:31 Beijing time (5:31

Greenwich); sunset was at 22:24 Beijing

time (18:24 Greenwich). We chose a 75 x

75 m plot that had both open and brush

microhabitats (Fig. 4). The plot was on a

sloping hill, with rocky, sandy clay

substrate. Using the line-intercept method

(Brower and Zar 1984), we sampled the

flora along two 75 m transects that trisected

the plot, north to south. The transects were

broken up into 15 m intervals. From 7

Sept., 1987 to 11 Sept., 1987, we

repeatedly and systematically transected the

site and sampled P. axillaris and E. velox.

The lizards were noosed and their cloacal

temperature recorded with a Fluke 25 heat-

resistant multimeter with thermocouple

transducer while still suspended from the

noose and with hind leg grasped (Avery

1982) if no longer than 30 seconds had

elapsed from the time of first sighting.

They were sexed, measured, uniquely toe

clipped and paint marked on the dorsal

surface with nail polish, and released. We

recorded the ground and air temperature,

and the distance to the nearest bush from

the area where the individual was first

sighted. Daily recapture data allowed us to

calculate a rough density estimate for both

species, using the Schnabel method of

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 7

FIG. 1 Desert regions of western China.

mark-recapture sampling (Giles 1971;

Schnabel 1938).

Results

1. Habitat.

Alhagi sparsiflora (Leguminosae)

(Fig 5) was the only plant sampled on a

150 m line transect. Its frequency among

fifteen 10 m intervals was 0.93. The

density and coverage were 0.31 plants/m

and 0.21, respectively. A plant of Nitraria

sp. (Zygophyllacae) was rare on the site.

Daytime air temperatures ranged from 24 to

45 °C.

2. Species interactions and behavior.

Eremias velox frequently move

from bush to bush and forage most often in

the shade. Phrynocephalus axillaris emerge

from their burrows, which seem to all be in

open ground, and remain motionless unless

approached. Small rocks allow the lizards

to orient with respect to the sun.

Phrynocephalus axillaris.do not seem to

make any use of the shade provided by the

bushes. We observed intraspecific

agonistic behavior in E.velox. In one case,

an individual pursued another for over 20

m. Intra- and interspecific interactions

were not evident in P. axillaris.

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 8

FIG. 3. Eremias velox at Turpan, Xinjiang Uygur Autonomous Region, China.

May 1988 Chinese Herpetological Research WolwZ- No mseap

FIG. 4. (ABOVE) Study site at Turpan, Xinjiang Uygur Autonomous Region, China.

FIG. 5. (BELOW) Alhagi sparsiflora, the dominant plant at the Turpan study site, Xinjiang Uygur

Autonomous Region, China.

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 10

Psammophis lineolatus were active in the

early morning and late afternoon in the low,

spiny branches of A. sparsiflora. One

individual stayed motionless, without any

contact with the substrate. This snake was

very wary and retreated to its burrow when

approached. The stomach contents of

another specimen captured in a bush

included a juvenile E. velox. Teratoscincus

scincus were present in great number on the

site. After sunset, these lizards emerge

from their extensive burrows, contrary to

Pianka's (1986) prediction that nocturnal

lizards should not occur at such high

latitudes. We also observed Teratoscincus

przewalskii only after dark in the

Taklimakan. Both species spend much of

their time motionless, but frequently move

several meters at a time, presumably

foraging for invertebrates.

3. Habitat utilization and thermoregulation.

The mean distance to the nearest

plant was significantly greater (t = 8.65,

DF = 87, a = 0.05) in P. axillaris (2.17 m,

std. dev. 2.00) than in E. velox (0.04 m,

std. dev. 0.10). There is not a significant

linear relationship between body

temperature and distance to the nearest plant

in either P. axillaris or E. velox (Fig. 6&7).

P. axillaris were active (on clear days) from

1:29 to 3:39 after sunrise and from 5:30 to

1:45 before sunset (11:00 to 13:10 and

from 16:50 to 20:35 Beijing time). At the

point of sighting, air temperatures ranged

from 26.6 to 41.7 °C, and ground

temperatures ranged from 29.6 to 42.0 °C.

Body temperature vs. air temperature can

be approximated by the linear equation, BT

= 16.78 + 0.60AT (R2 = 0.58, t= 7.54, a

= 0.05) (Fig. 8). Mean body temperature

in P. axillaris was 36.8 °C (std. dev. 3.58)

and ranged from 29.7 to 42.8 °C. Eremias

velox were active (on clear days) from 1:24

after sunrise to 1:35 before sunset (10:55 to

20:45 Beijing time). Air temperatures

ranged from 24.8 to 42.0 °C , and ground

temperatures ranged from 24.7 to 44.7 °C.

Body temperature vs. air temperature can

be approximated by the linear equation, BT

= 19.94 + 0.53AT (R2 = 0.67, t = 11.66,

a = 0.05) (Fig. 9). Mean body temperature

in adult E. velox was 38.4 °C (Std. Dev

2.94), and ranged from 26.3 (at

emergence) to 43.5 °C. Juvenile E. velox

seem to spend most of their time under

dense cover; we were not able to effectively

sample the juveniles.

4. Density.

Rough density estimates on the site

are 66+11.49 (117/hectare) (a = 0.05) for

adult E.velox, and 18+2.56 (32/hectare) (a

= 0.05) for P. axillaris. _Psammophis

lineolatus were sighted 3 times during the

study. A very rough density estimate

obtained by eyeshining T. scincus is 30

individuals on the site.

5. Body size and evidence of predation.

Mean snout-vent length was 43 +14

mm, and mean tail length was 66 +21 mm

in P. axillaris; 36% of these fall in the 45-

49 mm range (Fig. 10). Mean snout-vent

length was 59 +12 mm, and mean tail

length was 97 +51 mm in E.velox; over

71% of these fall in the 55-64 mm range

(Fig. 11). Twenty-nine out of 59 E. velox

showed evidence of tail regeneration, with

a mean regenerated tail length of 90 +58

mm. There was no evidence of tail loss in

P. axillaris.

Discussion

While our results are not

unexpected, it is interesting to note that

Phrynocephalus does not shun bushes

throughout its range. In Korla, Xinjiang

Uygur Autonomous Region (south of

Turpan in the Taklimakan Desert),

Dunhuang, Gansu Province (the extreme

eastern Taklimakan Desert), and Shapotou,

Ningxia Hui Autonomous Region (Central

Gobi Desert), Phrynocephalus is found

very Close to and under bushes. Eremias

were very rare in the area we sampled near

Korla and it is possible that a greater

microhabitat diversity may be available to

Phrynocephalus in the absence of Eremias.

The temperature, however, was much

cooler when we were in Korla and the

significance of a comparison of P. axillaris

between Turpan and Korla is questionable

at best. P. axillaris seems similar in form

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 11

Dist to Bush (m)

>

0

25 30 35 40 45

Body Temp (C)

FIG. 6. Distance to the nearest bush vs. air

temperature in Phrynocephalus axillaris at Turpan,

Xinjiang Uygur Autonomous Region, China. P.

axillaris were not found closer than 0.25m to a

bush. There is no significant linear relationship

between body temperature and distance to bush.

Body Temp (C)

25 30 35 40 45

Alr Temp (C)

FIG. 8. Body temperature vs. air temperature in

Phrynocephalus axillaris at Turpan, Xinjiang

Uygur Autonomous Region.

to Callisaurus draconoides (Iguanidae) and

in behavior to Phrynosoma platyrhinos

(Iguanidae) of North America. We

observed another distinct behavioral type

(that we term the "Callisaurus" type) in

Phrynocephalus przewalskii at Wuwei,

Gansu Province. These lizards were very

wary and would flee for long distances if

approached. They seemed to have a greater

capacity for sustained activity than the P.

axillaris of Turpan and Korla. E. velox

seems similar to small Cnemidophorus

0.8

0.6

0.4

Dist. to Bush (m)

0.2

0.0

25 30 35 40 45

Body Temp (C)

FIG. 7. Distance to the nearest bush vs. air

temperature in Eremias velox at Turpan, Xinjiang

Uygur Autonomous Region, China. E. velox were

most often found in or close to bushes. There is

no significant linear relationship between body

temperature and distance to bush.

Body Temp (C)

25 30 35 40 45

Alr Temp (C)

FIG. 9. Body temperature vs. air temperature in

Eremias velox at Turpan, Xinjiang Uygur

Autonomous Region, China. Arrow indicates an

emergence temperature.

(Teiidae) of North America. Mean body

temperatures in E. velox (38.4 °C) are

similar to those observed in the closely

related Pedioplanis lineoocelata (38.4 °C)

and Pedioplanis namaquensis (38.5 °C),

which were formerly placed in the genus

Eremias (Brattstrom 1965).

The activity period of P. axillaris is

restricted to mornings and afternoons, and

we conclude that this is related to the

lizards’ preference for open microhabitat.

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 12

No. of Individuals

30-34 35-39 40-44 45-49 50-54 55-59

Snout-Vent (mm)

FIG. 10. Snout-vent frequency in P. axillaris at

Turpan, Xinjiang Uygur Autonomous Region,

China.

Average Precipitation (mm)

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

FIG. 12.

Xinjiang Uygur Autonomous Region, China.

Eremias velox is able to actively

thermoregulate and forage throughout the

day. This lizard is a wide-forager, may be

territorial, maintains a high density, and

experiences a high degree of predation.

Psammophis lineolatus and T. scincus are

probably the primary predators, the latter

foraging nocturnally above ground, and

diurnally in burrows. Psammophis

lineolatus probably preys more intensely on

E. velox than on P. axillaris since it is

associated with bushes, and since E. velox

is a widely-foraging lizard and is more

likely to be encountered by a sit-and-wait

snake (Huey and Pianka 1981) The diet

analysis of Psammophis lineolatus and T.

scincus (J. R. Macey and Y. Z. Wang 1988

in prep.), as well as the small size of both

predators suggest that most of the

30

ct}

5 20

bs

=

wo

p}

~~

San0

3

Zz

30-34 35-39 40-49 45-49 50-54 55-59 60-64 65-69

Snout-Vent (mm)

FIG. 11. Snout-vent frequency in E. velox at

Turpan, Xinjiang Uygur Autonomous Region,

China.

Average Temperature (C)

Jan FebMar Apr May Jun Jul Aug Sep Oct Nov Dec

Month

Average monthly rainfall at Turpan, FIG. 13. Solid bars denote average daily air

temperature for each month of the year at Turpan,

Xinjiang Uygur Autonomous Region, China.

Vertical lines above 0°C denote maximum average

temperature; lines below 0°C denote minimum

average temperature.

predation pressure is on juvenile E. velox.

The lack of tail loss in P. axillaris is

inconclusive since agamids do not easily

autotomize their tails.

Mean annual rainfall (Fig. 12) at

Turpan is 16.6 mm (Anon. undated),

which contributes to the extremely low

floral species diversity. Mean temperature

exceeds 20°C only during a five-month

period from April to September (Fig. 13).

This probably limits the activity period of

reptiles to five months or less.

We plan to expand our study to

three sites in 1988: Shapotou, Ningxia Hui

Autonomous Region in the Gobi Desert, a

Taklimakan Desert site, as well as the

Turpan site. This will allow us to compare

May 1988 Chinese Herpetological Research Vol. 2, No. 1, p. 13

the thermal niche breadth and microhabitat

selection in a manner similar to Grant and

Dunham (1988) in Eremias-

Phrynocephalus communities across the

Chinese deserts. We also plan to

investigate the behavioral differences in

Phrynocephalus at the various regions in a

physiological context.

Acknowledgements

We would like to thank Ermi Zhao

of the Chengdu Institute of Biology

(Academia Sinica, China) for support of the

project. We also thank Theodore J.

Papenfuss and J. Robert Macey of the

Museum of Vertebrate Zoology (Univ.

Calif. Berkeley) for editing, assistance in

the field, and use of specimens, Margaret

Fusari of the Univ. Calif. Santa Cruz for

invaluable advice, Jonathan Losos of the

Museum of Vertebrate Zoology (Univ.

Calif. Berkeley) for last minute editing,

Terry Horn of Scimitar Meats for the

donation of gourmet beef jerky, and Zhang

Zemin of the Chengdu Institute of Biology

(Academia Sinica, China) for making our

expedition run smoothly.

Literature Cited

ANONYMOUS. undated. Turpan brochure.

Promotion Dept. of the National Tourism

Admin. of the People's Republic of China,

China Travel and Tourism Press, Turpan,

Xinjiang Uygur Autonomous Region, China.

AVERY, R. A. 1982. Field studies of body

temperatures and thermoregulation. In Biology

of the reptilia vol. 12, pp.941-166. Academic

Press, London.

BRATTSTROM, B. H. 1965. Body temperatures of

reptiles. Amer. Midl. Nat. 73(2):377-422.

BROWER, J. E. AND J. H. ZAR. 1984. Field and

laboratory methods for general ecology, 2nd ed.

William C. Brown Pub., Dubuque, Iowa.

HUEY, R. B. AND E. R. PIANKA. 1981.

Ecological consequences of foraging mode.

Ecology 62(4):991-999.

GILES, R. H. 1971. Wildlife Management

Techniques. Wildlife Society, Washington DC.

GRANT, B. W. AND A. E. DUNHAM. 1988.

Thermally imposed time constraints on the

activity on the desert lizard Sceloporus merriami.

Ecology 69(1):167-176.

PIANKA, E. R. 1986. Ecology and natural history

of desert lizards. Princeton Univ. Press,

Princeton, New Jersey.

SCHNABEL, Z. E. 1938. Estimation of the total

fish population of a lake. Amer Math. Monthly.

45:348-352.

XI, Y. (ed.). 1985. Atlas of the paleogeography of

China, compiled by Institute of Geology,

Chinese Academy of Geological Sciences and

Wuhan College of Geology. Cartographic

Publishing House, Beijing, China.

oN 5

as

} oe aie E

j Eee ma at hors

a

i

i i i j

oP Ai

5 2 \ i ae

x cu

i wees

2 ’ ; it £15*

a j

j x

Z

CHINESE

[ALTERPETOLOGICAL

IRESEARCH

Volume 2, No. 2 April, 1989

CONTENTS

WEN, YETANG. A New Species of the Genus Paramesotriton (Amphibia: Caudata) from

Guangxi and a Comparison with P. QUAN QXIENSIS..............cccecceecececcennecceennees 15

GREENE, HARRY W. Defensive Behavior and Feeding Biology of the Asian Mock Viper,

Psammodynastes pulverulentus (Colubridae), a Specialized Predator on Scincid

Estat sists Mindins Slosjce Wale ciclo caietinaitie nena melee ne ae eae tate a cle eae cna eR elscaie see ic eee 21

TAN, ANMING, GUANFU WU, AND ERMI ZHAO. The Karyotype of the Treefrog

Rhacophorus reinwardtii (Boie) from Xishuangbanna Daizu Autonomous Prefecture,

Yunnan: Provinces Chinai ac eee seo a aE I 33

ZHENG, XIAOMAO, AND GUANFU WU. Cytotaxonomical Studies on Chinese Pelobatids

V. The Karyotypes, C-bands and Ag-NORs of Megophrys omeimontis and

OR COLAIAK. ) SCRMIULI Gs 2 aoe a See ROO EOL ee 37

YANG, YUHUA, FUJI ZHANG, AND ERMI ZHAO. Karyotypic Analyses of Four Species in

Hourn,Genera of -Colubrid i Snlakess execs cso ee on eee ae ee ane ena 46

YANG, YUHUA, ZHENGFA GAO, AND ERMI ZHAO. Karyotypic studies of

Sphenomorphus indicus (Scincidae) and Takydromus septentrionalis

(IE aCertidae ce Eee ea OS Ne a ON HO RM Or 55

AUTUMN, KELLAR, AND BATUR HAN!. Mimicry of Scorpions by Juvenile Lizards,

Neratoscincus: roborowskit \ (GeKkonidae) ia st eee 60

BUSKIRK, JAMES R. New Locality Records for Chinese Non-Marine Chelonian......... 65

ZHANG, ZHENGDONG. A Major Research Achievement in Captive Reproduction of

Chinese wAN Batons. Men Mais. OMe Meee Date CO dee meh A uel ue aw en AS 69

1 Sino-Soviet-American Arid Asian Desert Regions Research Paper no. 4.

A New Species of the Genus Paramesotriton (Amphibia: Caudata) from

Guangxi and a Comparison with P. guangxiensis

YETANG WEN!

Department of Biology, Guangxi Medical College, Nanning, Guangxi, China

Abstract. -A new species of Paramesotriton is described from Guangxi Zhuang Autonomous Region.

This species is characterized by tips of the fore limbs which exceed the anterior margin of the eyes to a

greater extent and granular warts that are much more dense than observed in P. guangxiensis.

Key Words: Amphibia, Caudata, China, Guangxi, Paramesotriton.

FIG. 1. Holotype of Paramesotriton fuzhongensis, GMC 81-021 from Gupo Hill, Wanggao (24° 35'N

111° 25'E), Guangxi Autonomous Region, China.

Paramesotriton fuzhongensis

sp. nov.

Holotype: GMC 81-021 (Fig. 1), an

adult male from Gupo Hill, Wanggao

(24°35'N 111°25'E), Zhongshan Xian

(county), Guangxi Autonomous Region,

China, altitude 400 m. The specimen was

collected on August 12, 1981 by Chaoliang

Lai and is deposited in Guangxi Medical

College collection (GMC).

Allotype: GMC 81-022 an adult male

was collected with the holotype.

Paratype: Two males GMC 86-006 &

86-009 and three females GMC 86-004,

86-005, & 86-007 were sent by Fuchwan

County Science and Technology

Committee. The specimens were collected

from Xilin Hill, Fuchwan Xian (county)

(QiacS OuNGee lle oslGgE Pee Gilanicexe

Autonomous Region, China, altitude

500m. The exact date and collector are

unknown.

Diagnosis: This new species closely

resembles Paramesotriton guangxiensis

Huang, Tang and Tang, but differs from

the latter in the following ways: 1) When

the fore limbs are drawn forward, their tips

exceed the anterior margin of the eyes to a

greater extent. 2) When the fore and hind

limbs are drawn simultaneously along the

flank toward the middle, the palm and

Vol. 2, No. 2, p. 16 Chinese Herpetological Research April 1989

TABLE. 1. Measurements of the Holotype, Allotype, and Paratypes of Paramesotriton fuzhongensis. The

mean given is only for the Paratypes

Paratype Paratype Paratype Paratype

Holotype = Allotype | Paratype

Mal Male Female Female Female

le Male Male

GMC 81-021 81-022

86-009 86-004 86-005 86-007

Snout-vent length 88 75 &2 81 80 fo) B 77.0 -

Head length 27 24 2B 23 34 21 21 22.3 0.29

Head width 20 17 16 17 16 15 14 15.4 0.20

Snout length 09 08 07 08 08 07 08 07.5 0.10

Internasal space 06 (0's) it'.) 6 0 04 04 04.4 0.06

Diameter of eye 06 05 05 it's) 05 04 065 04.3 0.06

Interorbital space 08 07 07 07 07 07 07 06.9 0.09

Axilla-groin 39 32 35 34 37 33 34 34.6 0.45

Fore limb length 29 25 25 27 26 20 2 24.0 0.31

Hind limb length 30 25 27 28 27 2 23 25.4 0.33

Tail length 78 58 66 B 79 65 65 70.6 0.92

Tail-base width 10 07 0 09 10 08 09 08.9 0.12

Tail height 14 12 12 12 11 10 10 10.9 0.14

tarsus are overlapping. 3) The granular

warts have a higher frequency and density.

4) The coloration differs between the two

species.

Description of holotype: Total length

166 mm, snout-vent length (SVL) 88 mm;

head depressed slightly, longer than broad;

slightly ladder shaped in dorsal aspect, and

hind region wider than fore; snout

apparently longer than diameter of eye,

with tip even and slightly bent, projecting

far beyond anterior margin of lower jaw,

with canthus rostalis prominent; loreal

region slopes somewhat outward; top of

head has two ridges behind eyes, reaching

back of jugular plica and has nostrils

lateral, on tip of snout that are not seen in

dorsal view; oral gap exceeds posterior

margins of eyes; upper labial fold is

prominent and more developed under eyes;

vormerine teeth V-shaped; tongue

ovalform, lateral margins free, adhering to

floor of mouth; lengths of fore and hind

limbs nearly equal, with hindlimbs stouter

and tip of fore-limb reaching midway

between nostril and eye; adpressed limbs

overlap along flank palm and tarsus;

holotype has four fingers and five toes,

very expanded and unwebbed, with blunt,

round tips; first finger and toe very small;

tail is shorter than SVL with thick base that

gradually becomes laterally compressed,

nearly a thin sheet at end; cloacal walls

swollen and protuberant with many layers

of papillae.

Skin very rough with prominent,

protuberant dorsal ridge, anterior end

separates and reaches posterior margin of

the eyes; irregular costal grooves present on

flanks and anterior part of tail; dense

granules or warts cover dorsum of head,

loreal region, throat, dorsum of body,

flank, anterior part of tail and dorsal

surfaces of limbs; large dorsolateral warts

which form two longitudinal ridges; labial

folds, belly, ventral surfaces of limbs,

fingers, toes, palms, and tarsus smooth.

Measurements of the new species are listed

in Table 1.

In life, dorsum of head, back, lateral

area of body and dorsal surface of limbs

olive or have small black spots; anterior

part of tail light brown, fading at end; sides

have grayish-white stripes or various black

spots; throat and belly light pale with

irregular reddish orange spots that are

smaller and more dense on throat; ventral

caudal fin from tip of tail, to and including

vent, and ventral surfaces of limbs reddish

orange. In preservative color fades to

whitish-gray.

Tail of female longer and lower than

male; reddish orange spots on throat larger

than male; vent of female shorter than that

of male, not swollen, no papillae.

Habitat: The new species is restricted to

streams at mid-slope where a broad leafed

forest is present. Adults are usually found

under rocks, and sometimes ashore. This

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 17

@ Paramesotriton fuzhongensis

<= indicates type locality

A Paramesotriton guangxiensis

FIG. 2. Distribution of Paramesotriton fuzhongensis and Paramesotriton guangxiensis in Guangxi

Autonomous Region China.

species is sympatric with P.

caudopunctatus on Xilin Hill, Fuchuan

Xian (County).

Distribution: Paramesotriton

guangxiensis is known only from the type

locality of 478 m, Paiyang mountain,

Mingjiang (22°09'N 107°12'E), Ningming

County, Nanning Prefecture, Guangxi

Zhuang Autonomous Region, China. The

new species P. fuzhongensis ranges from

Fuchwan (24°50'N 111°16'E) south by

southeast to Wanggao (24°35'N

111°25'E). Both localities are in Wuzhou

Prefecture, Guangxi Zhuang Autonomous

Region, China. Mingjiang, where P.

guangxiensis occurs, is in the extreme

southwestern portion of Guangxi

Autonomous Region. In contrast, the two

sites where the new species is found are in

the northeastern part of Guangxi

Autonomous Region. These two areas are

separated by the Xi River system which

drains most of Guangxi Zhuang

Autonomous Region (Fig.2).

Comparisons: The new species differs

from P. guangxiensis in a number of

morphological traits besides those listed in

the diagnosis. I have made additional

comparisons as follows:

The head of the new species is longer

and narrower (head width/head length =

69.5 %), and broader in P. guangxiensis

(head width/head length = 78.2 %) [Fig.

3]. The tail of the new species is longer and

Vol. 2, No. 2, p. 18 Chinese Herpetological Research April 1989

Ler

HIG. 3. X-Ray of Paramesotriton fuzhongensis (above) and Paramesotriton guangxiensis (below).

FIG. 4. Skulls of Paramesotriton fuzhongensis

Paramesotriton guangxiensis (right).

lower (tail length/SVL = 91.7 %, tail

height/SVL = 14.0 %), and is shorter and

higher in P. guangxiensis (tail length/SVL

= 82.5 %, tail height/svl = 19.3 %). The

axilla to groin ratio is longer in the new

(left) with slender fronto-squamosal arch, and

species (axilla to groin/SVL = 44.9 %) than

in P. guangxiensis (axilla to groin/SVL =

41.0 %). The data for P. guangxiensis are

according to Huang, Tang and Tang

(1983).

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 19

FIG. 5. Skulls of Paramesotriton fuzhongensis (left) with maxillary separated from the pterygoid, and

Paramesotriton guangxiensis (right).

The fronto-squamosal arch of the new

species is slender and the outer edge is

almost a straight line. It is larger and stout,

with the outer edge nearly a right angle, in

P. guangxiensis (Figs. 3 & 4). The

maxillary of the new species is separated

from the pterygoid by a large interval,

while it almost touches the anterior tip of

the pterygoid in P. guangxiensis (Fig. 5).

The notch of nares internus on the vomer in

the new species is shallow and vertical. It is

5mm

FIG. 6. Hyoid aparatus of Paramesotriton

fuzhongensis (left), and Paramesotriton

guangxiensis (right). Arrows show ceratohyal.

deeper and transverse in P. guangxiensis

(Fig. 5). The trunk vertebrae of the new

species are slender and the ribs direct more

backward; the trunk vertebra are stout and

the ribs direct more laterally in P.

guangxiensis (Fig. 3). The seventeenth

vertebra (third caudal vertebra) begins to

have a hemal canal in the new species,

instead of the sixteenth vertebra (second

caudal vertebra) in P. guangxiensis. The

hyoid apparatus in the two species differs

in appearance, the ceratohyal being

apparently shorter in the new species (Fig.

6).

Literature Cited

CHANG, M. 1955. The amphibia of China. In

Fudan Univ. Series. pp. 1-75. (In Chinese.)

CHANG, T., AND A. M. BORING. 1934-35.

Studies in variation among the Chinese

amphibia. Peking Nat. Hist. Bull. 9(4):327-

358.

HU, S., E. DJAO!, AND C. LIU. 1973. A

survey of amphibians and reptiles in Kweichow

Province, including a herpetofaunal analysis.

Acta Zoologica Sinica 19(2):149-178. (In

Chinese.)

1 Currently, the name E. Djao is written E. Zhao

(ed.).

Vol. 2, No. 2, p. 20 Chinese Herpetological Research April 1989

HUANG, Z., Z. TANG, AND Z. TANG. 1983. A

new species of the genus Trituroides from

Guangxi, China. Acta Herpetologica Sinica

1983-2(2):37-39. (In Chinese.)

TIAN, W., AND Y. JIANG, (eds.) 1986.

Identification handbook of Chinese Amphibia

and Reptilia. Science Press, Beijing. pp.1-41.

(in Chinese.)

ZHAO, E., AND Q. HU. 1984. Studies on

Chinese tailed amphibians. Sichuan Scientific

and Technical Publishing House. pp. 1-50. (In

Chinese.)

Defensive Behavior and Feeding Biology of the Asian Mock Viper,

Psammodynastes pulverulentus (Colubridae),

a Specialized Predator on Scincid Lizards

HARRY W. GREENE!

1Museum of Vertebrate Zoology and Department of Zoology, University of California,

Berkeley, California , USA, 94720

Abstract. -Mock vipers react to simulated predation by immobility, locomotor escape, striking, and

biting; they resemble some sympatric, venomous pit vipers in color pattern, enlarged front teeth, and head

shape. One hundred and thirteen prey items included lizards, primarily scincids (70.8%); frogs (23.0%); and

colubrid snakes (6.1%). There is no evidence of ontogenetic dietary variation and slight evidence of

geographic dietary variation. Prey is subdued with venom and constriction, then swallowed head-first.

Skinks are difficult for some predators to grasp, because teeth and talons slide on their hard, shiny scales and

because their trunk musculature permits vigorous axial bending. Enlarged anterior maxillary and dentary

teeth in Psammodynastes pulverulentus and several other genera of snakes evidently concentrate forces,

function as pawls on a ratchet, and prevent scincid lizards from rolling off the tooth rows. Enlarged anterior

teeth and a diet of skinks perhaps have evolved several times in colubrid snakes, but phylogenctic

information is not available to resolve historical patterns of adaptation, exaptation, and convergence for

multiple biological roles. The evolutionary success of scincids might result in part from decreased

vulnerability to unspecialized lizard predators.

Key words: Snakes, Psammodynastes, Defense, Diet, Dentition, Adaptation.

Introduction

Much of the (morphological

diversification of snakes probably is related

to defense against predators and to feeding,

especially within the Caenophidia (or

Colubroidea, "advanced snakes"; e.g.,

Gans 1961, Savitzky 1980, 1983,

Ananjeva and Orlov 1982, Cundall and

Greene 1982, Greene 1983, 1988a,

Cundall 1987). Nevertheless, despite

longstanding interest in that topic, few of

the more than 2000 species have been

subjected to detailed studies. The need for

additional information is most acute for the

Colubridae (sensu lato), because that

largest of snake families is especially

diverse in terms of cephalic morphology,

defense, and diet (e.g., Malnate 1960,

Savitzky 1983, Greene 1988a).

This paper summarizes the antipredator

behavior and feeding biology of

Psammodynastes pulverulentus, a small,

rear-fanged, tropical forest snake that

occurs at low to moderate elevations over

much of southeastern Asia and the Indo-

Australian archipelagos (Wall 1910a, Smith

1943, Rasmussen 1975, Leviton 1983). A

case study of this species is of interest for

three reasons, the first two of which are

addressed in the Discussion section:

(i) Psammodynastes pulverulentus has

an unusual and narrow diet that is

functionally correlated with grooved,

posterior fangs and enlarged anterior teeth.

Its response to predators includes biting

and overall resemblance to a venomous pit

viper, such that the derived dentition might

fill multiple biological roles.

Understanding the relationships among

morphology and behavioral ecology in

mock vipers will be relevant to future,

synthetic considerations of snake evolution,

especially if the fangs of highly venomous

proteroglyphous snakes (Elapidae) are

indeed derived from enlarged anterior teeth

(e.g., Hoffstetter 1939, Smith 1952,

Rasmussen 1985, McDowell 1986).

(ii) Unlike many other lizards, scincids

are relatively safe from generalized

predators. Nevertheless, mock vipers eat

mainly skinks, suggesting that both might

have been involved in a coevolutionary

"predator-prey race". With additional

information on population biology and

phylogenetic relationships, these taxa might

prove useful in elucidating circumstances

Vol. 2, No. 2, p. 22 Chinese Herpetological Research April 1989

under which that phenomenon can occur

(see Mitter and Brooks 1983, Abrams

1986, Greene 1988a).

(iii) Psammodynastes pulverulentus is a

widespread and sometimes common snake,

judging from museum collections and

comments in the literature (e.g., Wall

1910a, Rasmussen 1975), so it might play

an important role in local ecosystems as

predator and prey of other vertebrates (cf.

Inger and Colwell 1977, Brown and Alcala

1986, Greene 1988b).

Methods

I examined all specimens of

Psammodynastes pulverulentus in the

American Museum of Natural History

(AMNH); California Academy of Sciences;

Field Museum of Natural History; Museum

of Vertebrate Zoology, University of

California, Berkeley (MVZ); National

Museum of Natural History; and

Rijksmuseum van Naturlijke Historie

(RMNH). Only stomach contents were

examined, thereby avoiding bias from

differential digestability of prey (see

Savitzky 1981). Direction of ingestion

(inferred from orientation in the gut),

identification, and linear dimensions of

prey were recorded whenever possible.

Intact prey (or a reference specimen of

comparable size) and the predator snakes

were weighed after blotting and draining

them briefly on paper towels.

Abbreviations refer to snout-vent length

(SVL) and prey/predator mass ratio (MR).

My results also incorporate literature

records (Wall 1908, 1909, 1910a, 1910b,

1910c, 1926, Pope 1929, Mertens 1930,

Smedley 1931, Brongersma 1934, Taylor

and Elbel 1958, Saint Girons 1972,

Auffenberg 1980, Leviton 1983), and I

took care to account for redundancy among

them and museum specimens (e.g.,

Brongersma 1934 and RMNH 6221).

Behavioral observations were made on an

adult male (SVL 325 mm, 16 g, now MVZ

172412) from an unknown locality.

Results

Morphology and Systematics

Psammodynastes pulverulentus is a

small snake (maximum SVL of females ca.

630 mm, Wall 1910a). There is sexual

dichromatism, but the dorsal color is

always some shade of brown or gray. In

general, snakes from the mainland and Java

have lower ventral and caudal scale counts

and shorter tails than do those from islands,

differences that are perhaps correlated with

increased arboreality in the latter

(Rasmussen 1975).

Beginning anteriorly, the first 2-3

maxillary teeth of Psammodynastes

pulverulentus are small, followed by one or

two markedly enlarged teeth, a diastema, 4-

12 small teeth, and two enlarged, grooved

fangs; the large anterior and posterior teeth

are approximately twice as long as the

intervening small teeth. Secretions of the

well-developed Duvernoy's glands

presumably are conducted by the grooved

posterior maxillary fangs. There is

geographic variation in the number of small

maxillary teeth, with a mode of eight or

nine on the Philippine Islands, Sulawesi,

Flores, Komodo, and Enggano, and a

mode of five to seven elsewhere in the

range. The anterior dentary teeth are

enlarged and followed by smaller, widely

spaced teeth (based on West 1895, Wall

1910a, Taub 1967, Gabe and Saint Girons

1971, Rasmussen 1975, and AMNH

27782; see Figs. 1, 2).

The only other species in the genus,

Psammodynastes pictus, has dentition

similar to that of the high tooth count

populations of P. pulverulentus. Grossly

similar dentition, with abruptly enlarged

anterior or middle maxillary and dentary

teeth are found in Old World colubrids of

the genera Ahaetulla, Cyclocorus,

Lepturophis, Lycodon, Lycophidion,

Macroprotodon Mehelya, and Psammophis

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 23

FIG. 1. Maxillae and dentition of (A) Psammo-

dynastes pulverulentus, (B) Lycodon aulicus, and

(C) Ahaetulla nasuta (adapted from Smith 1943).

Anterior to right; not to scale.

(Boulenger 1893, West 1895, Smith 1943,

Grandison 1972, Marx and Rabb 1972,

Broadley 1983, Savitzky 1981), and in the

Round Island bolyeriids Bolyeria and

Casarea (Cundall and Irish 1989).

However, in some Ahaetulla the anterior

teeth are grooved (Hoffstetter 1939,

Underwood 1967), in Lycophidion and

Mehelya the duct of Duvernoy's gland

opens near the enlarged anterior teeth

(Savitkzy 1981), and in bolyeriids the

disparity in tooth length is functionally

enhanced by an intramaxillary joint

(Cundall and Irish 1989). In Ditypophis

vivax of Sokotra, the maxillary teeth

increase in length from anterior to

posterior, followed by a diastema, a single

small tooth, and two enlarged grooved

fangs (Parker 1949). Pythonodipsas

carinatus, of southern Africa, is a rear-

fanged colubrid with enlarged anterior

palatine and dentary teeth that might be

functionally equivalent to those described

above (Marx et al. 1982).

The relationships among Old World

colubrid genera exhibiting the

heterogeneous dentition described above

have not been resolved. Smith (1943)

grouped Psammodynastes and

Psammophis, but noted that the former

seemed distant from all other colubrids.

Parker (1949:90) viewed the maxillae of

Ditypophis and Psammodynastes as "very

similar," and thought those genera as well

as Pythonodipsas are closely related.

Underwood (1967) placed Ahaetulla and

Psammophis in the Colubridae;

Cyclocorus, Lycodon, Lycophidion, and

Mehelya in the Lycodontinae of the

Dipsadidae; and Ditypophis,

Macroprotodon, and Psammodynastes in

the Boiginae of the Homalopsidae. Recent

biochemical approaches to snake

systematics have not included those genera

(Dowling et al. 1983, Dessauer et al. 1987,

Cadle 1988). Irregardless of relationships

among Psammodynastes and other

colubrids exhibiting enlarged anterior or

middle maxillary teeth, it is clear that

condition is uncommon and derived relative

to other advanced snakes (Marx and Rabb

1972). Lacking outgroup comparisons,

Rasmussen's (1975) conclusion that a high

number of small maxillary teeth is primitive

for Psammodynastes seems premature.

Defensive Behavior and Mimicry

Many snakes are eaten by a variety of

other reptiles, birds, and mammals (Greene

1988a); the only definite record of

predation on Psammodynastes

pulverulentus is RMNH 6221, from

Sulawesi, found in the stomach of a

colubrid snake (Boiga dendrophila,

Brongersma 1934). Mock vipers exhibit a

hierarchy of defensive reactions in response

to simulated predation by humans,

beginning with immobility and then

vigorous attempts at locomotor escape

("even indulging in a series of leaps," Wall

1910a:75). When cornered or restrained,

they typically assume a striking coil, with

head erect and forebody retracted into

sigmoid curves, and bite readily (Wall

1910a, Wall 1910c, Schmidt 1927, Pope

Vol. 2, No. 2, p. 24

Chinese Herpetological Research

April 1989

FIG. 2. Lateral view of live Psammodynastes pulverulentus holding a prey lizard (Anolis carolinensis).

Note buccal tissue surrounding the enlarged anterior maxillary and dentary teeth of the snake.

1929, Mertens 1930, Frith 1977, pers.

obs.). Smith (1915:174) saw one that

"...was Shy and very active, but made no

attempt to bite when handled." The tissue-

covered, enlarged anterior teeth are readily

visible when the snake's mouth is opened

(Schmidt 1927, Pope 1929, cf. Fig. 3),

and the effect is reminiscent of the open-

mouthed threat of some pit vipers. I noted

no effects, other than brief bleeding, from a

bite on the finger by MVZ 172412.

According to Wall (1910a:73), the

similarity between Psammodynastes

pulverulentus and a Himalayan pit viper,

Agkistrodon himalayanus, "...is especially

striking...and I know of no more

remarkable resemblance between any two

snakes of different families...The short and

rather stout body, contracted tail, flattened

head [there is a distinct canthus rostralis],

swollen lips, large eye with vertical pupil,

lustreless dorsal scales, and highly polished

ventral plates are all very characteristic

viperine traits, but the resemblances do not

stop here, for its attitude of menace is very

like that of vipers, added to which it is

viviparous in habit."

The resemblance of Psammodynastes

pulverulentus to dangerously venomous

snakes (also noted by Soderberg 1967 and

Saint Girons 1972) is subjectively

impressive and merits further study (see

Pough 1988). However, the mimicry

hypothesis is complicated by the fact that

the color pattern is cryptic in the snake's

natural habitat (Wall 1910a), that the

resemblance to sympatric pit vipers is

probably less precise in many parts of its

range than described by Wall (1910a; cf.

Smith 1943, Lim 1979), and that the

defensive posture is widespread among

other colubrid snakes and thus need not be

explained by mimicry (Kroon 1975,

Greene 1988a, Pough 1988).

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 25

Diet

One hundred and nine Psammodynastes

pulverulentus from throughout the species’

range contained 113 prey items

(x=1.03/snake): 65 scincid lizards (57.5%;

including 1 "young" Eumeces

quadrilineatus, 1 Lygosoma spp., 3

Sphenomorphus florensis, 2 S. indicus, 1

S. schlegeli; most of the rest appeared to be

lygosomines), 1 limbless anguid lizard

(0.9%; Ophisaurus gracilis), 2 lacertid

lizards (1.8%; 1 Tachydromus sp., 1 T.

Sauteri), 5 gekkonid lizards (4.5%, 1

Cyrtodactylus darmandvillei, 1

Hemidactylus sp.), 7 agamid lizards (6.3%;

including 1 Calotes jerdoni,1C.

mystaceus, 1 Ptyctolaemus gularis), 26

frogs (23.2%; including 1 Oreophryne

jeffersoniana, Microhylidae; 1 Rana

limnocharis, 1 R. microdisca, Ranidae),

and 7 colubrid snakes (6.3%; including 3

cf. Calamaria sp.). The frogs, Calamaria,

O. gracilis, and lygosomine skinks are

mainly terrestrial, whereas the geckos,

agamids, and 7. sauteri are at least partly

arboreal (e.g., Schmidt 1927, Pope 1929,

Smith 1935, Inger 1954, Auffenberg 1980,

Lue et al. 1987). If the anguid and snakes

are combined with skinks, elongate

vertebrates with shiny scales constitute ca.

65% of the prey.

Most mock vipers had empty stomachs

or contained a single item; stomachs with

multiple items contained three skinks (once)

and a skink and a colubrid snake (twice).

There is no evidence of ontogenetic

variation in diet: eight juveniles (SVLs

140-198 mm, cf. Wall 1910a) had eaten six

skinks, one agamid (Calotes mystaceus),

and one frog. Given small sample sizes,

the data provide only a hint of significant

geographic dietary variation. The

incidences of lizards, frogs. and snakes,

respectively, are as follows: 52, 15, and 5

versus 26, 11, and 2 for the grouped low

versus high tooth count populations

(Rasmussen 1975, see above); 14, 2, and 4

versus 38, 13, and 1 for Java versus

grouped other low tooth count populations;

45, 9, and 4 versus 33, 17, and 3 for short-

tailed versus long-tailed populations

(Rasmussen 1975, see above); and 31, 7,

and 0 versus 47, 19, and 7 for grouped

mainland versus island populations. If

lizards and snakes are lumped (to provide

minimum expected values for chi square

analysis), the only significant difference in

predation on reptiles versus frogs is for

short-tailed vs. long-tailed groups (p<.05).

These findings and those of Rasmussen

(1975) suggest that snakes on the mainland

and Java might have shorter tails, have

fewer pre-diastemal small teeth, and

consume relatively more reptiles than frogs

compared to those from other islands. All

records of predation on snakes are from

islands.

MR values (0.06-0.26, x=0.13, N=12)

are not especially large relative to other

snakes (cf. Voris and Moffett 1981, Greene

1983, 1984, Seib 1984, 1985, Jayne et al.

1988). Mean MRs for seven prey lizards

(0.13), three prey snakes (0.14), and two

prey frogs (0.14) are so similar to the

overall mean that simple frequencies

accurately represent the importance of each

prey type in the diet (see Greene 1986,

Losos and Greene 1988).

Feeding Behavior

Individual and geographic variability

remain to be assessed, but the feeding

behavior of my captive mock viper was

consistent during at least 12 complete

sequences. Prey was always approached to

within ca. 5 cm and seized with a rapid

strike to the torso, neck, or snout.

Constricting coils were applied by winding

anterior loops without a twist, an action

pattern common to many other colubrids

(Willard 1977, Greene and Burghardt

1978; Fig. 3). Skinks were typically

pinned behind the enlarged, anterior

maxillary and dentary teeth during

constriction, and even large prey

(maximum MR=0.31) never escaped

despite vigorous struggles (Fig. 3).

Venom clearly affected small iguanid and

Vol. 2, No. 2, p. 26

Chinese Herpetological Research

April 1989

FIG. 3. Mock viper, Psammodynastes pulverulentus, constricting and envenomating a scincid lizard,

Eumeces tetragrammus. Note enlarged anterior maxillary and dentary teeth restraining the skink, and

posterior maxillary fang penetrating the prey's nostril.

scincid lizards, as they were limp and

immobile during ingestion (Shaw [cited in

Smith 1943] and Gabe and Saint Girons

[1971] noted rapid mortality of lizards and

a snake bitten by this species).

Prey were always manipulated in the

jaws without release, then usually

swallowed head-first by the captive mock

viper, as is typical of many snakes (Greene

1976, 1983, 1984, Ananjeva and Orlov

1982, Voris and Voris 1983, Seib 1984,

1985). Among 74 natural prey items for

which direction of ingestion was

determined, 46 lizards, 12 frogs, and seven

snakes were swallowed head-first; three

lizards and two frogs were swallowed bent-

double; and three lizards and one frog were

swallowed tail-first (p < .01, chi square

test, for head-first versus not head-first).

Times from seizing 4 and 5 g prey lizards

(Eumeces sp.) until completion of ingestion

were 24.7 and 29.8 minutes, respectively,

for the captive snake.

Information on Other Taxa

Species of Ahaetulla (Smith 1943, Lim

1956, Leviton 1968, Henderson and

Binder 1980, pers. obs.), Cyclocorus

(Taylor 1922, Leviton 1965a), Lepturophis

(Lim and Kamarudin 1975), Lycodon

(Smith 1943, Leviton 1965b, pers. obs.),

Lycophidion (Branch 1976, Savitzky 1981,

Broadley 1983), Macroprotodon (Ferrand

de Almeida and Ferrand de Almeida 1986),

Mehelya (Savitzky 1981, Broadley 1983),

and Psammophis (Smith 1943, Broadley

1983) feed frequently on lizards,

especially, at least in some cases, on

skinks. Relatively large skinks are trapped

behind and/or impaled on the enlarged

anterior teeth of Lycophidion (Broadley

1983, Savitzky pers. comm.), but I am not

aware of observations on feeding in the

other genera with respect to use of the

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 27

anterior teeth. Nothing is known about the

diet of Ditypophis vivax. Pythonodipsas

carinatus eats skinks and geckos (Marx et

al. 1982). A bolyeriid, Casarea

dussumieri, eats skinks and geckos in

nature, and its enlarged anterior teeth and

divided maxillae might function

analogously to those of Psammodynastes

pulverulentus during feeding (Cundall and

Irish 1989).

Skinks are occasionally taken by a

variety of reptiles, birds, and mammals

throughout the world (e.g., Taylor 1952,

Fitch 1954, Greene 1988a, Losos and

Greene 1988), but observations on captive

predators indicate that they might often

have trouble handling these lizards. Vitt et

al. (1977:329) noted that "Smooth scales

and rolling movements allowed some

skinks [Eumeces sp.] to escape by breaking

the grip...[of snakes, Lampropeltis sp. and

Diadophis punctatus, both of which lack

enlarged anterior teeth]". A rear-fanged

snake, D. punctatus, lost two of five small

skinks (Eumeces sp., MRs ca. 0.10) when

they rolled out of its grip (pers. obs.). In

captivity a skink escaped from the grasp of

a Casarea dussumieri, “owing to the glossy

nature of its scales" (Vinson 1975:64). A

pigmy owl, Glaucidium minutissimum,

readily ate Anolis lizards, but each of four

times "only a slight wiggle caused...[a

skink's] extremely smooth body to squirt

out of her talons" (Janzen and Pond

1976:73). Some gekkonid lizards might

pose similar problems for predators, due to

their fragile, easily torn skin (Greene

1988a, Bauer et al. 1988, Branch 1988:Pl.

84).

Discussion

Psammodynastes pulverulentus

resembles many other small, venomous,

constricting snakes in some aspects of

defensive behavior and feeding biology

(e.g., sigmoid threat posture and biting,

prey size, head-first ingestion), but is

highly unusual in terms of dental

morphology, diet, and perhaps resemblance

to sympatric pit vipers. Three concepts that

describe historical patterns among

phenotypic attributes, performance

advantage, and ecology are relevant here:

A feature is an adaptation for a particular

task if its evolutionary origin was

associated with a performance advantage at

that task. A feature is an exaptation if its

origin preceded an association with that

performance advantage. Convergence is

independently derived similarity, and might

reflect adaptive, exaptive, or non-functional

resemblance. Untangling these

relationships requires information on

phylogenetic relationships, natural history,

and performance capabilities (for further

discussion and references, see Gould and

Vrba 1982, Greene 1986, Schaefer and

Lauder 1986).

Three lines of evidence suggest that

markedly enlarged anterior maxillary and

dentary teeth in Psammodynastes

pulverulentus and certain other snakes are

functionally correlated with increased

effectiveness at capturing skinks:

(i) Insofar as known, most snakes with

that dentition emphasize skinks in the diet.

Examples include several genera of

colubrids and possibly two genera of

bolyeriids; the only clear exceptions are two

genera of arboreal boids, Chondropython

and Corallus, that prey mainly on mammals

(McDowell 1975, Groves and Greene,

unpubl.), and the former does eat skinks.

(ii) Direct observations of Lycophidion

sp. and Psammodynastes pulverulentus

(see above) suggest that the enlarged

anterior teeth function like pawls on a

ratchet, viz., by trapping slippery, writhing

prey in the mouth. A single observation of

Casarea dussumieri (Vinson 1975) is

contradictory.

(iii) Preliminary observations on a

captive mock viper and on other predators

lacking the specialized dentition indicate a

difference in handling ability consistent

with that hypothesis. Those observations

also support the hypothesis that skinks

escape from generalized predators by virtue

of their powerful trunk musculature and

shiny, impenetrable scales.

The evidence is consistent with an

Vol. 2, No. 2, p. 28 Chinese Herpetological Research April 1989

adaptive relationship between enlarged

anterior teeth and a diet of skinks in certain

colubrids, as is the order of their respective

origins in the Mesozoic (Estes 1983) and

Cenozoic (Cadle 1987). However, the

functional relationship might be exaptive,

as implied by the fact that some species

with enlarged anterior teeth might feed

frequently on frogs and/or non-scincid

lizards (e.g., some Ahaetulla, Cyclocorus,

Lepturophis, Psammodynastes, and

Psammophis, such that a diet of skinks

might not be primitive), and by the

possibility of a defensive role for the

enlarged teeth in Psammodynastes

pulverulentus. Lacking information on the

relationships of genera with enlarged

anterior teeth to each other and to taxa

lacking that attribute, we cannot distinguish

between homologous and convergent

similarity--adaptive or otherwise--in

morphology and ecology among the

colubrids. Surely the similar dentitions of

bolyeriids and skink-eating colubrids are

independently derived (Cundall and Irish

1989).

Thus far, most limbless or near-limbless

squamates that eat mainly skinks fall into

two groups. Pygopodid lizards, xenopeltid

snakes, and several genera of colubrids

have small hinged teeth that impinge on the

trailing edges of skinks' scales (Savitzky

1981, 1983; Patchell and Shine 1986a,

1986b, 1986c); in at least one colubrid,

Scaphiodontophis, ingestion is extremely

rapid (Henderson 1984). Bolyeriids,

Psammodynastes pulverulentus, and

several other genera of colubrids use

enlarged anterior teeth to restrain a

struggling skink's body, and ingestion is

not particularly rapid. African colubrids of

the genera Lycophidion and Mehelya

possess both tooth types on a single

maxillary bone (Savitzky 1981). Whether

those differences reflect chance,

morphological constraints among different

lineages, or ecological differences is an

interesting and unanswered question.

Venom aids Psammodynastes

pulverulentus and perhaps other snakes that

eat skinks in subduing potentially difficult

and even dangerous prey, but it also might

be involved in the antipredator response of

mock vipers. However, as yet neither of

those roles can be applied to questions

about the adaptive origin of rear fangs

(e.g., Kardong 1979) because we lack

phylogenetic information on the origin of

venom injection in the history of mock

vipers (cf. Cadle 1983).

Skinks are widespread, morphologically

and ecologically diverse, and often

abundant components of lizard faunas;

more than 1000 extant species have been

described worldwide (Greer, 1976).

Observations on the diets and behavior of

lizard predators (see above) suggest that the

impressive evolutionary success of this

family might reflect, in part, a decreased

vulnerability to unspecialized predators.

Although a variety of lizard-eating reptiles,

birds, and mammals occasionally eat skinks

(Greene 1988a), every species of snake and

pygopodid lizard known to prey frequently

on them in Africa and the Indo-Australian

region exhibits morphological

specializations for that diet (see above and

Savitky 1981, 1983, Patchell and Shine

1986a, 1986b, 1986c, Cundall and Irish

1989). Putative antipredator mechanisms

cannot entirely explain the success of

skinks, because shiny, hard scales

(underlain by osteoderms) and powerful

trunk musculature probably facilitate a

variety of limbless locomotor responses

(Gans 1975), and because each of those

traits is primitive for a higher taxon within

which other families have sustained only

modest radiations (Estes 1983, Estes et al.

1988).

Acknowledgments

I thank R. C. Drewes, M. S.

Hoogmoed, A. E. Leviton, H. Marx, C.

W. Myers, and G. R. Zug for permission

to examine specimens in their care; J. E.

Cadle, D. Cundall, and F. J. Irish for

helpful discussions; A. H. Savitzky for

critical review of the manuscript; and the

American Museum of Natural History

(Theodore Roosevelt Memorial Fund),

Field Museum of Natural History (Karl P.

Schmidt Fund), Fourth Bremen

Symposium on Biological Systems Theory,

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 29

and National Science Foundation (BNS 76-

19903, BSR 83-00346) for financial

support.

Literature Cited

ABRAMS, P. A. 1986. Adaptive responses of

predators to prey and prey to predators: the

failure of the arms-race analogy. Evolution

40:1229-1247.

ANANJEVA,N. B., AND N. L. ORLOV. 1982.

Feeding behaviour of snakes. Vertebrata

Hungarica 21:25-31.

AUFFENBERG, W. 1980. The herpetofauna of

Komodo, with notes on adjacent areas. Bull.

Florida St. Mus. 25:39-156.

BAUER, A. M., A. P. RUSSELL, AND R. E.

SHADWICK. 1988. Morphological and

mechanical aspects of integumentary autotomy in

the gecko Ailuronyx. Amer. Zool. 28:194A.

BOULENGER, G. A. 1893. Catalogue of snakes in

the British Museum (Natural History). Vol. 1

Brit. Mus. (Nat. Hist.), London.

BRANCH, W. R. 1976. The wolf snakes,

Lycophidion capense and Lycophidion

variegatum (Reptilia, Serpentes, Colubridae) in

South Africa. J. Herpetol. 10:1-11.

BRANCH, W.R. 1988. Field guide to the snakes

and other reptiles of southern Africa. Ralph

Curtis Books, Sanibel Island, Florida.

BROADLEY, D. R. 1983. FitzSimons' snakes of

southern Africa. Delta Books, Johannesburg.

BRONGERSMA, L. D. 1934. Contributions to

Indo-Australian herpetology. Zool. Meded.

Rijksmus. Nat. Hist. 17:161-251.

BROWN, W.C., AND A. C. ALCALA. 1986.

Comparison of the herpetofaunal species richness

of Negros and Cebu Islands, Philippines.

Silliman J, 33:74-86.

CADLE, J. E. 1983. Problems and approaches in

the interpretation of the evolutionary history of

venomous snakes. Mem. Inst. Butantan 46:255-

274.

CADLE, J. E. 1987. Geographic distribution:

problems in phylogeny and zoogeography. Pp.

77-105 in R. A. Seigel, J. T. Collins, and S. S.

Novak (eds.), Snakes: ecology and evolutionary

biology. Macmillan Publ. Co., New York.

CADLE, J. E. 1988. Phylogenetic relationships

among advanced snakes: a molecular perspective.

Univ. California Publ. Zool. 119:1-77.

CUNDALL, D. 1987. Functional morphology.

Pp. 106-140 in R. A. Seigel, S. S. Novak, and

J. T. Collins (eds.), Snakes: ecology and

evolutionary biology. Macmillan Publ. Co.,

New York.

CUNDALL, D., AND H. W. GREENE. 1982.

Evolution of the feeding apparatus in

Alethinophidian snakes. Amer. Zool. 22:924.

CUNDALL, D., AND F. J. IRISH. 1989. The

function of the intramaxillary joint in the Round

Island boa, Casarea dussumieri. J. Zool.,

London, in press.

DESSAUER, H. C., J. E. CADLE, AND R.

LAWSON. 1987. Patterns of snake evolution

suggested by their proteins. Fieldiana, Zool.

(new ser.) 34:1-34.

DOWLING, H. G., R. HIGHTON, G. C. MAHA,

AND L. R. MAXSON. 1983. Biochemical

evaluation of colubrid snake phylogeny. J.

Zool., London 201:309-329.

ESTES, R. 1983. The fossil record and early

distribution of lizards. Pp. 365-398 in A.

Rhodin and K. Miyata (eds.), Advances in

herpetology and evolutionary biology: essays in

honor of Ernest E. Williams. Mus. Comp.

Zool., Cambridge, Massachusetts.

ESTES, R., K. DE QUEIROZ, AND J. GAUTHIER.

1988. Phylogenetic relationships within

Squamata. Pp. 119-281 in R. Estes and G.

Pregill (eds.), Phylogenetic relationships of the

lizard families: essays commemorating Charles

L. Camp. Stanford Univ. Press, Stanford,

California.

FERRAND DE ALMEIDA, N. AND F. FERRAND

DE ALMEIDA. 1986. On the occurance and

feeding habits of the false smooth snake

Macroprotodon cucullatus (Geoffroy, 1827) in

Portugal (Serpentes: Colubridae). Amphibia-

Reptilia 7:75-81.

FITCH, H. S. 1954. Life history and ecology of

the five-lined skink, Eumeces fasciatus. Univ.

Kansas Publ. Mus. Nat. Hist. 8:1-156.

FRITH, C. B. 1977. A survey of the snakes of

Phuket island and the adjacent mainland of

Vol. 2, No. 2, p. 30 Chinese Herpetological Research April 1989

peninsular Thailand. Nat. His. Bull. Siam Soc.

26:263-316.

GABE, M., AND H. SAINT GIRONS. 1971.

Particularites histologiques des formations

glandulaires du vestibule buccal de Calamaria

pavimentata Dumeril et Bibron, 1854 et

Psammodynastes pulverulentus (Boie, 1827);

(Colubridae, Serpentes). Ann. Sci. Natur.,

Zool., Ser. 12, 13:149-162.

GANS, C. 1961. The feeding mechanism of snakes

and its possible evolution. Amer. Zool. 1:217-

227.

GANS, C. 1975. Tetrapod limblessness:

evolutionary and functional corollaries. Amer.

Zool. 15:455-467.

GOULD, S.J., AND E. VRBA. 1982. Exaptation--

a missing term in the science of form.

Paleobiology 8:4-15.

GRANDISON, A. G. C. 1972. The Gunong Benom

Expedition 1967. 5. Reptiles and amphibians of

Gunong Benom with a description of a new

species of Macrocalamus. Bull Brit. Mus. (Nat.

Hist.) 23:45-101.

GREENE, H. W. 1976. Scale overlap, a directional

sign stimulus for prey ingestion by

ophiophagous snakes. Z. Tierpscyhol. 41:113-

120.

GREENE, H. W. 1983. Dietary correlates of the

origin and radiation of snakes. Amer. Zool.

23:431-441.

GREENE, H. W. 1984. Feeding behavior and diet

of the eastern coral snake, Micrurus fulvius.

Spec. Publ. Mus. Nat. Hist. Univ. Kansas

10:147-162.

GREENE, H. W. 1986. Diet and arboreality in the

emerald monitor, Varanus prasinus, with

comments on the study of adaptation. Fieldiana,

Zool. (new ser.) 31:1-12.

GREENE, H. W. 1988a. Antipredator mechanisms

in reptiles. Pp. 1-152 in C. Gans and R. B.

Huey (eds.), Biology of the Reptilia, Vol. 16,

Ecology B, Defense and life history. Alan R.

Liss Inc., New York.

GREENE, H. W. 1988b. Species richness in

tropical predators. Pp. 259-280 in F. Almeda and

C. M. Pringle (eds.), Tropical rainforests:

diversity and conservation. California Acad. Sci.

and Amer. Assoc. Adv. Sci., San Francisco.

GREENE, H. W., AND G. M. BURGHARDT. 1978.

Behavior and phylogeny: constricting in ancient

and modern snakes. Science 200:74-77.

GREER, A. E. 1976. A most successful invasion,

the diversity of Australia's skinks. Austral. Nat.

Hist. 18:428-433.

HENDERSON, R. W. 1984. Scaphiodontophis

(Serpentes: Colubridae): natural history and test

of a mimicry-related hypothesis. Univ. Kansas

Mus. Nat. Hist. Spec. Publ. 10:185-194.

HENDERSON, R. W., AND M. H. BINDER. 1980.

The ecology and behavior of vine snakes

(Ahaetulla, Oxybelis, Thelotornis, Uromacer): a

review. Contr. Biol. Geol., Milwaukee Publ.

Mus. 37:1-38.

HOFFSTETTER, R. 1939. Contribution a l'etude

des Elapidae actuels et fossiles et de l'osteologie

des ophidiens. Archs. Mus. Hist. Nat. Lyon

15:1-78.

INGER, R. F. 1954. Systematics and

zoogeography of Philippine Amphibia.

Fieldiana: Zool. 33:181-531.

INGER, R. F., AND R. K. COLWELL. 1977.

Organization of contiguous communities of

amphibians and reptiles in Thailand. Ecol.

Monogr. 47:229-253.

JANZEN, D. H., AND C. M. POND. 1976. Food

and feeding behavior of a captive Costa Rican

least pigmy owl Glaucidium minutissimum

rarum Griscom (Aves: Strigidae). Brenesia 9:71-

80.

JAYNE, B. C., H. K. VORIS, AND K. B. HEANG.

1988. Diet, feeding behavior, growth, and

numbers of a population of Cerberus rynchops

(Serpentes: Homalopsinae) in Malaysia.

Fieldiana, Zool. (new ser.) 50:1-15.

KARDONG, K. V.

evolution of snake fangs.

443.

1979. "Protovipers” and the

Evolution 33:433-

KROON, C. 1975. A possible Muellerian mimetic

complex among snakes. Copeia 1975:425-428.

LEVITON, A. E. 1965a. Contributions to a review

of Philippine snakes, IX. The snakes of the

genus Cyclocorus. Philippine J. Sci. 94:519-

532.

LEVITON, A. E. 1965b. Contributions to a review

of Philippine snakes, VIII. The snakes of the

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 31

genus Lycodon H. Boie. Philippine J. Sci.

94:117-140.

LEVITON, A. E. 1968. Contributions to a review

of Philippine snakes, X. The snakes of the

genus Ahaetulla. Philippine J. Sci. 96:73-90.

LEVITON, A. E. 1983. Contributions to a review

of Philippine snakes, XIV. The snakes of the

genera Xenopeltis, Zaocys, Psammodynastes, and

Myersophis. Philippine J. Sci. 112:195-223.

LIM, B. L. 1956. The natural food of some

Malayan snakes. Malay. Nat. J. 10:139-143.

LIM, B. L. 1979. Poisonous snakes of peninsular

Malaya. Malay. Nat. Soc., Kuala Lumpur.

LIM, B. L. AND M. S. KAMARUDIN. 1975.

Notes on new locality records of some rare

snakes in peninsular Malaysia. Malay. Nat. J.

29: 23-27.

LOSOS, J. B.. AND H. W. GREENE. 1988.

Ecological and evolutionary implications of diet

in monitor lizards. Biol. J. Linn. Soc. 35: 379-

407.

LUE, K.-Y., S.-H. CHEN, Y.-S. CHEN, AND S.-L.

CHEN. 1987. Taiwan lizards. Taiwan Prov. Ed.

Ministry. (In Chinese)

MALNATE, E. V. 1960. Systematic division and

evolution of the colubrid snake genus Nairix,

with comments on the subfamily Natricinae.

Proc. Acad. Nat. Sci. Philadelphia 112:41-71.

MARX, H., AND G. B. RABB. 1972. Phyletic

analysis of fifty characters of advanced snakes.

Fieldiana: Zool. 63:1-321.

MARX, H., G. B. RABB, AND S. J. ARNOLD.

1982. Pythonodipsas and Spalerosophis,

colubrid snake genera convergent to the vipers.

Copeia 1982:553-561.

MCDOWELL, S. B. 1975. A catalogue of the

snakes of New Guinea and the Solomons, with

special reference to those in the Bernice P.

Bishop Museum. II. Anilioidea and Pythoninae.

J. Herpetol. 9:1-80.

MCDOWELL, S. B. 1986. The architecture of the

comer of the mouth of colubroid snakes. J.

Herpetol. 20:353-407.

MERTENS, R. 1930. Die Amphibien und

Reptilien der Inseln Bali, Lombok, Sumbawa

und Flores (Beitraege zur Fauna der Kleinen

Sunda-Inseln. I.). Abh. Senckenberg.

Naturforsch. Ges. 42:115-344.

MITTER, C., AND D. R. BROOKS. 1983.

Phylogenetic aspects of coevolution. Pp. 65-98

in D. J. Futuyma and M. Slatkin (eds.),

Coevolution. Sinauer Assoc., Sunderland,

Massachusetts.

PARKER, H. W. 1949. The snakes of Somaliland

and the Sokotra Islands. Zool. Verh. 6:1-115.

PATCHELL, F. C., AND R. SHINE. 1986a. Food

habits and reproductive biology of the Australian

legless lizards (Pygopodidae). Copeia 1986:30-

39.

PATCHELL, F. C., AND R. SHINE. 1986b.

Hinged teeth for hard-bodied prey: a case of

convergent evolution between snakes and legless

lizards. J. Zool., London (A) 208:269-275.

PATCHELL, F.C., AND R. SHINE. 1986c.

Feeding mechanisms in pygopodid lizards: how

can Lialis swallow such large prey? J. Herpetol.

20: 59-64.

POPE, C. H. 1929. Notes on reptiles from Fukien

and other Chinese provinces. Bull. Amer. Mus.

Nat. Hist. 58:335-487.

POUGH, F. H. 1988. Mimicry and related

phenomena. Pp. 153-234 in C. Gans and R. B.

Huey (eds.), Biology of the Reptilia, Vol. 16,

Ecology B, Defense and life history. Alan R.

Liss Inc., New York.

RASMUSSEN, J. B. 1975. Geographical variation,

including an evolutionary trend, in

Psammodynastes pulverulentus (Boie, 1827)

(Boiginae, Homalopsidae, Serpentes). Vidensk.

Meddr. Dansk. Naturh. Foren. 138:39-64.

RASMUSSEN, J. B. 1985. A re-evaluation of the

systematics of the African rear-fanged snakes of

Bogert's groups XIII-XVI, including a discussion

of some evolutionary trends within Caenophidia.

Pp. 531-548 in K.-L. Schuchmann (ed.),

Proceedings of the International Symposium on

African Vertebrates: systematics, phylogeny and

evolutionary ecology. Zool. Forschungsinst.

Mus. Alexander Koenig, Bonn.

SAINT GIRONS, H. 1972. Les serpents du

Cambodge. Mem. Mus. Nat. Hist. Natur.,

nouv. ser. 74:1-170.

SAVITZKY, A. H. 1980. The role of venom

delivery strategies in snake evolution. Evolution

Vol. 2, No. 2, p. 32 Chinese Herpetological Research April 1989

34:1194-1204.

SAVITZKY, A. H. 1981. Hinged teeth in snakes:

an adaptation for swallowing hard-bodied prey.

Science 212:346-349.

SAVITZKY, A. H. 1983. Coadapted characer

complexes among snakes: _ fossoriality,

piscivory, and durophagy. Amer. Zool. 23:397-

409.

SCHAEFER, S. A., AND G. V. LAUDER. 1986.

Historical transformation of functional design:

evolutionary morphology of feeding mechanisms

in loricarioid catfishes. Syst. Zool. 35:489-508.

SCHMIDT, K. P. 1927. The reptiles of Hainan.

Bull. Amer. Mus. Nat. Hist. 54:395-465.

SEIB, R. L. 1984. Prey use in three syntopic

neotropical racers. J. Herpetol. 18:412-420.

SEIB, R. L. 1985. Euryphagy in a tropical snake,

Coniophanes fissidens. Biotropica 17:57-64.

SMEDLEY, N. 1931. Amphibians and reptiles

from the Cameron Highlands, Malay Peninsula.

Bull. Raffles Mus. 6:105-123.

SMITH, H. M. 1952. A revised arrangement of

maxillary fangs of snakes. Turtox News

30:214-218.

SMITH, M. A. 1915. The snakes of Bangkok. J.

Nat Hist. Siam. 1:173-178.

SMITH, M. A. 1935. The fauna of British India,

including Ceylon and Burma. Reptilia and

Amphibia. Vol. II. Sauria. Taylor and Francis,

London.

SMITH, M. A. 1943. The fauna of British India,

Ceylon, and Burma, including the whole of the

Indo-Chinese sub-region. Reptilia and Amphibia.

Vol. HI. Serpentes. Taylor and Francis,

London.

SODERBERG, P. 1967. Notes on a collection of

herpetological speciment recently donated to the

Centre for Thai National Reference Collections.

Nat. Hist. Bull. Siam. Soc. 22:151-166.

TAUB, A. M. 1967. Comparative histological

studies on Duvernoy's gland of colubrid snakes.

Bull. Amer. Mus. Nat. Hist. 138:1-50.

TAYLOR, E. H. 1922. The snakes of the

Philippine Islands. Bureau of Printing, Manila.

TAYLOR, E. H. 1952. Third contribution to the

herpetology of the Mexican state of San Luis

Potosi. Univ. Kansas Sci. Bull. 34:793-815.

TAYLOR, E. H., AND R. E. ELBEL. 1958.

Contributions to the herpetology of Thailand.

Univ. Kansas Sci. Bull. 38:1033-1189.

UNDERWOOD, G. 1967. A contribution to the

classification of snakes. Brit. Mus. (Nat. Hist.),

London.

VINSON, J. 1975. Notes on the reptiles of Round

Island. Mauritius Inst. Bull. 8:49-67.

VITT, L. J.. J. D. CONGDON, AND N. A.

DICKSON. 1977. Adaptive strategies and

energetics of tail autotomy in lizards. Ecology

58:326-337.

VORIS, H.K. AND M. W. MOFFETT. 1981. Size

and proportion relationships between the beaked

sea snake and its prey. Biotropica 13:15-19.

VORIS, H. K. AND H. H. VORIS. 1983. Feeding

strateegies in marine snakes: an analysis of

evolutionary, morphological, behavioral, and

ecological relationships. Amer. Zool 23: 411-

425.

WALL, F. 1908. Notes on a collection of snakes

from the Khasi Hills, Assam. J. Bombay Nat.

Hist. Soc. 18:312-338.

WALL, F. 1909. Notes on snakes from the

neighborhood of Darjeeling. J. Bombay Nat.

Hist. Soc. 19:337-357.

WALL, F. 1910a. A popular treatise on the

common Indian snakes. Part XIII. J. Bombay

Nat. Hist. Soc. 20:65-79.

WALL, F. 1910b. Further notes on snakes from

the Chin Hills. J. Bombay Nat. Hist. Soc.

20:770-775.

WALL, F. 1910c. Notes on snakes collected in

upper Assam. Part II. J. Bombay Nat. Hist.

Soc. 19:825-845.

WALL, F. 1926. Snakes collected in Burma in

1925. J. Bombay Nat. Hist. Soc. 31:558-566.

WEST, G. S. 1895. On the buccal glands and teeth

of certain poisonous snakes. Proc. Zool. Soc.

London 1895:812-826.

WILLARD, D. E. 1977. Constricting methods of

snakes. Copeia 1977:379-382.

The Karyotype of the Treefrog Rhacophorus reinwardtii (Boie) from

Xishuangbanna Daizu Autonomous Prefecture, Yunnan Province, China

ANMING TAN!:2, GUANFU WU!, AND ERMI ZHAO!

1Chengdu Institute of Biology, Academia Sinica, Chengdu, Sichuan,China

2(Present address) Museum of Vertebrate Zoology, University of California, Berkeley, California, USA

Abstract. -The Chinese Flying Treefrog, Rhacophorus reinwardtii, has a diploid chromosome number of

26, with 5 pairs of large and 8 pairs of small chromosomes. A secondary constriction occurs in the

paracentric region of the short arm of the first chromosome pair. Submetacentric chromosomes occur on

large pairs 2, 3, and 4. No other rhacophorid frogs are known to have these karyotypic features. The

significance of these findings for chromosomal evolution in rhacophorid frogs is considered.

Key words: Anura, Rhacophoridae, Rhacophorus, Cytotaxonomy, Yunnan, China.

Introduction

Chinese rhacophorid frogs are numerous

(about 40 species) and their taxonomic

position, especially their supraspecific

classification, is controversial (Liu and Hu

1961, Liem 1970, Jiang et al. 1987). In

recent years we have been attempting to use

cytotaxonomical methods and data to clarify

the situation.

Karyotypes have been reported for only

a few of the many species of Chinese

rhacophorid frogs. Gao et al. (1985)

reported the karyotype and its C-banding of

Rhacophorus dennysi. Yang and Wu

(1986) reported the karyotype of a new tree

frog, Rhacophorus gongshanensis. Tan et

al. 1987a, reported the karyotype, C-

banding and Ag-NORs of Rhacophorus

chenfui. Tan (1987) reported an

extraordinary condition in which different

diploid numbers (2n=26 and 2n=16) were

found in single individuals of the small

treefrog, Philautus doriae. This paper

reports the karyotype of Rhacophorus

reinwardti.

Methods

In a survey of the herpetofauna of

Xishuangbanna Daizu Autonomous

Prefecture, Yunnan Province, China,

during May and June, 1986, over 50

specimens of the Chinese Flying Treefrog,

Rhacophorus reinwardtii, were collected in

the vicinity of Mengleng, at Buyuan River,

19 km WNW of Mengxing (21° 55'N

101° 22' E). Fifteen of these specimens

were used to examine its karyotype. The

reproductive habits of this treefrog were

observed and described in another paper

(Tan et al. 1987b).

Mitotic chromosomes were prepared

from bone marrow tissues using a modified

direct chromosome-making method. An

injection of 0.2 ml of a 0.1% solution of

colchicine was injected intraperitoneally,

and after 10 hours the specimens were

sacrificed. The long bones were dissected

from the four limbs. The two ends of the

bones were removed and the bone marrow

cells were washed directly onto the slides

and fixed by using a vapor of ethanol,

acetic acid, and distilled water in a ratio of

1:2:3. After being fixed for 2 hours, the

slides were stained in a 2% Giemsa

solution (pH 6.8-7.0), and photographed

using a camera fixed on a Nikon

microscope. For detailed information, refer

to Tan et al.(1987a). Terminology was

adopted from Levan et al. (1964).

Results

More than 50 metaphase chromosome

spreads were observed and counted.

Rhacophorus reinwardtii has a diploid

chromosome number of 26. The karyotype

Vol. 2, No. 2, p. 34 Chinese Herpetological Research April 1989

TABLE. 1. Proportional data from 10 well spread metaphase chromosomes.

1 14.83 + 0.63

2 12.14 + 0.50

3 11.14 + 0.55

4 10.69 + 0.44

5 09.61 + 0.24

6 06.32 + 0.28

7 05.89 + 0.24

8 05.78 + 0.20

9 05.40 + 0.23

10 04.96 + 0.31

11 04.61 + 0.28

12 04.28 + 0.30

04.02 + 0.35

of Rhacophorus reinwardtii consists of 5

pairs of large (relative length larger than

9%) and 8 pairs of small (relative length

smaller than 7%) chromosomes (Table 1).

Discussion

Rhacophorus reinwardtii has a karyotype

of 2n=26 with 5 large and 8 small pairs of

chromosomes. This is often seen in the

Rhacophoridae and may be either a

primitive retention, or a synapomorphy.

Many ranid frogs which are thought to be

sister groups of the family Rhacophoridae

(Inger 1967, Duellman 1988) have a similar

karyotype.

As we have pointed out previously, the

arm ratios of the chromosomes are useful in

differentiating closely related species (Zhao

et al. 1987). Moreover, Morescalchi

(1973) proposed that there may be a

relationship between how primitive the

chromosomes are and the chromosomal

arm ratios: the more primitive species tend

to have more chromosomes with

submetacentric, subtelocentric and

telocentric centromeres. This hypothesis

has been corroborated in most of the

groups we have studied (Tan et al 1987a).

This rather specialized and advanced

Chromosome no. Relative Length Arm Ratio Centromeric Type

1.30 + 0.15 metacentric

1.89 + 0.22 submetacentric

1.83 + 0.22 submetacentric

1.73 + 0.19 submetacentric

1.26 + 0.07 metacentric

1.39 + 0.15 metacentric

1.40 + 0.17 metacentric

1.27 + 0.09 metacentric

1.44 + 0.21 metacentric

1.23 + 0.17 metacentric

1.19 + 0.13 metacentric

1.15 + 0.10 metacentric

1.2 metacentric

FIG. 1. Metaphase chromosomes of Rhacophorus

reinwaratii. Arrows indicate the secondary

constrictions.

treefrog (Jiang et al. 1987) with only 3

pairs of submetacentric chromosomes,

supports this hypothesis. Exceptions to

this generalization also have been observed

in some groups (refer to Zhao et al. 1987).

Attention should be paid to different

members of closely related groups when

April 1989

Chinese Herpetological Research

Vol. 2, No. 2, p. 35

3 BA BS MS RB wa aan xx

RX AR NB RK BA

FIG. 2. Karyotype of metaphase chromosomes from figure 1.

studying chromosomal evolution and

phylogeny.

The position of the secondary

constriction on the chromosomes is another

important criterion in cytotaxonomy. As

shown in figures 1 and 2, Rhacophorus

reinwardtii has a prominent secondary

constriction in the paracentric regions of the

short arms of the largest no. 1

chromosome. This is presumed to be the

Nuclear Organizing Regions (NORs). In

contrast, Rhacophorus’ chenfui

(=Polypedates chenfui) has a prominent

satellite on the terminal of the long arm of

no. 11 chromosome, which has been

shown to be the NORs.

To date several authors have reported on

the classification of the Rhacophoridae

which has a distribution limited to the Old

World. Liem's family revision (1970)

divided it into 14 genera in the Orient,

Madagascar and the African tropics. Frost

(1985) recoginized 2 subfamilies

(Philautinae and Rhacophorinae) and 10

genera. Jiang et al. (1987) studied 14

species of Chinese Rhacophoridae and

identified 5 genera (Buergeria, Philautus,

Polypedates, Rhacophorus and Chirixalus)

instead of the 2 previously recognized

genera (Rhacophorus and Philautus). The

karyotypes of Polypedates dennysi

(=Rhacophorus dennysi), P. leucomystax

(=Rhacophorus leucomystax), P. chenfui

(=Rhacophorus chenfui), Chirixalus doriae

(=Philautus doriae) and Rhacophorus

reinwardtii have been reported. Further

studies should be carried out in order to

give convincing evidence for the

taxonomical position and phylogeny of the

Chinese rhacophorid frogs.

Acknowledgments

The senior author would like to express

his sincere gratitude to T. J. Papenfuss and

J. R. Macey for their encouragement to

finish this paper, and to D. B.Wake for

reviewing and improving the manuscript.

Literature Cited

DUELLMAN, W. E. 1988. Evolutionary

relationship of the Amphibia. Pp. 13-34 in

Fritzsch, B. (ed.) The evolution of the

Amphibian auditory system. John Wiley &

Son Inc., New York.

FROST, D. R. 1985. Amphibian species of the

world. Allen Press and The Association of

Systematics Collections, Lawrence, Kansas.

832 pp.

GAO, J., B. GENG, AND X. CHEN. 1985.

Analyses on the karyotype and C-banding

pattern of Rhacophorus dennysi. Acta

Herpetologica Sinica 1985-4(1):1-4. (In

Chinese.)

INGER, R. F. 1967. The development of a

phylogeny of frogs. Evolution 21:369-384.

JIANG, S., S. HU, AND E. ZHAO. 1987. The

approach of the phylogenetic relationship and

the supraspecific classification of 14 Chinese

species of treefrogs (Rhacophoridae). Acta

Herpetologica Sinica 1987-6(1):27-42. (In

Chinese.)

LEVAN, A., K. FREGDA, AND A. A.

SANDBERG. 1964. Nomenclature for

centromeric position on chromosomes.

Hereditas, 52: 201-220.

Vol. 2, No. 2, p. 36 Chinese Herpetological Research April 1989

LIEM, S. S. 1970. The morphology, systematics,

and evolution of the Old World treefrogs

(Rhacophoridae and Hyperoliidae). Fieldiana

Zoology 57:1-145.

LIU, C., AND S. HU. 1961. Tailless amphibians

in China. Scientific Press, Peking. 364 pp. (In

Chinese.)

MORESCALCHI, A. 1973. Amphibia. Pp. 233-

348 in Chiarelli, A. B., and C. Capanna (eds).

Cytotaxonomy and vertebrate evolution.

Academic Press, New York.

TAN, A. 1987. A rare case of karyotype in Anura

-- A preliminary study on the karyotype of

Philautus doriae (Boulenger) with different

diploid numbers of 26 and 16. Chinese

Herpetological Research 1987: 12-16.

TAN, A., E. ZHAO, AND Z. WU. 1987a. Studies

on the karyotype, C-bands and Ag-NORs of

Rhacophorus chenfui. Acta Zoologica Sinica

33(2):105-109. (In Chinese.)

TAN A., X. ZHENG, AND Y. CHU. 1987b.

Preliminary observations of the reproductive

habits of Rhacophorus reinwardtii

(Rhacophoridae). Acta Herpetologica Sinica

1987-6(2):71. (In Chinese.)

YANG, W., AND G. WU. 1986. The study on the

karyotype of Rhacophorus gongshanensis. Acta

Herpetologica Sinica 1986-5(3):225-226. (In

Chinese.)

ZHAO, E., A. TAN, AND G. WU. 1987.

Karyotypes of Chinese species of Occidozyga

(family Ranidae), with discussion on the

taxonomic status of O.laevis martensi. Chinese

Herpetological Research. 1987:7-11.

April 1989 inese Herpetological Researc Vol. 2, No. 2, pp. 37-45

Cytotaxonomical Studies on Chinese Pelobatids V. The Karyotypes, C-

bands and Ag-NORs of Megophrys omeimontis and Oreolalax schmidti

XIAOMAO ZHENG! AND GUANFU WU!

1Chengdu Institute of Biology, Academia Sinica, Chengdu, Sichuan,China

Abstract. -Chromosome preparations were successfully stained for C-bands and Ag-NORs in two Chinese

pelobatids, Megophrys omeimontis and Oreolalax schmidti. It is thought that karyotypes of Chinese

pelobatids have two formulae: 5+6 and 6+7. Most of them have the secondary constriction on 6q always

associated with the C-band, and in these two species, associated with the NORs.

Key words: Anura, Pelobatidae, Megophrys, Oreolalax, Cytotaxonomy, Sichuan, China.

TABLE 1. The diploid chromosome numbers of Megophrys omeimontis and Oreolalax schmidti.

Sex

Megophrys omeimontis Female

Female

Oreolalax schmidti

Introduction

The karyotypes of pelobatids have been

reviewed by Morescalchi (1973). The

karyotypes of 13 species of pelobatids in

China have been investigated (four species

of the genus Vibrissaphora, Zhao et al.

1983; Scutiger boulengeri, Wu 1984;

Brachytarsophrys carinensis, Tan et al.

1987; Vibrissaphora ailaonica, Wu and

Zhao 1987; Megophrys lateralis and

Atympanophrys shapingensis, Wu 1987;

and the four species of the genus Oreolalax,

Wuetal. 1988). Until now the majority of

comparative chromosomal investigations on

pelobatids have been performed by

conventional staining methods. Two

instances of banding analyses on Chinese

pelobatids were on Atympanophrys

shapingensis (Wu 1987) and the four

species of Oreolalax (Wu et al. 1988). In

this paper both C-bands and Ag-NORs

No. of cells

2n=24 2n=25 2n=26 2n=27 2n=28

were used for analysis of the karyotypes.

The results from C-banding and Ag-NORs

banding in pelobatids will provide evidence

for discussions on the evolution in the

family Pelobatidae.

Methods

Two species of Pelobatidae, Megophrys

omeimontis and Oreolalax schmidti were

collected at Mt. Emei (29°32'N 103°21'E),

Sichuan Province, China in May 1988.

Both ends of the femur, tibiofibula,

humerus, ilium, and coccygeum were cut

off and marrow cells were washed out with

0.4M KCL for chromosome preparation by

a centrifugal air-drying method (Wu et al.

1981). The samples were stained with 2%

Giemsa (diluted by PBS, PH 6.8-7.0). C-

banding was prepared following Sumner

(1972). Ag-NORs banding was made by a

developed Ag-NORs banding making

Vol. 2, No. 2, p. 38 Chinese Herpetological Research April 1989

TABLE. 2. Proportional data of the karyotypes from Megophrys omeimontis and Oreolalax schmidti.

Megophrys omeimontis Megophrys omeimontis Megophrys omeimontis

Arm Ratio Centromere Position

OCMOAIAAMKHAP WN

OPAAIRAMNRWNDY

18.24 + 1.33

15.15 + 0.89

13.40 + 0.95

11.22 + 0.78

9.90 + 0.62

6.97 + 0.56

4.80 + 0.80

4.22 + 0.38

4.02 + 0.36

3.49 + 0.30

3.24 + 0.28

2.81 + 0.28

2.53 + 0.29

Oreolalax schmidti

17.52 + 0.70

13.96 + 0.86

12.56 + 0.78

11.84 + 0.64

10.35 + 0.39

6.24 + 0.59

5.11 + 0.47

4.77 + 0.37

4.32 +041

3.95 + 0.43

3.53 + 0.43

3.04 + 0.34

2.57 + 0.30

1.13 + 0.16

1.73 + 0.19

1.85 + 0.18

1.64 + 0.34

1.57 + 0.17

1.37 + 0.18

2.42 + 0.80

1.50 + 0.31

1.29 + 0.26

1.16 + 0.22

2.36 + 1.01

Oreolalax schmidti

1.36 + 0.10

1.63 + 0.12

1.71 + 0.10

1.93 + 0.20

1.51 + 0.10

1.53 + 0.30

1.50 + 0.32

1.54 +0.11

1.60 + 0.28

1.34 + 0.33

1.27 + 0.38

1.06 + 0.14

metacentric

metacentric, submetacentric

submetacentric

metacentric

metacentric

metacentric

submetacentric

metacentric

telocentric

metacentric

metacentric

submetacentric

telocentric

Oreolalax schmidti

metacentric

metacentric

submetacentric

metacentric

metacentric

metacentric

metacentric

metacentric

metacentric

metacentric

metacentric

metacentric

telocentric

technique (Tan et al.1986).

Results

Chromosome numbers for the two

species studied are shown in table 1. We

found that the diploid chromosome number

of Megophrys omeimontis was 26.

However, two of the three specimens of

Oreolalax schmidti were 28 and one was

26. Variation in the number of diploid

chromosomes was also found in another

species of pelobatid, Brachytarsophrys

carinensis (Tan et al. 1987). We assume

the standard diploid chromosome number

of O. schmidti is 26. There was a pair of

extra small chromosomes (Table 1).

The karyotypes in which the diploid

chromosome numbers are 26 were

measured for each species (Table 2). The

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 39

: ae

FIG. la. Spread of metaphase chromosomes from Oreolalax schmidti, a, b, and c showing c-bands and

Ag-NORs 2n = 26; d, e, and f showing c-bands and Ag-NORs 2n = 28.

Vol. 2, No. 2, p. 40

mews

ae

ae

Chinese Herpetological Research

April 1989

ag 20 a6 A Cb AG eG BH

» # :

th fia My ta te % ae ao

HA Go @ a La em ‘ .

FIG. 1b. Karyotype of metaphase chromosomes from Oreolalax schmidti, a, b, and c showing c-bands and

Ag-NORs 2n = 26; d, e, and f showing c-bands and Ag-NORs 2n = 28.

karyotype of M. omeimontis was 2n=26

with seven pairs of metacentric

chromosomes (nos. 1, 4, 5, 6, 8, 10, and

11); three pairs of submetacentric

chromosomes (nos. 3, 7, and 12); two

pairs of telocentric chromosomes (nos. 9

and 13); the centromere type of the no. 2

chromosome was between metacentric and

submetacentric.

The chromosomes of O. schmidti were

metacentric except for no. 2 which was

submetacentric and no. 13 which was

telocentric.

The secondary constriction can be found

in both species. It is in a pericentric

position on the long arm of chromosome

no. 6. This secondary constriction was not

conspicuous in a few cells from both

species. It is impossible to know whether

sex chromosomes are present because only

male M. omeimontis and female O.

schmidti were looked at.

The results of C-banding and Ag-NOR

banding are presented in table 3 and figures

1 and 2. In both O. schmidti and M.

omeimontis there was a strongly C-band-

positive associated with the Ag-NORs

located in the position of the no. 6

chromosome long arm. In addition, the

strongly C-band-positive was in a centric

position on each pair of the M. omeimontis

chromosomes. Neither the C-banding nor

the Ag-NORs banding technique

demonstrated heteromorphism between

homologous heterochromatic regions in the

two species.

On the chromosomes of O. schmidti,

whose diploid chromosome number is 28,

C-banding and Ag-NORs were successful,

but these techniques did not reveal

heterochromatic regions on the extra pair of

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 41

as

a6 7) ue oe 8 “ : — .

2 “- 6% 4“ vA uke eo Be ey ae 8 #3

b ] Ee # ¢ # 4s Pe ‘ ‘ * g 42 é

. _ y

& e te g g * ee 2 *

& ae ¥ as % ae ie Oe: ie

c : sf boo § ne OR: ae «CO

eS “ 4 3 4

FIG. 2. Spread and karyotypes of metaphase chromosomes from Megophrys omeimonitis, a, b, and c

showing c-bands and Ag-NORs 2n = 26.

Vol. 2, No. 2, p. 42 Chinese Herpetological Research

April 1989

TABLE. 3. The karyotypes of Chinese pelobatid species that have been investigated.

Megophrys lateralis

Megophrys omeimontis

Oreolalax omeimontis

Oreolalax pingti

Oreolalax rugosa

Oreolalax schmidti

Scutiger boulengeri

Vibrissaphora ailaonica

Vibrissaphora boringti

Vibrissaphora liui

chromosomes.

Discussion

1. The chromosomal polymorphism of O.

schmidtii.

From the results, it is surprising that the

diploid chromosomal number is 28 in two

specimens and 26 in the third individual

(Fig.1).

The former have an extra pair of smaller

chromosomes. Moreover, heterochromatin

was not shown on any segments of this

extra small pair of chromosomes by C-

banding and no NORs were discovered

(Fig.1). A similar situation was found in

Atympanophrys shapingensis

Brachytarsophrys carinensis

Vibrissaphora leishanensis

Vibrissaphora yaoshanensis

Brachytarsophrys carinensis which were

also obtained from the Heng Duan

Mountains (Tan et at. 1987). They

examined 12 specimens and found that the

diploid chromosome number was 26 in

eight individuals, 27 in three, and 28 in

one. In 2n=27, the single extra

chromosome was the same as the extra pair

of chromosomes in 2n=28. As in O.

schmidti no C-band-positive or NORs were

found. Since B-chromosomes are

consistently associated with

heterochromatin (Schmid 1978b), they

consider the extra 1-2 chromosomes not to

be B-chromosomes, but a chromosomal

polymorphism.

All except one species of Chinese

pelobatid is distributed in the Heng Duan

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 43

TABLE. 4. C-banding and Ag-NORs position of Chinese pelobatid species that have been investigated.

all 13 centric

1q, 2q, 7q (pericentric or proximal)

Megophrys lateralis

1 - 5q (terminal)

1p (interstitial)

all 13 centric

6q (interstitial)

Oreolalax pingii all 13 centric

1 - 5q (terminal)

6q (interstitial)

Megophrys omeimontis

Oreolalax rugosa all 13 centric

6q (interstitial)

Mountains region. In this unusual

geographic region, anura often have special

chromosomal karyotypes. In addition to

uncommon karyotypes reported in Ranidae

(Wu and Zhao 1984a,b) unusual

karyotypes were found in pelobatids where

one of four Atympanophrys shapingensis

examined was found to be triploid and the

karyotype of the single female Megophrys

lateralis studied was 2n+1 (Wu 1987).

2. Karyotypes.

Morescalchi (1973) concludes that two

main karyotype formulae are found in the

family Pelobatidae. The first consists of 6

pairs of large and 7 pairs of small

C-band hetero-morphism Position of Ag-NORs

Oreolalax schmidti 6q (interstitial) 6q (interstitial)

Oreolalax popei all 13 centric

all 13 terminal

6q (interstitial)*herozygesity

1q (telomeric)

6q (pericentric or proximal)

chromosomes. The second consists of 5

pairs of large and 8 pairs of small

chromosomes. Chinese pelobatids are

divided into two subfamilies: Oreolalaxinae

and Megophryinae. The majority of the

species studied in the Oreolalaxinae have

the first formula (6+7), and most of the

chromosomes are 2n=26 with metacentric,

submetacentric, or subtelocentric

chromosomes (NF=52)[see Table 3]. Only

O. schmidti reported on in this paper has

the second formula (5+8) and NF=50. In

the Megophryinae, the karyotype is variable

in the four species studied (Table 3).

Atympanophrys_ shapingensis has a

karyotype formula of 5+8 with NF=52, B.

carienensis has a formula of 5+8 with

NF=52, M. lateralis has 6+7 with NF=52,

Vol. 2, No. 2, p. 44 Chinese Herpetological Research April 1989

and M. omeimontis has 5+8 with NF=48.

It is widely believed that species with fewer

numbers of NF (possessing more pairs of

homologous with telocentric and small

chromosomes) are more primitive (Schmid,

1978a). For this reason, we speculate that

M. omeimontis and Brachytarsophrys

carinensis are more primitive in karyotype

evolution.

In the evolution of Amphibia, the rate of

gene mutation is more rapid than karyotype

mutation. The form of the gene mutation is

revealed by the secondary constriction

(SC). Compared with the more advanced

groups like ranids and hylids, the number

of SC in pelobatids is fewer, only one or

two. In 10 of 14 species listed in table 3,

the SC is found on the no. 6 chromosome

long arm. We consider the frequent

presence of the SC on the long arm of no. 6

chromosome an indication of genetic

stability in Pelobatidae. Furthermore the

SC which. is on no. 5p in M. lateralis, 9q

in Brachytarsophrys carinensis,1q in

Atympanophrys shapingensis, and 2p in

Scutiger boulengeri probably forms from

pericentric inversions of chromosomes, or

chromosome segment translocation. We

speculate these species are more developed

in genetic phylogeny.

3. C-bands and NORs.

Not only are the pelobatids classified as

primitive through morphological taxonomy,

but they are also ancestral in the character

of the constitutive heterochromatin which is

shown by C-bands (see Table 4). The C-

bands, which are found mainly at

centromeres and telomeres, are lower in

number in pelobatids than in the more

advanced ranids and hylids. In M.

omeimontis there is a centric C-band on

every pair of chromosomes in addition to

one inter-C-band on the 6q. However, O.

schmidti has only one inter-C-band on the

6q and no bands on the others, indicating

it's genetic stability.

It is speculated that karyotypes with a

single large NOR pair are more ancestral

than those in which the NORs are

distributed over several chromosomes (Hsu

1975). Some species of ranids have

several additional small NORs instead of a

large NOR (Schmid 1978b). Schmid

suggests that the single large NOR which

appears ancestral would have been

transferred to other chromosomes as

complete packages, rather than as multiple

smaller units distributed throughout the

genome by translocations and inversions

(Schmid 1978a).

M. lateralis (Wu 1987), M. omeimontis

and O. schmidti (in this paper) all have

only a large NOR. This suggests that

pelobatids are more primitive that ranids.

As illustrated in tables 3 and 4, most of the

14 species of Chinese pelobatids that have

been studied have the conspicuous SC on

the no. 6q, along with the strongly C-band-

positive. In addition, this is associated

with the NORs in M. omeimontis and O.

schmidti. We suggest that this position of

the constitutive heterochromatin must have

been primitive. It is a character of genetic

stability in the pelobatids. It is indicated

that pelobatids without this character have

genetic constriction altered during the

evolutionary prosses.

M. omeimontis and O. schmidti appear

to be primitive both by C-banding and Ag-

NORs banding. Although the

morphological characters are different, both

species possess extremely similar

karyotypic characters.

Literature Cited

HSU, T. C. 1975. A possible function of

constitutive heterochromatin: the bodyguard

hypothesis. Genetics 79:137-150.

MORESCALCHI, A. 1973. Amphibia. Pp. 233-

348 in Chiarelli, A. B., and C. Capanna (eds).

Cytotaxonomy and vertebrate evolution.

Academic Press, New York.

SCHMID, M. 1978a. Chromosome banding in

Amphibia. I. Constitutive heterochromatin and

nucleolus organizer regions in Bufo and Hyla.

Chromosoma (Berl.) 66:361-388.

SCHMID, M. 1978b. Chromosome banding

Amphibia. II. Constitutive heterochromatin and

nucleolus organizer regions in Ranidae,

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 45

Microhylidae and Rhacophoridae. Chromosoma

Gerl.) 68:131-148.

SUNMER, A. T. 1972. A simple technique for

demonstrating centromeric heterochromatin.

Exp. Cell Res. 75:304-306.

TAN, A., Z. WU, AND E. ZHAO. 1986. Studies

of the Karyotype, C-bands and Ag-NORs of

Rana limnocharis (Bioe). Acta Herpetologica

Sinica 1986-5 (3):176-180. (In Chinese.)

TAN, A., X. ZENG, G. WU, AND E. ZHAO.

1987. Cytotaxonomical studies on Chinese

pelobatids I. A preliminary study on the

karyotype of Brachytarsophrys carinensis and the

variation in their chromosome number. Acta

Herpetologica Sinica 1987-6(2):1-4. (In

Chinese.)

WU, G. 1981. 3. A preliminary observation on

karyotype of Vibrissaphora liui (Pope) studies

on genus Vibrissaphora (Amphibia:

Pelobatidae). Acta Herpetologica Sinica 1981-

5(22):139-142. (In Chinese.)

WU,G. 1984. Karyotypes of Scutiger boulengeri

(Pelobatidae) of Sichuan and Altirana parkeri

(Ranidae) of Xizang. Acta Herpetologica Sinica

1984-3(1):33-36. (In Chinese.)

WU, G. 1987. Cytotaxonomical studies on

Chinese pelobatids III. The analysis of the

karyotypes of Megophrys lateralis and

Atympanophrys shapingensis. Acta

Herpetologica Sinica 1987-6(3):45-48. (In

Chinese.)

WU,G., AND E. ZHAO. 1984a. A rare karyotype

of anurans, the karyotype of Rana phrynoides.

Acta Herpetologica Sinica 1984-3(1):29-32. (In

Chinese.)

WU,G., AND E. ZHAO. 1984b. Two rare

karyotypes of anurans, the karyotypes of

Staurois mantzorum and S. liangshanensis.

Acta Herpetologica Sinica 1984-3(4):5-10. (In

Chinese.)

WU, G., AND E. ZHAO.1987. Cytotaxonomical

studies on Chinese pelobatids II. The karyotype

‘of Vibrissaphora ailaonica, with a discussion on

the synonomy of V. liui and V. yaoshanensis.

Acta Herpetologica Sinica 1987-6(3):42-44. (In

Chinese.)

WU,G., A. TAN, AND E. ZHAO. 1988

Cytotaxonomical studies on Chinese pelobatids

IV. The karyotype and C-bands of four species

in the genus Oreoolalax. Acta Herpetologica

Sinica 1988-6(1):1-4. (In Chinese.)

ZHAO, E., G. WU, AND W. YANG. 1983.

Studies on genus Vibrissaphora (Amphibia:

Pelobatidae) 5. A comparitive study of the

karyotypes of the genus Vibrissaphora. Acta

Herpetologica Sinica 1983-2(1):15-20. (In

Chinese.)

Vol. 2, No. 2, pp. 46-54

April 1989

Karyotypic Analyses of Four Species in Four Genera of Colubrid Snakes

YUHUA YANG!, FUJI ZHANG2, AND ERMI ZHAO2

1Department of Bioengineering, Sichuan University, Chengdu, Sichuan, China

2Chengdu Institute of Biology, Academia Sinica, Chengdu, Sichuan, China

Abstract. -The karyotypes are presented for four species, belonging to different genera in the snake

family Colubridae. Pseudoxenodon macrops sinensis has 2n=38 with 8 pairs of macrochromosomes and 11

pairs of microchromosomes. The diploid number of Dinodon rufozonatum is 2n=46 with 8 pairs of

macrochromosomes and 15 pairs of microchromosomes. Elaphe mandarina has 2n=38 with 8 pairs of

macrochromosomes and 11 pairs of microchromosomes. Zaocys nigromarginatus has 2n=34 with 8 pairs

of macrochromosomes and 9 pairs of microchromosomes. The relationship between karyotype and

phylogeny is discussed.

Key words: Cytotaxonomy, Colubridae, Pseudoxenodon, Dinodon, Elaphe, Zaocys, China

TABLE. 1. The diploid chromosome number of the four colubrid snake species.

No. of Cells Observed

Dinodon rufozonatum

Elaphe mandarina

Pseudoxenodon macrops sinensis

Zaocys nigromarginatus

Introduction

The family Colubridae is a large and

phylogenetically complex group of

serpents, including 37 genera and 144

species in China. Various karyotypic data

on Colubridae have been reported. This

paper presents the karyotypes of four

species in the family: Pseudoxenodon

macrops sinensis Boulenger, Dinodon

rufozonatum (Cantor), Elaphe mandarina

(Cantor), and Zaocys nigromarginatus

(Blyth).

Methods

Pseudoxenodon macrops sinensis (2

males), Dinodon rufozonatum (1 female),

Elaphe mandarina (2 males) and Zaocys

nigromarginatus (4 males) were purchased

% of 2n Cells

in July, 1986 from Miyi County (26° 50' N

102° 03' E), Dukou Municipality, Sichuan

Province, China. The specimens were

sacrificed after intraperitoneal injection of

lmg/ml colchicine (Smg/gr body weight).

Partial spines and ribs were taken out, from

which the bone marrow was flushed with

1% sodium citrate and the cell suspension

was treated hypotonically (0.4% KCL) for

half an hour at room temperature, fixed in a

fresh fixative (ethanol:glacial acetic acid =

3:1) after centrifuging for 7 min. at 1000

rpm, and stored in the fixative over night at

4°C. Then the chromosome preparations

were made using an air-drying technique,

stained for 30-40 min in 10% Giemsa, and

observed under an oil objective for

counting and microphotographing the

chromosomes. The relative length and arm

April 1989 Chinese H erpetological Research Vol. 2, No. 2, p. 47

ratio of chromosomes were calculated

according to the method proposed by Singh

(1972). Classification of chromosomes

follows Levan et al. (1964).

Results

The chromosome data, including diploid

number, and morphology, relative length

and arm ratio for the macrochromosomes of

the four species are shown in Table 1 and

Table 2.

Pseudoxenodon macrops sinensis. The

diploid number is 38 with 8 pairs of

macrochromosomes and 11 pairs of

microchromosomes. There are 5 pairs of

metacentric chromosomes (nos. 1-5) and 3

pairs of submetacentric chromosomes

(Nos. 6-8) among the macrochromosomes.

Dinodon rufozonatum. The diploid

number is 46 with 8 pairs of

macrochromosomes and 15 pairs of

microchromosomes. Among the

macrochromosomes, pair 1 _ is

submetacentric, and the heteromorphic third

pair should probably be considered to be

sex elements. However this can't be

proven since only one female and no males

were examined. The Z chromosome is

metacentric and the W chromosome is

submetacentric, with the Z longer than the

W. The rest of the macrochromosomes are

acrocentric (Fig. 2).

Elaphe mandarina. The diploid number

is 38, and the karyotype is readily divided

into 8 pairs of macrochromosomes and 11

pairs of microchromosomes. For the

macrochromosomes, Pairs 1-4 are

metacentric, pairs 5 and 6 submetacentric

and pairs 7 and 8 acrocentric (Fig. 3).

Zaocys nigromarginatus. The diploid

number is 34 with 8 pairs of

macrochromosomes, in which nos. 1, 3-4,

and 6 are metacentric and the rest

submetacentric. There are 9 pairs of

microchromosomes (Fig. 4).

Discussion

The diploid chromosome number in the

family Colubridae ranges from 24 to 50.

The most commonly observed karyotype is

2n=36 with 8 pairs of macrochromosomes

and 10 pairs of microchromosomes. There

is a distinction between macro- and

microchromosomes (Gorman, 1973). In

general, the difference in diploid number

might be caused by a variation in number of

microchromosomes. However, there are

exceptions. For instance, Amphiesma

Stolata has 7 pairs of macrochromosomes

(Singh, 1972), Natrix erythrogaster has 16

pairs, and both N. harteri and N.

rhombifera have 17 pairs (Baker et al.

1972). Based on the Robertsonian

translocation about chromosome evolution,

the karyotypes with a large chromosome

number, more microchromosomes and

acrocentric chromosomes, are often

considered to be primitive ones

(Morescalchi et al. 1979). Some species

of Natrix (sensu lato), which have more

macrochromosomes and_ fewer

microchromosomes, might be more

advanced species.

A number of karyotypes of Elaphe have

been reported. A karyotypic study of 3

species of Chinese Elaphe was done by

Yang et al. (1986). Most species examined

have a karyotype of 2n=36., however, the

Chinese E. bimaculata has 2n=34, and the

North American E. subocularis has 2n=40

(Baker et al. 1972). This paper reports the

karyotype of E. mandarina_ which has

2n=38, including 8 pairs” of

macrochromosomes and 11 pairs of

microchromosomes. The karyotype differs

from other studied species of Elaphe, not

only in number of microchromosomes (22,

instead of 20), but also in the morphology

of the macrochromosomes (2 pairs of

acrocentric chromosomes, not 1 pair)

[Table 3]. As described above, there are

interspecific differences in Elaphe. One of

the present authors (Zhang 1988) divided

the genus into three groups based on the

morphological characters of the skeleton,

muscles, and hemipenes. E. carinata

differs from the other species of Elaphe in

these morphological characters but not in its

karyotype. On the contrary, there are

karyotypical differences between E.

mandarina and other species of Elaphe,

Vol. 2, No. 2, p. 48

Chinese Herpetological Research

April 1989

TABLE. 2. Macrochromosome data of the four colubrid snake species.

Dinodon rufozonayum __Dinodon rufozonayum _ Dinodon rufozonayum

SCIDANAHN—

30.30 + 1.41

17.94 + 1.11

16.66 + 1.19

13.95 + 0.73

15.26 + 1.11

11.06 + 1.00

10.10 + 0.51

8.54 + 0.70

6.81 + 0.61

25.36 + 1.42

20.11 + 0.91

15.41 + 1.05

11.50 + 0.60

7.88 + 0.71

7.14 + 0.68

6.92 + 0.54

5.65 + 0.69

22 OEEMESS

19.04 + 1.24

14.60 + 1.33

11.01 + 1.01

9.87 + 0.44

8.48 + 0.45

7.49 + 0.60

6.82 + 0.59

23.04 + 1.00

18.93 + 1.25

14.64 + 1.10

10.76 + 0.63

9.91 + 0.33

8.86 + 0.44

7.34 + 0.87

6.52 + 0.69

Arm Ratio

1.91 + 0.22

1.63 + 0.24

2.27 + 0.41

1.15 £0.12

1.62 + 0.20

1.20 + 0.20

1.32 + 0.16

2o3\) a= i105)

1.94 + 0.26

1.16 + 0.09

1.61 0.18

1.30 + 0.16

1.31 + 0.12

1.44 + 0.29

1.73 + 0.41

1.88 + 0.40

1.17+0.11

1.72 + 0.15

1.25 + 0.09

1.32 + 0.14

2.10 + 0.30

1.24 + 0.15

MoT) a8 OAD)

173 10:36

Centromere Position

submetacentric

telocentric

metacentric

submetacentric

telocentric

telocentric

telocentric

telocentric

telocentric

metacentric

metacentric

metacentric

metacentric

submetacentric

submetacentric

telocentric

telocentric

Pseudoxenodon macrops Pseudoxenodon macrops Pseudoxenodon macrops

metacentric

metacentric

metacentric

metacentric

metacentric

submetacentric

submetacentric

submetacentric

metacentric

submetacentric

metacentric

metacentric

submetacentric

metacentric

submetacentric

submetacentric

April 1989 Chinese Herpetological Research

BY XS wa ee as

Ge ae oe ee ee

7 8 8 10 11

&e *¢ ** Me & es t+

13 14 16 16 17 18

—______ ok

FIG. 1. Karyotype of Pseudoxenodon macrops, Female.

Vol. 2, No. 2, p. 49

o@

12

19

Vol. 2, No. 2, p. 50 Chinese Herpetological Research April 1989

9 i0 {1 12 13 14 15

ao & ay ow o &% we _ a»

16 17 18 19 20 21 22 23

FIG. 2. Karyotype of Dinodon rufozonayum, Female.

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 51

Ss

1 8

~~ ee oe oe ow

3 10 1 12 13

«~* o* ee te #o *”

14 15 16 W% 18 13

FIG. 3. Karyotype of Elaphe mandarina, Male.

Vol. 2, No. 2, p. 52 Chinese Herpetological Research April 1989

FIG. 4. Karyotype of Zaocys nigromarginatus, Female.

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 53

TABLE. 3. Karyotypic data of some species in the genus Elaphe, from China.

2n Macrochromosome Microchromosome Reference

Elaphe rufodorsata

Elaphe taeniura

Elaphe mandarina

16(7M, 7S, &2A)

16(7M, 7S, &2A)

16(7M, 7S, &2A)

16(7M, 7S, &2A)

16(7M, 7S, &2A)

16(7M, 7S, &2A)

16(8M, 4S, &4A)

SSS888a

Xie et al. 1983

Xie et al. 1983

Yang etal. 1986

Yang etal. 1986

Yang etal. 1986

Xie et al. 1983

here reported

while there are similarities in morphological

characters between it and E. rufodorsata as

well as E. dione.

We report the karyotype of Dinodon

rufozonatum, which is consistent with that

of the same species reported by Nakamura

(1935) in diploid number and the numbers

of macro- and microchromosomes (2n=46

with 8 pairs of macrochromosomes and 15

pairs of microchromosomes). However,

there are differences in the morphology of

the macrochromosomes. We found 6 pairs

of acrocentric, 1 pair of submetacentric,

and 1 pair of metacentric chromosomes.

Nakamura found 7 pairs of acrocentric and

1 pair of metacentric chromosomes. The

sex chromosomes in the majority of species

of colubrids are the fourth largest pair

(Yang et al. 1983; Zie et al. 1983; Baker et

al. 1972). A female specimen of D.

rufozonatum was studied by the present

authors and clear heteromorphism was

detected in the third largest pair, which

might indicate a sex chromosome. Within

Colubridae, Clelia occipitolutea has 2n=50,

including 6 pairs of acrocentric, and 1 pair

of metacentric chromosomes, Oxyrophus

petolaris has 2n=46, with 5 pairs

acrocentric, 2 pairs submetacentric, and 1

pair metacentric. Macropisthodon rudis has

2n=46 with 7 pairs acrocentric and one pair

metacentric. These karyotypes suggest

that the three species above and D.

rufozonatum may be primitive species in

the family Colubridae.

We found that the karyotype of Zaocys

nigromarginatus is 2n=34, comprised of 8

pairs of macrochromosomes and 9 pairs of

microchromosomes, while the karyotype of

the same species, as reported by Nakamura

(1935), had 2n=36 with 8 pairs of

macrochromosomes and 10 pairs of

microchromosones. More studies have to

be completed in order to determine if these

karyotypic differences show geographic

patterns. There is a different diploid

number between Z. dhumnades (2n=36,

Yang et al. 1986) and Z. nigromarginatus

(2n=34), but the morphology of the

macrochromosomes is similar except for

pairs 4 and 5. The fourth pair is

metacentric and the fifth pair is

submetacentric in Z. dhummnades,

whereas the fourth pair is submetacentric

and the fifth pair is metacentric in Z.

nigromarginatus.

The karyotype of Pseudoxenodon

macrops sinensis is 2n=38, with one more

pair of microchromosomes than the typical

karyotype of colubrid snakes (2n=36).

There are 5 pairs of metacentric and 3 pairs

of submetacentric macrochromosomes . It

does not appear to be a primitive species.

Literature Cited

BAKER, J. R., G. A. MENGDEN, AND J. J. BULL.

1972. Karyotypic studies of thirty-eight species

of North American snakes. Copeia (2):257-265.

GORMAN, G. C. 1973. The chromosomes of the

Reptilia, a cytotaxonomic interpretation. pp.

349-424 in Cytotaxonomy and Vertebrate

Evolution. (A.B. Chiarell and E. Capama Eds.).

Vol. 2, No. 2, p. 54 Chinese Herpetological Research April 1989

Academic Press, New York.

LEVAN, A., K. FREGDA, AND A. A. SANDBERG.

1964. Nomenclature for centromeric position on

chromosomes. Hereditas 52:201-220.

MORESCALCHI, A., G. ODIERNA, AND E. OLMO.

1979. Karyology of the primitive salamander

family Hynobiidae. Experientia 35:1434-1436.

NAKAMURA, K. 1935. Studies on reptilian

chromosomes. VI. Chromosomes of some

snakes. Mem. Coll. Sci. Kyoto Imp. Univ.

B10:361-402.

SINGH, L. 1972. Evolution of karyotypes in

snakes. Chromosoma 38:185-236.

XIE, X., Y. QU, Y. YANG, AND M. HUANG. 1983.

A preliminary observation on the karyotype of

three species of genus Elaphe. Acta

Herpetologica Sinica 2(3):33-36. (in Chinese.)

YANG, Y., M. HUANG, Y. QU, AND X. XIE.

1986. A comparative study on the karyotypes of

four species in Colubrinae. Acta Herpetologica

Sinica 5(1):30-33. (in Chinese.)

ZHANG, F. 1988. Preliminary studies on the

phylogeny of Elaphe, Zaocys, Ptyas, and

Entechinuss Acta Herpetologica Sinica 1988-

6(2):103-111. (in Chinese.)

hinese Her,

Karyotypic studies of Sphenomorphus indicus

etolog

ical Research Vol. 2, No. 2, pp. 55-59

(Scincidae) and Takydromus

septentrionalis (Lacertidae)

YUHUA YANG! ZHENGFA GAO2, AND ERMI ZHAO?

1Department of Bioengineering, Sichuan University, Chengdu, Sichuan, China

2County Middle School of Anxian, Sichuan , China

3Chengdu Institute of Biology, Academia Sinica, Chengdu, Sichuan , China

Abstract. -Sphenomorphus indicus (Gray) has a diploid number of 28 with 14 metacentric

macrochromosomes and 14 microchromosomes. The NF value is 42. Takydromus septentrionalis

(Guenther) has a karyotype with 36 acrocentric macrochromosomes and only 2 microchromosomes (2n=38).

The NF value is 38. These karyotypic data are compared with related species.

Key words: Sauria, Scincidae, Lacertidae, Sphenomorphus, Takydromus, Cytotaxonomy, Sichuan, China.

80

40

Frequency

20

23 24 25 26 27 28 29 30 31 32

Diploid Number

FIG. 1. Sphenomorphus indicus has a diploid

chromosome number of 28. This was determined

by counting the mitotic metaphase chromosomes

of 100 individual cells.

Introduction

The karyotypes of two species of

lizards, Sphenomorphus indicus (Gray) in

the family Scincidae and Takydromus

septentrionalis in the family Lacertidae are

reported in this paper.

Methods

Sphenomorphus indicus (1 male and 1

female) and Takydromus septentrionalis (1

male and 1 female) were collected in June,

1978 from An Xian County (31° 40' N

104° 26' E), Mianyang Prefecture, Sichuan

Province, China.

The specimens were _ injected

intraperitoneally with 0.01 ml colchicine

solution (3mg/ml) per gram of body

weight. Sixteen hours after the injection,

he specimens were killed. The bone

marrow Cells were flushed with 1% sodium

citrate, the testes were removed, and cell

suspensions were prepared. Then the

chromosome preparations were made using

an air-drying technique.

The diploid number (2n) was determined

by observing the preparations under an oil

objective. Ten selected cells were

photographed, enlarged, and measured for

calculating the chromosome parameters.

The chromosomes were classified with the

standard proposed by Levan et al. (1964).

The abbreviations for the morphology of

chromosomes are as follows: "M" is used

for a macrochromosome, "m" for a

microchromosome, "V" for a metacentric

chromosome and "I" for an acrocentric

chromosome. The "Nombre Fondamental"

(NF) is the total number of arms in the

karyotype. A metacentric (V) has two

arms, an acrocentric (I) has one arm, and a

microchromosome is also considered to

have one arm regardless of its morphology.

Results

Sphenomorphus indicus (Gray). The

diploid number is 28 consisting of

14V+14m, NF=42 (Table 1, Figs. 1 and

2). In addition to mitotic metaphase cells,

numerous diakineses were seen in the

males, having 7 macrobivalents and 7

Vol. 2, No. 2, p. 56 Chinese Herpetological Research April 1989

oe

] 2 3 4 5 6 7

te Me & a% te Sl

& 9 10 tt 12 13 14

rt :

16u *

FIG. 2. The chromosomes of Sphenomorphus indicus. There are seven pairs of metacentric

macrochromosomes and seven pairs of microchromosomes.

80

40

Frequency

20

Sos) SIS SG 37 9B SIO) 0)

—_ &@

Lia. Diploid Number

FIG 3. Numerous diakineses are present in the | FIG. 4. Takydromus septentrionalis has a diploid

testes cells of male Sphenomorphus indicus. chromosome number of 38. This was determined

by counting the mitotic metaphase chromosomes

of 100 individual cells.

April 1989 Chinese Herpetological Research

Vol. 2, No. 2 p. 57

TABLE 1.

Data on chromosome number, relative length, arm ratio, and centromere position for

Sphenomorphus indicus (Scincidae) and Takydromus septentrionalis (Lacertidae).

Relative Length

Chromosome no.

Sphenomorphus indicus

. 21.83 + 1.22

19.14 + 0.77

16.19 + 0.44

15.32 + 0.65

11.62 + 1.34

08.98 + 0.85

06.83 + 1.01

Sphenomorphus indicus

Arm Ratio Centromere Position

Sphenomorphus indicus

Takydromus septentrionalis Takydromus septentrionalis Takydromus septentrionalis

9.43 + 0.80

8.69 + 0.58

8.15 + 0.38

7.56 + 0.46

6.71 + 0.32

6.43 + 0.29

6.23 + 0.26

5.76 + 0.37

5.55 + 0.34

5.09 + 0.31

4.78 + 0.36

4.46 + 0.37

4.12 + 0.21

3.97 + 0.21

3.65 + 0.24

3.39 + 0.21

3.28 + 0.16

2.80 = 0.29

1

2

3

4

5

6

7

8

9

microbivalents (n=14) [Fig. 3].

Takydromus septentrionalis (Guenther).

The diploid number is 38 with 361 and 2m,

NF=38 (Table 1, Figs. 4 and 5). The

diakineses were observed in males,

containing 18 macrobivalents and only one

microbivalent (Fig. 6).

Discussion

Sphenomorphus indicus and

Takydromus septentrionalis belong to two

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

Telocentric

different families. and there are distinct

differences between their karyotypes. The

former has 14V+14m with the ratio of M:m

being 14:14, while the latter has 361+2m

with the ratio of M:m being 36:2. It is

obvious that their karyotypic difference is

consistent with the distance in their

phylogenies.

King (1981) suggested that some groups

of lizards are chromosomally conservative,

whereas others are extraordinarily diverse.

Vol. 2, No. 2, p. 58

13 i4 15

Chinese Herpetological Research

April 1989

16 il 42

ae ae «*

16 Wy 18 19

FIG. 5. The chromosomes of Takydromus septentrionalis. There are eighteen pairs of acrocentric

macrochromosomes and one pair of microchromosomes.

The karyotype data from 7 genera, 19

species of Scincidae (Gorman, 1973) show

that the diploid numbers vary, including

24, 26, 28, 30, and 32 with the ratio of

M:m variable ranging from 8:20 to 12:12.

Additionally, the karyotype is not readily

divisible into macro- and

microchromosomes for some species. The

variation is found not only at the species or

genus level, but also in different local

April 1989

FIG. 6. Only one microbivalent chromosome

(shown by the arrow) is present in the testes of

male Takydromus septentrionalis.

populations of the same species. For

example, Makino and Momma (1949)

reported that the karyotype of

Sphenomorphus indicus is 2n=28 with

10V+18m and the ratio of M:m being

10:18. The karyotype of the same species

reported by the present authors has a

diploid number of 28 with 14V+14m and

the ratio of M:m is 14:14. This appears to

be an unreported karyotype pattern in the

Scincidae. On the other hand, the

chromosomal data of 7 genera, 32 species

or Lacertidae (Gorman 1973) reveal that the

family's karyotypes are conservative,

having a diploid number of 38 with 36I and

2m, except for a few species. The

karyotype of Takydromus septentrionalis is

2n=38 comprised of 36I and 2m, which

agrees with that of the T. septentrionalis

reported on by Nakamura (1935). It

appears that there is no geographical

variation between the two populations of T.

septentrionalis.

Lizards vary markedly in their

karyotypic evolution (Gorman 1973, King

1981) and consisted of 331 genera and

3105 species as of Underwood (1957). It

will reveal valuable insights into the

evolution of lizards to study their

karyotypes deeply and widely.

Chinese Herpetological Research

Vol. 2, No. 2 p. 5

Literature Cited

GORMAN, G. C. 1973. The chromosomes of the

Reptilia, a cytotaxonomic interpretation. Pp.

349-424 in Chiarell, A. B. and E. Caponna, eds.

Cytotaxonomy and Vertebrate Evolution.

Academic Press, New York.

KING, M. 1981. Chormosome change and

speciation in lizards. Pp. 262-285 in Atchley,

W.R. and D. S. Woodruff, eds. Evolution and

Speciation. Cambridge University Press.

LEVAN, A., K. FREGDA, AND A. A. SANDBERG.

1964. Nomenclature for centromeric position

on chromosomes. Hereditas 52:201-220.

MAKINO, S., AND E. MOMMA. 1949. An

idiogram study of the chromosomes in some

species of reptiles. Cytologia 15: 96-108.

NAKAMURA, K. 1935. Studies on reptilian

chromosomes. VI. Chromosomes of some

snakes. Mem. Coll. Sci. Kyoto. Imp. Univ.

Ser. B 10:361-402.

UNDERWOOD, G. 1957. On lizards of the family

Pygopodidae. A contribution to the

morphology and phylogeny of the Squamata.

Journal of Morphology 100:207-268.

9

inese Herp

Vol. 2, No. 2, pp.

Mimicry of Scorpions by Juvenile Lizards, Teratoscincus roborowskii

(Gekkonidae)

KELLAR AUTUMN! AND BATUR HAN2

Dept. of Environmental Studies, University of California, Santa Cruz, California, 95064, USA

2Urumgi Institute of Biology, Academia Sinica, Urumqi, Xinjiang Uygur Autonomous Region, China

Abstract. -Teratoscincus roborowskii is a large nocturnal gecko that occurs only in the Turpan Depression

of northwestern China. The striking similarity of dorsal coloration, escape behavior, and body size of

juvenile T. roborowskii and a species of scorpion in the genus Mesobuthus (Bathidae) indicates probable

Batesian mimicry by these lizards.

Key Words: Batesian mimicry, lizards, Gekkonidae, Teratoscincus, scorpions, China.

Introduction

Batesian mimicry is a well-documented

phenomenon in both invertebrates and

vertebrates (See Pough 1988a, 1988b for

review of vertebrate mimicry). In the

simplest case, three organisms are included

in a Batesian mimetic relationship: 1) a

noxious or potentially dangerous MODEL;

2) an edible and relatively harmless MIMIC;

and 3) a potential predator of both model

and mimic, the DUPE (Pasteur 1982, Pough

1988a, 1988b). Rejection of the model as a

prey item confers an adaptive advantage to

the dupe, which manifests itself in a learned

or instinctive avoidance of the model. A

superficial similarity to the model should be

adaptive to other organisms preyed upon by

the dupe; hence a mimetic relationship

evolves (Bobisud and Potratz 1976, Turner

1988).

Mimicry in lizards is poorly documented

and largely anecdotal (Pough 1988a,

1988b, 1989 pers. comm., Parker and

Pianka 1974, Huey and Pianka 1977). Our

account is anecdotal as well, and we hope

that as more examples of lizard mimicry

surface, solid scientific studies will follow.

We present a case of probable Batesian

mimicry between juvenile Teratoscincus

roborowskii (Gekkonidae) and a scorpion

in the genus Mesobuthus (Bathidae).

Teratoscincus roborowskii occurs only in

the second lowest spot on Earth (-154 m.),

the Turpan Depression, Xinjiang Uygur

Autonomous Region, China (taxonomy

after Macey et al. in prep). Sympatric with

T. roborowskii is the scorpion,

Mesobuthus sp. It was the superficial

similarity between Mesobuthus sp. and

juvenile T. roborowskii, and indeed our

care in making a proper identification when

collecting the lizards, that first suggested

Methods

During the 1987 and 1988 cooperative

Chengdu Institute of Biology, University

of California, California Academy of

Sciences herpetological expeditions to the

deserts of western China, we observed and

collected Teratoscincus roborowskii in the

Turpan area. The study area was 4.4 km

west of the Main Mosque in Turpan

(42°58'N 89°10'E), on the Turpan-Jiaohe

road, Turpan Prefecture, Xinjiang Uygur

Autonomous Region, China. This area

consists of a series of rocky hills with

sparse vegetation -primarily Alhagi

sparsiflora (Leguminosae). This was the

same study area that was used by Autumn

and Wang (1988). The scorpion,

Mesobuthus sp., is common on the site and

can be easily collected by turning rocks.

The scorpion specimens were donated to

the California Academy of Sciences. The

lizard specimens were donated to the

Museum of Vertebrate Zoology and to the

California Academy of Sciences.

In order to compare T. roborowskii with

the other Teratoscincus species, we

April 1989 Chinese H erpetolo gical Research Vol. 2, No. 2, p. 61

examined T. przewalskii from Dunhuang,

Gansu Province, China, T. bedriagai from

the Siesitan Basin, Iran, and 7. microlepis

and T. scincus from the Helmud Basin,

Afghanistan. None of these species are

sympatric with T. roborowski. These

specimens were provided by the Museum

of Vertebrate Zoology, and by the

California Academy of Sciences.

Results

Teratoscincus roborowskii was

extremely common on the study site.

While holding a bright flashlight at

shoulder height and turning in a full circle,

we often observed over 40 eyeshines at a

single spot. The geckos remain motionless

on open ground, a short distance from their

burrows. They seem to be solitary, and

adults and juveniles were never observed

together.

The similarity between juvenile T.

roborowskii and Mesobuthus sp. is evident

in the dorsal coloration, size, defensive

posture, and escape behavior of the two

animals:

1. Dorsal coloration.

Juvenile T. roborowskii are banded

from head to tail (Fig. 1). These bands are

dark, thin, and are only present on the

dorsal surface. This banding is lost in the

adult. The number of bands on the body

ranges from 5 to 8, with a mode of 7. The

number of bands on a non-regenerate tail

ranges from 4 to 6, with a mode of 5.

Similarly, Mesobuthus sp. has 7 plates on

the body and 5 segments on the tail

(excluding the stinger, which is folded

back) [Fig. 2, Fig. 3]. When viewed under

moonlight, the patterns of the two animals

are extremely difficult to differentiate.

DSIZE:

The mean snout-vent length of juvenile

T. roborowskii is 42 mm (N = 43, SD =

4.1, range: 35-53 mm). The mean non-

regenerate tail length is 27 mm. (N = 39,

SD = 2.1, range: 23-37 mm). The mean

total length is 69 mm. (N = 39, SD = 5.65,

range: 58-90 mm.) The largest

Mesobuthus sp. we collected had a body

length of 30 mm and a tail length of 37

mm., for a total length of 67 mm.

3. Defensive posture.

When provoked (by humans), juvenile

T. roborowskii stiffen the body and raise

the tail in an arc. This is in contrast to the

adults, which adopt an arched posture and

wave the tail from side to side, producing

hissing noise from the specialized caudal

scales, as demonstrated by Werner (1967).

The defensive posture of Mesobuthus sp. is

typical of most scorpions. The legs and

pedipalps are extended and the tail is held in

an arc above the abdomen. This arc is

similar in form to that seen in the juvenile

T. roborowskiii

4. Escape behavior.

When approached, juvenile T.

roborowskii scuttle quickly away -usually

in a straight line- and stop a short distance

away. The defensive posture mentioned

above is usually maintained throughout

flight. In contrast, adult T. roborowskii

run for much longer distances, in a zig-zag

pattern, and the body is thrown from side

to side during flight. Both adults and

juveniles will hide under Alhagi sparsiflora

clumps or retreat into burrows if pursued.

Mesobuthus sp. also run in approximately

straight lines, for short distances.

The juvenile T. scincus we examined

had 5 dorsal bands on the body and 5 thin

dorsal bands on the tail. These specimens

were the most similar in dorsal coloration to

juvenile T. roborowskii and to Mesobuthus

sp. The other Teratoscincus species

specimens either had bands that were too

broad or too few in number to accurately

resemble scorpions. The juvenile T.

Vol. 2, No. 2, p. 62 Chinese Herpetological Research April 1989

FIG. 1. Juvenile Teratoscincus roborowskii assuming a defensive posture while being photographed at

night in the Turpan Depression, Xinjiang Uygur Autonomous Region, China, Sept. 10, 1988. This

individual has 13 dorsal bands.

FIG. 2, Mesobuthus sp. assuming a derensive posture while being phoweraphrdla at night in the Turpan

Depression, Xinjiang Uygur Autonomous Region, China, Sep 10, 1988. This species has 13 obvious

body and tail segments.

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 63

WB Body Bands

EE) Tail Bands

Ci a a a ae a A

SNYNNNNNNS

Frequency

Number of Bands

FIG. 3. Number of body and tail bands occurring

in Teratoscincus roborowski in the Turpan Basin.

The mode of the body bands is 7 and the mode of

the tail bands is 5. Similarly, the scorpion

Mesobuthus sp., has 7 body segments and 5 tail

segments.

przewalskii had 6-7 dorsal bands on the

body and 2-4 broad dorsal bands on the

tail. The juvenile T. bedriagai were

unevenly banded on the body, and had 2

broad bands on the tail. Juvenile T.

microlepis were mottled, with light bands

on the body, and 7 light, uneven bands on

the tail.

Discussion

Since tail-raising and arching is a

common defensive behavior in other

Teratoscincus (Mebs 1966), and other

gekkonids as well (Bustard 1967, Johnson

and Brodie 1974, Parker and Pianka 1974,

Marcellini 1977, Vitt, et al 1977, Greene

pers. comm., Huntley pers. comm), it may

be an ancestral trait shared by most

gekkonids, rather than a mimetic derivation

in juvenile Teratoscincus roborowskii.

Tail-raising and the possibly conspicuous

tail-banding may misdirect the predator's

attention to the readily autotomized tail

(Pough 1988a, 1988b, Johnson and Brodie

1974, Parker and Pianka 1974, Vitt et al.

1977, Dial and Fitzpatrick 1981, Greene

1988 and Arnold 1988 for reviews).

However, 1) the behavioral differences

of juvenile T. roborowskii, when compared

with adults, 2) their similarity in dorsal

coloration, size, defensive posture, and

escape behavior to Mesobuthus sp., and 3)

their relative availability to potential

predators strongly indicates a mimetic

relationship. Juvenile T. roborowskii

differ greatly from other Teratoscincus

juveniles in dorsal coloration. Only T.

roborowskii and T. scincus possess thin

dorsal bands that resemble scorpion

segments. Furthermore, the kinematics of

tail raising in juvenile T. roborowskii is

not the same as that in Coleonyx

variegatus, C. brevis (see Vitt, et al. 1977,

p.328, and Dial and Fitzpatrick 1981 p.311

for figures), adult T. roborowskii. and

adult T. scincus (Autumn, pers. obs.,

Mebs 1966 p. 17, 19 for figures). While

juvenile T. roborowskii maintain a

relatively rigid posture with the tail raised

and parallel to the body, Coleonyx and

adult Teratoscincus adopt an arched

posture and wave the tail. It is plausible

that the evolution of scorpion mimicry in

juvenile T. roborowskii was made possible

by the ancestral banding and postural

characters.

Candidates for the dupe include the little

owl, Athene noctua (de Schauensee 1984),

the red fox, Vulpes vulpes (Tate 1947), the

sand boa, Eryx tatarticus, and adult

Teratoscincus roborowskii. Macey and

Papenfuss (1986 pers. comm) reported that

several adult T. roborowskii had

regurgitated undigested juvenile T.

roborowskii while in a sack. Adults and

juveniles were kept together and it is likely

that predation took place in the sack.

Juvenile T. roborowskii may not able to

function as mimics throughout their activity

period. Unless the mimicry is tactile as

well, the similarity should only be present

while the lizards are out of their burrows

and are in sufficient light, as is present

during moonlit hours. However, the

selective advantage should still be

substantial if moonlight provides the

greatest mimetic protection during a period

when predation pressure by visual

predators is likely to be the highest.

Vol. 2, No. 2, p. 64 Chinese Herpetological Research April 1989

Acknowledgements

We thank E. Zhao for making our

research possible, R. Macey and T.

Papenfuss for encouragement and

companionship in the field, H. Greene and

H. Pough for valuable advice, S. Williams

for scorpion identification, and V.

Friedman, S. Autumn, and M. Fusari for

editorial assistance. Our research was

funded in part by the California Academy

of Sciences.

Literature Cited

ARNOLD, E. N. 1988. Caudal Autotomy as a

defense. Pp. 235-273 in C. Gans, and R. B.

Huey, eds. Biology of the reptilia, vol. 16,

ecology b. Allen R. Liss, Inc., New York. 659

Ppp.

AUTUMN, K. AND Y. Z. WANG. 1988.

Observations on the ecology of Phrynocephalus

axillaris and Eremias velox in the Turpan Basin,

Xinjiang Uygur Autonomous Region, China.

Chinese Herp. Res. 2(1):6-13.

BOBISUD, L. E. AND C. J. POTRATZ. 1976. One-

trial learning versus multi-trial learning for a

predator encountering a model-mimic system.

Am. Nat. 110:121-128.

BUSTARD, R. H. 1967. Defensive display behavior

of the Australian gecko, Nepherurus asper.

Herpetologica 23(2):126-129.

CUSHING, B. S. AND A. MATHERNE. 1980.

Stinger utilization and predation in the scorpion

Paruroctonus boreus. Great Basin Nat. June

1980: 193-195.

DIAL, B. E. AND L. C. FITZPATRICK. The

energetic costs of tail autotomy to reproduction

in the lizard Coleonyx brevis

(Sauria:Gekkonidae). Oecoligia (Berl) 51:310-

317.

GREEN, H. W. 1988. Antipredator mechanisms in

reptiles. Pp. 1-152. in C. Gans, and R. B.

Huey, eds. Biology of the reptilia, vol. 16,

ecology b. Allen R. Liss, Inc., New York. 659

pp.

HUEY, R. B. AND E. R. PIANKA. 1977. Natural

selection for juvenile lizards mimicking noxious

beetles. Science 195:201-203.

JOHNSON, J. A. AND E. D. BRODIE, JR. Defensive

behaviour of the western banded gecko, Coleonyx

variegatus.. Anim. Behav. 22:684-687.

MARCELLINI, D. 1977. Acoustic and visual

display behavior of Gekkonid lizards. Amer.

Zool. 17:251-260.

MEBS, D. 1966. Studien zum aposematischen

verhalten von Teratoscincus scincus.

Salamandra. 1(2):16-20.

PARKER, W. S. AND E. R. PIANKA. 1974.

Further ecological observations on the western

banded gecko, Coleonyx variegatus. Copeia

1974:528-531.

PASTEUR, G. 1982. A classificatory review of

mimicry systems. Annu. Rev. Ecol. Syst.

13:169-199.

POUGH, F. H. 1988a. Mimicry of vertebrates: are

the rules different? Pp. 67-102 in L. P. Brower,

ed. Mimicry and the evolutionary process.

University of Chicago Press, Chicago.

POUGH, F. H. 1988b. Mimicry and related

phenomena. Pp. 155-234 in C. Gans, and R. B.

Huey, eds. Biology of the reptilia, vol. 16,

ecology b. Allen R. Liss, Inc., New York. 659

pp.

DE SCHAUENSEE, R. M. 1984. The birds of

China. Smithsonian Institution Press,

Washington, D. C. 602 pp.

TATE, G. H. H. 1947. Mammals of eastern Asia.

Macmillan Co., New York. 363 pp.

TURNER, J. R. G. The evolution of mimicry: a

solution to the problem of punctuated

equilibrium. Pp. 42-66 in L. P. Brower, ed.

Mimicry and the evolutionary process.

University of Chicago Press, Chicago.

VITT, L. J.. J. D. CONGDON, AND N. A.

DICKSON. 1977. Adaptive strategies and

energetics of tail autotomy in lizards. Ecology

58(2):326-337.

WERNER, Y. L. 1967. Regeneration os

specialized scales in tails of Teratoscincus

(Reptilia: Gekkonidae). Seneck. Biol.

48(2):117-124.

Chinese Herp

Vol. 2, No. 2, pp. 65-68

New Locality Records for Chinese Non-Marine Chelonians

JAMES R. BUSKIRK!

1413] Terrace St., Oakland, California , USA 94611

Introduction

In the past decade field biology has

enjoyed a renaissance in China.

Distributional information concerning a

wide range of reptiles and amphibians in

the People's Republic of China has been

updated and recorded. Prior geographical

distribution data often predate the Second

World War, and the current generation of

Chinese herpetologists has published their

findings mostly in Chinese language

journals with limited overseas circulation.

Even in those cases where an English

summary is provided, the translated

information is usually less complete than

the original Chinese. Changes in the

English rendition of localities in China has

been an additional problem. What follows

are several new localities published since

1973 for 12 species of Chinese turtles

representing 4 families.

Classification is in accordance with the

latest revisions (see Iverson 1986, and

Iverson and McCord 1988). Where

controversy exists, the classification given

in the Chinese journal is presented

parenthetically following the current

approved classification. Province (in

capitals) and locality are given following

each species. The rendering of Chinese

localities and authors’ names is in

accordance with the current Pinyin

standard; therefore, hyphens have been

deleted from names and localities prior to

1983. The nature of the record (literature,

collection, purchase) is indicated along with

other parameters, if given. The

geographical significance of the new

records is discussed, and habitat

information, if given in the original paper,

is presented. In 2 cases (Testudo

horsfieldii and Manouria impressa )

specific locality information lacking in the

original texts was provide via personal

communication from Ermi Zhao The order

of species follows Iverson (1986).

Family Emydidae

Chinemys reevesii: SHAANXI:

Shangnan. Song and Fang, 1982. Iverson

(1986) shows 2 localities in central

Shaanxi, one of which is apparently Xi'an.

Shangnan is about 110 km SW Xi'an.

ANHUI: Changling Commune, Jinzhai Co.

and Puoliangtin Commune, Shu shuong

Co., Dabie Shan. Zou, 1983. Lovich et

al. (1985) list 28 Anhui specimens in 2

American museums. The Dabie Shan

localities are more than 100 km W of the

Anhui localities plotted by Iverson (1986),

and are the first for that province north of

the Chang Jiang (Yangtze River).

JIANGXI: Lu shan (literature record);

Nanchang (1 specimen collected by author);

Anfu (specimen documented but not

located). Zhong and Wu, 1981. Lovich et

al. (1985) refer to 13 specimens from

Jiansu (sic) in 4 American museums.

ZHEJIANG: coastal isles and part of

Zhoushan Qundao, Dinghai Co. Gu et al,

1982. Lovich et al. (1985) cite 13

specimens from Zhejiang in 2 American

museums. Additionally, they list 4

specimens (CAS) from Chusan Island

(Zhoushan Qundao), which they list as a

province.

Chinemys nigricans (Chinemys

kwangtungensis): GUANGXI: Nanning

and Liuzhou. Zong and Ma, 1985. Eleven

specimens in the Shanghai Museum of

Natural History. Carapace lengths (CL)

range from 123 to 257 mm. Two black and

white figures showing head, plastrons. The

Guangxi localities represent range

extensions of approximately 320 and 130

km W and WNW Wuzhou, heretofore the

only published Guangxi locality. Iverson

and McCord (1988) show C.

kwangtungensis to be an invalid taxon.

Vol. 2, No. 2, p. 66 Chinese Herpetological Research April 1989

Cistoclemmys flavomarginata (Cuora

flavomarginata): GUANGXI: Wuzhou.

Wen, 1983. Guangxi Medical College

81014. An adult of unspecified gender

whose CL is 165 mm. First provincial

record. Extends range approximately 180

km W Guangzhou, Guangdong (Iverson

1986).

Cistoclemmys galbinifrons (Cuora

hainanensis): HAINAN: Dali of Diaulo

Shan; Nanxi of Diaulo Shan, and Chien

Fung Ling. Hu et al., 1975. Nine

specimens (4 males, 5 females) collected

between 1957 and 1964, of which museum

numbers are given for only 2 specimens,

64 III 6110 (Sichuan Biol. Res. Inst., now

Chengdu Inst. Biol.), and FU 200.

Altitudes given for Dali (200m) and Nanxi

(82 m). First record for China. The brief

description of "Cuora hainanensis " and

comparison with C. flavomarginata in

Hu's paper was misleading to Western

readers, who concluded erroneously that it

was a subspecies of the latter. Chinese

herpetologists at the time were unfamiliar

with C. galbinifrons, with which C.

hainanensis is presently being synonymized

(Zhao, in press). GUANGXI: Xiaodong

village, Qin Prefecture. Liu and Zhang,

1987. Two female specimens, Chengdu

Inst. Biol. 8601 and Guangxi Aquatic

Products Inst. 8602, purchased in Nanning

farmer's market on 3/19/86 and 8/22/86,

respectively. Identified by Ermi. Zhao.

Several dimensions and mass of these

specimens given. Another 22 specimens,

whose fate is not specified, were purchased

in the farmer's market. All were said to

have been found in a ravine near Xiaodong

village. First provincial record and first

record for mainland China. Until 1975 (Hu

et. al) this species was known from only 3

localities in northern Vietnam (Petzold

1965), the nearest of which, Tam Dao, is

approximately 380 km WSW Xiaodong

village.

Cuora pani: SHAANXI: Xujiaba,

Pingli Co. Song, 1984. Holotype Shaanxi

Inst. Zool. 80170 (male), allotype SIZ

80171 (female). New species, and a NE

range extension for the genus by

approximately 900 km (Yunnan Fu,

Yunnan, for C. yunnanensis). Two figures

(showing carapace and plastron), several

dimensions and ratios are given in this

description which contrasts the new taxon

with C. yunnanensis without mention of

C. trifasciata to which pani bears a

stronger resemblance. Several specimens

of box turtles identified as C. pani have

entered the United States and Europe via

the pet trade, their geographical origin a

mystery to the purchaser. In his

redescription of C. pani, Ernst (1988)

used15 pet trade animals of unstated origin,

and relied upon an incomplete translation of

Song's paper. The plastral pattern of the

turtle in Ernst's figure 3 is remarkably

different from that of the holotype (Song

1984). It is possible that future research

will reveal that C. pani and other new taxa

of Chinese box turtles being described from

a very small sample are no more different

from C. trifasciata than are the races of

Terrapene carolina from one another.

Mauremys mutica (Clemmys mutica):

ZHEJIANG: coastal isles and part of

Zhoushan Qundao, Dinghai Co. Gu et al.,

1982. Zhoushan Qundao (Chusan Island)

is the type locality for this species.

GUANGXI: Tian Deng and Da Xin. Lin,

1985. Four unnumbered male specimens,

2 captured on 10/4/83 and 2 on 8/16/84.

Dimensions and mass recorded. First

provincial record, and first record between

Guangzhou, approximately 600 km to the E

(Iverson 1986) and Ha Tinh, Vietnam,

approximately 350 km WSW (Petzold

1965).

Ocadia sinensis: GUANGXI: commune

near Wuming. Lin, 1984. A dead male

turtle weighing 1300 g was found on

4/13/83 and deposited in the Guangxi

College Trad. Chin. Med. First provincial

record, extending range approximately 450

km W Guangzhou.

Sacalia bealei (Clemmys quadriocellata):

JIANGXI: Jiulanshan in Longnan Xian.

Zhong, 1981. Two juveniles, Jiangxi

Medical College 00183-4, collected in

September 1979. Dimensions given. First

provincial record, extending range

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 67

approximately 170 km NE Guangzhou

(Iverson 1986).

Family Platysternidae

Platysternon megacephalum: JIANGXI:

Qianshan (3 specimens collected by

author); Pingxih; Longnan; Anfu;

Pingxiang (specimens documented but not

located); Xuniao (literature record). Zhong

and Wu, 1981. First provincial records.

Less than 200 km from known localities in

neighboring Fujian, Guangdong, and

Hunan.

Family Testudinidae

Testudo horsfieldii: XINJIANG: Lower

Ili and Kax valleys near Yining,

approximately 550 km W Urumgi. Zhao,

1973. Dimensions are given for SBRI

(CIB) 625145-6, a male and female,

respectively. First record for China. This

tortoise may be present in other valleys

along China's western frontier as there are

several records for the adjacent portion of

Soviet Kazakhstan (Iverson 1986).

Manouria impressa (Testudo impressa,

Geochelone impressa): HAINAN: no

locality given. Zong and Ma, 1974. A

specimen obtained at the Wildlife Trading

Bureau in Haikou in 1964 is preserved in

the Shanghai Museum of Natural History.

First record for China. YUNNAN:

Xishuangbanna. Zhao, 1986. This record

is based on an unspecified number of the

shells of butchered specimens. First record

for mainland China. HUNAN: Shaoyang,

at junction of Shao and Yangtze Rivers.

Zhao, X., 1986. A photograph and

measurements accompany this largely

descriptive account of a tortoise found in

mid-January, 1986. First provincial

record. The remoteness of Shaoyang from

the known distribution of this species in

extreme southern China, and its having

been captured in midwinter argue strongly

that the tortoise was an introduction.

Family Trionychidae

Trionyx sinensis: LIAONING:

Zhuanghe; Ingkou; Sueizhueng (1 female

each); Linguan (2 males, 4 females). Zhao

and Huang, 1982. Measurements, dates of

capture, and contents of reproductive

organs are given for 3 female specimens,

CIB 806001-2 and 806016, but which

locality they came from is not indicated, nor

is what became of the remaining 6 turtles.

Iverson (1986) records the species from

Shenyang, Liaoning. This species was

recorded from the vicinity of Harbin and

the Amur River basin in Heilongjiang

(Manchuria) by Pavlov (1926).

SHAANXI: Luonan. Song and Fang,

1982. Foping Nature Conservation Area:

between 610-780 m. Yuan, 1985. The

species is well represented in southern and

central Shaanxi (Iverson 1986). ANHUI:

Puoliangtin Commune, Shushuong Co.

Zou, 1983. The lower Yangtze basin is

well represented by records for this taxon

(Iverson 1986). ZHEJIANG: coastal isles

and part of Zhoushan Qundao. Gu et al.,

1982. These records are apparently the

first for the offshore archipelago. There are

numerous records from the adjacent

mainland (Iverson 1986). JIANGXI:

Anfu; Pingxiang; Longnan (one specimen

at each locality collected by the author); Lu

shan (literature record). Zhong and Wu,

1981. The literature record, apparently

known to Iverson (1986) is hundreds of

km N of the other 3 localities.

Acknowlegments

I wish to thank Ermi Zhao for providing

most of the original papers and for

encouragement. The superb translation

skills of Eileen Tang, Kinji Hayashi, and

Paul Triolo were instrumental in

accomplishing this project. John Iverson

and Ted Papenfuss provided valuable

editorial assistance.

Literature Cited

ERNST, C. H. 1988. Redescriptions of two

Chinese Cuora (Reptilia: Testudines: Emydidae).

Proc. Biol. Soc. Wash 101(1):155-161.

GU, H., Y. JIN, AND J. GENG. 1982. A

herpetological survey of the isles belonging to

Dinghai County, Zhejiang. Acta Herpetologica

Sinica 1982-1(1):89-91. (In Chinese.)

Vol. 2, No. 2, p. 68 Chinese Herpetological Research April 1989

HU S., E. ZHAO, AND C. HUANG. 1975. Three

new species of reptiles from Hainan Island,

Guangdong Province. Acta Zoologica Sinica

21(4):379-384. (In Chinese.)

IVERSON, J. B. 1986. A checklist with

distribution maps of the turtles of the world.

Richmond, IN. 282 pp.

IVERSON, J. B., AND W. P. MCCORD. 1988 (in

press). The proper taxonomic allocations of

Emys nigricans Gray, Emys muticus Cantor, and

Geoclemys kwangtungensis Pope. Amphibia-

Reptilia.

LIN, L. 1984. A new record of Testudinata of

Guangxi--Ocadia sinensis (Gray). Acta

Herpetologica Sinica 1984-3(1):14. (In

Chinese.)

LIN, L. 1985. Mauremys mutica (Cantor)--a

record new to Guangxi. Acta Herpetologica

Sinica 1985-4(1):49. (In Chinese.)

LIU, Z., AND T. ZHANG. 1987. Cuora

hainanensis --a record new to Guangxi. Acta

Herpetologica Sinica 1987-6(2):77. (In

Chinese.)

LOVICH, J. E., C. H. ERNST, AND S. W. GOTTE.

1985. Geographic variation in the Asiatic turtle

Chinemys reevesii (Gray) and the status of

Geocemys grangeri Schmidt. Journal of

Herpetology 19(2):240-245.

PAVLOV, P. A. 1926. Zhivotnyi mir

Man'chzhurii po Kollektsiiam Muzeia

Obshchestva Izucheniia Man'chzhurskogo Kraia

(presmykaiushchiesia i zemnovodnye) (Wildlife

of the Manchurian Union in the collection of the

Public Museum for Studies of the Land of

Manchuria [reptiles and amphibians]. Harbin. 48

pp.

PETZOLD, H. G. 1965. Cuora galbinifrons und

andere siidostasiatische Schildkréten im Tierpark

Berlin. Aq. u. Terr. Z. 18(3/4)87-91, 119-121.

SONG, M. 1984. A new species of the turtle

genus Cuora (Testudoformes: Testudinidae).

Acta Zootax. Sinica 9(3):330-332. (In Chinese.)

SONG, M., AND R. FANG. 1982. On collections

of amphibians and reptiles from Shangnan-

Luonan area, Shaanxi. Acta Herpetologica

Sinica 1982-1(1):88-89. (In Chinese.)

WEN, Y. 1983. Several new records of reptiles

from Guangxi. Acta Herpetologica Sinica 1983-

2(3):72-74. (In Chinese.)

YUAN, H. 1985. Herpetological survey of Foping

Natural Conservation, Shaanxi. Acta

Herpetologica Sinica 1985-4(1):50-51. (In

Chinese.)

ZHAO, E. 1973. A new record of Chinese land

tortoise from Sinkiang--Testudo horsfieldii

Gray. Acta Zoologica Sinica 19(2):198. (In

Chinese.)

ZHAO, E. 1986. A revised catalogue of Chinese

tortoises. Acta Herpetologica Sinica 1986-

§(2):145-148. (In Chinese.)

ZHAO, E., AND K. HUANG. 1982. A survey of

amphibians and reptiles in Liaoning Province.

Acta Herpetologica Sinica 1982-1(1):1-23. (In

Chinese.)

ZHAO, X. 1986. Geochelone impressa (Guenther)

discovered from Shaoyang, Hunan. Acta

Herpetologica Sinica 1986-5(4):313. (In

Chinese.)

ZHONG, C. 1981. Two new records of reptiles

from Jiangxi Province. Acta Herpetologica

Sinica 1981-5(15):95-98. (In Chinese.)

ZHONG, C., AND G. WU. 1981. A preliminary

list and its geographical distribution of reptilia of

Jiangxi Province. Acta Herpetologica Sinica

1981-5(16):99-110. (In Chinese.)

ZONG, Y., AND J. MA. 1974. A new record of

Chinese land tortoise from Hainan--Testudo

impressa (Giinther). Acta Zoologica Sinica

20(3):108. (In Chinese.)

ZONG, Y., AND J. MA. 1985. Studies on the

genus Chinemys in China. Proc. Sino-Jap.

Herpetologica Symp. presented by Chinese

herpetologists. Special publ. Acta Herpetologica

Sinica 1985-5(2):145-148. (In Chinese.)

ZOU, S. 1983. A survey of amphibians and

reptiles in Huaibei and Dabie Shan. Acta

Herpetologica Sinica 1983-2(3):74-76. (In

Chinese.)

April 1989

Chinese Herpetological Research

Vol. 2, No. 2, pp. 69-71

A Major Research Achievement in Captive Reproduction of Chinese

Alligators

ZHENGDONG ZHANG!

1Anhui Research Center of Chinese Alligator Reproduction, Xuancheng, Anhui, China

FE a. \

FIG. 1. Hatchling Chinese Alligator leaving its shell.

The Chinese Alligator (Alligator

chinensis) is the only species of crocodilian

found in China. For various reasons,

alligators nearly went extinct in China. By

1979, their range had been reduced to five

counties in Xuancheng Prefecture, Anhui

Province and the total population was less

than 500.

The Anhui Research Center of Chinese

Alligator Reproduction was established in

1979 to study the preservation of this

species through captive reproduction. In

1981, with the help of Prof. Bihui Chen,

we had mastered the techniques of artificial

incubation and in 1982 we developed

methods for feeding young alligators.

By 1983, only four years after the center

opened, both the number of eggs hatching

and the survival rate of young alligators

reached about 95%. The nesting rate of

adult female alligators, which are housed is

designed dens, had increased from 14% to

67% with about 90% of the eggs laid being

fertile. The male/female sex ratio is 1:6.

We are now hatching about 800 alligators

each year, and the size of the captive

population is increasing rapidly.

Our final goal with the captive breeding

program was to complete the egg to egg

live cycle at the center. Due to time

limitations we were not able to accomplish

this goal for a long time. For the last few

years, we have paid much attention to the

feeding and care of the young alligators

hatched in 1982. Special diets were

provided and natural habitats were

incorporated into the design of the pen.

Finally on July 5, 1988, the first nest of 25

eggs was was produced by alligators that

had hatched in 1982. After an incubation

period of 54 days, a second generation of

Vol. 2, No. 2, p. 70 Chinese Herpetological Research April 1989

EN EA AE EI eg ee

aS

FIG. 2. A nest of Chinese Alligator eggs.

FIG.3. Adult and hatchling Chinese Alligators basking.

April 1989 Chinese Herpetological Research Vol. 2, No. 2, p. 71

FIG. 4. Four year old Chinese Alligator feeding on a Rana.

TABLE 1. Length, weight, and sex of 1982's alligators.

Weight (kg) Range (kg) Length (cm) Range (cm)

Male 10.62 + 2.01 8.5 - 13.5 139.75 + 6.38 132 - 147

Female 35 8.65 + 2.06 4.5 - 12.5 130.17 + 8.10 112 - 146

22 Chinese Alligators hatched.

On September 20, 1988 we

examined a sample of 39 alligators from the

1982 population in order to measure their

weight and size and to determine sex ratio

(Table 1). Chinese Alligators grow slowly

and there is considerable variation in weight

and size. As in the wild population, the

frequency of males is very low. Since only

one next was produced this year, it takes

at least six years for Chinese Alligators to

reach sexual maturity. Most will take seven

years and some maybe even longer. In

order to produce more second generation

alligators, we are going to reduce the

density in the rearing areas and remove

some smaller individuals.

cone

MACEY, J. ROBERT, THEODORE J. PAPENFUSS, AND ERMI ZHAO.

Ningxia Hui Autonomous Region as an Indication of a Baie:

Phrynocephalus axillaris and Eremias velox in the ean Dep

Autonomous). Region; China... 20... coves Cast cece cster nee

Psammodynastes pulverulentus (Colubridae), a Specialized Predator C

ARATHS colleen stwadiare daauaee seeeaeeeeaeeneuseesentecenesesaanseanastens

TAN, ANMING, GUANFU WU, AND ERMI ZHAO. The Karyotype of the T

Rhacophorus reinwardtii (Boie) from eemsuaaerscsr: Daizu Autonom mous |

Yunnan Province, China........ citiusandevekedceuecane eee ar naee

YANG, YUHUA, ZHENGFA GAO, AND ERMI ZHAO, Karyotypic studies of on :

Sphenomorphus ndicus (Scincidae) and Takydromus os

Teratoscincus roborowskii patieneone Oc poses na ventsssnieouibuinenGheees

BUSKIRK, JAMES R. New Locality Records for Chinese Non-Marine Chelonians..

ZHANG, ZHENGDONG. *A ae Research Achievement in Captive Reproduction ae

Chinese Alligators......... exnanenunagien osaaneosedoqedsaitelg magn ete Gna hale Mem OnIn iar Coat nna