Hemidactylus varadgirii, Chaitanya & Agarwal & Lajmi & Khandekar, 2019

Chaitanya, R., Agarwal, Ishan, Lajmi, Aparna & Khandekar, Akshay, 2019, A novel member of the Hemidactylus brookii complex (Squamata: Gekkonidae) from the Western Ghats of Maharashtra, India, Zootaxa 4646 (2), pp. 236-250 : 239-246

Hemidactylus varadgirii sp. nov.

Figures 3–6 ; Tables 1–3 .

Holotype. BNHS 2377 , adult male, from Amboli (15.9590° N 73.9970° E; ca. 740 m asl), Maharashtra state, India, collected by RC and Varad Giri on 19 May 2018.

Paratypes. BNHS 2375 , BNHS 2378 , adult males, BNHS 2374 , BNHS 2376 adult females, collected on 15 February 2019, other collection details same as those of holotype.

Etymology. The specific epithet is a Latinized patronym in honour of Varad Giri for mentoring the authors in gecko taxonomy. Varad has made sizeable and significant contributions to the taxonomy of reptiles in India and has also been working tirelessly for over 15 years towards the conservation of the Amboli landscape, the type locality of this species.

Suggested Common Name. Giri’s ‘brookiish’ gecko or Amboli ‘ brookiish ’ gecko.

Diagnosis. A medium sized Hemidactylus , SVL to at least 62 mm (n=5). Dorsal pholidosis composed of granular scales that are heterogeneous in size and shape throughout, intermixed with enlarged, conical, keeled tubercles that are heterogeneous in size, arranged in 17 or 18 fairly regular longitudinal rows at midbody, extending from posterior part of the head to groin; tubercles in parasagittal rows are smaller, keeled and more rounded; approximately six rows of tubercles on either side of parasagittal tubercle rows are highly enlarged, remarkably conical and strongly keeled; ventrolateral folds indistinct; about 30–34 scale rows across the venter. Supralabials up to jaw and midorbital position, 11–13 and eight or nine respectively; 8–11 infralabials. Digits with enlarged scansors, lamellae in straight transverse series, all divided except the apical and a few basal, seven or eight (manus & pes) lamellae beneath fourth digit and five or six (manus & pes) beneath first digit. Males with nine or ten femoral pores (rarely eight) on each side separated by four or five poreless scales. Ventral scales on tail enlarged, imbricate, with a single median row of roughly rectangular, enlarged subcaudal plates covering almost entire tail base, including regenerated tails.

Comparison with Indian congeners from the brookii group. Hemidactylus varadgirii sp. nov. differs from all other Hemidactylus based on phylogenetic position ( Figure 1 ) and is therefore compared herein only with members of the brookii group. It is 11.2% –16.3% divergent based on uncorrected ND2 sequence divergence from all other members of the H. murrayi clade ( Table 2 ). It can be diagnosed from the ground-dwelling clade within the brookii group by the presence of divided sub-digital lamellae versus undivided or notched sub-digital lamellae in all species in the ground-dwelling clade. Further, it can be diagnosed from the ground-dwelling members based on the enlarged, remarkably conical, strongly keeled tubercles on the dorsum (as opposed to): enlarged tubercles absent from dorsum in H. imbricatus Bauer, Giri, Greenbaum, Jackman, Dharne & Shouche, 2008 ; tubercles on dorsum only slightly larger than surrounding granular scales in H. albofasciatus Grandison & Soman, 1963 , H. sataraensis Giri & Bauer, 2008 and H. gracilis Blanford,1870 ; tubercles on dorsum rounded, only slightly larger than surrounding granules in H. reticulatus Beddome, 1870 and H. vijayaraghavani Mirza, 2018 . The other scansorial congeners from the brookii group can be diagnosed from H. varadgirii sp. nov. based on the following characters: 9 or 10 femoral pores (rarely 8) on each side separated by 4–6 poreless scales, and seven or eight lamellae under digit IV of pes (conflicting characters in parentheses): H. brookii sensu stricto (12 or 13 femoral pores separated by a single poreless scale), H. chipkali (seven femoral pores on each side separated by eight poreless scales), H. gleadowi (12 or 13 femoral pores separated by a single poreless scale), H. kushmorensis (femoral pores separated by two or three poreless scales, ten lamellae under digit IV of pes), H. malcolmsmithi (femoral pores separated medially by 1–3 poreless scales, 9–11 lamellae beneath digit IV of pes), H. murrayi Gleadow, 1887 (6–8 femoral pores, never more than eight, separated by 5–7 poreless scales), H. parvimaculatus (11–17 femoral pores separated by 1–3 poreless scales), H. treutleri (seven femoral pores on each side separated by seven poreless scales), H. sankariensis (15 femoral pores on each side) and H. chikhaldaraensis (six or seven femoral pores on either side separated by seven poreless scales).

Description of holotype. The holotype is in good condition except for a small vertical incision on the venter (7.1 mm) made to extract tissue, slightly curved towards the right, a strong fold of skin between forelimb and hindlimb insertions on the left flank, hemipenis on the right, partially everted—all artefacts of preservation, final third of tail regenerated. Adult male, SVL 55.4 mm. Head short (HL/ SVL 0.28), slightly elongate ( HW /HL 0.66), slightly depressed (HH/HL 0.36), distinct from neck ( Figure 3A ). Loreal region slightly inflated, canthus rostralis indistinct ( Figure 4C ). Snout short ( SE /HL 0.43); slightly shorter than eye diameter ( ED / SE 0.47); scales on snout, canthus rostralis, forehead and inter-orbital region heterogeneous, mostly granular and conical; scales on the snout and canthus rostralis much larger than those on occipital, forehead and inter-orbital regions ( Figure 4A ). Eye small ( ED /HL 0.20); pupil vertical with crenulated margins; supraciliaries small, mucronate, gradually decreasing in size posteriorly. Ear opening roughly circular (greatest diameter 1.3 mm); eye to ear distance greater than diameter of eye ( EE / ED 1.42). Rostral much wider than deep ( RD / RW 0.44), partially divided dorsally by a weakly developed rostral groove; two supranasals, larger compared to surrounding scales, separated from each other by a much smaller scale; single, large, postnasal, much smaller than supranasal, on either side; rostral in contact with nasal scale, supralabial I, supranasals and the small scale separating the supranasals; nostril oval shaped, covers the nasal scale on either side; nasal surrounded by supranasal, rostral, supralabial I and single postnasal on either side; two rows of scales separate orbit from supralabials. Mental triangular; two well-developed postmentals, the inner pair much shorter (1.0 mm) than the mental (1.8 mm), and in strong contact with each other (0.6 mm) behind mental; outer pair slightly shorter (0.7 mm) than inner pair and separated from each other by four gular scales ( Figure 4B ). Inner postmental bordered by mental, infralabial I and two gular scales on either side, outer postmentals; a small scale separating inner postmental from infralabial II on right side; outer postmentals bordered by inner postmentals, eight gular scales on left and six on right and the anterior-most of a series of flattened, enlarged scales bordering the infralabials; A row of enlarged gular scales below infralabials II to V, the ones from below infralabial III, subimbricate. Supralabials 12 on right and 13 on left, nine (on both sides) to midorbital position; infralabials ten on right and 11 on left.

Body relatively stout, not very elongate ( TRL / SVL 0.42). Dorsal pholidosis composed of granular scales that are heterogeneous in size and shape throughout, intermixed with enlarged, conical, keeled tubercles that are heterogeneous in size, arranged in 18 fairly regular longitudinal rows at midbody, extending from posterior part of head to groin; tubercles in parasagittal rows are smaller, keeled and more rounded; approximately six rows of tubercles on either side of parasagittal rows highly enlarged, remarkably conical and strongly keeled; tubercles on flanks slightly larger than those on parasagittal rows but much smaller than tubercles on rest of dorsal aspect of trunk. Enlarged tubercles surrounded by a rosette of approximately 10–12 granules; enlarged tubercles on nape and shoulder smaller than those on trunk, keeled, those on occiput and temporal region slightly smaller, smooth.

Ventral scales homogeneous, larger than dorsal granular scales, smooth, imbricate, getting slightly smaller in the pectoral region; midbody scale rows across belly 33 or 34; gular region with much smaller, flat, subimbricate scales, becoming slightly larger and juxtaposed on anterior aspect. Palmar and plantar scales, granular; scales on dorsal aspect of upper arm and forearm much larger than granules on dorsum, flat, weakly pointed, smooth and subimbricate, intermixed with a few scattered enlarged tubercles that are slightly more conical and distinct on forearm than upper arm; scales on dorsal part of thigh and shank are granular, except those on knee that are flat and imbricate, intermixed with enlarged, conical, rounded tubercles, much larger on thigh compared to shank; anterior aspect of thigh with flatter scales, posterior aspect with granular scales. Nine femoral pores on left and 10 on right, separated medially by a diastema of four poreless scales ( Figure 4D ).

Fore and hind limbs relatively short, stout; forearm short ( FL / SVL 0.13); tibia short (CL/ SVL 0.15); digits moderately short, strongly clawed; indistinct webbing between all digits of manus and digits I–IV of pes; terminal 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 a straight transverse series, divided except for distal and two or three basal scansors on digit I and one or two in other digits: 5–7–7–8–7 (left manus), 5–7–7–8–8 (right manus), 6–8–8–7–7 (left pes), 6–8–8–7–8 (right pes) ( Figures 4E & 4F ). Relative length of digits (measurements in mm in parentheses): IV (4.7)> III (4.4)> V (4.2)> II (3.8)> I (3.0) (right manus); IV (6.2)> V (6.0)> III (5.4)> II (4.3)> I (3.1) (right pes).

Tail depressed, flat beneath, verticillate, with indistinct median furrow, final third regenerated; tail, including regenerated portion, as long as snout to vent length (TL/ SVL 1.01). Scales on dorsal aspect of tail flat, subimbricate, smooth, slightly larger than granules on dorsum, with a series of eight much enlarged, strongly pointed tubercles on the first whorl and five or six in the rest; tubercles on the first to sixth whorls keeled, rest smooth; tubercles on the last row on ventrolateral aspect of tail, broader than long and extends down to the tenth whorl. Ventral scales on tail enlarged, imbricate, with a single median row of roughly rectangular, enlarged subcaudal plates covering almost entire tail including regenerated part. First scale (closest to vent) in enlarged median row largest ( Figure 3D ). A pair of much enlarged postcloacal spurs on both sides, anterior spur of similar in size to the largest of dorsal tubercles and slightly larger than anterior.

Colouration in life based on paratype BNHS 2374. Dorsal ground colour of head, body, limbs and tail greyish-brown ( Figure 6 ). Dorsal aspect of head with a scattering of dark brown and white spots, snout intermixed with darker spots; labials paler than rest of head; a faint post-orbital streak extends until ear opening; eye golden brown with darker venal colouration; Head, body, limbs and original portion of tail with dark brown blotches, some tubercles on occiput and dorsum white. A dark circle in the occipital region followed by six dark rectangular patches along the vertebral column up to tail base. Flanks with dark markings, a streak in the nuchal region followed by six blotches up to the hindlimb insertion on each side; tail with faint, alternating dark and light bands in the original portion; regenerated portion of tail chiefly ground colour with indistinct dark spots. Venter off-white, immaculate.

Variation and additional information from type series. Supranasals in contact in BNHS 2374 . Seven gular scales bordering inner postmentals in BNHS 2378 . Inner postmentals in contact with infralabial II in BNHS 2374 . Outer postmentals bordered by six gular scales on either side in BNHS 2376 , BNHS 2374 & BNHS 2375 . Right outer postmental in BNHS 2378 bordered by three gular scales. Outer postmentals bordered by infralabials I & II on both sides in BNHS 2378 & BNHS 2375 . Outer postmentals bordered by infralabials I & II on right and infralabial II on left in BNHS 2376 and outer postmentals bordered only by infralabial II and separated from infralabial I on either side in BNHS 2374 . All other meristic and mensural variations are listed in Table 3 .

Distribution and Natural history. Hemidactylus varadgirii sp. nov. is known with surety only from the type locality, Amboli in Maharashtra state at elevations of 650–750 m asl. Predominant parts of the Amboli landscape are covered by basaltic floors, commonly called the Deccan Traps, and are thought to have formed during the upper cretaceous to the lower Eocene ( Punekar et al. 2014). The vegetation types here can be classified as mid-elevation wet evergreen and southern tropical semi-evergreen in the comparatively less disturbed areas ( Kanade et al. 2008). Similar morphotypes to H. varadgirii sp. nov. were encountered in Rajgoli village, Maharashtra state that lies ~ 50 km east in straight line distance from Amboli. This species potentially inhabits a wider region in the Western Ghats of southern Maharashtra, Goa and northern Karnataka. The first individuals were encountered during the day, resting under rocks in Amboli. They were later spotted using eyeshine at night, occupying lateritic walls and on the plateaus or ‘sadas’ in Amboli. They seem to be reluctant climbers as they were always found not more than five feet from the ground. They also occasionally slough their skin in defence when caught. Sympatric lizards at the type locality include Hemidactylus prashadi Smith, 1935 , Cnemaspis amboliensis Sayyed, Pyron and Dileepkumar,2018 , Monilesaurus rouxii (Duméril & Bibron, 1837) , Calotes versicolor (Daudin, 1802) and Eutropis carinata (Schneider, 1801) .

Figures

FIGURE 1. A Maximum Likelihood (ML) phylogeny of the members of the brookii group sensu Bauer et al. (2010) showing the position of Hemidactylus varadgirii sp. nov. Black circles at nodes indicate bootstrap support> 95 and Grey circles,> 85, the branches they correspond to.

FIGURE 3. Hemidactylus varadgirii sp. nov. (holotype, BNHS 2377), dorsal view (A), ventral view (B), dorsal view of tail (C), and ventral view of tail (D). Scale bars 10 mm.

FIGURE 4. Hemidactylus varadgirii sp. nov. (holotype, BNHS 2377), head in dorsal view (A), ventral view (B), lateral view (C), view of femoral region showing femoral pores (D), ventral view of right manus (E), and ventral view of right pes (F). Scale bars 5 mm.

FIGURE 5. Details of dorsal pholidosis at midbody (A) and details of ventral scales at midbody (B) of (holotype, BNHS 2377) Hemidactylus varadgirii sp. nov. Scale bars 5 mm.

FIGURE 6. Hemidactylus varadgirii sp. nov. (paratype, BNHS 2374) in life.

Tables

TABLE 2. Pairwise sequence divergence (%) between members of the H. murrayi clade. Numbers along diagonal in bold represent within species divergence.

Species	Voucher	Genbank Accession Number	Locality
Hemidactylus varadgirii sp. nov.	Unvouchered	MK569850	India, Maharashtra, Kolhapur, Amboli
Hemidactylus varadgirii sp. nov.	VG349	MK569849	India, Maharashtra, Kolhapur, Amboli
Hemidactylus varadgirii sp. nov.	AQ473	MK569848	India, Maharashtra, Kolhapur, Amboli
Hemidactylus sp.	CAS252886	KM975948	United States, Louisiana
Hemidactylus sp.	VG156	MK569851	India, Tripura,

TABLE 1. Sequences used in this study. Museum abbreviations as follows:ADS, Anslem de Silva field series; ADS, Anslem de Silva field series; AMB, Aaron M. Bauer field series (now in National Museum, Colombo, Sri Lanka); AQ/NCBS (National Centre for Biological Sciences, Bangalore); CAS, California Academy of Sciences; CES/ CESG/ CESL/ ESV, Centre for Ecological Sciences; IAG, Ishan Agarwal field series; JS, Jay Sommers personal collection; LSUHC, La Sierra University Herpetological Collection; USNM, National Museum of Natural History; ZRC, Zoological Reference Collection (ZRC) of the National Museum of Singapore; VG, Varad Giri field series.

Species	Voucher	Genbank Accession Number	Locality
Hemidactylus chikhaldaraensis Hemidactylus chipkali Hemidactylus chipkali Hemidactylus frenatus Hemidactylus frenatus Hemidactylus frenatus Hemidactylus cf. gleadowi Hemidactylus cf. gleadowi Hemidactylus cf. gleadowi Hemidactylus imbricatus Hemidactylus malcolmsmithi Hemidactylus malcolmsmithi Hemidactylus cf. malcolmsmithi Hemidactylus cf. malcolmsmithi Hemidactylus murrayi	NCBS-BH684 CESG322 CESG323 AMB 7420 CAS 229633 LSUHC 4871 CES11014 IAG058 IAG059 JS11 CES11065 IAG061 CAS 252886 CAS 252886 CAS 206638	MK569807 MK569808 MK569809 EU268359 HM559629 GQ458049 MH454761 MK569811 MK569810 EU268353 MH454765 MK569842 KM975948 KM975948 GQ458054	India, Maharashtra, Amravathi, Gawilgarh Fort India, Madhya Pradesh, Hoshangabad, Pachmarhi India, Madhya Pradesh, Hoshangabad, Pachmarhi Sri Lanka, Anuradhapura District, Ritigala Myanmar, Tanintharyi Div., Kaw Thaung Dist Malaysia, Pahang, Bukit Bakong India, Karnataka, Bagalkot, India, Maharashtra, Buldhana, Near Lokhanda India, Maharashtra, Amravathi, Chichghat Pakistan, pet-trade India, Himachal Pradesh, Hamirpur, Sujanpur India, Maharashtra, Amravathi, Chikhaldara USA, Louisiana, New Orleans USA, Louisiana, New Orleans Myanmar, Mandalay Division
Hemidactylus murrayi	CAS 208159	GQ458052	Myanmar, Yangon
Hemidactylus murrayi	CAS 229632	GQ458051	Myanmar, Tanintharyi Division
Hemidactylus murrayi Hemidactylus murrayi	LSUHC 6754 LSUHC 6755	EU268365 EU268366	Malaysia, Pulau Pinang, Empangon Air Hitam, Malaysia, Pulau Pinang, Empangon Air Hitam,
Hemidactylus murrayi	USNM 579430	KM975941	Timor-Leste, Com
Hemidactylus murrayi	USNM 579441	KM975942	Timor-Leste, Com
Hemidactylus murrayi	USNM 579442	KM975943	Timor-Leste, Com
Hemidactylus murrayi	USNM 579443	KM975944	Timor-Leste, Com
Hemidactylus murrayi	USNM 579728	KM975945	Timor-Leste, Dili
Hemidactylus murrayi	USNM 579729	KM975946	Timor-Leste, Dili
Hemidactylus murrayi	USNM 581298	KM975947	Timor-Leste, Dili
Hemidactylus murrayi	ZRC 2.6167	GQ458050	Borneo, Sarawak, Loagan Bunut National Park
Hemidactylus parvimaculatus Hemidactylus parvimaculatus Hemidactylus parvimaculatus	ADS 36 AMB7466 AMB7475	GQ458053 GQ458056 GQ458055	Sri Lanka, Kartivu Sri Lanka, Mampuriya, Sri Lanka, Kandy,
Hemidactylus parvimaculatus	CES14174	MH454766	India, Andhra Pradesh, Visakhapatanam, Araku forest guest house
Hemidactylus reticulatus Hemidactylus sankariensis Hemidactylus treutleri	CES13062 NCBS-BH682 CES14216	MH454767 MK569844 MH454770	India, Karnataka, Chamrajnagar, Kollegal India, Tamil Nadu, Erode, Sankari India, Gaganpahad, Hyderabad, Andhra Pradesh

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