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Accepted by S. Carranza: 12 Jan. 2011; published: 15 Feb. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2765: 21–37 (2011)
www.mapress.com/zootaxa/Article
21
A new cryptic rock-dwelling Hemidactylus (Squamata: Gekkonidae)
from south India
ISHAN AGARWAL1,4, VARAD B. GIRI2 & AARON M. BAUER3
1Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India. E-mail: ishan.agarwal@gmail.com
2Bombay Natural History Society, Hornbill House, S.B. Singh Road, Mumbai 400 023, Maharashtra, India.
E-mail:varadgiri@gmail.com
3Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085 USA.
E-mail: aaron.bauer@villanova.edu
4Corresponding author
Abstract
A new species of gecko, Hemidactylus graniticolus sp. nov. is described from Karnataka state, south India. This large-
sized (SVL to at least 110.6 mm), rupicolous gecko differs from congeners in having 16–18 longitudinal rows of fairly
regularly arranged, subtrihedral, weakly keeled, striated tubercles at midbody; 9–11 and 12–13 subdigital lamellae on the
first and fourth digits, respectively, of both manus and pes; tail with transverse series of four enlarged tubercles on each
tail segment; 23–28 femoral pores on each side separated by 1–3 poreless scales; 12–14 supralabials and 9–11 infralabials.
Molecular data support the distinctiveness of the new species and its affinities with large-bodied, tuberculate Hemidacty-
lus spp. from India and Sri Lanka.
Key words: Hemidactylus graniticolus sp. nov., H. maculatus, H. hunae, H. prashadi group, cryptic species, India
Introduction
Hemidactylus is the most species-rich Indian gekkonid genus with at least 25 described species (Giri and Bauer
2008; Mahony 2009; Bauer et al. 2010a). Though dominant across the subcontinent, Hemidactylus geckos remain
poorly known, with a number of recent descriptions and taxonomic works significantly altering the view of both
regional and local diversity (Zug et al. 2007; Giri 2008; Giri and Bauer 2008; Giri et al. 2009; Mahony 2009; Bauer
et al. 2010a,b).
Hemidactylus maculatus Duméril & Bibron is one of the largest geckos in the Indian subcontinent, and is
widely distributed, with confirmed reports from localities across the Western Ghats of Gujarat, Maharashtra, Ker-
ala and Tamil Nadu (Smith 1935; Tikader and Sharma 1992; Das 2002; Sharma 2002; personal observations).
There are stray reports of H. maculatus from Madhya Pradesh (Sanyal and Dasgupta 1990; Ingle 2003), and
Andhra Pradesh (McCann 1945). Both these localities lie outside the Western Ghats, in central and eastern India
respectively. There has been no detailed study on these specimens, and our observations on McCann’s specimen
from Vishakhapatnam, Andhra Pradesh show that it differs from true H. maculatus.
Deraniyagala (1937) recorded Hemidactylus maculatus from Sri Lanka based on a female specimen collected
from Okanda, Eastern Province. He described this as a new form, H. maculatus hunae Deraniyagala, designating
the Sri Lankan specimen as the type for this subspecies; with the material examined and assigned to this subspecies
including specimens from South India (Malabar, Tinnevelly and Salem). Deraniyagala (1937) restricted the nomi-
nate race H. m. maculatus to northern localities. More recently, Bauer et al. (2010a) recognized Hemidactylus
hunae as a distinct species, within the brookii group and sister to the Indian endemic H. prashadi Smith.
Based on our observations of specimens in the collection of the Natural History Museum, London, the material
referred to Hemidactylus maculatus hunae by Deraniyagala (1937) is composed of three different forms, which
include H. hunae, represented by the type specimen from Sri Lanka; typical H. maculatus, represented by three
AGARWAL ET AL.
22 · Zootaxa 2765 © 2011 Magnolia Press
specimens from Tinnevelly (Tirunelveli, Kerala) and a juvenile specimen from ‘Malabar’; and an unnamed taxon,
represented by six specimens from Salem and a specimen from ‘Malabar’.
The unnamed taxon is a cryptic species, morphologically very similar to H. hunae. Bansal and Karanth (2010)
included this species under the name H. maculatus in their recent phylogeny of Indian Hemidactylus, where it fell
within the H. prashadi group. We take this opportunity to describe this new species based on recently collected
material from near Bangalore, Karnataka, south India. We also present molecular data to support the distinctiveness
of the new species from closely related congeners, as well as to elucidate its phylogenetic position.
Material and methods
Morphological data. The following measurements were taken with an Aerospace digital caliper (to the nearest 0.1
mm): snout-vent length (SVL; from tip of snout to vent), trunk length (TRL; distance from axilla to groin measured
from posterior edge of forelimb insertion to anterior edge of hindlimb insertion), body width (BW; maximum width
of body), crus length (CL; from base of heel to knee); tail length (TL; from vent to tip of tail), tail width (TW; mea-
sured at widest point of tail); head length (HL; distance between retroarticular process of jaw and snout-tip), head
width (HW; maximum width of head), head height (HH; maximum height of head, from occiput to underside of
jaws), forearm length (FL; from base of palm to elbow); orbital diameter (OD; greatest diameter of orbit), nares to
eye distance (NE; distance between anteriormost point of eye and nostril), snout to eye distance (SE; distance
between anteriormost point of eye and tip of snout), eye to ear distance (EE; distance from anterior edge of ear
opening to posterior corner of eye), internarial distance (IN; distance between nares), interorbital distance (IO;
shortest distance between left and right supraciliary scale rows). Scale counts and external observations of mor-
phology were made using a Wild M5 dissecting microscope. Comparisons were made with representatives of most
South Asian species based on material in the collection of the Bombay Natural History Society, Mumbai (BNHS),
Natural History Museum, London (BMNH), Centre for Ecological Sciences, Bangalore (CES), and Aaron M.
Bauer field series (AMB; accession in National Museum of Sri Lanka, Colombo pending). Material examined of
the most similar species to the new form is listed in Appendix 1. Information on morphological characters of spe-
cies that could not be examined, as well as supplemental data for all Hemidactylus was obtained from the literature
(Deraniyagala 1937, 1953; Smith 1935; Giri and Bauer 2008; Mahony 2009; Somaweera and Somaweera 2009).
Supplementary information on life colouration and certain morphological details of Hemidactylus hunae were
based on photographs and data provided by R. Somaweera and M. Wickramasinghe.
Molecular data. As mentioned earlier, the new species, misidentified as Hemidactylus maculatus, was
included in a recent phylogeny of Indian Hemidactylus, falling in the H. prashadi group (Bansal and Karanth
2010). The H. prashadi group of Bansal and Karanth (2010) clearly corresponds to the subclade of “strictly South
Asian endemics” within the brookii group in the phylogeny of South Asian Hemidactylus presented by Bauer et al.
(2010a). Thus, besides sequences of the new species, we chose to include sequences of congeners that included all
the species from the H. prashadi group (Bansal and Karanth 2010) as well as species from the corresponding sub-
clade of the brookii group (Bauer et al. 2010a); with representatives of the other subclades of the Tropical Asian
clade (Bauer et al. 2010a) used to root the tree.
Sequences of the mitochondrial gene cyt b (239 bp) and the nuclear genes PDC (395 bp) and RAG1 (1044 bp)
were downloaded from Genbank (accession numbers listed in Appendix 2). Uncorrected sequence divergence was
calculated for the cyt b gene to assess genetic divergence of the new species from closely related congeners.
Downloaded sequences were aligned with default gap penalties using CLUSTALW (Thompson et al. 1994) in
the program MEGA 4 (Tamura et al. 2007), and protein coding genes were translated to amino acids in MEGA to
check the reading frame and for premature stop codons. Phylogenetic relationships were assessed with combined
data from PDC and RAG1 using parsimony and likelihood criteria. Both Maximum parsimony (MP) and Maxi-
mum Likelihood (ML) analyses were conducted in PAUP*4.0b10 (Swofford 2002). The Akaike Information Crite-
rion (AIC) in ModelTest 3.7 (Posada and Crandall 1998) was used to find the model of evolution that best fit the
data for ML analyses. ML and MP trees were constructed through heuristic searches with the following conditions:
100 random addition replicates, tree bisection-reconnection (TBR) branch swapping, zero-length branches col-
lapsed to yield polytomies, and gaps treated as missing data. Clade support in the resulting trees was assessed by
nonparametric bootstrap (1000 pseudoreplicates) with 100 random addition replicates per pseudoreplicate for the
MP trees and 10 random addition replicates for the ML trees.
Zootaxa 2765 © 2011 Magnolia Press · 23
NEW CRYPTIC INDIAN HEMIDACTYLUS
Systematics
Hemidactylus graniticolus sp. nov.
Figs. 1–7
Holotype. Bombay Natural History Society (BNHS) 1850, adult female; collected from hills near Harohalli, Ban-
galore Rural District, Karnataka, India (12°40’57.97’’N, 77°29’21.30’’E, 913 m asl) on 9 June 2008. Collected by
I. Agarwal, V. Mistry, N. Page and P. Chanchani.
Paratypes. BNHS 1859, adult male, BNHS 1858, adult female and BNHS 1860, juvenile female, collected
along with holotype; BNHS 1826, adult female, collected from adjacent hill on 11 November 2007 by I. Agarwal
and V. Mistry. BNHS 2016, adult female, collected from near Yercaud, Salem District, Tamil Nadu (11° 45’ 3.02’’
N 78° 11’ 18.78’’ E, 606 m asl) on 5 December 2009 by I. Agarwal and A. D. Roy; BMNH 1946.8.23.70–71 and
BMNH 1946.8.23.73–75, four adult females, and BMNH 1946.8.23.72, adult male, collected from Salem District,
Tamil Nadu, India by R. H. Beddome; BMNH 1946.8.23.76, adult male, collected from ‘Malabar’, India by R. H.
Beddome.
Diagnosis. A large sized Hemidactylus, snout-vent to at least 110.6 mm. Dorsal pholidosis heterogeneous, with
16–18 fairly regularly arranged longitudinal rows of subtrihedral, striated tubercles at midbody (Fig. 1). First
supralabial in broad contact with nasal. Two well-developed pairs of postmentals, the inner pair longer than the
outer pair and mental, and in broad contact behind the mental. Ventrolateral folds distinct, about 40–46 scale rows
across venter. 12–13 enlarged, divided scansors beneath fourth digit and 9–10 (rarely 11) beneath first digit of both
manus and pes. 23–28 femoral pores on each side separated by one to three poreless scales in males. Original tail
depressed, oval in transverse section with a median dorsal furrow; scales on the tail slightly larger than dorsals of
body, striated, imbricate, with a longitudinal series of two enlarged, weakly keeled, striated, flattened tubercles on
either side of the median dorsal furrow. Body dorsum with a series of pale saddles from occiput to sacrum, tail with
distinct alternating light and dark bands.
Hemidactylus graniticolus sp. nov. may be distinguished from all other Indian and Sri Lankan congeners on
the basis of (taxa with differing or non-overlapping character states indicated parenthetically): dorsum with coni-
cal, granular, striated scales intermixed with enlarged, fairly regularly arranged, longitudinal rows of 16–18 sub-tri-
hedral, weakly keeled, striated tubercles (dorsum with small granules usually intermixed with scattered, rounded,
feebly keeled or conical tubercles in H. frenatus Schlegel; enlarged dorsal tubercles usually few or sometimes
absent in H. leschenaultii Duméril & Bibron; dorsum with very few enlarged tubercles, more often absent alto-
gether in H. flaviviridis; small, uniform, granular dorsal scales except along the sides where they may form a single
line of larger rounded tubercles in H. garnotii Duméril & Bibron; no enlarged dorsal tubercles in H. anamallensis
(Günther), H. giganteus Stoliczka, H. platyurus (Schneider) and H. aquilonius McMahan & Zug; dorsum with uni-
form, imbricate, scales in H. scabriceps (Annandale) and H. imbricatus (Bauer, Giri, Greenbaum, Jackman, Dharne
& Shouche), 23–28 femoral pores on each side in males (9–13 precloacal pores in H. persicus Anderson; angular
series of 6 precloacal pores in H. porbandarensis Sharma; 6 precloacal pores in H. gracilis Blanford; 6-12 precloa-
cal pores in H. reticulatus Beddome; 7–10 precloacal pores in H. albofasciatus Grandison & Soman; and 4–6 pre-
cloacal pores in H. sataraensis Giri & Bauer, 7–16 femoral pores on each side in H. brookii Gray and
parvimaculatus Deraniyagala; 6–14 femoral pores on each side in H. lankae Deraniyagala, H. subtriedrus Jerdon
and H. triedrus (Daudin); 14 femoral pores in H. treutleri Mahony; 12–14 femoral pores on each side in H. gujarat-
ensis Giri, Bauer, Vyas & Patil; 16–19 femoral pores on each side in H. depressus Gray).
Hemidactylus graniticolus sp. nov. may also be distinguished from all the above mentioned species on the
basis of its large size (adult SVL to at least 110.6 mm). The only four Indian and Sri Lanka congeners that grow
beyond 100 mm SVL are H. aaronbaueri Giri, H. prashadi, H. maculatus and H. hunae.
H. aaronbaueri can be distinguished from Hemidactylus graniticolus sp. nov. by the presence of 15–19 femo-
ral pores on each side separated by 6 poreless scales in males (vs. 23–28 femoral pores on each side separated by
1–3 poreless scales in males) and 18–20 rows of irregularly arranged, enlarged, rounded and feebly keeled tuber-
cles on dorsum (vs. 16–18 rows of fairly regularly arranged sub-trihedral, weakly keeled, striated tubercles on dor-
sum). H. prashadi can be distinguished from the new species by the presence of 17–20 femoral pores separated by
3 poreless scales in H. prashadi (vs. 23–28 femoral pores on each side separated by 1–3 poreless scales in males)
and lamellae beneath first toe 8 and fourth toe 10 (vs. lamellae beneath first toe 9–11 and fourth toe 12–13).
AGARWAL ET AL.
24 · Zootaxa 2765 © 2011 Magnolia Press
FIGURE 1. Dorsal view of holotype (BNHS 1850, adult female) of Hemidactylus graniticolus sp. nov.
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NEW CRYPTIC INDIAN HEMIDACTYLUS
FIGURE 2. Details of dorsal pholidosis. A) Hemidactylus graniticolus sp. nov. holotype BNHS 1850, B) H. hunae type
BMNH 1946.8.23.77, C) H. maculatus BNHS 1836.
FIGURE 3. Dorsal view of tail. A) Hemidactylus graniticolus sp. nov. holotype BNHS 1850, B) H. hunae type BMNH
1946.8.23.77, C) H. maculatus BNHS 1836.
The new species is similar in size and general appearance to Hemidactylus maculatus, but differs with respect
to (H. maculatus versus H. graniticolus sp. nov.): dorsal pholidosis (back with small juxtaposed, conical, granular
scales and large trihedral tubercles arranged in 20 fairly regular longitudinal rows versus back with conical, granu-
lar, striated scales intermixed with enlarged, fairly regularly arranged, longitudinal rows of 16–18 sub-trihedral,
weakly keeled, striated tubercles (Fig. 2)); dorsal pholidosis of tail (small, irregular, more or less pointed, keeled
scales and a series of six or eight large, keeled, trihedral tubercles versus small, imbricate, striated scales and a
series of four enlarged, keeled and weakly striated and flattened tubercles (Fig. 3)); femoral pores in males (16–19
femoral pores on each side with a gap of 5 to 9 scales versus 23–28 femoral pores on each side separated by 1–3
scales (Fig. 4)); colouration (brown above with darker spots, dorsal pattern with a series of dark, transverse undu-
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26 · Zootaxa 2765 © 2011 Magnolia Press
lating crossbars versus dorsal pattern with a series of alternating broad pale saddle-markings and narrower, darker,
interspaces; no scattered dark spots.
The new species most closely resembles the Sri Lankan Hemidactylus hunae, but differs with respect to (H.
hunae versus H. graniticolus sp. nov.): dorsal pholidosis of tail (a series of six large, keeled, pointed/recurved
tubercles, dorsolateral rows largest versus a series of four enlarged, keeled, weakly striated and flattened tubercles,
dorsolateral row absent; femoral pores in males (22–24 femoral pores on each side with a gap of 3–6 scales versus
23–28 femoral pores on each side separated by 1–3 scales).
Description. The holotype is generally in good condition with some minor exceptions (Fig. 1). The body shape
is somewhat dorsoventrally flattened. A fold of skin is present on the right side of the gular region, running from
below the eye to the axilla, an artefact of preservation. The tail is partially regenerated and the tail tip is missing
(stored for phylogenetic analysis in the personal collection of IA).
FIGURE 4. Pericloacal region of adult male Hemidactylus graniticolus sp. nov. (BNHS 1859) showing a series of 25 femoral
pores on left and 27 on right side separated mesially by one scale.
FIGURE 5. Head of Hemidactylus graniticolus sp. nov. holotype, BNHS 1850. A) Dorsal view, B) Lateral view, C) Ventral
view.
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NEW CRYPTIC INDIAN HEMIDACTYLUS
FIGURE 6. A) Ventral view of right manus of Hemidactylus graniticolus sp. nov. holotype, BNHS 1850, B) Ventral view of
right pes of H. graniticolus sp. nov. holotype, BNHS 1850.
Head short (HL/SVL ratio 0.28), slightly elongate (HW/HL ratio 0.71), not strongly depressed (HH/HL ratio
0.41), distinct from neck (Fig. 5 A). Loreal region slightly inflated, canthus rostralis not prominent. Snout short
(SE/HL ratio 0.43); slightly longer than eye diameter (OD/SE ratio 0.52); scales on snout, canthus rostralis, fore-
head and between eyes homogenous, juxtaposed, and weakly pointed; scales on snout, canthus rostralis and fore-
head twice the size of those on the occipital and interorbital region, canthus rostralis with slightly enlarged patch of
scales (Fig. 5 B). Eye small (OD/HL ratio 0.22); pupil vertical with crenulated margins; supraciliaries small,
pointed, those at the anterior end of orbit slightly larger. Ear opening oval (greatest diameter 2.5 mm); eye to ear
distance slightly greater than diameter of eye (EE/OD ratio 1.22). Rostral wider (4.1 mm) than deep (2.3 mm),
incompletely divided dorsally by weakly developed rostral groove; two internasals, enlarged, in contact anteriorly,
posteriorly separated by a single scale, one supranasal on each side which is smaller than internasal, one pair of still
smaller postnasals; rostral in contact with nostril, supralabial I, and internasal; nostrils large (1.1 mm), circular,
each surrounded by supranasal, internasal, rostral, supralabial I and postnasal; 3–4 rows of scales separate orbit
from supralabials. Mental triangular, two well developed postmentals, the inner pair slightly longer (4.1 mm) than
mental (3.8 mm), and in extensive contact with each other (2.5 mm) behind mental, outer pair about half the size of
the inner pair, separated from each other by inner pair (Fig. 5 C). Inner postmental bordered by mental, infralabial
I, outer postmental and three gular scales; outer postmental bordered by infralabials I and II, inner postmental, and
6 gular scales of which the outer 2 are enlarged and continue as a single row of enlarged scales below infralabials.
Infralabials bordered by a single row of enlarged scales, about 2 to 8 rows of scales below infralabials III to VIII
are enlarged and weakly imbricate. Supralabials (to midorbital position) 9 (right) – 9 (left); supralabials (to angle of
jaw) 13 (right) – 12 (left); infralabials (to angle of jaw) 9 (right) – 9 (left).
Body relatively stout, not elongate (TRL/SVL ratio 0.40), with ventrolateral folds without denticulate scales.
Dorsal pholidosis heterogeneous, composed of conical, granular, striated scales intermixed with enlarged, fairly
regularly arranged, longitudinal rows of 16–18 subtrihedral, weakly keeled, striated tubercles at midbody, extend-
ing from occipital region to tail, each enlarged tubercle roughly two to three times longer than adjacent granules,
surrounded by rosette of 10–13 small granules, 2–5 granules between two adjacent enlarged tubercles; enlarged
tubercles on back smallest on two most medial parasagittal rows, increasing in size toward the flanks, the last row
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FIGURE 7. Photographs of Hemidactylus graniticolus sp. nov. in life. A) Adult colouration, light phase (holotype BNHS
1850, adult female), B) Adult colouration, dark phase (specimen not collected), C) juvenile colouration (paratype BNHS 1860,
juvenile female). Note the prominent, contrasting patterning in the juvenile, which fades in adult specimens.
Zootaxa 2765 © 2011 Magnolia Press · 29
NEW CRYPTIC INDIAN HEMIDACTYLUS
on flank smallest after medial rows; enlarged tubercles more strongly keeled and slightly larger on flanks and close
to the tail than on the dorsum; enlarged tubercles on nape, shoulder small and pointed, those on occipital, temporal
region still smaller, strongly pointed. Ventral scales larger than dorsal, smooth, imbricate, slightly larger on preclo-
acal and femoral region than on chest and abdominal region; midbody scale rows across belly 42–46; gular region
with still smaller, sub-imbricate scales, those on lateral aspect of neck granular, anterior gular scales slightly larger
than the rest.
Scales on the palm and sole smooth, imbricate, rounded; scales on dorsal aspect of upper arm larger than gran-
ules on dorsum, subimbricate and striated, dorsal aspect of forearm with smaller, striated, conical and granular
scales, intermixed with a few enlarged conical tubercles; those on dorsal part of thigh and shank conical, granular,
striated, intermixed with enlarged, striated, sub trihedral tubercles, which are numerous on shank compared to ante-
rior aspect of thigh, posterior aspect of thigh lacks enlarged tubercles.
Fore- and hind limbs relatively short, stout; forearm short (FL/SVL ratio 0.14); tibia short (CL/SVL ratio
0.14); digits moderately short, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; ter-
minal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than
half as long as associated toepad; scansors beneath each toe in straight transverse series, divided except for distal
and three to four basal scansors on digit I and one or two in all digits that are single; scansors from proximal most
at least twice diameter of palmar scales to distalmost single scansor: 10-13-13-13-13 (right manus; Fig. 6 A), 9-13-
13-13-13 (right pes; Fig. 6 B). Relative length of digits (measurements in mm in parentheses): III (8.9) > IV (8.8) =
V (8.8) > II (8.7) > I (7.0) (right manus); II (10.0) > III (9.9) > V (9.6) > IV (9.4) > I (7.6) (right pes).
Tail depressed, flat beneath, verticillate, with well defined median furrow; length of the partially regenerated
and slightly broken tail less than snout-vent length (TL/SVL ratio 0.73); tail covered above with small (slightly
larger than those on the dorsal granules), posteriorly-pointed, imbricate, striated scales and a series of two enlarged,
keeled and weakly striated, posteriorly pointed and flattened tubercles on either side of the median furrow (Fig. 3
A); ventral scales larger, imbricate, median row (subcaudal plates) slightly broader, about twice as broad as adja-
cent scales, not extending across width of the tail proximally, but distally they extend almost across the width of the
tail. Coloration (in preservative). Dorsum grayish- brown with a vague, pale beige vertebral stripe and a series of
narrow, wavy, mid-brown transverse markings defining the edges of somewhat paler saddles (indistinct saddle on
occiput, one across shoulders, two between fore- and hindlimb insertions, and one on sacrum). Dorsolateral tuber-
cle rows mostly dark brown, those on flanks predominantly ashy to white. Crown of head with scattered vague
mid-brown markings. A cream stripe, bordered above and below by a narrow (one scale wide) dark brown margin,
extending from nostril to midorbital rim, through eye, and on to temporal region, confluent with posterior edge of
pale occipital marking. Infralabials and posterior supralabials pale, anterior supralabials with scattered dark pig-
ment, anterior three scales darkest. Limbs pale, yellowish-gray with vague irregular dark markings, more distinctly
alternating with pale interspaces distally, especially on metapodial and phalangeal segments. Original tail mostly
gray-brown with lighter mottling; distal portion of original tail with alternating vague lighter and darker bands.
Regenerated portion of tail unmarked. Venter cream with scattered patches of diffusely pigmented scales, particu-
larly under thighs; palms and soles grayish. Ventrolateral surfaces of tail darkly pigmented, with scattered speck-
ling on proximal portions of midventral scutes.
Coloration (in life). In life dorsal markings much more evident, especially in light phase, although vertebral
stripe not visible. Saddle-shaped markings ashy with dark brown edges. Interspaces between saddles with diffuse
orangey-brown highlights. Tail distinctly banded, with alternating light and dark bands similar to trunk dorsum in
colour; darker bands somewhat wider than pale bands. Mottled pattern of limb bases and banding of distal limbs
clearly demarcated. In dark phase dorsal colours much less contrasting; paler saddles light brown, darker inter-
spaces a dark, slightly reddish-brown (Fig. 7 A). Iris marbled greenish-silver suffused with brown-rust veination
that is most prominent near the pupil.
Etymology. The species is named for the conspicuous granite rock formations upon which it lives. The specific
epithet is a masculine adjective.
Variation and additional information from type series. Mensural data for the type series and additional
material is given in Table 1. There is one male, and four females including a juvenile, ranging in size from 39.2 mm
to 110.6 mm. All paratypes resemble the holotype except as follows: internasals separated by two (BNHS 1858) or
three (BNHS 1826, 1859, 1860) scales. Range of supralabials is from 12–14 (9–10 below eye) and infralabials is
from 9–11. The scales across belly also range from 40–46 in the paratypes. Males have a series of 23–28 femoral
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30 · Zootaxa 2765 © 2011 Magnolia Press
pores separated mesially by 1-3 poreless scales (BNHS 1859 – 25/27 pores on left/right side separated by a single
poreless scale, BMNH 1946.8.23.72 – 27/28 pores separated by 2 scales, BMNH 1946.8.23.76 – 25/23 pores sepa-
rated by 3 scales).
Colouration. Adult colouration variation primarily due to relative prominence of the dorsal pattern elements.
Light and dark phases physiological so may be seen in the same individual (Fig. 7 A, B). Juvenile pattern very
bold, brighter than in adults, with very sharp edges to dorsal markings (Fig. 7 C). Light markings on head, edges of
dorsal pale saddles, and lateral tubercles pale yellowish. Darker portions of trunk and limbs purplish-brown. Crown
distinctly lighter than nape and temporal region. Tail very boldly banded with alternating dark chocolate and ashy
(proximal) to white (distal) bands; 10 pale bands from tail base to tip. Venter in dark phase heavily mottled with
brown.
TABLE 1. Measurements (mm) and scale counts for type series of Hemidactylus graniticolus sp. nov. Abbreviations as in
Materials and Methods. * indicates tail is incomplete, - measurement not taken.
Holotype Paratypes
BNHS 1850 BNHS 1826 BNHS 1858 BNHS 1859 BNHS 2016
Sex F F F M F
SVL 103.8 82.5 101.5 108.5 81.0
TRL 41.4 37.1 42.6 46.6 34.5
BW 23.4 19.6 25.5 23.5 -
CL 14.9 14.3 17.2 17.8 13.1
TL 75.5* 67.2* 105.6 107.8 65.0*
TW 13.5 10.2 13.6 14.6 -
HL 29.0 23.3 27.9 30.6 22.8
HW 20.7 16.8 20.2 21.2 16.8
HH 11.9 8.7 11.9 12.5 10.3
FL 15.0 11.6 14.9 16.0 11.6
OD 6.4 5.2 6.2 5.9 -
NE 9.7 8.1 9.8 10.4 7.6
SE 12.4 9.9 12.1 12.6 9.8
EE 7.8 6.2 7.3 8.2 6.6
EL 2.5 2.4 2.8 3.3 -
IN 2.9 2.5 3.0 2.9 2.4
IO 8.7 8.5 10.7 9.8 9
Lamellae
L Manus 9-11-12-12-11 10-12-12-12-12 11-12-12-14-13 10-12-13-13-12 10-10-12-12-13
R Manus 9-12-12-12-12 10-12-12-12-12 11-12-12-13-13 10-10-13-13-13 10-12-12-12-13
L Pes 9-14-13-12-12 9-13-13-13-12 10-13-13-13-13 10-13-13-13-13 10-13-13-13-13
R Pes 9-13-13-12-12 9-13-13-12-13 10-13-13-13-12 10-13-13-13-13 10-13-12-13-12
Supralabials
L 12 (9) 14 (10) 12 (9) 12 (10) 13 (9)
R 13 (9) 13 (9) 14 (10) 12 (9) 12 (8)
Infralabials
L91010911
R 1010101010
Zootaxa 2765 © 2011 Magnolia Press · 31
NEW CRYPTIC INDIAN HEMIDACTYLUS
continued.
Distribution. Hemidactylus graniticolus is known based on voucher specimens from the following localities
(Fig. 8), in Karnataka state: Bangalore Rural District (Harohalli), Ramnagaram District (Ramnagaram); and Tamil
Nadu state: Nilgiri District (Masanagudi-Ooty road), Salem District (near Yercaud). This species has also been
visually observed in the following localities (Fig. 8), in Andhra Pradesh: Chittoor District (Kangundi); Karnataka:
Mysore District (Chamundi Hills, Kollegal); and Tamil Nadu: Vellore District (near Vellore), Villupuram District
(near Gingee).
Natural history. The type locality is a rocky hillock in the outskirts of Bangalore, Karanataka, India (Fig. 9).
These broken hills are a conspicuous feature of this landscape. Hemidactylus graniticolus appears to be strictly
rupicolous, recorded only from boulders. The boulder habitats of this species are both in hills and rocky outcrops in
the plains; localities around Bangalore lie on the Mysore plateau and may be above 1000 m asl while localities fur-
ther south such as in Salem District are as low as 200 m. These areas are characterized by sparse scrub vegetation
and few trees or occasionally deciduous forests. The average annual rainfall at the type locality is about 889 mm
(Hegde et al. 2008), and the altitude is 830 m.
Paratypes
1946.8.23.70 1946.8.23.71 1946.8.23.72 1946.8.23.73 1946.8.23.74 1946.8.23.75 1946.8.23.76
Sex F F M F F F M
SVL 101.5 88.1 76.5 86 93.5 94 99.5
TRL 47.5 36 31.6 34.8 38.5 44.9 44.1
BW 20 16.7 16.4 18.2 19.1 20 24.7
CL 16.9 14.9 12.3 14.8 15.6 14.7 16.2
TL 90* 90* 85 90* *- 75* *-
TW 11.3 8.6 8.2 7.8 *- 9.5 *-
HL 30.1 25.6 22.5 25.1 28.1 27.4 29.7
HW 22.1 18.1 16.4 18 20.4 19.3 22.9
HH 12.1 10.9 9 10.5 12 10.9 13.5
FL 14.6 14.3 11.4 12.8 13.3 13.9 13.5
OD 7.1 5.6 4.1 5.7 6.2 5.8 5.6
NE 10.5 8.2 8 9.1 9.5 9.9 9.8
SE 12.9 11.5 9.1 11.4 12 12.6 12.7
EE 7.5 6.5 6.5 6.8 7.9 8 8.7
EL ----- - -
IN 2.9 2.8 2.6 2.7 2.6 2.6 3
IO 8.7 7.1 6.6 6.8 8.4 6.9 8.5
Lamellae
L Manus 10-13-13-13-13 10-12-13-13-13 10-12-12-12-13 11-13-13-13-13 10-13-12-13-13 10-12-13-13-13 12-12-12-13-13
R Manus 10-13-13-13-13 10-11 -13-12-14 10-12-13-12-12 11-12-13-13-13 10-12-12-12-13 10-13-13-14-13 11-13-14-12-13
L Pes 10-13-12-13-13 10-13-13-13-13 9-13-13-13-14 10-13-13-12-13 10-13-13-13-13 10-15-14-13-13 10-14-13-12-13
R Pes 10-13-13-13-13 10-13-14-14-13 9-13-13-13-13 10-13-13-13-13 9-13-13-13-12 10-14-13-13-13 10-12-13-13-13
Supralabials
L 13 (11) 10 (9) 10 (11) 12 (10) 10 (8) 12 (10) 11 (10)
R 11 (9) 10 (8) 10 (11) 13 (10) 12 (10) 12 (10) 11 (10)
Infralabials
L 9 9109 81010
R98101091010
AGARWAL ET AL.
32 · Zootaxa 2765 © 2011 Magnolia Press
FIGURE 8. The known distribution of Hemidactylus graniticolus sp. nov. in South India: 1, near Harohalli, Karnataka (type
locality); 2, Ramnagaram, Karnataka; 3, Nilgiri District, Tamil Nadu; 4, near Salem, Tamil Nadu; 5, Kangundi, Andhra
Pradesh; 6, Chamundi Hills, Karnataka; 7, near Kollegal, Karnataka; 8, near Vellore, Tamil Nadu, and 9, near Gingee, Tamil
Nadu. Triangles indicate localities with voucher specimens, squares represent visual observations.
The species is active after dark on rock faces and occasionally culverts, in the day they may be found in crev-
ices among boulders. These fast-moving geckos are relatively abundant and widespread within their range, found
on even isolated outcrops. H. graniticolus is the most commonly seen gecko at the type locality. These large geckos
have fragile skin that is easily damaged by handling. Juveniles have been observed in June.
Sympatric congeners at the type locality include Hemidactylus cf. brookii, H. frenatus, and H. triedrus. The
ground-dwelling H. reticulatus occurs in the same habitats at various other localities; and the large rupicolous H.
giganteus is sympatric with H. graniticolus at Ramnagaram (Karnataka) and Kangundi (Andhra Pradesh).
Discussion
The phylogenetic relationships of Indian and South Asian Hemidactylus spp. have been elucidated by recent
molecular studies (Bansal and Karanth 2010; Bauer et al. 2010a,b). Based on morphological affinities, Hemidacty-
lus graniticolus is likely to be a member of the broad brookii group (Bauer et al. 2010a; = Indian clade/Tropical
Asian clade 2 of Bansal and Karanth 2010), and more specifically the H. prashadi group (Bansal and Karanth
Zootaxa 2765 © 2011 Magnolia Press · 33
NEW CRYPTIC INDIAN HEMIDACTYLUS
2010; = subclade of “strictly South Asian endemics” Bauer et al.2010a). In fact, Bansal and Karanth (2010) used
sequences of this new species under the name H. maculatus (from Karnataka, specimen number CES/07/005; and
Tamil Nadu CES/08/026), and these fell in the H. prashadi group. Data from the cyt b gene reveals that Hemidac-
tylus graniticolus is genetically divergent from other members of the H. prashadi group (Bansal and Karanth 2010)
and members of the subclade of “strictly South Asian endemics” within the brookii group (Bauer et al. 2010a) as
well as H. aaronbaueri, the only other large, superficially similar species, with an uncorrected sequence divergence
of 14–22% (Table 2). The two specimens of Hemidactylus graniticolus are similar, with 5.0% sequence diver-
gence.
TABLE 2. Uncorrected pairwise sequence divergence (%) for the cyt b gene (239 bp) between Hemidactylus graniticolus sp.
nov. (n=2) and closely related congeners (H. depressus (n=2), H. hunae (n=1), H. lankae (n=1), H. maculatus (n=1), H. pra-
shadi (n=5), H. triedrus (n=5) and H. aaronbaueri (n=2). Ranges are presented, details of specimens used in the analysis listed
in Table A2. The divergence between both H. graniticolus samples was 5.0 %.
FIGURE 9. Type locality of Hemidactylus graniticolus sp. nov. near Harohalli, Bangalore Rural District, Karnataka, India.
The holotype (BNHS 1850) and paratypes (BNHS 1858–1860) were collected among the boulders in this habitat.
Species Sequence divergence
Hemidactylus aaronbaueri 19.1–23.1
Hemidactylus depressus 17.1–18.1
Hemidactylus hunae 17.6–19.1
Hemidactylus lankae 15.6–17.1
Hemidactylus maculatus 14.1
Hemidactylus prashadi 13.6–15.6
Hemidactylus triedrus 16.1–22.1
AGARWAL ET AL.
34 · Zootaxa 2765 © 2011 Magnolia Press
FIGURE 10. Maximum-likelihood phylogram of species of the H. prashadi group based on combined nuclear data (PDC +
RAG1, 1439 bp). Numbers adjacent to nodes indicate parsimony and likelihood bootstrap support values respectively.
The MP and ML phylogenies derived from PDC and RAG1 data (Fig. 10) strongly support the monophyly of
the H. prashadi group (parsimony, likelihood bootstrap support, BS, 99%), which includes all the large-bodied
Indian and Sri Lankan Hemidactylus species that have trihedral or subtrihedral dorsal tubercles – H. depressus, H.
graniticolus, H. hunae, H. lankae, H. maculatus, H. prashadi and H. triedrus (Fig. 10). Within this clade, H. gra-
niticolus forms a polytomy (BS 87, 75%) with H. prashadi, H. hunae, and (H. maculatus (H. triedrus, H. lankae),
with H. depressus basal to this unresolved group.
Hemidactylus graniticolus shows close morphological affinities to H. hunae from Sri Lanka, and they are
likely to be sister species, although cyt b divergence between the species was substantial and these two taxa are no
more similar than other members of the H. prashadi group (Table 2). Both species are large-bodied and tuberculate,
and are known to live among granite boulders. While H. graniticolus appears fairly widespread across semi-arid
areas of Southern India with the appropriate boulder habitat, H. hunae is restricted to the dry zone of Uva and East-
ern provinces in southeast Sri Lanka (Somaweera and Somaweera 2009). Until 2010, these two species had been
lumped together as H. maculatus hunae, a subspecies of H. maculatus (Bauer et al. 2010a). H. maculatus, however,
inhabits relatively wet areas across the Western Ghats from Gujarat, south to Tamil Nadu.
Recent species descriptions have revealed previously undocumented diversity in Indian Hemidactylus geckos
(Giri 2008; Giri and Bauer 2008; Giri et al. 2009; Mahony 2009). While many of these descriptions were of com-
pletely new forms, the description of this new species and a cryptic species from within the H. brookii complex
(Mahony 2009), as well as high genetic divergence between species from India and Sri Lanka that were previously
considered conspecific (Bauer et al. 2010a, b) is indicative of undocumented cryptic diversity within many of the
widely distributed Hemidactylus species in India.
Zootaxa 2765 © 2011 Magnolia Press · 35
NEW CRYPTIC INDIAN HEMIDACTYLUS
Acknowledgments
At BNHS we are thankful to A. Rahmani, Director and A. Kothari, Honorary Secretary for supporting this
research, V. Hegde for curatorial assistance and K. Gaikwad for assistance with data collection. At CES, P. Karanth
and R. Bansal provided data and access to specimens, J. Joshi provided useful comments and help with the map
and phylogenetic analyses, and A. Datta-Roy provided additional distributional and natural history data. V. Mistry
photographed the specimens, provided inputs on the draft and helped collect the first specimen. N. Page, A.
Krishna, P. Chanchani, S. Khan assisted in collecting specimens. M. Wickramasinghe and R. Somaweera provided
data and photographs of H. hunae. VG’s visit to the BMNH, London was supported by a Scholarship and Travel
Award from the Society of Systematic Biologists and a grant from the BMNH. VG thanks M. Wilkinson, D. Gower
and C. McCarthy. AMB was supported by grants DEB 0515909 and DEB 0844523 from the National Science
Foundation of the United States.
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APPENDIX 1. Material examined.
Institutional abbreviations are given in materials and methods. Old BNHM numbers are listed in Table A1.
Hemidactylus aaronbaueri. Holotpye BNHS 1739 (male), near Ghatghar, District Pune, Maharashtra, India.
Hemidactylus hunae: Type specimen BMNH 1946.8.23.77 (female), Okanda, Eastern Province, Sri Lanka; AMB 7416, Pita-
kumbura, Sri Lanka (07°13’22”N, 81°18’25”E).
Hemidactylus maculatus. BNHS 1184 and BNHS 1903 (males), Matheran, District Raigad, Maharashtra, India; BNHS 1255
(male), Malabar, Kerala; BNHS 1026 (male), Poombarai, Kodaikanal, Tamil Nadu, India; BNHS 846 (male), Courtallam,
Tamil Nadu, India. BNHS 648 (female), Ahwa Dangs, Gujarat, India; BMNH 1946.8.23.67 (female),BMNH 1946.8.23.68
(male), BMNH 1946.8.23.69 (female), Tinnevelly, Tamil Nadu, India; BMNH 1946.8.20.3 (juvenile female), ‘Malabar’,
India.
Hemidactylus prashadi. BNHS 1540 (male), BNHS 1577 (male) and BNHS 1491-1494 (females), Amboli, Sindhudurg Dis-
trict, Maharashtra, India; BNHS 146 (one male and two females) and BNHS 146/2 (female), Gersoppa Falls, North Kanara
District, Karnataka, India.
Hemidactylus graniticolus. Holotype BNHS 1850 (female), near Harohalli, Bangalore Rural District, Karnataka, India. Para-
types BNHS 1859 (male), BNHS 1826 (female), BNHS 1858 (female), BNHS 1860 (juvenile female), near Harohalli,
Bangalore Rural District, Karnataka, India; BNHS 2016 (female), near Yercaud, Salem District, Tamil Nadu, India;
BMNH 1946.8.23.70 (female), BMNH 1946.8.23.71 (female), BMNH 1946.8.23.72 (male), BMNH 1946.8.23.73
(female), BMNH 1946.8.23.74 (female), BMNH 1946.8.23.75 (female), Salem District, Tamil Nadu, India; BMNH
1946.8.23.76 (male), ‘Malabar’, India. Additional material: BNHS 1861 (male), near Harohalli, Bangalore Rural District,
Karnataka, India, CES/07/005 (tail only), CES/07/041 (male), Ramnagaram District, Karnataka, India; CES/08/026
(female), Nilgiri Hills, Nilgiri District,Tamil Nadu, India.
Hemidactylus sp. BNHS 89 (female), Lamarsinghe, Vishakhapatnam District, Andhra Pradesh, India.
TABLE A1. New BNHM specimen numbers and corresponding old BNHM numbers cited in Deraniyagala (1937)
Old number New number
1936.6.7.7 1946.8.23.177
74.4.29.1183–1188 1946.8.23.70–75
74.4.29.1049–1051 1946.8.23.67–69
83.9.5.1 1946.8.23.76
74.4.29.1411 1946.8.20.3
Zootaxa 2765 © 2011 Magnolia Press · 37
NEW CRYPTIC INDIAN HEMIDACTYLUS
APPENDIX 2. List of samples used for genetic comparisons
TABLE A2. List of specimens corresponding to sequences used for genetic comparisons with museum numbers, locality and
Genbank accession numbers.
Sample Museum No. Locality Genbank Accession Numbers
cyt bRAG1 PDC
Hemidactylus aaronbaueri CES/08/022 Pune, near Ghatgar, Maha-
rashtra, India HM595640 - -
Hemidactylus aaronbaueri CES/08/016 Peth Fort, Raigadh Dist.,
Maharashtra, India HM595641 - -
Hemidactylus brookii CAS 229632 Tanintharyi Division, Myan-
mar - GQ375313 GQ375307
Hemidactylus depressus 1 AMB 7481 Matale, Sri Lanka HM559593 HM559691 HM559658
Hemidactylus depressus 2 AMB 7524 Galle, Sri Lanka HM559594 HM559692 HM559659
Hemidactylus frenatus AMB 7420 Rathegala, Sri Lanka - EU268298 EU268328
Hemidactylus garnotii CAS 223286 Taung Gok Township, Rakh-
ine State, Myanmar - EU268302 EU268332
Hemidactylus giganteus CES/08/013 Hampi, Karnataka, India - HM622357 HM622372
Hemidactylus graniticolus
1CES/08/026 Nilgiri Hills, Tamil Nadu,
India HM595664 HM622361 HM622375
Hemidactylus graniticolus
2CES/07/005 Ramnagar, Karnataka, India HM595663 - -
Hemidactylus hunae AMB 7416 Pitakumbura, Sri Lanka HM559606 HM559706 HM559673
Hemidactylus lankae AMB 7453 near Medavachchiya, Sri
Lanka HM559615 HM559714 HM559681
Hemidactylus maculatus BNHS 1516 Zirad, Raigad Dist.,Maha-
rashtra, India HM559607 HM559707 HM559674
Hemidactylus prashadi 1 JB 30 India (captive specimen) HM559609 HM559709 HM559676
Hemidactylus prashadi 2 JB 02 India (captive specimen) HM559608 HM559708 HM559675
Hemidactylus prashadi 3 CES/07/040 Castle Rock, Karnataka,
India HM595668 HM622364 HM622378
Hemidactylus prashadi CES/07/037 Ratnagiri, Maharashtra, India HM595666 - -
Hemidactylus prashadi CES/06/170 Seeta Nadhi, Udupi, Karnat-
aka, India HM595667 - -
Hemidactylus triedrus 1 JB 08 Pakistan (captive specimen) HM559617 HM559716 HM559683
Hemidactylus triedrus 2 JB 09 India (captive specimen) HM559616 HM559715 HM559682
Hemidactylus triedrus 3 CES/07/007 Ramnagar, Karnataka, India HM595673 HM622365 HM622379
Hemidactylus triedrus CES/07/011 Devanahalli, Karnataka,
India HM595674 - -
Hemidactylus triedrus CES/07/023 Atigulipura, Talvadi, Karnat-
aka, India HM595675 - -
