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A New Species of Amolops (Anura: Ranidae) from Southwest China

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DOI: 10.1643/0045-8511(2007)7[913:ANSOAA]2.0.CO;2

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 Copeia, 2007(4), pp. 913–919

A New Species of Amolops (Anura: Ranidae) from Southwest China

DING-QI RAO AND JEFFERY A. WILKINSON

A new species of Amolops is described from a mountainous area of southern Yunnan

Province, China. The species is unique in having a dark purple dorsum with small light
yellow spots. The spots are smaller than the smallest finger disk. Other characters that
distinguish this species from other species of Amolops include smooth skin (lacking
tubercles) over the entire body and lack of dorsolateral folds, transverse bars on the
legs, and a visible pineal body.

ROGS of the genus Amolops, commonly
F referred to as cascade or torrent frogs, are
distributed in mountain streams of India, South-
east Asia, and southern China. Typically these
frogs can be distinguished from other species of
Asian ranids by the presence of enlarged disks on
the fingers and toes, non-glandular skin, absence
of humeral glands, presence of gular pouches in
males, dorsoventrally flattened bodies, tadpoles
with poison glands and a gastromyzophorous
adhesive disk (abdominal sucker), and inhabit-
ing swift torrents and splash zones (Inger, 1966;
Yang, 1991). However, several frogs in the
nominal genera Hiua, Hylarana, Odorrana, Mer-
istogenys, Rana, and Staurois also contain combi-
nations of these characters.
Recent molecular phylogenetic studies by Chen
et al. (2005), Frost et al. (2006), Matsui et al.
(2006), and Ngo et al. (2006) support Amolops,
with the exception of A. chapaensis, as a mono-
phyletic group. Amolops chapaensis was shown to be
more closely related to species of Odorrana and
thus reassigned to that genus by Ngo et al. (2006),
while Frost et al. (2006) assigned all species of
Odorrana, including O. chapaensis, to the genus
Huia based on H. nasica being nested within an
Odorrana clade (a result similar to Chen et al.
[2005] and Ngo et al. [2006]). However, they
recognize that this taxonomy is problematic and
the assigned name may be inappropriate, as H.
nasica may not be closely related to other species
of Huia (not included in their analysis). Matsui et
al. (2006) indicate that Huia (H. cavitympanum
and H. sumatrana in their study) may be more
closely related to Meristogenys, though they did not
include H. nasica or species of Odorrana in their
study. Because of these discrepancies and in-
adequacies in taxon sampling between the studies,
we still maintain Odorrana separate from Huia, but
with O. nasica and O. chapaensis within Odorrana as
in Chen et al. (2005) and Ngo et al. (2006).
Though these studies confirmed Amolops as
a monophyletic group, they did not define
Amolops based on morphological characters be-
yond what were presented in Yang (1991).
# 2007 by the American Society of Ichthyologists and Herpetologists
Of the 35 known species of Amolops, over half
are from mainland China (D. F. Frost, Amphib-
ian species of the world: an online reference,
v4.0, http://research.amnh.org/herpetology/
amphibia/index.php [American Museum of Nat-
ural History, 17 August 2006]). Some species
have only recently been discovered from south-
west China (Liu and Yang, 2000; Liu et al., 2000;
Zhao et al., 2005). We believe the lateness of
these findings is partly due to inadequate
sampling of remote areas within the north–south
aligned Trans-Himalayas (Hengduangshan
mountain range) and their southern extensions
in this region. Zhao (1999) considers these high
mountain ranges with deep valleys a center of
speciation for some amphibians including Amo-
lops due to the vertical zonation of climate and
vegetation that provides a wide range of environ-
ments. He bases this argument on the high
degree of diversity and specific endemism found
in this region (highest among his regions in
mainland China). The results of Matsui et al.
(2006) partially support this claim with different
origins of southwestern and southern lineages of
Chinese Amolops.
Specimens of a previously undescribed species
of Amolops were photographed from Mount
Huanglianshan, Yunnan Province, on the Viet-
nam border. The specimens were found in the
spring of 2002 by local people who were
participating in a gibbon survey with the staff of
the Kunming Institute of Zoology (KIZ). Sub-
sequently, the first author and staff of the
Huanglianshan National Nature Reserve
(HNNR) were able to collect voucher specimens
of this species. This species is described herein,
based on those specimens.
MATERIALS AND METHODS
All specimens were collected by hand and
euthanized. Tissue samples were removed from
the paratypes (except for KIZ-Luchun04A016)
and all specimens were fixed in 10% formalin and
later stored in 75% ethanol. All specimens are

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914 COPEIA, 2007, NO. 4

TABLE 1. MEASUREMENTS OF THE TYPE SPECIMENS OF Amolops caelumnoctis (IN MM). See text for abbreviations.

KIZ-2003 KIZ-Luchun KIZ-Luchun KIZ-Luchun KIZ-Luchun KIZ-Luchun KIZ-Luchun KIZ-Luchun

Specimen A018 04A005 04A004 04A006 04A001 04A002 04A003 04A016
Sex/size male male female female female female female juvenile

SVL 71.3 73.7 78.0 86.0 90.6 88.4 81.0 39.2

HL 25.3 27.3 29.8 31.3 32.0 31.5 29.0 13.8
HW 24.8 24.4 29.3 31.2 32.3 31.3 28.0 13.3
SL 10.5 11.2 10.7 12.5 12.9 12.8 11.5 5.7
IOD 9.0 9.3 9.8 9.7 10.6 10.2 9.5 4.5
ED 8.8 8.4 9.5 10.3 10.9 10.0 8.9 5.0
TD 3.1 3.2 3.2 3.3 3.2 3.3 3.1 1.6
FAL 38.2 39.5 48.6 51.6 51.6 51.2 46.1 21.6
HaL 27.0 24.6 31.9 32.8 34.1 33.4 30.0 14.4
HLL 127.2 129.5 153.2 156.0 161.5 165.0 154.2 71.6
TL 41.0 38.5 48.7 48.6 50.6 50.0 47.6 23.2
FL 39.6 40.5 47.2 45.5 50.0 48.8 42.5 21.8

housed at KIZ. The following measurements were
obtained (Table 1): snout–vent length (SVL),
head length (HL), head width (HW), snout
length (SL), interorbital distance (IOD), eye
diameter (ED), tympanum diameter (TD), fore-
arm length (FAL), hand length (HaL), hind limb
length (HLL), tibia length (TL), and foot length
(FL). All measurements were taken using digital
calipers to the nearest 0.1 mm. The specimens
were compared with available specimens (see
Material Examined) and original descriptions of
the following currently recognized species of
Amolops from China and neighboring South and
Southeast Asian countries (D. F. Frost, Amphibian
species of the world: an online reference, v4.0,
http://research.amnh.org/herpetology/amphibia/
index.php [American Museum of Natural History]
17 August 2006): A. aniqiaoensis, A. bellulus, A.
chakrataensis, A. daiyunensis, A. cremnobatus, A.
gerbillus, A. jansauri, A. kangtingensis, A. kaulbacki,
A. larutensis, A. liangshanensis, A. loloensis, A.
longimanus, A. medogensis, A. mengyangensis, A.
nepalicus, A. panhai, A. spinapectoralis, A. tormotus,
A. tuberodepressus, A. viridimaculatus, and A. wuyien-
sis. These descriptions were supplemented with
works by Fei (1999), Fei et al. (2005), and Yang
(1991) that included descriptions of A. chunga-
nensis, A. formosus, A. granulosus, A. hainanensis, A.
himalayanus, A. hongkongensis, A. jinjiangensis, A.
lifanensis, A. mantzorum, A. marmoratus, A. monti-
cola, A. ricketti, and A. torrentis. In addition, the
specimens were compared with original descrip-
tions and works by Fei (1999) and Fei et al.
(2005) for the following species: Huia absita, H.
melasma, Hylarana erythraea, H. macrodactyla, Odor-
rana andersonii, O. anlungensis, O. archotaphus, O.
aureola, O. bacboensis, O. banaorum, O. bolavensis, O.
chapaensis, O. chloronota, O. compotrix, O. cucae,
O. daorum, O. exiliversabilis, O. gigatympana, O.
grahami, O. graminea, O. hainanensis, O. heatwolei, O.
hejiangensis, O. hmongorum, O. hosii, O. indeprensa, O.
iriodes, O. jingdongensis, O. junlianensis, O. khalam, O.
kuangwuensis, O. leporipes, O. livida, O. lungshengen-
sis, O. margaretae, O. megatympanum, O. monjerai, O.
morafkai, O. nasica, O. nasuta, O. orba, O. schmackeri,
O. sinica, O. tabaca, O. tiannanensis, O. trankieni, O.
versabilis, O. vitrea, and O. wuchuanensis.
Amolops caelumnoctis, new species
Figures 1–3
Holotype.—KIZ-2003A018, adult male, China,
Yunnan Province, Honghe Prefecture, Luchun
County, Qimaba Township, collected from
a mountain stream in the forest of Mount
Huanglianshan (HNNR), 2400 m, approximately
12.4 km NNE of Ha-Zha Village, 22u549360N,
102u149240E, datum: WGS84, 18 May 2003, D.-Q.
Rao (geocoordinates were added retrospectively
and should not be considered original data
supplied by the collector).
Paratypes.—1 male (KIZ-Luchun04A005), 5 fe-
males (KIZ-Luchun04A001, 002–004, 006), and 1
juvenile (KIZ-Luchun04A016), China, Yunnan
Province, Honghe Prefecture, Luchun County,
Qimaba Township, collected in streams near Ha-
Zha Village, March 2004, staff of HNNR.
Diagnosis.—The new species differs from all other
species of this genus by a combination of the
following characters: numerous small round light
yellow spots, irregularly distributed, on a dark
purple background on dorsum, head, limbs,
fingers, and toes; smooth skin (lacking tubercles)
over entire body; lack of dorsolateral folds; lack
of transverse bars on limbs; and lack of visible
pineal body on top of head.

Copeia cope-07-04-13.3d 10/11/07 13:13:11 914 Cust # CH-06-248R

 RAO AND WILKINSON—NEW SPECIES OF AMOLOPS 915

Fig. 1. Holotype (KIZ-2003A018) of Amolops caelumnoctis: (A) dorsal view and (B) ventral view (scale 5
10 mm).

Copeia cope-07-04-13.3d 10/11/07 13:13:12 915 Cust # CH-06-248R

916 COPEIA, 2007, NO. 4
Fig. 2. Holotype (KIZ-2003A018) of Amolops
caelumnoctis: (A) lateral view of the head (scale 5
5 mm), (B) palmar view of the left hand, and (C)
plantar view of the left foot (scale 5 5 mm).
Description of holotype.—Adult male, SVL 71.3 mm
(see Table 1 for additional measurements); body
depressed and elongate; width of head less than
length, greater than width of body. Head dorso-
ventrally flattened; snout sloping from anterior of
eye to tip, projecting only slightly beyond lower
jaw; canthus rostralis rounded; loreal region
concave, not oblique, almost vertical; nostril
dorsolaterally located midway between tip of
snout and eye; top of head oblique from occipital
to frontal, interorbital and frontal areas concave;
snout rounded in dorsal view; pineal body not
visible; eyes large and anterolaterally protuberant,
almost as long as snout; temporal region not
swollen; tympanum small, distinct; vomerine teeth
strongly converging backward between choanae;
choanae mere slits, laterally located and ob-
structed by lingual shelf in ventral–lateral view;
tongue deeply notched, posterior 1/3 free; vocal
sac and vocal apertures absent.
Arm slightly enlarged and elongate; fingers
slender and long, when adpressed, finger formu-
la 3–4–2–1, third almost as long as forearm; tips
of fingers 2 through 4 expanded to disks, disks at
fingertips with circummarginal grooves, disk of
third finger largest, larger than tympanum;
webbing and lateral fringes on hand absent;
subarticular tubercles present on midventral
Copeia cope-07-04-13.3d 10/11/07 13:13:17 916
ridge, large, oval or almost round, obviously
bulging, formula 1,1,2,2; supernumerary tubercle
present at base of each finger, indistinct; inner
and outer metacarpal tubercles small, narrow
and flat; velvety nuptial pad, brownish black
ventrally, grayish white dorsally, extending medi-
ally from base of first finger to distal edge of first
subarticular tubercle.
Legs long and slender; heels overlap when legs
at right angles to body; tibial–tarsal articulation
reaches tip of snout; tibia approximately 57%
length of body; toes fully webbed, webbing
formula I1–1KII1–2III1K–2IV2K–1KV follow-
ing Myers and Duellman (1982) as modified by
Savage and Heyer (1997) with distinct fringes
reaching to disks; disks of toes smaller than those
of fingers with circummarginal grooves; oval
subarticular tubercles present on all toes, sub-
articular tubercle formula 1,1,2,3,2; inner meta-
tarsal tubercle moderate, narrow and elongate,
slightly bulging, less than half of length of first toe;
outer metatarsal tubercle absent. Skin dorsally
and ventrally smooth, including throat, chest,
abdomen, ventral surface of limbs, and flanks; no
dorsolateral fold; temporal fold distinct.
Coloration in life.—Dorsum and flanks dark
purple, irregularly interspersed with round light
yellow spots (Fig. 3); spots in three distinct size
classes, spots on flanks slightly larger than those
on dorsum; eyes dark brown with scattered yellow
spotting and yellow ring around iris; canthus
rostralis light brown, dark nictitating membrane,
eyelids dark purple as with dorsum of body;
throat and chest mostly light blue or white with
some black mottling; anterior abdomen light
gray, posterior abdomen with dark gray mottling;
groin dark gray with small white irregular
spotting; dorsal aspect of limbs lack transverse
bands; ventral aspect of arms light blue or white;
ventral aspect of thighs light blue with black
reticulation; ventral aspect of tibia and tarsus
light blue with tiny light yellow spots; ventral
aspect of hands including tubercles dark gray;
ventral aspect of feet including tubercles, disks of
digits, and both inner and outer metacarpal
tubercles and inner metatarsal tubercle dark
gray; webbing between toes yellow.
Coloration in preservative.—Dorsum black with small
white spots. Ventrum light brown, white, or gray.
Variation.—Snout–vent length varies from 71.3 to
73.7 mm in the two males and 78.0 to 90.6 mm
in the five adult females. Coloration and size of
spots are same between the seven adults and one
juvenile, but number of spots increases with body
size so that density of spots is similar. KIZ-
Cust # CH-06-248R
 RAO AND WILKINSON—NEW SPECIES OF AMOLOPS
Fig. 3. Photograph in life of the holotype (KIZ-2003A018) of Amolops caelumnoctis.
Luchun04A002–004 and KIZ-Luchun04A006
have several relatively large spots on shoulder;
KIZ-Luchun04A001 and 002 have longitudinal
dark mottling and bars on light blue ventrum,
whereas the mottling and bars are absent on the
remaining specimens; the two males, juvenile,
and KIZ-Luchun04A006 have darker gray ventra.
Dimorphism.—Males smaller than females (Ta-
ble 1); males have velvety nuptial pads on base
of first fingers making first fingers appear short
and stumpy; males with relatively thicker but
shorter arms, and shorter hands, while females
have more slender and longer arms and hands,
with more slender first fingers; inner metacarpal
tubercle of males much larger than outer, not so
in females. Males without vocal sac or gular
pouches. Dorsal spots of males slightly smaller
than females.
Distribution.—Known only from the localities of
the holotype and paratypes (Fig. 4). According
to interviews with the local people of Jinping
Copeia cope-07-04-13.3d 10/11/07 13:13:23 917
917
County, this species may be distributed in the
mountainous areas of Jinping County, which is
a border region with Vietnam. Its range, there-
fore, possibly extends to northern Vietnam.
Ecology and habitat.—The new species lives in
cascading mountain streams of heavily forested
mountains, 2400 m elevation. The streams have
sandy bottoms and large boulders beside or
within the stream. Flow was high at the time
collections were made.
Amolops tuberodepressus, Nanorana unculuanus,
and a tadpole of Vibrissaphora sp. were collected
during the survey in March of 2004. The new
species was observed breeding in the early spring,
differing from other species of Amolops. The
breeding season appears to be comparatively
short, lasting only one month as most specimens
were found during this period even though
surveys continued through April. In contrast,
the sympatric Amolops tuberodepressus breeds later
in June and July. No tadpoles or calls were
collected.
Cust # CH-06-248R
918 COPEIA, 2007, NO. 4
Fig. 4. Type locality of Amolops caelumnoctis
(indicated by star). Yunnan Province is solid white.
Huanglianshan National Nature Reserve is outlined
in white (taken from WDPA Consortium ‘‘World
Database on Protected Areas’’ 2006—Copyright
World Conservation Union [IUCN] and UNEP–
World Conservation Monitoring Centre [UNEP–
WCMC]. Electronic database accessible at http://
www.unep-wcmc.org/wdpa).
Comparisons.—Because members of the genus
Amolops share many of the adult characters
mentioned in the introduction with species of
other ranid genera, the presence of the abdom-
inal sucker in the tadpole is considered the
defining character for the genus, as it is present
in all species for which the tadpole is known and
absent in all species of Huia, Hylarana, and
Odorrana in China and Southeast Asia (Cambo-
dia, Laos, Thailand, and Vietnam) for which the
tadpole in known (see Frost et al., 2006 for
a discussion of the dubiousness of the abdominal
sucker in the tadpole of O. nasica). Because
the tadpole for the new species is unknown but
the adult characters are mostly consistent with
the characters for Amolops, we place the new
species in this genus.
Several species of Amolops are known from
India, Southeast Asia, and southern China.
Amolops caelumnoctis differs from all of these
species by its unique dorsal color pattern of
small light yellow spots interspersed on a dark
purple background. In addition to the dorsal
color pattern, A. caelumnoctis differs from A.
Copeia cope-07-04-13.3d 10/11/07 13:13:33 918
aniqiaoensis, A. bellulus, A. chunganensus, A.
cremnobatus, A. gerbillus, A. granulosus, A. jinjiang-
ensis, A. liangshanensis, A. longimanus, A. monticola,
A. nasicus, and A. tormotus by the absence of
dorsolateral folds; from A. hainanensis, A. hong-
kongensis, A. marmoratus, A. nepalicus, A. ricketti, A.
spinapectoralis, A. tuberodepressus, and A. wuyiensis
by smooth skin lacking any tubercles; from A.
daiyunensis, A. larutensis, A. lifanensis, A. loloensis,
A. macrorhynchus, A. mantzorum, A. mengyangensis,
A. panhai, A. torrentis, and A. viridimaculatus by
the absence of a visible pineal body; and from A.
chakrataensis, A. jaunsari, A. kaulbacki, A. kangting-
ensis, and A. medogensis by the absence of trans-
verse bars on the hind limbs.
Amolops caelumnoctis also differs from the
several species of Huia, Hylarana, and Odorrana
from this region again by a smooth dorsum with
the unique pattern of small light yellow spots
interspersed on a dark purple background and
a lack of transverse bars on the hind limbs. All of
these species have yellow or olive to green or
brown dorsa with or without brown or black
spotting. All but two (O. leporipes and O. livida)
have transverse bars on the hind limbs; several
have dorsolateral folds and shagreened or
tuberculate dorsal skin.
Etymology.—The specific name is derived from
Latin for the sky of the night. The name refers to
the numerous tiny rounded irregularly arranged
light yellow spots on a dark purple dorsal
background resembling stars in the night sky.
MATERIAL EXAMINED
Amolops tuberodepressus. KIZ821090, KIZ821122–
821125, KIZ821130, China, Yunnan Province, Dali
Prefecture, Yangbi County; KIZ581719, China,
Yunnan Province, Simao Prefecture, Jingdong
County.
Amolops loloensis. KIZ820286, China, Sichuan
Province, Ya-an Prefecture, Shimian County,
altitude 2150 m; KIZ820263–820264, China, Si-
chuan Province, Ganzi Prefecture, Kangding
County, altitude 1550–1650 m.
Amolops viridimaculatus. KIZ75II0132, 0136,
0138, 0139, 0141, 1043, 0189, KIZ75II0343,
0356, KIZ75II0383, 0386–0388, 0390, 0391,
0393, and KIZ75III345, China, Yunnan Province,
Simao Prefecture, Jingdong County, Modaohe
and Dingpa, altitude 1800–2000 m; KIZ820339,
0341, 0343, 0344, 0355, 0356, 0357, 0358–0360,
China, Yunnan Province, Baoshan Prefecture,
Longling County; KIZ79I236, 373, China, Yun-
nan Province, Lincang Prefecture, Yongde Coun-
ty, altitude 1600–2100 m; KIZ74II0259–0268,
0270, 0271, China, Yunnan Province, Baoshan
Cust # CH-06-248R
 RAO AND WILKINSON—NEW SPECIES OF AMOLOPS
Prefecture, Tengchong County, Datang Town-
ship, altitude 2060 m.
ACKNOWLEDGMENTS
We appreciate the staff of the Huanglianshan
National Nature Reserve and the Forest De-
partment of Honghe Prefecture for providing
permission to collect specimens and help in the
field. We thank X. Ma who brought the initial
pictures of this species to D. Rao. This work was
supported by the Chinese Academy of Sciences,
Key Laboratory of Cell and Molecular Evolution
at KIZ, a National Science Foundation Grant
(DEB-0103795) to J. Slowinski and P. Fritsch, and
funds from the Lakeside Foundation of the
California Academy of Sciences to D. Rao. The
paper was completed while D. Rao was visiting
the Herpetology Department of the California
Academy of Sciences and profited from the
assistance of colleagues from that department.
We thank D. Lin for providing the photographs
for Figures 1 and 2 and L. Irving for providing
Figure 4. We also thank E. Muenk and E. Arnold
for assisting with the naming of the species and
J. Vindum and H. Brignall for critically reading
and editing the manuscript.
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mous Region, 2. Medog. Sichuan Journal of
Zoology 24:250–253.
(DQR) KUNMING INSTITUTE OF ZOOLOGY, CHINESE
ACADEMY OF SCIENCES, KUNMING, YUNNAN,
650223, CHINA ; AND (JAW) DEPARTMENT OF
HERPETOLOGY, CALIFORNIA ACADEMY OF SCIENCES,
875 HOWARD STREET, SAN FRANCISCO, CALIFORNIA
94103. E-mail: (DQR) raodq@mail.kiz.ac.cn;
and (JAW) jwilkinson@calacademy.org. Send
reprint requests to JAW. Submitted: 13 Oct.
2006. Accepted: 8 Feb. 2007. Section editor:
M. J. Lannoo.
Cust # CH-06-248R