Postilla

Published by the

PEABODY MUSEUM OF NATURAL HISTORY
YALE UNIVERSITY
New Haven, CT 06520-8118

Number 217
9 April 1999

Reptiles and Amphibians
from Nan Ao Island, China

James Lazell

Curatorial Affiliate for Recent Vertebrates
Peabody Museum of Natural History
Yale University

and

The Conservation Agency

6 Swinburne Street

Jamestown, RI 02835

6661 90 030

Jonathan Kolby
The Conservation Agency

Yong-Mu Lin (deceased), late of
Science and Technology Bureau of Nan Ao County
Guangdong 515920, China

Dong-Hong Zhuang

Department of Biology

Santou University

Santou, Guangdong 515063, China

Wenhua Lu
The Conservation Agency

Postilla

Published by the
PEABODY MUSEUM OF NATURAL HISTORY
YALE UNIVERSITY

Curatorial Editor-in-Chief
Jacques Gauthier

Executive Editor
Lawrence F. Gall

Publications Editor
Rosemary Volpe

Published at irregular but frequent intervals since 1950, Postilla presents short papers based on
original scientific research by Yale Peabody Museum staff and research associates, and their
colleagues, in the natural science disciplines represented by the collections of the Yale Peabody
Museum’s curatorial divisions.

All manuscript submissions are sent out for external peer review by one or more referees, as well as
being reviewed internally at the Peabody Museum by the Executive Editor in consultation with the
Curatorial Editor-in-Chief and one or more members of the appropriate curatorial division.
Interested authors should contact the Executive Editor at the address below before submitting
manuscripts.

Other titles published by the Yale Peabody Museum are the Bulletin of the Peabody Museum of Natural
History and the Yale University Publications in Anthropology. A complete list of titles, availability and
ordering information as well as submission guidelines for contributors can be obtained from the Yale
Peabody Museum website at http://www.peabody.yale.edu/publications/ or requested from the
Publications Office at the address below.

Address correspondence to:

Publications Office
Peabody Museum of Natural History
Yale University

170 Whitney Avenue Phone: (203) 432-3786
P.O. Box 208118 Fax: (203) 432-9816
New Haven, CT 06520-8118 USA E-mail: peabody.publications@yale.edu

Copyright © 1999 Peabody Museum of Natural History, Yale University. All rights reserved.
No part of this publication, except brief quotations for scholarly purposes, may be reproduced
without the written permission of the Director, Peabody Museum of Natural History.

ISBN 0-912532-48-3

Printed in the U.S.A.
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

Postilla

Number 217
9 April 1999

Reptiles and Amphibians
from Nan Ao Island, China

James Lazell
Jonathan Kolby
Yong-Mu Lin
Dong-Hong Zhuang
Wenhua Lu

(Received 2 July 1998; revised and accepted !0 November | 998)

Abstract

Specimens of 18 native reptile and
amphibian species were collected for the
Department of Biology, Santou University,
on Nan Ao Island, Guangdong Province,
China. Four of five lizard species appear
to be new records: Gekko subpalmatus
(major range extension), Ateuchosaurus
chinensis (we present life history data),
Eumeces chinensis and Sphenomorphus
indicus. All five snake species appear to be
new records: Typhlops (““Ramphotyphlops’’)
braminus, Elaphe radiata, Natrix
(“Xenochrophis”) piscator, Oligodon
formosanus and Trimeresurus cf. albolabris.
Five frogs and toads are species already
recorded, but Rhacophorus leucomystax
(Polypedates megacephalus) and Microhyla
ornata are newly recorded.A single Bufo
specimen may represent a new species.
Rana catesbeiana, introduced from
America, is now established in the wild.
The native Rana rugulosa was not found.
Geological history, ecological conditions
and habitat alterations are discussed.

Key words

Biogeography, Squamata, Gekko
subpalmatus, Ateuchosaurus chinensis,
Trimeresurus, Bufo, habitat alteration.

Introduction

Nan Ao is the largest island on the conti-
nental shelf of Asia between Taiwan and
Hong Kong. Variously spelled “Nanngau”’
or “Namoa,” and translated to mean
“south crescent beach” or “south bay,”
the island figured prominently in the his-
tory of western exploits in China
(Williams 1895). However, little has been
published on its natural history. In 1996,
the Department of Biology, Santou Uni-
versity, began an exchange program with
The Conservation Agency involving per-
sonnel, publications and specimens, many
of which were deposited at South China
Normal University, the University of
Hong Kong, the Yale Peabody Museum
(YPM) and the Museum of Comparative
Zoology (MCZ). Here we report on re-
cent reptile and amphibian collections
made on the island.

Nan Ao is circa 106 square kilometers,
with two rugged massifs on its eastern and
western portions connected by an alluvial
plain largely filled by the city of Houzhai.
The western massif rises to 587 meters at
Da Jian Shan, the high point of the island,
and the larger eastern massif rises to 576
meters. Nan Ao is on the Tropic of Cancer
at the eastern edge of Guangdong

4 Postilla 217

Reptiles and Amphibians
from Nan Ao Island, China

Figure |

A portion of South China showing
the location of Nan Ao (N), Hong
Kong (V), and an estimate of the
range of Gekko subpalmatus on the
mainland (shaded). Dots are locality
records from Pope (1935) in the
provinces of Sichuan (S), Guangdong
(G) and Fujian (F); he also notes
Zhejiang (Z) without precise locality.
Zhao and Adler (1993) add the
provinces of Guizhou (U), Guangxi
(K) and Jiangxi (J). We are unaware
of any record from Hunan (H).
Lower right bar represents 100 km.

Province, and was historically portioned
between Guangdong and Fujian, the neigh-
boring province to the east (Figs. | and 2).
The island is a remnant of the Zhe Min
Oldland and has been largely above sea
level since the Early Jurassic (Wang 1985),
unlike the adjacent mainland, which is
nearly flat and has been repeatedly sub-
merged (as recently as the Sangamon in-
terglacial, some 120,000 years ago).

Nan Ao’s climate is monsoonal, resem-
bling that of Hong Kong to the west
(Dudgeon and Corlett 1994). The average
temperature is 21.5°C. January is the
coolest month, at the height of the
northerly dry monsoon, with lows of
2.9°-14°C. August is the warmest month,
with highs of 27.5°—-36.5°C. Rainfall aver-
ages |,348 mm per year, with most falling
during the southerly wet monsoon from
June through September. It rains an aver-
age of 2,290 hours per year, and the is-
land is struck by an average of 19 cy-
clonic storms annually (YL, unpubl. data),
notably more than Hong Kong (Dudgeon
and Corlett 1994). The island was origi-
nally covered in evergreen monsoon for-
est, but only tiny remnants remain as
fungshui woods on the edges of coastal
towns, or as patches in the steeper inland

ravines. Even these remnants are selec-
tively cut for valuable wood products and
attain little more than shrub stage today.
There are some tree plantations, largely
of exotic Eucalyptus and American Pinus.
Prior to 1950 Nan Ao had extensive
mangrove swamps, but these have been
converted to commercial salt works or
filled for urban development.

Materials and Methods

We collected 267 specimens on 25 July
1996 and 13-23 July 1997 on Nan Ao and
Wu Yu islands (Fig. 2). Our collecting was
highly selective; we did not attempt to
amass large series. Ve attempted to col-
lect specimens that vouchered particular
habitat types, individual variation, sizes
and sexes. We did record in a bound field
journal (JL) the presence of species seen
but not collected. In 1996 we collected
only on the Houzhai plain for one
evening. In 1997 we spent four days can-
vassing the island for the most productive
collecting sites (see Discussion below).
Efforts to document the herpetofauna of
South China and the islands of the South
China Sea span two decades (Lazell and
Liao 1986, Lazell 1988a, and works cited

LAZELL

LAZELL

Reptiles and Amphibians
from Nan Ao Island, China

Postilla 217 5

therein and herein). Specimens were pre-
served in plastic trays and injected with a
solution of one part stock formalin
(37-41% formaldehyde in water, buffered)
in 15 parts 70% isopropanol. They were
maintained wet in the trays in 70% iso-
propanol (without formaldehyde) until
firm, then transferred for storage to 60%
isopropanol (contra Simmons in Zhao and
Adler 1993).All specimens were tagged in
the field and thus end up with two tags
affixed, the second reflecting museum ac-
quisition. Ventral scale counts on snakes
were made by the method of Dowling
(1951) so as to reflect number of trunk
vertebrae. We also examined comparative
material in the American Museum of Nat-
ural History (AMNRH).

Gekko subpalmatus (Guenther)

This species was quite unexpected on
Nan Ao (Fig. 2:2, 5 and 7), and especially
on Wu Yu (Fig. 2:8). Zhao and Adler
(1993) include Guangdong in the range
but we have seen no mainland specimens.
Pope (1935) provides details of distribu-
tion indicating an upland and inland range
(Fig. 1). In view of the apparent range dis-

junction, the paucity of available informa-
tion on this species and subtle differences
between populations, we provide a de-
tailed description. We compared our
specimens with AMNH 23586, 33018-21,
33025, 35157, 35160 and 39970, plus one
untagged, from “mountains near Yeng
Ping,’ and with MCZ 45960—| from
“Chungan Hsien,” all inland Fujian
Province (Fig. |). For other species, we
used the accounts of Zhou et al. (1989)
and Ota et al. (1995), and examined 14
MCZ specimens of G. chinensis (Gray)
from Hong Kong and mainland Guang-
dong. Data for all 21 specimens from Nan
Ao (YPM 9094-7, 9105, 9158 and
9160-2) and Wu Yu (YPM 9111-22) are
combined in the description below.

Ten males measured 48-63 (average
59) mm SVL and || females measured
46-79 (average 64) mm SVL. Head width
measured 10.1—13.0 (average 12.2) mm
for males and 9.7—16.0 (average 13.1) mm
for females. Eye diameter measured
2.7—3.8 (average 3.2) mm for both sexes.
In males, 7-10 (average 9) preanal pores
are present, as well as a large cloacal spur
on each side of the base of the tail.
Supranasal scales are always in contact or

Figure 2

Nan Ao and adjacent islets.
Contours are at 300 m and
500 m. Bar represents | km.
T, right, indicates the Tropic
of Cancer. Localities are:

|, Gui Chen; 2, Huanghua
Shan Reservoir; 3, Houzhai
city; 4, Bai Sha Wan; 5, Shen
Ao; 6, Wu Ping Zai; 7, Tai Zhu
Ao; 8, Wu Yu. Inset shows the
position of Nan Ao (arrow)
relative to the Chinese
mainland (C), Hainan Dao
(H), Taiwan (T) and Luzon,
Philippines (L).

6 Postilla 217

Reptiles and Amphibians
from Nan Ao Island, China

separated by a minuscule granule. There
are no significant differences in the num-
ber of middorsal scales in eye diameter
among populations, but there is a weak
modal difference between the inland Fujian
specimens and those from Nan Ao and
Wu Yu. In the latter there are 12-21 (aver-
age 16) scales in eye diameter, while those
from inland Fujian have 14-27 (average 19)
dorsal scales in eye diameter. In the Nan
Ao and Wu Yu specimens, especially those
from Shen Ao, the head is tapered and
somewhat pointed, rather than rounded as
in material from inland Fujian (Fig. 3). The
rostral is quadrangular and about twice as
wide as high. In YPM 9097, from Shen Ao,
there are scales present within the rostral
borders anterior to the supranasals (Fig.
3); this individual is unique in this respect
among all examined.

All mainland specimens we have seen
are patterned in shades of brown, de-
spite decades in alcohol. The first live
Nan Ao G. subpalmatus we observed
(YPM 9094-7) were gray dorsally, dirty
yellowish abdominally, and with a “lad-
der” pattern especially well developed on
the nape and shoulders (Fig. 4). On Wu
Yu, live G. subpalmatus (YPM 91 | 1-22)
were mossy green to olive, banded with
chocolate brown to tan.Abdominal yel-
low was reduced or absent, but throats
often had yellow tones. The dorsal band-
ing was regular, without the longitudinal
“ladder” components. These specimens
had a prominent cephalic “W” mark (Fig.
4), and could turn pale ashy gray and tan
when captured.

G. subpalmatus from other Nan Ao lo-
calities were intermediate. For example,
YPM 9158 from Huanghua Shan Reser-
voir approached Wu Yu specimens in pat-
tern, with the cephalic “Wy,” but with the
dorsal transverse bars less regular and
more broken. There were no longitudinal

nape or shoulder bars.When first found,
this individual was gray brown with con-
spicuous ash white dots, but after capture
it turned shades of warm brown and the
dots disappeared. At Tai Zhu Ao, YPM
9162 was colored in life in shades of lead
to slate gray like the Shen Ao specimens.
It lacked both the ladder pattern and
cephalic “WV.” Three more individuals
(YPM 9105, 9160—1) showed varying de-
grees of both pattern elements, but little
or no brown coloration.

Hemidactylus bowringi (Gray)

This is the abundant, edificarian gecko of
South China (Karsen et al. 1986), with nu-
merous specimens in institutional collec-
tions from many localities, including small
islands (Lazell 1988b, Lazell et al. 1997).
We examined females YPM 9090 (49 mm
SVL) and MCZ 181800 (50 mm SVL),
both from Houzhai, and YPM 9093 (47
mm SVL) from Shen Ao.We saw these
geckos at several other localities (Fig. 2:2
and 7). The Wu Yu series (YPM 9153-6)
includes males 47 and 49 mm SVL, a 46
mm SVL female with two large, shelled
eggs, and a 22 mm SVL juvenile (Fig. 2:8).

Ateuchosaurus chinensis Gray

This is an uncommon lizard over most of
its range (Karsen et al. 1986), but it
sometimes occurs in numbers on small
islands (Lazell 1994).All of our collected
specimens (YPM 9101—2, 9108-10 and
9152) were found beneath rocks and logs
at Huanghua Shan Reservoir (Fig. 2:2), but
one individual was observed basking on
14 July 1997. Several were turned out
from cover submerged in streams.A fe-
male, YPM 9108 (58 mm SVL), contained
remnants of a grasshopper (Orthoptera:
Acrididae).A male, YPM 9110 (61 mm

Reptiles and Amphibians
from Nan Ao Island, China

Postilla 217 7

Figure 3

Gekko subpalmatus. A, MCZ 45960 from Fujian. B, YPM 9097 from Nan Ao. C, right pes, YPM 9097.

Ilustration © 1999 by Jonathan Kolby.

SVL), contained orthopteran eggs.

All three males (YPM 9102, 9110 and
9152) have 30 scale rows at midbody.
One female (YPM 9109) has 30, and two
(YPM 9101 and 9108) have 28. Midven-
trals ranged 61—64 (average 62) from
just posterior to the mental to the edge
of the vent in both sexes. Subdigital
lamellae under the fourth (longest) toe
of the pes were |6—18 (average 17) in
both sexes. In life these A. chinensis were
rich chestnut to mahogany brown dor-
sally with nearly black lateral fields con-
taining bold, nearly white spots, with
vinaceous tones anterolaterally on the
head and neck. In males, the vinaceous
color was especially bright, the throats
bright orange and the bold light dots
pale lime green to cream yellow.

In addition to the six from Nan Ao,
we have available 14 specimens from
seven islands and the mainland of the
Hong Kong region (MCZ 112194,
172612—4, 172447-70, |76214-6,
176651, 177076 and 179514—5). Nan Ao
female YPM 9109, collected 20 July 1997,

contained three shelled eggs measuring
12-14 by 7.5-8 mm. These eggs were
arranged in the body cavity lengthwise,
occupying (with the oviduct between
them) more than 42 mm within a skink
75 mm SVL.The anterior egg thus ex-
tended far into the thoracic area. The vis-
cera were much reduced in size and the
abdominal muscle wall was noticeably
thickened compared to other specimens.
Both smaller females YPM 9101 (54 mm
SVL) and YPM 9108 (58 mm SVL) con-
tained ovarian eggs approaching | mm in
diameter when collected |7 and 20 July
1997, respectively.

The ontograph for A. chinensis (Fig. 5)
indicates year classes, implying seasonal
reproduction (Auffenberg and Auffenberg
1989). The pattern, however, does not fit
skinks from this region documented by
Lazell et al. (1997) or Lazell and Ota
(1998). The wide seasonal span shown by
Okada et al. (1992) might account for this
pattern, but we hypothesize that females
spend the wet, warm monsoonal season
yolking up eggs that hatch late in that

8 Postilla 217

season. If our seasonal hypothesis is cor-
rect, females would probably not lay eggs
until about 23 months of age.

Eumeces chinensis (Gray)

Our sole voucher (YPM 9100) is an adult
female 97 mm SVL collected in a wet
paddy in the early evening of 16 July 1997
(Fig. 2:3). In life the specimen was similar
to Hong Kong specimens, as in the pho-
tograph provided by Karsen et al. (1986):
muddy gray brown with yellowish brown
sides and rusty lateral blotches. However,
juveniles of E. chinensis may be quite dark,
nearly black, with bold light stripes and
bright blue tails (e.g, MCZ 176225-9,
Hong Kong region). We saw skinks that
appeared to be typical adult E. chinensis
(as YPM 9100 above) and that were dark,
striped and blue tailed at Shen Ao on 20
July 1997.We saw several other dark,
striped, blue-tailed skinks we believed
might be E. quadrilineatus (Blyth) at
Huanghua Shan Reservoir, on |7 July
1997, and twice near Tai Zhu Ao, on 16
and 17 July 1997.

Sphenomorphus indicus (Gray)

Our larger specimen, YPM 9103 (72 mm
SVL), was dark brown with a ragged
nearly black lateral zone and similarly
blackish blotches in the brown middorsal
field. The chin and throat were nearly
black and white. This bold pattern faded
posteriorly along the ventral surface to
coppery light yellow on the sides of the
abdomen and around the vent. The un-
derside of the tail was golden. There was
a prominent series of light brown dash
marks forming a dorsolateral stripe. Our
smaller specimen, YPM 9107 (63 mm
SVL), was similar but had what appeared
to be solid deposits, bright yellow in

Reptiles and Amphibians
from Nan Ao Island, China

color, at the elbows, ankles and in the dig-
its. S. indicus occurs on Hainan and Taiwan
(Zhao and Adler 1993) and on the main-
land in the Hong Kong region, but not on
the larger islands of Hong Kong and Lan-
tau (Karsen et al. 1986). It has been
recorded from small islands of the Matsu
Group (Ota and Lin 1997). Ours were
collected in shady woods at Huanghua
Shan Reservoir (Fig. 2:2).

Typhlops (‘‘Ramphotyphlops”’)
braminus (Daudin)

Our two vouchers of this partheno-
genetic species (Fig. 2:1 and 2) have the
typical 20 midbody scale rows; YPM 9091
is 122 mm SVL, tail 3 mm, with 315 ven-
trals in midline, chin to vent. YPM 9104 is
135 mm SVL, tail 3.5 mm, with 336 ven-
trals. This is an abundant species on is-
lands of tropical China (Karsen et al.
1986, Lazell 1988b, 1994).

We follow Hahn (1977) in using Ty-
phlops rather than “Ramphotyphlops,” and
note that Wallach (1993) describes a con-
tinuum of morphology. The genus “Ram-
photyphlops” was diagnosed solely on the
hemipenes, structures of males only, so
not present in T. braminus.

Elaphe radiata (Schlegel)

Our voucher (YPM 9159) is an adult male
circa |.7 m in life, prepared as a skin and
complete skeleton. There are the normal
21 scale rows at the nape, 19 at midbody
and |7 just anterior to the vent. There are
227 ventrals and 95 subcaudals (tail per-
fect). These counts are low compared to
those given by Pope (1935): 228-231 and
95-101, respectively, for males, or to ven-
tral counts (225—246) given by Hu et al.
(1980). However, Wu et al. (1985) provide
a table with several males having fewer

Reptiles and Amphibians
from Nan Ao Island, China

Figure 4

Postilla 217 9

4),
“e

aay ers

&

&

Gekko subpalmatus dorsal patterns.A, MCZ 45960 from Fujian. B, YPM 9120 from Wu Yu. C, YPM
9097 from Shen Ao, Nan Ao. Illustration © 1999 by Jonathan Kolby.

than 227 ventrals, including one with 221.
The subcaudal count is near their median,
86-101. In life YPM 9159 was somber,
dark gray brown posteriorly, striped with
dull gray tan anteriorly. There were slate
black blotches in the dorsolateral fields
trimmed with cream white to ash gray.
The venter was pink anteriorly, with gray
increasingly encroaching ventrolaterally,
and cloudy gray posteriorly. This highly ac-
tive, diurnal snake is known as the “cop-
perhead racer” in the Hong Kong region
and is common on the larger islands like

Hong Kong itself and Lantau (Karsen et al.

1986). We trapped ours at Tai Zhu Ao
using paddy frogs for bait (Fig. 2:7).

Natrix (‘‘Xenochrophis’’)
piscator (Schneider)

All of our specimens (YPM 9092, 9099 and
9157) are males (Fig. 2:1, 2 and 3). Two are
certainly young of the year:YPM 9099, 127
mm SVL (tail 51 mm), collected 16 July
1997 at Houzhai; and YPM 9157, 131 mm
SVL (tail 55 mm), collected 21 July 1997 at
Huanghua Shan Reservoir. However, nei-
ther of these small specimens shows a
yolk sac scar, so we do not refer to them
as hatchlings. Our Gui Chen specimen
(YPM 9092) was captured in the act of
swallowing a paddy eel, Fluta alba L., unfor-
tunately not measured or preserved.

10 Postilla 217

Reptiles and Amphibians
from Nan Ao Island, China

SVL:cms

All three have the normal dorsal scale
row counts: 19 nape, 19 midbody and |7
before the vent. Two have | 24 ventrals
(YPM 9092 and 9099); one has 125 ven-
trals. Subcaudal counts vary considerably:
YPM 9092 has 54, lower than the range of
67-88 given for males by Hu et al. (1980).
The tail is 107 mm, one third of SVL,
which is 324 mm; this is shorter than the
mode of circa 40% in most males. The tail
ends in a spine-like scale and does not ap-
pear damaged, but we opine it must have
been truncated. Subcaudals number 80 in
YPM 9099 and 73 in YPM 9157.

Zweifel (1960) noted that snake popu-
lations on the offshore islands of Pacific
Mexico had consistently higher ventral
counts (and therefore more trunk verte-
brae) than their mainland relatives. He
compared populations of 10 species, and
suggested that the constant shift was
“possibly not determined genetically.”
Zweifel (1960) did not speculate on what
environmental condition of those islands
might cause high ventral counts. We note
that ventral counts of our N. piscator, like
that of Elaphe radiata above, are near the
low extreme for males of their species,

Figure 5

Ontograph of Ateuchosaurus. Solid
dots, Nan Ao; the circled dot is
gravid female YPM 9019. Hong Kong
region: squares, Lantau Island; circles,
Peng Chau; triangles, Ma Wan Island;
X, Ngong Shuen Chau (Stonecutters
Island); asterisk, Lam Chau (now
destroyed); star, Ping Chau; H, Hong
Kong Island; S, Sai Kung Peninsula.

122—138 (Hu et al. 1980). We suggest that
cold ocean currents in the eastern Pacific
may lower the average incubation tem-
peratures on Zweifel’s Mexican islands,
while the warm waters of the South
China Sea might warm island tempera-
tures along the coasts of tropical China.

Oligodon formosanus (Guenther)

An adult male (YPM 9098) was captured
alive on 16 July 1997, and an adult female
was found dead on a road on 22 July
1997, both at Tai Zhu Ao (Fig. 2:7). The
male has |9 dorsal rows at nape and mid-
body, |5 before the vent; the female has
19 at nape and midbody, but |7 before
the vent. The male has |6I ventrals and
51 subcaudals; ranges for males from Hu
et al. (1980) are 159-167 and 46-55, re-
spectively. The female has |71 ventrals and
46 subcaudals; ranges for females from Hu
et al. (1980) are 156-173 and 39-5I, re-
spectively. Both snakes were patterned in
somber shades of gray brown. The male in
life was bright rose pink on the posterior
venter; this color faded to nearly white
anteriorly and under the tail. The female

Reptiles and Amphibians
from Nan Ao Island, China

had only a faint tint of posteroventral
pink, possibly as a result of postmortem
fading. Pink posterior ventral color is typi-
cal (Hu et al. 1980). The male had a yel-
lowish chin and throat. O. formosanus is a
regular inhabitant of tropical China’s is-
lands such as Hong Kong and Lantau
(Karsen et al. 1986), and even some no
larger than Ping Chau, circa 0.7 square
kilometers (Lazell 1994).

Trimeresurus cf. albolabris (Gray)

This is our most notable snake, YPM
9151, found alive on the road, but mor-
tally wounded, at 2000 hrs, 22 July 1997,
at Wu Ping Zai (Fig. 2:6). The specimen is
a subadult male, SVL 360 mm, tail 95 mm;
the hemipenis is partially everted on the
right and the left has been dissected.
There is an anomalous half ventral just
before the anal plate. Table | lists some
characters of this individual, compared to
T. albolabris and T. stejnegeri Schmidt, the
other two green “bamboo” vipers of
South China.As in both T. albolabris and
T. stejnegeri, YPM 9151 has the nasal scale
in broad contact with the plate forming
the anterior border of the pit, and a sep-
aration of the supraoculars by 12
smooth, subimbricate scales (variation
8—|3, both species). The dorsals are
keeled, and there are the usual 21 rows
at midbody. There are 72 subcaudals
(male range: 65—73 in T. albolabris, 68—74
in T. stejnegeri). The hemipenes are not
fully developed, but the sulcus is already
prominent. There are three large, al-
though still soft, spines at the base of the
fork on each member.

In life this snake was bright green with
a brilliant yellow belly and supralabials;
there were very faint, shadow-like dark
transverse markings, but these appear
more prominent in preservative. The

Postilla 217 1 |

upper half of the first dorsal scale on
each side was bold lime white, forming a
nearly continuous stripe along the side.
The tail top was russet. The morphology
accords well with T. albolabris from
Guangdong and Hong Kong. However, in
life the lower half of the first dorsal scale
on each side was brick red. The iris was
similarly brick red, notably darker than in
T. albolabris. In both color characters YPM
9151 agrees with T. stejnegeri. The pres-
ence of developing spines on the imma-
ture hemipenes is unlike T. albolabris and
accords with T. stejnegeri (Pope 1935).

Bufo melanostictus Schneider

This is the abundant toad of South China
and nearby coastal islands, documented
on Nan Ao by Li and Pan (1995).We took
a large female, YPM 5471 (89 mm SVL), as
a voucher at Tai Zhu Ao, 23 July 1997, and
note MCZ 125859 from Houzhai. Sight
records are from Fig. 2:2 and 5.

Bufo sp. indet.

We here call attention to a peculiar
subadult toad (Fig. 6) from Houzhai (Fig.
2:3), MCZ 125860, 35 mm SVL, that was
exchanged as Bufo gargarizans Cantor on
geographic grounds. It certainly is not
that species. In life it was patterned en-
tirely in soft browns, without dark spines
or crest edges, or any blackish coloration
dorsally, but with dark ventral spotting.
The cranial crests, albeit weakly devel-
oped, are unlike either the South China
B. gargarizans or B. melanostictus. Profes-
sor Li Zhen-chang visited MCZ on 19 Au-
gust 1998 and compared this individual
directly to the Chinese Bufo and the sev-
eral American species sympatric with
Rana catesbeiana. We suspected it might
have been introduced inadvertently with

12 Postilla 217

eggs. We are satisfied, however, that it dif-
fers strikingly from all these forms. We
await collection of additional specimens.

Rana catesbeiana Shaw

American bullfrogs, vouchered by YPM
5521-3 and collected in the wild at
Houzhai on |5 July 1997, and at Bai Sha
Wan on 23 July 1997, are now well estab-
lished on Nan Ao (Fig. 2:3 and 4). They
were introduced as a commercial venture
for food.We were informed locally that
three “kinds” have been imported: “Ameri-
can bullfrogs, Cuban bullfrogs and river
frogs.” We have seen only R. catesbeiana,
but that species is notorious for eating
other frogs and is implicated in the decline
and local extirpation of several species na-
tive to areas where exotic R. catesbeiana
has been introduced and become estab-
lished (Hayes and Jennings 1986).

Rana guentheri Boulenger

This species is common on South China
Sea islands like Hong Kong and Lantau
(Karsen et al. 1986) and documented on
Nan Ao by Li and Pan (1995). We col-
lected YPM 5526-7 at Houzhai, YPM
5529-32 at Shen Ao, and YPM 5528 at Tai
Zhu Ao between | 3-20 July 1997 (Fig. 2:3
and 7). These frogs are somewhat differ-
ent from those we are accustomed to
elsewhere in Guangdong and in the Hong
Kong region. They are noticeably less red
brown, tending to dark mustard or olive
brown. They have lustrous, metallic look-
ing platinum to gold upper lips and folds
from tympanum to shoulder, much more
prominent than usual elsewhere. Four
males are 65—74 (average 68) mm SVL
and three females are 61—74 (average 69)
mm SVL. Males are easily recognized by
their wrinkled lateral throat vocal sacs.

Reptiles and Amphibians
from Nan Ao Island, China

Rana limnocharis Boie

The paddy frog is the most abundant frog
at present on Nan Ao. We put up 12
voucher specimens (YPM 5549 and
5551-61) and recorded them at six local-
ities (Fig. 2: western tip, |, 2, 3,5 and 7).
We used this species frequently for bait.
It is common on South China Sea conti-
nental shelf islands, including some rather
small ones (Lazell |988b), and is docu-
mented on Nan Ao by Li and Pan (1995).
Our specimens measured 28-49 (average
38) mm SVL; only the largest appeared
reproductive, a gravid female. Of a pair of
adults from Houzhai collected 25 July
1996, the male (MCZ 125857) measured
32 mm SVL, and the female (MCZ
125858) measured 41 mm SVL.The sexes
are easily distinguished by the sooty black
patches on the throat sides of males.

Three striking pattern variants of R. lim-
nocharis vary geographically in frequency:
unstriped, narrow striped and broad
striped middorsally (Ota and Lin 1997). Of
14 from Nan Ao, seven are unstriped, four
have narrow stripes, two have broad
stripes and one is anomalous. On the an-
terior half of the trunk of YPM 5556 the
middorsal stripe is broad; at midbody
there is a sharp bend to the left, the stripe
makes an acute angle, and resumes its
middorsal position as narrow on the pos-
terior trunk. Only color pattern seems
anomalous in this individual. There is no
obvious scar tissue and the longitudinal ru-
gosities are symmetrical and appear nor-
mal, even within the bent stripe area.

Rhacophorus leucomystax
(Gravenhorst) or Polypedates
megacephalus Hallowell

This brown tree frog was not recorded
on Nan Ao by Li and Pan (1995). Taxo-

Reptiles and Amphibians

from Nan Ao Island, China

Table |

Nan Ao
Supranasal and nasal fused
Internasals in contact
Dorsal rows at nape 21
Dorsal rows at vent 17
Ventrals 154.5
Belly color yellow
First dorsal color white/red
Transverse bars present
Hemipenial spines 3
Hemipenial sulcus prominent
Iris color brick red

Postilla 217 13

T. albolabris T. stejnegeri
fused separate
in contact separate
21-27 2\—23
15 15
153-162 161-171
yellow green
white/green white/red
present absent
none many
prominent weak
pale red brick red

Some characteristics of male South China green “bamboo” vipers. Data from Pope (1935) or
Hu et al. (1980) except for the Nan Ao specimen, YPM 9151.

nomic uncertainty, from family through
species level, is noted by Lazell et al.
(1997). Our vouchers, YPM 5604-6 and
5608-9, from Houzhai and Tai Zhu Ao, re-
spectively (Fig. 2:3 and 7), measured
47-66 (average 54) mm SVL; the largest is
a gravid female, YPM 5604, collected in
amplexus with YPM 5605, 13 July 1997.
Brown tree frogs are abundant on Hong
Kong (type locality of P. megacephalus),
Lantau and many smaller coastal islands
(Karsen et al. 1986, Lazell 1988b, Lazell et
al. 1997).

Kaloula pulchra Gray

First recorded on Nan Ao in July 1996
(Lazell and Lu 1996), the “painted frog,”
“Malaysian narrowmouth toad” or “un-
derground ox” is abundant in the remain-
ing wetlands on the Houzhai plain, but we
did not encounter it elsewhere (Fig. 2:3).
We have noted the ability of these frogs
to survive urbanization, living in gardens
and parks and breeding in storm drains in

Houzhai and Hong Kong. Although their
occurrence is spotty, they inhabit some
small islands (Lazell et al. 1997). In addi-
tion to the vouchers in China and MCZ,
we have YPM 5497,a 55 mm SVL male.

Microhyla pulchra (Hallowell)

This “pygmy” frog is one of the most in-
tricately patterned of anurans, colored in
shades of brown, typically with brilliant
yellow ventrally and on the concealed
surfaces of the hind limbs. Color pho-
tographs are provided by Karsen et al.
(1986) and a color illustration is in
Lazell (1989). Four adults (YPM 5514-7),
collected 16 and 21 July 1997, measured
29-31 (average 30) mm SVL.AIl are
from Houzhai (Fig. 2:3).A juvenile (YPM
5513), collected 13 July 1997, measured
16 mm SVL. This species is documented
from Nan Ao by Li and Pan (1995) and
on the larger coastal islands in the Hong
Kong region (Karsen et al. 1986). While
it does well in edificarian habitats such

14 Postilla 217

as the wet paddy, it does not seem to
survive urbanization.

Microhyla ornata
(Dumeril et Bibron)

This somber “pygmy” frog was not
recorded on Nan Ao by Li and Pan
(1995).We found it abundant at several
localities on the western part of the is-
land (Fig. 2:1, 2 and 3): YPM 5506-11. It is
known from some small islets in the
South China Sea (Lazell 1988b) and fre-
quents agricultural areas (Karsen et al.
1986). Our specimens ranged from 13-18
(average 16) mm SVL.

Discussion

Our total of only 19 species for a conti-
nental shelf island the size of Nan Ao is
low. Hong Kong island, much smaller in
area at 78 square kilometers, has at least
58 species of reptiles and amphibians
(Karsen et al. 1986). Further, Hong Kong
is massively urbanized with a nearly ring-
ing coastal metropolis from Victoria on
the north through Aberdeen in the west
to Stanley in the southeast. However,
much of the interior of Hong Kong island
is protected as watershed and country
parks, today largely covered in some of
the better (albeit mostly second growth)
monsoon forest remaining in tropical
China. In a very brief survey Ota and Lin
(1997) record 17 species from two is-
lands, totaling |7 square kilometers, on
the continental shelf.

Despite deforestation and urbaniza-
tion, we believe Nan Ao will prove to
support many more species than we
have recorded. We are confident that
pythons, kraits and cobras are present.
Indeed, we call attention to Pope’s
(1935:352) record of Naja atra Cantor

Reptiles and Amphibians
from Nan Ao Island, China

from “Lamock” island. The Le Men Lie
Dao, an archipelago within Nan Ao
County centered circa 10 kilometers
south of the eastern end of Nan Ao, has
at least four vegetated islets that might
support reptiles like cobras (and cer-
tainly geckos and skinks). Cobras have
been reported on Wu Yu, but a single in-
dividual may have been introduced there
to discourage sea bird eggers (YL, un-
publ. data). There also may be many more
species of colubrid snakes. We observed
two Coluber mucosus L. at a restaurant in
Houzhai, 25 July 1996, said to have been
captured locally. This snake is regularly
found on coastal islands (Lazell 1988b).
We saw a number of snakes we failed to
catch, and our funnel traps, baited with
frogs and geckos, were regularly raided,
even when the raider had to regurgitate
the bait to escape. On |5 July 1997 we
found a road killed skink, Mabuya longi-
caudata (Hallowell), at Shen Ao. The car-
cass was in two pieces and discarded. We
subsequently saw several more of these
large, conspicuous lizards alive, but failed
to catch one. We have noted above the
possibility that Eumeces quadrilineatus
may account for some of our sightings of
blue-tailed skinks. Local people speak of
a “fence dragon” (YL, unpubl. data). This
name refers to Calotes versicolor (Daudin)
(Agamidae) in Hong Kong.

There are two intriguing herpetologi-
cal legends on Nan Ao.The first describes
a small frog that lived only around one
temple in the hills. Its persistent, noctur-
nal calling so irritated the monks that
they cut its throat. Thereafter it was the
only mute species of frog on the island
and could be recognized by the white
scar-like marking across its throat. We
cannot guess this species. The second
concerns the name of Gui Chen, “turtle
flat” (Fig. 1:1). The site was originally a

Reptiles and Amphibians
from Nan Ao Island, China

Pally Up

2

€

Figure 6

WHAAHHAL

Si
“3 |
SY

LUUATUIUUYIIL ALU TTD

|

Postilla 217 15

in

Bufo sp. indet., MCZ 125860, Houzhai, Nan Ao. Photographed by William K. Sacco.

marsh with ponds inhabited by the
“golden coin turtle,’ now believed extir-
pated. This name applies to Sacalia quadri-
ocellata (Siebenrock), a species well
known from southern Guangdong and
Hainan Dao, where it inhabits streams at
100-400 m elevation (Pope 1935, as
Clemmys quadriocellata). The name derives
from the prominent, light-colored, dark-
centered ocellate spots (two on each
side of the head) that are rich golden yel-
low in young specimens.

On Nan Ao there are few natural
wetland habitats left. Prior to the con-
version of the mangrove swamps in the
1950s, there were several species of
aquatic marine snakes with great eco-
nomic value as delicacies. One, called the
“round-tailed sea snake,” fits the de-
scription of Enhydris bennetti (Gray), a

species we believe has been extirpated
from the entire coast of China east of
Hainan Dao, except at Mai Po in the
Hong Kong region (Karsen et al. 1986).
The effects of mangrove destruction on
Nan Ao’s fisheries were catastrophic.
Today, shrubs and small trees are cut to
provide long rows of stakes extending
into the bays at Houzhai and Shen Ao.
These make a sort of artificial mangrove
mimic for the cultivation of shellfish.
Cutting for this purpose further hastens
the deforestation of the uplands.
Although we visited more than 20 po-
tential collecting sites on Nan Ao, we ob-
tained specimens from only seven (Figs. 2
and 3). Of the total of 19 species col-
lected, |7 came from three sites: the
Houzhai plain, the Huanghua Shan Reser-
voir and Tai Zhu Ao. Of these 17, eight

16 Postilla 217

were found at only one of those sites.
The sites have disparate characteristics.
The Houzhai plain, near sea level and
with remnant freshwater wetlands, pro-
vided our only specimens of Eumeces chi-
nensis, Microhyla ornata, Kaloula pulchra
and the undetermined Bufo. The shady
ravine at Huanghua Shan Reservoir, circa
200 m above sea level, was our only site
for Ateuchosaurus chinensis and Sphe-
nomorphus indicus. Tai Zhu Ao is a coastal
site with an extensive paddy on the valley
floor and woodlands on the flanking
slopes; here we got Elaphe radiata and
Oligodon formosanus. We hunted fre-
quently at Shen Ao, for example, which is
coastal with considerable freshwater wet-
land and paddy, and with adjacent
wooded slopes, but got none of those
eight species or anything unique. No
doubt luck was a factor, but there may be
real, highly important intraisland habitat
differences too.

Plans to further develop and urbanize
Nan Ao were put forward in the early
1990s (Nan Ao County Geographical Bu-
reau 1994).A bridge, as yet incomplete, is
to connect the western end of Nan Ao to
the mainland. The land area is to be in-
creased by nearly one quarter, to circa
130 square kilometers, by filling the
largest bays, and four large coastal cities
are planned. Already most of the agricul-
tural area and wetland of the Houzhai
plain has been graded and filled. If these
plans are carried out, the impact on fauna
and flora will be incalculable. We herein
document several species as yet known
only from the Houzhai plain, surely once
the most extensive natural marshy part of
this island. Some of the anurans naturally
abundant in this habitat, like Rana guen-
theri and R. limnocharis, are desirable food
items of economic value elsewhere in
South China. Introduction and cultivation

Reptiles and Amphibians
from Nan Ao Island, China

of exotic bullfrogs, like staking the bays,
seems a poor substitute for the native
ecosystems and species lost. We did not
find Rana rugulosa Wiegmann, a large,
highly edible native species recorded by Li
and Pan (1995). Its large freshwater habi-
tats may already have been destroyed.A
major effort to document the fauna and
flora of Nan Ao, and the more than 20
smaller islands in Nan Ao County, is
clearly called for (Qian and Lu 1994).

Acknowledgments

The late Lin Yong-Mu, our co-author, pro-
vided all our physical and most of our
ecological data on Nan Ao and Wu Yu. We
dedicate this paper to his memory. We
are indebted to Jiang Tong, Cai Fa-sheng,
Hong Sheng-gin and Zeng Li-rong (San-
tou University). Huang Ruo-kun, Yang Si-
chang (Nan Ao County) and Li Zhen-
chang (South China Normal University)
provided logistical assistance to our on-
going exchange program. In the field we
were ably assisted by Ben Calvi, Shannon
Corliss, Phaedra Demers, Peter Lynch,
Andrea Scott, Fred Sibley, Tom Smith, Car-
olyn Tate, Bonnie Welcker, Wu Shao Ping
and Adam Young. This research was sup-
ported by Santou University, Yale Univer-
sity, the University of Hong Kong, the Ex-
plorers Club, the Gailer School, the
Hayes Foundation, the Science and Tech-
nology Bureau of Nan Ao County and
The Conservation Agency.

Reptiles and Amphibians Postilla 217 17
from Nan Ao Island, China

Literature Cited

Auffenberg, W. and T. Auffenberg. 1989. Reproductive patterns in sympatric Philippine
skinks (Sauria: Scincidae). Bull. Florida State Mus. Biol. Sci. 34:201—247.

Dowling, H. G. 1951.A proposed standard system for counting ventrals in snakes. Brit.
J. Herpetol. 1:184—185.

Dudgeon, D. and R. Corlett. 1994. Hills and streams: an ecology of Hong Kong. Hong
Kong: Hong Kong Univ. Press. 234 pp.

Hahn, D. E. 1977. Review: revision systematique des Typhlopidae d’Afrique, Reptilia—
Serpentes. J. Herpetol.Assoc.Africa |5:34-39.

Hayes, M. P. and M. R. Jennings. 1986. Decline of ranid frog species in western North
America: are bullfrogs responsible? J. Herpetol. 20:490—-509.

Hu, B., M. Huang, J. Zhe, E. Zhao, Y. Jiang and X. Huang. 1980. Chinese snakes illustrated.
Shanghai: Sci. Technol. Press. 166 pp, | 12 plates.

Karsen, S.,M. Lau and A. Bogadek. 1986. Hong Kong amphibians and reptiles. Hong
Kong: Urban Council. 136 pp.

Lazell, J. 1988a. Herpetology in South China. Herpetol. Rev. 19(3):49-51.
1988b. Soko Islands, South China Sea. Explorers J. 66(2):80—85.
1989. Hong Kong frogs. Earthwatch 9(1):9-I 1.

1994.A brief herpetological reconnaissance of Ping Chau, Mirs Bay. Porcupine,
Univ. Hong Kong 11:17.

Lazell, J. and W. Liao. 1986. Contribution to the herpetofauna of Dinghushan,
Guangdong. Acta Herpetol. Sinica 5(1):70—71.

Lazell, J. and W. Lu. 1996. Geographic distribution. Kaloula pulchra pulchra (Asiatic
painted frog). Herpetol. Rev. 27:209.

Lazell, J.. M. Lau and W. Lu. 1997.A brief herpetological excursion to Wai Ling Ding,
Wanshan Islands, South China Sea. Asiatic Herpetol. Research 7:80—84.

Lazell, J. and H. Ota. 1999. Aspects of the life history, ecology, and distribution of the
Asiatic four-lined skink, Eumeces quadrilineatus, in China. Breviora. In press.

Li, Z. and J. Pan. 1995. Guangdong Province amphibians: faunal regions and
biogeography. Chinese amphibian biogeography | 6. Herpetol. Series 8:125—1 30.

Nan Ao County Geographical Bureau. 1994. Travel and tourism map. Guangzhou:
Guangdong Cartographic Press. | sheet.

Okada, S., H. Ota, M. Hasegawa, T. Hikida, H. Miyaguni and J. Kato. 1992. Reproductive
traits of seven species of lygosomine skinks (Squamata: Reptilia) from East Asia. Nat.
Hist. Research 2:43—52.

Ota, H. and J. Lin. 1997. On the herpetofauna of the Matsu Group—I. Reptiles and
amphibians recorded from Nankan and Peikan Islands. J. Taiwan Mus. 50:93—105.

Ota, H., M. Lau, T. Weidenhofer, Y. Yasukawa and A. Bogadek. 1995. Taxonomic review of
the geckos allied to Gekko chinensis Gray 1842 (Gekkonidae: Reptilia) from China and
Vietnam. Trop. Zool. 8:181—196.

18 Postilla 217 Reptiles and Amphibians
from Nan Ao Island, China

Pope, C. H. 1935. The reptiles of China. New York: Amer. Mus. Nat. Hist. 604 pp.

Qian, H. and B. Lu. 1994. Establishment of Nan Ao Island as a synthetic oceanic natural
reserve and its implication. Ecolog. Sci. 1994:138—142 [in Chinese].

Wang, H. 1985.Atlas of the paleogeography of China. Beijing: Cartographic Publ.
House. |43 pp.

Wallach, V. 1993.A new species of blind snake, Typhlops marxi, from the Philippines.
Raffles Bull. Zool. 41:263—278.

Williams, S$. W. 1895. The middle kingdom. New York: C. Scribners Sons. 836 pp, 775 pp.
Wu, L., D. Li and J. S. Liu. 1985. Guizhou fauna: Reptilia. Guiyang, China: Guizhou
People’s Publisher. 434 pp [in Chinese].

Zhao, E. and K.Adler. 1993. Herpetology of China. Oxford, Ohio: Soc. Study
Amphibians and Reptiles Contrib. Herpetol. 10.522 pp.

Zhou, K.,Y. Liu and D. Li. 1989. Three new species of Gekko and remarks on Gekko
hokouensis (Lacertiformes, Gekkonidae). Smithsonian Herpetol. Info. Svc. 77:1—12.

Zweifel, R. G. 1960. Results of the Puritan-American Museum of Natural History
expedition to western Mexico 9. Herpetology of the Tres Marias Islands. Bull. Amer.
Mus. Nat. Hist. 119:1 12-115.

“iii

3 9088 01206 5223