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ACKNOWLEDGEMENTS. We thank Director, WIHG, Dehradun for
technical and administrative support, Director, BSIP, Lucknow for 14C
dating of the samples; the DST for financial support under Kangra pro-
ject. We acknowledge the support provided by Prof. Vinod Nautiyal,
Prof. R. C. Bhatt, Dr P. M. Saklani, Dr Y. S. Farswan and the support
staff of the Department of History, Ancient Indian History and Culture
and Archaeology of the H.N.B. Garhwal University, Srinagar for ar-
chaeological excavation and study. Dr N. R. Phadatare provided useful
Received 19 March 2009; revised accepted 22 March 2010
A ground-dwelling rhacophorid frog
from the highest mountain peak of
the Western Ghats of India
S. D. Biju1,*, Yogesh Shouche2, Alain Dubois3,
S. K. Dutta4 and Franky Bossuyt5,*
1Systematics Lab, Department of Environmental Studies, University of
Delhi, Delhi 110 007, India (e-mail: firstname.lastname@example.org)
2National Centre for Cell Science, Pune University Campus,
Pune 411 007, India (e-mail: email@example.com)
3Reptiles et Amphibiens, UMR 7205 OSEB, Département de
Systématique and Evolution, Muséum National d’Histoire Naturelle,
CP 30, 25 rue Cuvier, 75005 Paris, France (e-mail: firstname.lastname@example.org)
4Department of Zoology, North Orissa University, Takatpur,
Baripada 400 023, India (e-mail: email@example.com)
5Amphibian Evolution Lab, Biology Department, Unit of Ecology and
Systematics, Vrije Universiteit Brussel, Pleinlaan 2, B-1050 Brussels,
Belgium (e-mail: firstname.lastname@example.org)
A new rhacophorid frog is described from Eravikulam
National Park in the Western Ghats of India. The
species is morphologically dissimilar from any known
member of this family in having a bright orange to
reddish colouration, multiple macroglands on the body
and extremely short limbs. Phylogenetic analyses of
mitochondrial genes indicate that this new frog is nested
in a radiation of shrubfrogs that had its origin on the
Indian subcontinent, and which is here recognized as
a distinct genus, Raorchestes gen. nov. The new species,
Raorchestes resplendens sp. nov. is likely restricted to
less than 3 sq. km on the summit of Anamudi, and
deserves immediate conservation priority.
Keywords: Endotrophic development, macroglands,
mitochondrial phylogeny, Raorchestes gen. nov., Raor-
chestes resplendens sp. nov., Western Ghats.
THE Rhacophoridae constitute a radiation of about 300
species of frogs1 with a distribution in the tropical regions
of Asia and Africa. A large part of their taxonomy has
traditionally been based on the variety of reproductive
strategies, such as the use of foam nests or development
without going through a free-swimming tadpole stage.
Molecular evidence has recently solved the phylogenetic
position of many rhacophorid species2–7 and has resulted
in the erection of new genera, such as Gracixalus3, Fei-
hyla4, Liuixalus5 and Ghatixalus6. Although rhacophorids
find breeding microhabitats from the soil to the forest
canopy, calling males are often found in vegetation
layers8, and the family as a whole is therefore usually
considered arboreal. Here we describe an exceptional
glandular rhacophorid with extremely short limbs from
Anamudi summit (2695 m asl) in the Western Ghats of
India. We performed molecular analyses to infer its phylo-
genetic position within the family and studied its repro-
duction and development.
CURRENT SCIENCE, VOL. 98, NO. 8, 25 APRIL 2010
We constructed a data matrix of 3370 bp mtDNA con-
taining a part each of 12S rRNA, tRNAVAL, 16S rRNA,
ND1, tRNAILE, tRNAGLN and tRNAMET genes for a variety
of rhacophorid frogs to infer the phylogenetic position of
the new species within the family (GenBank accession
number GU808563 for BNHS 5301). Maximum parsi-
mony (MP) and maximum likelihood (ML) analyses were
executed with the software PAUP* 4.0b109, Bayesian
analyses and calculation of posterior probabilities were
performed with MrBayes10. Our results largely agree with
previous studies6,11,12 and show that frogs traditionally
classified in the genus Philautus1 do not form a mono-
phyletic group. Consistent with recent analyses5,12, they
form a clade with a main distribution on the Sunda shelf
(Philautus sensu stricto), and a group with its centre of
diversification on the Indian subcontinent. MP, ML and
Bayesian analyses all place the new species in the latter
clade of direct developing frogs, a group that in itself
consists of two reciprocally monophyletic clades (Figure
1, left): the largest part of direct developing frogs form a
distinct radiation with a notably large diversity in the
Western Ghats. Its sister clade is a mainly Sri Lankan
radiation, which has known a single dispersal event to the
Indian mainland, resulting in few extant species in the
Western Ghats11. Although the name Kirtixalus has been
proposed for the latter clade13, the older name Pseudo-
philautus Laurent 1943 is available for this island radia-
tion14,15 and its nested Indian members11. The type
species of Pseudophilautus, Ixalus temporalis Günther
1864, is considered to be extinct16, but clearly is a mem-
ber of the Sri Lankan radiation of direct developing frogs.
Pseudophilautus can be defined in a phylogenetic frame-
work as the most inclusive clade including the species
Ixalus temporalis Günther 1864 and excluding the species
I. glandulosus Jerdon 1854. To distinguish species of the
mainland radiation, we here suggest recognizing this
remarkable and fairly large radiation (Figure 1, left) as a
Raorchestes gen. nov. Type species, Ixalus glandulosus
Etymology: The generic epithet is derived from the name
‘Rao’ and the scientific nomen ‘Orchestes’. The former is
chosen in honour of C. R. Narayan Rao, in recognition of
his contribution to Indian batrachology. The latter is
based on the first generic nomen coined for frogs of the
Philautus group, Orchestes Tschudi 1838.
Diagnosis: We distinguish different kinds of taxog-
noses17 and define the new genus according to two of
them: Idiognosis (brief characterization of a taxon through
characters): relatively small frogs, adult snout-vent length
between 15 and 45 mm; vomerine teeth absent; large gular
pouch transparent while calling; largely nocturnal species;
direct development without free-swimming tadpoles in all
species for which the development is known. Entexognosis
(characterization of a taxon in a phylogenetic framework,
through inclusion and exclusion of taxa or organisms): the
most inclusive clade including Ixalus glandulosus Jerdon
1854 and excluding I. temporalis Günther 1864.
Distribution: Most species are from the Western
Ghats of India, but recent molecular phylogenetic analyses
supporting a nested position for ‘Philautus’ menglaensis,
longchuanensis and gryllus in this clade12,15 indicate that
the range of Raorchestes extends at least to southern
China, Laos and Vietnam1. Raorchestes is absent from
Figure 1. Left, Maximum likelihood tree based on a 3370 bp data matrix of mtDNA, showing the nested phylogenetic position of Raorchestes
resplendens in the genus. Right, Maximum likelihood tree for known Raorchestes species (all except R. ochlandrae) of the Western Ghats, based
on a 1488 bp data matrix of mtDNA. Black circles indicate Bayesian posterior probabilities above 95%.
CURRENT SCIENCE, VOL. 98, NO. 8, 25 APRIL 2010 1121
Table 1. Morphometric measurements (all in mm; range, mean and standard deviation) of Raorchestes resplendens sp. nov. Abbreviations are
defined in the text
Male (N = 5) Female (N = 2)
BNHS BNHS BNHS BNHS ZSIC BNHS BNHS
4087 5301 5302 5303 A9140 Range Mean SD 5304 5305 Range Mean SD
SVL 24.0 24.5 22.7 24.0 24.5 22.7–24.5 23.9 0.7 28.3 25.2 25.2–28.3 26.8 2.2
HW 9.4 9.6 8.6 9.2 9.5 8.6–9.6 9.3 0.4 10.6 10.2 10.2–10.6 10.4 0.3
HL 9.1 8.9 8.1 8.7 9.1 8.1–9.1 8.8 0.4 10.1 9.8 9.8–10.1 10.0 0.2
SL 3.0 3.3 3.0 3.4 3.4 3.0–3.4 3.2 0.2 3.6 3.6 3.6 3.6 0.0
EL 2.4 2.6 2.7 2.6 2.6 2.4–2.7 2.6 0.1 2.7 2.8 2.7–2.8 2.6 0.1
IUE 2.7 3.1 2.9 2.9 2.8 2.7–3.1 2.9 0.1 3.3 3.4 3.3–3.4 3.4 0.1
UEW 1.6 1.3 1.3 1.5 2.1 1.3–2.1 1.6 0.3 1.3 1.3 1.3 1.3 0.0
FLL 4.8 5.0 4.7 4.6 5.0 4.6–5.0 4.8 0.2 5.4 5.3 5.3–5.4 5.4 0.1
HAL 6.0 6.3 5.8 6.6 6.3 5.8–6.6 6.2 0.3 6.9 6.6 6.6–6.9 6.8 0.2
ShL 8.3 8.2 7.8 8.2 8.3 7.8–8.3 8.2 0.2 9.8 8.7 8.7–9.8 9.3 0.8
TL 8.8 9.1 8.3 9.0 9.4 8.3–9.4 8.9 0.4 10.6 9.1 9.1–10.6 9.9 1.1
TFOL 12.9 12.6 11.8 12.8 13.2 11.8–13.2 12.7 0.5 14.7 12.4 12.4–14.7 13.6 1.6
FOL 8.5 9.1 7.0 8.5 8.9 7.0–9.1 8.4 0.8 1.1 9.1 9.1–10.1 9.6 0.7
Figure 2. Raorchestes resplendens sp. nov. holotype. a, Dorsal view;
b, Ventral view.
Raorchestes resplendens sp. nov. Holotype: BNHS 4087
(Figures 2, 3a left animal), an adult male collected by
SDB at an altitude of 2695 m asl, at Anamudi summit,
10°10′N 77°05′E, Idukki District, Eravikulam National
Park, Kerala, India. This specimen is kept in the Bombay
Natural History Society (BNHS), Mumbai, Maharashtra,
India. Paratypes: four adult males, BNHS 5301, 5302,
5303 and ZSIC A9140 (Kolkata, West Bengal, India);
two adult females, BNHS 5304 and 5305. All were col-
lected along with the holotype (Table 1).
Etymology: The species epithet ‘resplendens’ is a
Latin term meaning ‘bright-coloured, glittering, resplen-
dent’. It refers to the bright red or orange colouration.
Suggested vernacular name: ‘resplendent shrubfrog’.
Diagnosis: Raorchestes resplendens can be distin-
guished from all members of the genus by its multiple
prominent glandular swellings; laterally behind the eyes,
on the side of the dorsum, on the anterior side of the vent,
on the dorsal side of the forearms and shanks, and on the
posterior side of tarsus and metatarsus (Figures 2 and 3).
Additionally, the colour of the iris is invariably bright red
(Figure 3) and the legs are extremely short (Figure 4).
Phylogeny: To infer a more precise phylogenetic po-
sition of R. resplendens, we constructed a 1488 bp data
matrix covering part of the 16S rRNA, part of the ND1,
the complete tRNAILE and tRNAGLN, and part of the
tRNAMET genes for all known extant species (except for
R. ochlandrae) from the Western Ghats2. MP, ML and
Bayesian are in agreement with previous analyses2. Addi-
tionally, R. resplendens comes out as a close relative of
R. beddomii, R. munnarensis and R. dubois (Figure 1,
right). The low support for relationships within this clade
however indicates that either of the latter three species
may be the sister taxon of R. resplendens.
Description: The description (all measurements in
mm) of the holotype follows terminology used else-
where2,18–20. Medium sized frog (SVL 24); head slightly
wider than long (HL 9.1; HW 9.4; MN 7.4; MFE 6.6;
MBE 4.1); outline of snout in dorsal view nearly
rounded, its length (SL 3.0) larger than the horizontal di-
ameter of the eye (EL 2.4); canthus rostralis indistinct;
loreal region vertically acute; distance between posterior
margins of eyes (IBE 8.0) 2.1 times in distance between
anterior margins of eyes (IFE 3.8); interorbital area (IUE
2.7) larger than upper eyelid (UEW 1.6); tympanum (TYD
1.3) rather distinct, rounded; vomerine ridge absent;
CURRENT SCIENCE, VOL. 98, NO. 8, 25 APRIL 2010
Figure 3. Raorchestes resplendens sp. nov. in life. a, Holotype (BNHS 4087, left-hand side); paratype (BNHS
5302, right-hand side); b, Paratype (BNHS 5304), profile; c, Paratype (BNHS 5304), ventral side, showing granu-
Figure 4. Plot of relative thigh length (x-axis, TL/SVL) versus rela-
tive shank length (y-axis, ShL/SVL) for 200 males of Indian Raor-
chestes and Pseudophilautus species. The plot shows a wide ecomorph
diversity, likely corresponding to different microhabitat preferences.
The extremely short legs of Raorchestes resplendens reflect its terres-
tongue cordate, without a papilla; supratympanic fold
from posterior corner of upper eyelid to base of forelimb.
Forelimb length (FLL 4.8) shorter than hand (HAL 6.0);
tips of fingers with discs, with distinct circummarginal
grooves; webbing on fingers absent; fingers without lat-
eral dermal fringe; subarticular tubercle indistinct, pre-
pollex oval; supernumerary tubercles absent, nuptial pads
absent. Hind limbs moderately short; shank about 3 times
longer (ShL 8.3) than wide (ShW 2.6), shorter than thigh
length (TL 8.8), slightly shorter than distance from base
of inner metatarsal tubercle to tip of toe IV (FOL 8.5;
TFOL 12.9); tips of toes with discs and distinct circum-
marginal grooves; webbing reduced, reaching just below
the 2nd subarticular tubercle on inside of toe IV and up to
the 2nd subarticular tubercle on outside of toe IV; dermal
fringe along toe V absent; subarticular tubercles rather
indistinct; supernumerary tubercle absent; inner metatar-
sal tubercle indistinct.
Skin of snout smooth; between eyes shagreened; upper
eyelids sparsely granular; anterior and posterior part of
back and flanks with symmetrical glandular swellings: a
kidney-shaped gland behind eye, a rather long and wide
latero-dorsal gland all along upper part of flank, a small
rounded gland on anterior side of vent; dorsal parts of
limbs with glandular swellings: a long gland all along
dorsal part of forelimbs (from elbow to base of hand); a
rather long gland on dorsal part of shank, on posterior
CURRENT SCIENCE, VOL. 98, NO. 8, 25 APRIL 2010 1123
Figure 5. a–g, Raorchestes resplendens developmental stages in life. a, Eggs after 3 h; b, neural fold at 30 h; c, head, limb bud
and tail differentiation at 80 h and beginning of embryo rotation; d, hind limb differentiation at 140 h; e, elongation of toe and
forelimb differentiation at 230 h; f, froglet with reduced tail and mouth differentiated into upper and lower lips at 380 h and tail
and yolk considerably reduced in this stage; g, fully mobile froglets measuring SVL 4.5 ± 0.7 mm emerged after 540 h (about 22
side of tarsus, and on posterior side of metatarsus of toe
V; a few less prominent glands throughout the dorsal and
lateral parts of the body (Figure 3a, left); ventral side of
throat slightly granular, belly coarsely granular (oval to
hexagonal); thigh and lower arm sparsely granular.
Colour of holotype: In life (Figure 3a, left): The pre-
dominant colour of dorsum is deep brick red, which
sharply contrasts with the black filling irregular furrows
on the dorsum; the dorsal parts of limbs, fingers, toes and
tips are uniform deep brick red; glandular swellings are
surrounded with black patches; posterior lateral abdomi-
nal area, groin and inner side of the thighs are black; ven-
tral side light yellow, with irregular oval to hexagonal
granulation; iris bright red. In alcohol, the dominant brick
red colour changed into brownish grey which contrasts
with the black furrows; lateral side light yellowish; limbs
are reddish grey; ventral side light greyish white.
Breeding and development: Breeding observations
were conducted in the natural habitat without handling
the animals. During a late afternoon (16 h) in July, a pair
of R. resplendens in amplexus was located on the ground
in bamboo thickets. Initial amplexus was inguinal for a
period of about 3 h during which time the female was al-
most motionless. After that, the amplexus became axil-
lary, the female started to search for a suitable oviposition
site, and at 20 h, eggs were burrowed under the moss
covered forest floor, deep inside the base of bamboo
clumps. The male and female left the nest immediately
after completion of spawning. Several other clutches
were located within a radius of 7 m, and all were at the
base of bamboo clumps (number of eggs: 18–28, N = 7).
The female, male and egg clutches were collected and
developmental stages were observed under laboratory
conditions with a temperature difference from natural
conditions of maximum 3°C. As in other species of this
genus, R. resplendens undergoes direct development, i.e.
complete development in the eggs, without a free-
swimming larval stage. After oviposition (eggs: N = 24,
diameter 4.1 ± 0.4 mm), the female’s oviduct still con-
tained mature embryos (N = 18, diameter 3.8 ± 0.5 mm).
This indirectly suggests that the female may mate with
multiple males and/or may breed more than once in a single
season. The developmental time from embryo to hatch-
ling varied from 22 to 27 days (N = 7). Descriptions of
embryo morphology are based on observation of living
and preserved material. The major features of eight stages
are illustrated in Figure 5 a–g.
Comments on morphology: The combination of a
unique morphology and nested phylogenetic position of
the new species points to a quick morphological diver-
gence from congeners. The most striking character of R.
resplendens is the presence of numerous macroglands.
These are absent in all species of Philautus, Pseudo-
philautus and Raorchestes presently known and thus are
derived characters. The presence, distribution and size of
the macroglands shows no sexual dimorphism, so we
hypothesize that the primary function will not be
involved in reproduction, but this needs further testing.
The quick morphological evolution is also visible when
plotting relative limb length of Pseudophilautus and
Raorchestes species (Figure 4). These measurements in-
dicate that the new species has much shorter limbs than
CURRENT SCIENCE, VOL. 98, NO. 8, 25 APRIL 2010
Figure 6. Map showing the type locality of Raorchestes resplendens at Anamudi, in the Western Ghats. Map
modified from Inger et al.21. Shaded contours showing above 300 m asl (light) and above 1000 m asl (dark).
its congeners and members of the sister clade, pointing to
a rapid morphological evolution that is dissociated from
molecular evolution. These short limbs have resulted in a
much more pronounced crawling behaviour in this spe-
cies compared to its congeners.
Comments on conservation concern: R. resplendens
is known only from the type locality (Figure 6). The
highest temperature reported for the Anamudi summit is
30°C in May 1980 and the lowest is –3°C in June 1980
(ref. 22). The vegetation, climate and temperature of the
hills are different from other regions of the Western
Ghats, and share several floristic elements with Himala-
yas22–24. Most importantly, despite intensive searches in
suitable habitats close to the type locality, we were
unable to locate this species in any other place. It is there-
fore possible that the entire range of this species does not
exceed a few square kilometres. We suggest immediate
action to make this highly endangered species a top prio-
rity for conservation.
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ACKNOWLEDGEMENTS. We thank Kim Roelants, R. F. Inger,
Rohan Pethiyagoda, I. Das, Stephen Mahony and Rachunliu G. Kamei
for critical review of an early draft of the manuscript. SDB is supported
by a research grant from Department of Biotechnology, Govt of India
(DBT) and University of Delhi (Scheme to strengthen R&D doctoral
research programme). S.D.B. thanks T. M. Manoharan, Principal Chief
Conservator of Forests, Kerala for his constant source of inspiration
and financial support, and the Kerala Forest Department for giving
necessary permission to survey.
Received 17 September 2009; revised accepted 22 March 2010