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pp.85–92

A New Species of Rhacophorus (Anura: Ranidae) from China
YUN MING MO1, JIAN PING JIANG2,*, FENG XIE1 AND ANNEMARIE OHLER3
1Natural History Museum of Guangxi, Nanning 530012, China,
Chengdu Institute of Biology, The Chinese Academy of Science, Chengdu 610041, China,
3
Département Evolution et Systématique Muséum national d'Histoire naturelle, Paris 75005 France.
*Corresponding author E-mail: jiangjp@cib.ac.cn
2

Abstract.- A new brown tree frog species, genus Rhacophorus, is described on the basis of seven adult male specimens
collected from Cenwanglaoshan Nature Reserve, Guangxi, in southern China. This frog can be distinguished from allother Asian Rhacophorus Kuhl and van Hasselt, 1822 by the combination of: skin brown and smooth; Y-shaped cartilage visible dorsally on tips of fingers and toes; outer fingers one-third webbed; distinct dermal ridges present on
forearms, above vent, and calcars present on heels; anterior and posterior surface of thighs tangerine in color without
distinct dark or light spots; tympanum distinct and large, about 6.6% of SVL; dorsum brown with wide dark crossshaped mark.
Keywords.- Amphibia, Rhacophoridae, Rhacophorus, new species.

Introduction
The genus Rhacophorus Kuhl and van Hasselt, 1822, a
member of the family Rhacophoridae, contains approximately 60 species (sensu stricto) (Frost, 2007) that are
distributed in the tropical and temperate zones of East,
South, and Southeast Asia. Liem (1970) outlined a conservative definition of the genus (Rhacophorus sensu
stricto), and while some authors recognize the genus inthis sense (Jiang et al., 1987; Fei et al., 1990; Zhao andAdler, 1993; Inger et al., 1999; Malkmus et al., 2002;
Frost, 2007; Frost et al., 2006), others have adopted abroader definition (Rhacophorus sensu lato) that
includes the genus Polypedates (Tian and Jiang, 1986;
Dubois, 1987, 1992; Fei, 1999; Fei et al., 2005). In a
recent review of the Rhacophrinae, a new generic classification was proposed for the Rhacophorinae (Delorme
et al., 2005), where some members of Rhacophorus
were transferred to the new genus Aquixalus. This classification was adopted by Frost (2007) and Frost et al.
(2006) with some modification. In view of the various
interpretations of the Rhacophoridae, the classificationof the treefrogs should still be considered unstable(Wilkinson et al., 2002; Matsui and Orlov, 2004), and as
stressed by Frost et al. (2006), the boundaries of
Rhacophorus should be considered tentative.
All Chinese Rhacophorus (sensu lato) have beenfound in southern region north to Qinling, with most
species distributed in the tropical and subtropicalregions. Some new species were recently added to thegenus from the regions in and around southern China: R.
hainanus Zhao et al., 2005 from Hainan, R. minimus
Rao et al., 2006 from Guangxi, R. yinggelingensis Chouet al., 2007 from Hainan, and R. jarujini Matsui and© 2008 by Asiatic Herpetological Research

Panha, 2006 from eastern Thailand.
During the survey of the herpetofauna of
Cenwanglaoshan Nature Reserve, Guangxi, China, in
May 2004 and 2005, seven specimens of a small brown
Rhacophorus were collected (Fig. 1), which appeared tobe distinct from other congeners hitherto known from
China (Fei et al., 2005) and nearby countries (Bourret,1942), including Vietnam (Inger et al., 1999; Orlov etal., 2001), Thailand (Taylor, 1962) , Laos (Stuart, 1999),
India (Inger and Dutta, 1987), and Burma (Zug et al.,2003). These specimens, which resemble some southeastern Asia members of the genus, are described below
as a new species.

Materials and Methods
Morphological data.- The seven specimens included in
the new species were collected by hand in
Cenwanglaoshan Nature Reserve, Guangxi, China, in
May 2004 and 2005. Five were preserved in 10%
buffered formalin, and two were preserved in 70%
ethanol. Morphological information of related species
used for comparison was obtained from the literature
listed below.
Measurements.- Sixteen body measurements weremade using dial callipers to the nearest 0.1 mm: SVL =snout-vent length; HL = head length from tips of snout
to the commissure of the jaws; HW = head width at thecommissure of the jaws; SL = snout length from tip ofsnout to the anterior corner of the eye; INS = internarial
space; IOS = interorbital space, i.e., the smallest space
between the inner edge of upper eyelid; UEW = width of

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Figure 1. Localities recorded for the six species belonging to the Rhacophorus reinwardtii group in China, with thelocality recorded for R. hoanglinensis in Vietnam.

upper eyelid; ED = diameter of eye; TD = horizontal
diameter of tympanum; LAHL = length of lower arm
and hand; HAL = hand length; HLL = hindlimb length;
TL = tibia length; FTL = length of foot and tarsus; FL =foot length; and TFDD = third-finger disc transverse
diameter.
Analyses of advertisement calls.- We recorded theadvertisement calls of this new species using a
Panasonic SV-MP21V recorder (parameter set as22050Hz, 16 bit, monophone, wav file). Calls were analyzed using Cool Edit Pro V2.1 and BatSound V3.0.
Environmental parameters recorded during collection(2010–2340 h) were as follows: air temperature 22°C;
moisture 92%.
Museum acronyms.- CIB, Chengdu Institute of
Biology, Chinese Academy of Sciences; GXNM,
Natural History Museum of Guangxi, Nanning, China.

Taxonomy
Rhacophorus laoshan, new species
Fig. 2
Holotype.- Catalog number GXNM 2005081. Adult
male collected in Cenwanglaoshan Nature Reserve(Guangxi, China) on 19 May 2005 (106° 24′ 8.22′′ E,
24° 29′ 1.98′′ N) at 1389 m altitude by Yunming Mo.
Paratypes.- Six adult males, all from the same site as theholotype: CIB 2831k collected on 19 May 2004 by
Jianping Jiang, Annemarie Ohler, Yunming Mo, and
Feng
Xie;
GXNM2005079,
GXNM2005082,
GXNM2005095, CIB2005080, and CIB2005094 collected on 19–20 May 2005 by Yunming Mo.

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A.

B.

C.

D.

87

Figure 2. (A-D) Holotype of Rhacophorus laoshan sp. nov.; adult male GXNM2005081.

Diagnosis.- This new species Rhacophorus laoshan is asmall brown treefrog (SVL 35.1±1.3 mm; n = 7) that canbe distinguished from all other Asian Rhacophoridae by
the combination of the following characters: skin brownand smooth; Y-shaped cartilage visible dorsally on tipsof fingers and toes; outer two fingers one-third webbed;
dermal ridges present on forearm, above vent, and onheel calcars; anterior and posterior surfaces of thigh tangerine in color and without distinct dark or light spots;
tympanum large, about 6.6% of SVL; dorsum brownwith wide dark cross.
Description of holotype.- Body size small (SVL = 35.4mm) and moderately elongate. Head moderately compressed, dorsally flat, and wider (HW = 13.9 mm) thanlong (HL = 13.8 mm). Snout bluntly pointed, projecting,with length (SL = 6.4 mm) longer than horizontal diameter of eye (ED = 4.6 mm). Canthus rostralis distinct
with loreal region slightly concave. Nostril oval andcloser to tip of snout than eye; interorbital space almost
flat, and larger (IOS = 4.5 mm) than upper eyelid (UEW
= 3.9 mm) and internarial space (INS = 4.0 mm).
Tympanum rounded with diameter (TD = 2.3 mm) halfthat of eye; tympanum-eye distance about half tympanum diameter. Supratympanic fold distinct, present fromposterior corner of eye to above and behind insertion of

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arm. Vomerine ridges oblique, at an angle of 45° to bodyaxis, and closer to choanae than to each other. Vomerine
teeth on tongue pear-shaped, with deep notch on posterior end.
Forearm plus hand about half snout-vent length,with forearm (6.7 mm) shorter than hand (10.9 mm);forearm with distinct dermal ridge from elbow to wrist(Fig. 2); relative length of fingers: 3 > 4 > 2 > 1; fingers
with dermal fringes (Fig. 2); third and fourth fingers
with web on basal one-third; tips of all fingers with welldeveloped disks with distinct circummarginal grooves;discs relatively wide compared to fingers, except for
first finger; Y-shaped cartilage easily observable on
backs of fingers; subarticular tubercle developed; supernumerary tubercles below base of fingers distinct; inner
metacarpal distinct, large, flat, and oval.
Hind limbs rather long, with tibiotarsal articulation
reaching middle of eye when leg stretched forward;
heels strongly overlapping when limbs folded at rightangles to body; tibia (TL 18.0 mm) about half SVL andlonger than thigh (12.8 mm) and foot (FL 14.9 mm); relative length of toes: 4 > 3 = 5 > 2 > 1; tips of all toes withmoderately-sized disks (slightly smaller than those of
fingers except that of first finger); discs with distinct circummarginal grooves that are relatively wide compared
to toe width; Y-shaped cartilage easily observable on
backs of toes; web present, half developed, I 1 - 2 1/2 II 1
- 2 1/2 III 1 - 2 1/2 IV 2 - 1 V dermal fringe along toe V
distinct; subarticular tubercles prominent on all toes;
inner metatarsal tubercle distinct and oval; outer
metatarsal tubercle absent.
Dorsum smooth with granules scattered along sidesof body and head, along lower mandible, and back of
forearm; venter, head, and limbs covered with flat granules; outer edge of forearm and tarsus-metatarsus with
granulose ridge; dermal calcars present on heels; granules above vent forming transverse skin fold.
Color in life.- Dorsum chocolate brown; broad transverse strip present medially on upper lids and interorbital space; back with broad cross-shaped marking (Fig.
2); limbs with broad transverse stripes: 2 on forearms,
2–3 on carpals and fingers, 3 on thighs and tibiae, and
4–5 on feet and toes; anterior and posterior surfaces ofthighs tangerine in colour and usually without distinct
dark or light spots; inner surface of tarsus and foot tangerine; belly light gray-brown and without dark spots.
Male secondary sexual characters.- Adult males withnuptial pad on the base of first finger; internal subgular
vocal sacs present with two elongate openings; linea
masculina absent.
Variation.- Most variation was found in the appearance

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of the wide cross-shaped mark on the dorsum. Usually,
this mark was visible except for when the dorsum wasmore darkly-colored. The snout of some specimens was
green, and some specimens had a green spot on theshoulder; ovate yellow spots on the sides of the bodysometimes formed a line.
Etymology.- The species is named after the locality, i.e.,
Cenwanglaoshan Natural Preserve in northwestern
Guangxi, China, where it was found.
Measurements.- Sixteen body measurements are provided in Table 1.
Habitat and ecological notes.- This new species wasfound in a secondary broad-leaf forest with bamboo
undergrowth. Dense grass and deciduous leaves covered
the ground (Fig. 3). There were almost no perennial
streams in the region. During April and May, especiallyat night after rainfall, males were heard calling loudly inthe forest, with six to nine calls making up a chorus. The
holotype and paratypes were found on branches andleaves of trees one to three meters from the ground.
Advertisement calls.- The analyzed results of the callsby Cool Edit Pro 2.1 and BatSound indicated that calls
were emitted every 17–25 seconds and lasted about
3.6–4.6 seconds. The calls had 19-26 notes (6 individuals, 12 calls) (Fig. 4a), with a note interval of
0.171–0.267 seconds (mean ± SD: 0.189±0.0184, 6 individuals, 12 calls, 44 notes). The dominant frequency was2000 Hz and the second dominant frequency was 4000
Hz (Fig. 4b).
Comparisons.- The new species Rhacophorus laoshan
is most similar in appearance to R. hoanglienensis(Orlov et al., 2001) and R. verrucopus Huang, 1983. It
species can be distinguished from R. hoanglienensis by
a combination of the following characters: (1) anterior
and posterior surfaces of the thighs are orange-red andwithout distinct dark or light spots (black and white vermiculation is present in R. hoanglienensis); (2) bodysize smaller (SVL 35.1±1.3 mm; n = 7) than in R. hoanglienensis (SVL 43.2 mm; n = 1); (3) tympanum more
distinct and larger (TD = 2.3 mm, about 6.6% of SVL)
than that of R. hoanglienensis (TD = 2.1 mm, about
4.86% of SVL); (4) wide dark cross-shaped mark present on dorsum; (5) venter light gray-brown and without
spots (not creamy-white with small dark spots thatmerge in the distal parts of the fore and hind limbs).
Furthermore, the new species emits a call every 17–25
sec. (see above), while R. hoanglienensis calls every 3–5
min (Orlov et al., 2001), and white lines running fromthe supratympanic fold to tip of snout through eyelid and

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Table 1. Measurements (in mm) of Rhacophorus laoshan sp. nov. Variations (mean ± SD) are shown for paratypes andratios to SVL (%). Abbreviations as used in text.
Character
SVL
HL
HW
SL
INS
IOS
UEW
ED

Holotype

Paratypes (n = 6)

Character

Holotype

Paratypes (n=6)

35.4

33.2–36.7 (35.1±1.4)

TD

2.3

2.1–2.7 (2.3±0.2)-6.50%

6.6% (5.9–6.8)

13.8

12.2–13.4 (12.8±0.4)-39.00%36.5% (35.2–39.0)

13.9

12.8–13.5 (13.1±0.3)-39.30%37.3% (36.1-39.3)

6.4

5.5–6.2 (5.9±0.3)-17.10%16.7% (15.7–17.1)4

3.6–4 (3.8±0.1)-11.30%10.9% (10.4–11.7)

4.5

4.2–4.4 (4.3±0.1)-12.70%12.2% (11.7–13.3)

3.9

3.5–3.9 (3.7±0.2)-11.00%10.6% (10.5–11.0)

4.6

4.1–4.7 (4.5±0.2)-13.00%12.7% (12.3–13.0)

canthal ridge of female of R. hoanglienensis (Bain andTruong, 2004). This new species can be distinguishedfrom R. verrucopus by the latter lacking vocal sac, headlonger than width, outer fingers 1/2 webbed, the dermal
ridge on forearm and above vent more weak, and thedorsum without the wide dark cross mark present(Huang, 1983). Rhacophorus laoshan can be separated
from related species that also have dermal flaps on theforearms, tarsus, vent, or heel as follows. Rhacophoruslaoshan shows reduced toe webbing, which is more
extensive in R. annamensis, R. bipunctatus, R. exechopygus, and R. reinwardtii (Inger et al., 1999); complete webbing on the feet is seen in R. kio, R.
nigropalmatus, R. reinwardtii (Ohler and Delorme,
2006), R. maximus (Liu and Hu, 1961; Fei, 1999), R.
pardalis (Brown and Alcala, 1998; Inger, 1954, 1966;
Inger and Stuebing, 1997; Malkmus et al., 2002), R.
prominanus (Taylor, 1962; Inger, 1966), and R. robinsoni (Taylor, 1962; Inger, 1954) have. The absence of adark spot in temporal region distinguishes the new
species from R. cyanopunctatus (Malkmus et al., 2002);
and R. bipunctatus (Inger et al., 1999).
Rhacophorus baluensis can be separated from the
new species by being larger (male SVL = 50–55 mm)(Malkmus et al., 2002) and by having dark transversebars and irregular light or dark blotches on the dorsum(Inger and Stuebing, 1997; Malkmus et al., 2002).
Rhacophorus bimaculatus (Peters, 1867) is also larger insize (SVL = 65 mm) and has fingers that are almost fully
webbed (Fei, 1999). Almost fully-webbed fingers are

LAHL
HAL
HLL
TL
FTL
FL
TFDD

17.6

16.1–17.7 (17.1±0.5)-49.70%48.7% (47.1–51.2)

10.9

10.4–10.9 (10.6±0.2)-30.80%30.3% (29.4–31.9)

54.7

53.5–56.1 (54.7±1.1)-154.50%156.1% (154.0–169.0)

18

17.6–18.1 (17.9±0.2)-50.80%51.1% (49.3–53.9)

23.9

22.4–24.6 (23.7±0.7)-67.50%67.6% (63.1–72.3)

14.9

14.4–15.5 (15.1±0.4)-42.10%43.1% (41.4–45.5)

2.3

2.2–2.4 (2.3±0.1)-6.50%

6.6% (6.3–6.8)

also seen in R. dulitensis (Taylor, 1962; Inger andStuebing, 1997). Rhacophorus gauni can be separated
from the new species by having a conspicuous whitespot below the eye (Inger and Stuebing, 1997). The skinof R. kajau is leafy green dorsally and usually has
minute white spots scattered on the back, head, andexposed surface of the limbs (Inger and Stuebing, 1997;
Das, 2007). Rhacophorus rhodopus has more developed
finger and toe webbing, which is also red in color (Liuand Hu, 1959; Fei, 1999), not grey-brown. Rhacophorustranslineatus has dark transverse bars on dorsum(Sichuan Institute of Biology [Hu et al., 1977; Fei,1999]) while R. laoshan has a dark wide cross-shapedmark.
The new species can be distinguished from
Rhacophorus appendiculatus by having irregular low
ridges on the back, as well as a narrow flap on the heel
(Inger and Stuebing, 1997; Malkmus et al., 2002).
Rhacophorus calcaneus differs by having oval or rounddark brown spots on the dorsum, small enameled whitespots, or pinkish dorsolateral bands and a thin black network enclosing large white spots ventrolaterally (Ingeret al., 1999). Rhacophorus verrucosus has the vomerine
teeth absent (not present) and a low conical tubercle on
the heel (Inger et al., 1999). Rhacophorus bisacculus has
no prominent projections in the infra-anal area, but it
may have several short, pointed tubercles (Inger et al.,1999). Rhacophorus naso has a dorsolateral fold and askin projection on the tip of the snout (Fei, 1999).

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Frequency (kHz)

A.
4
3
2
1
0
0

B.

1

2

3
4
Time (seconds)

5

6

0

Intensity (dB)

100
800
60
40
20
0

Figure 3. Habitat of Rhacophorus laoshan sp. nov.

Remarks.- Those Rhacophorus with dermal flaps are
distributed in the tropical region of southeastern Asia,
extending north to the southern slopes of the eastern
Himalayas, the Hengduan Mountains, the YunnanGuizhou Plateau, and the Nanling Mountains. Within
this area, most species are distributed north of 20º N. Of
these taxa, R. bipunctatus, R. maximus, R. kio, R. hoanglienensis, R. laoshan, R, translineatus, and R. verrucopus, are distributed along the southern border of China(Fig. 1).
Rhacophorus bipunctatus (recorded as R. bimaculatus by Fei (1999) and Fei et al. (2005) in their list of
Chinese amphibians) has the widest Chinese distributionof these seven species (Fig. 1), and is also known from
Northeastern India, Myanmar, Thailand, Laos andVietnam. In China this species occupies the tropicalregion of the southern slopes of the Yunnan-Guizhou
Plateau, the Nanling Mountains, and southern Medog.
Rhacophorus maximus has and R. bipunctatus are
known from Yunnan and Tibet. The northern extent of R.
kio in China is the southern border regions of Yunnan
and Guangxi (Ohler and Delorme, 2006; Fei, 1999).
Rhacophorus translineatus and R. verrucopus are foundalong the southern slope of the eastern Himalayas in
Medog, while R. hoanglienensis and R. laoshan are
known from the southern slope of the Yunnan-Guizhou
Plateau.
In summary, there are only five Rhacophorus
species with dermal flaps along the southern slopes of
the Yunnan-Guizhou Plateau andthe Nanling
Mountains: R. maximus, R. kio, R. hoanglienensis, R.
laoshan, and R. bipunctatus. Of these, the new species

042Frequency (kHz)

Figure 4. Audio-spectrogram with FFT size 512 (A), andpower spectrum (B) of advertisement calls of the new
species Rhacophorus laoshan (Holotype)

R. laoshan is the most northern, R. bipunctatus the mosteastern and R. kio and R. hoanglienensis the most southern.
Lastly, only four rhacophorids were recorded in
Cenwanglaoshan Nature Reserve (Mo and Xie, 2005),
including the new species, suggesting that further surveys of the herpetofauna along the southern slopes of
Yunnan-Guizhou Plateau and the Nanling Mountains
should be carried out to further explore the little-known
biodiversity of this region. Such efforts may elucidate
more species or populations of Asian treefrogs or othernew species.

Acknowledgments
We are grateful to Prof. Fei Liang, Ye Changyaun (CIB),
and Prof. Masafumi Matsui (Kyoto University) and Dr.
R. Steven Wagner (Central Washington University) for
comments on the manuscript, Mr. Jian Li (CIB) for theline drawing, and Mr. Zhiming Xie (GXNM) for fieldassistance. This work was supported partially by NSFC
(No. 30000018, 30670245) and Life Science Special
Fund of Chinese Academy of Sciences (CAS) Supportedby the Ministry of Finance (STZ-01-19) to Jian-ping
Jiang.

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