A new species of rupicolus gecko of the genus Hemidactylus oken, 1817 (Reptilia: Squamata: Gekkonidae) from Maharashtra, Central India

June 15, 2017 | Autor: Dr. Ashish Tiple | Categoria: Herpetology, Biodiversity
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Russian Journal of Herpetology

Vol. 22, No. 3, 2015, pp. 233 – 240

A NEW SPECIES OF RUPICOLUS GECKO OF THE GENUS Hemidactylus OKEN, 1817 (REPTILIA: SQUAMATA: GEKKONIDAE) FROM MAHARASHTRA, CENTRAL INDIA Parag H. Dandge1 and Ashish D. Tiple2 Submitted June 6, 2014. The genus Hemidactylus Oken, 1817 has 26 species reported from the Indian region. Here we report a new species, Hemidactylus hemchandrai sp. nov. collected from the unprotected peripheral region of the Navegaon Bandh National Park, district Bhandara, eastern Maharashtra, Central India. The new species can be distinguished from other species of Hemidactylus, Hemidactylus hemchandrai having snout to vent length 63 to 86 mm, 12 to 15 rows of irregularly arranged flattened to weekly conical tubercles on the dorsum, 6 to 7 rows of tubercles on the first segment of the tail, 9 lamellae on first digit of pes, 14 lamellae on fourth digit of pes, 10 to 11 femoral pores in male separated by 5 to 6 scales, 11 to 12 supralabials and 8 to 10 infralabials. Keywords: Hemidactylus hemchandrai sp. nov.; Squamata; Reptilia; Central India; Navegaon National Park.

INTRODUCTION The genus Hemidactylus Oken 1817 is one of the most species rich genus of the family Gekkonidae and widely distributed of all reptile genera, been found in the tropical and subtropical regions of the world and hundreds of continental and oceanic islands. However, the great majority of Hemidactylus species has relatively small distributions confined to southern Asia and Africa. Despite having already 122 species, the number of species described in recent years is very high (Smid et al., 2013). 26 species reported from the Indian region (Giri and Bauer, 2008; Mahony, 2009; Bauer et al., 2010a; Agarwal et al., 2011). 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, 2010b; Agarwal et al., 2011). The distribution of genus Hemidactylus has been highly affected by repeated transmarine colonizations, human activity, spontaneous rafting, which have contributed significantly to the unusually wide distribution range (Smid et al., 2013). The new species described here, Hemidactylus hemchandrai was collected from the peripheral region of 1

2

Bahar Nature Foundation, Wardha, Maharashtra, 442001, India; e-mail: [email protected] Department of Zoology, Vidyabharti College, Seloo, Wardha, 442104, Maharashtra, India; e-mail: [email protected]

the Navegaon Bandh National Park, Central India. The new species differs from its allied species H. leschenaulti, H. giganteus, H. aaronbaueri, and H. gujaratensis in several characters and we also provide species variations, ecology and natural history of new species. MATERIAL AND METHODS We conducted herpetological field surveys in different regions of the Navegaon Bandh National Park, Central India since 2006 to 2014 along the reserve forest areas, buffer zone, lakes, rivers and surrounding area during the monsoon and post monsoon period. Specimens were hand collected and stored in 70% ethanol ~24 h after collection. All measurements were carried out using a Digital slide vernier caliper. Coloration in life was established with the use of digital photographs from a Canon XUS 115 HS, Canon 450 D DSLR. Morphometric abbreviations are as follows: SVL, snout to vent length; TL, tail length; TW, maximum tail width (excluding additional tubercle length); BW, maximum body width excluding additional tubercle length; HL, head length from posterior axis of the jaw to the tip of the snout; HW, head width at its widest point; HD, maximum head depth; ED, horizontal eye diameter; EL, maximum ear diameter; E-E, distance from posterior edge of eye to anterior edge of ear; E-N, distance from anterior edge of eye to posterior edge of nostril; E-S, anterior edge of eye to snout tip; IN, internarial distance; IO, transverse distance between anterior most supraciliaries; FAL, length

1026-2296/2015/2203-0233 © 2015 Folium Publishing Company

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Parag H. Dandge and Ashish D. Tiple

Fig. 1. Map showing type locality of Hemidactylus hemchandrai sp. nov.

Fig. 2. Hemidactylus hemchandrai sp. nov. from type locality, adult male (holotype).

from elbow to wrist; TBL, tibia length; and A-G, axilla to groin distance. Sex of the three adult types two males and one female were determined on the basis of the presence of slightly enlarged hemipenal swellings in male, complimented with the presence of femoral pores, considered indicative of males only. Comparative material was examined from the collection of the Zoological Survey of India, Kolkata (ZSI) and Jabalpur (ZSI). Data for the synonyms have been taken either from their original description or from Boulenger (1885); Amarsinghe et al. (2009) and Mahony (2009).

40 – 41 scale rows across venter between lowest rows of tubercles. 9 lamellae on first digit of pes, 14 lamellae on fourth digit of pes. 10 to 11 femoral pores on each thigh, separated by 5 to 6 scale, tail depressed, oval in transverse section, 6 to 7 tubercle present on the first segment of the tail, tail tuberculate with a median dorsal furrow; a single median row of transversely enlarged subcaudal plates started from second segment of the tail. Description of holotype. Adult male (Figs. 2, 5 – 10). Snout-vent length 86 mm. Head relatively short (HL/SVL 21.1%), head width (HW-SVL = 20.11%), not depressed (HD-HL = 55%), head distinct from neck, lores rounded, imbricate and interorbital region concave, forehead concave; snout relatively long (ES-HW ratio 0.66), longer than eye diameter (ED-ES = 49.52%); scales on snout and forehead rounded; scales on snout larger than those on occipital region; eyes large (ED-HL = 31.4%); canthus rostralis rounded; pupil vertically elliptical, with crenellated edges; 38 to 40 elongate supraciliaries, having spines, on anterior upto posterior half of orbit; ear-opening deep, roundish to oval, with major axis vertically (EL-ES = 24.8%); eye to ear distance greater than diameter of eyes (EE-ED = 1.3); rostral rectangular incompletely divided dorsally by a undeveloped rostral groove, Rostral over little more half as deep as wide (rostral width = 4.0 mm, depth = 2.2 mm; width depth ratio 1.81); connected to the nostril posteriorly; two enlarged supranasals separated by a internasal scale which is smaller than that of supranasal scales in size; rostral in contact with supralabial I, supranasals, internasal; nostrils rounded in shape and large in size; each surrounded by supranasal, rostral, first labial and two large postnasals, of which the posterior is larger than the remaining one, posterior nasal gives the appearance like of two fused postnasals; 3 rows of scales separate eyes from

SYSTEMATICS Hemidactylus hemchandrai sp. nov. Holotype (male) ZSI V-6218. India, Maharashtra, Bhandara: unprotected peripheral region of the Navegaon Bandh National Park (Fig. 1); 20°54¢55.5¢¢ N 80°06¢14.6¢¢ E (type locality); date of collection 15/V/2014. Leg. Mr. Parag H. Dandge. Specimen under present study will be deposited in National Zoological Collection of Zoological Survey of India, Central Zone Regional Centre, Jabalpur 482002, India. Paratype (1 male, 1 female) ZSI V-6219. Paratype collected from the same locality. Leg. Mr. Parag H. Dandge. Deposited in National Zoological Collection of Zoological Survey of India, Central Zone Regional Centre, Jabalpur-482 002, India. DIAGNOSIS A medium-sized Hemidactylus, SVL 63 to 86 mm. 12 to 15 rows of irregularly arranged, flattened to weakly conical tubercles. First supralabial contacting nostril narrowly. Two well-developed pairs of postmentals, the inner pair larger and in broad contact behind mental.

A New Species of the Genus Hemidactylus Oken, 1817 from Central India

Fig. 3. Adult male (not collected) from type locality, exhibiting typical coloration for the species.

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Fig. 5. Right lateral view of head of holotype of Hemidactylus hemchandrai sp. nov.

Fig. 4. Dorsal view of trunk of paratype of Hemidactylus hemchandrai sp. nov. male showing pholidosis.

supralabials; first supralabial in contacts nostril narrowly. mental subtriangular, little deeper than wide (3.4 mm wide, 3.5 mm deep, ratio 1.03); two paired postmentals, inner (2.9 mm deep, 1.3 mm wide) is larger than outer (1.9 mm deep, 1.1 mm) wide, respectively; outer pair of postmental is rounded posteriorly; first pair of postmental is contact in subequal size to mental and first infralabials broadly and in contact with second infralabial narrowly, outer pair postmental is in contact with second infralabial; row of enlarge scales bordering infralabial decreasing in size posteriorly, 8 scales ends on 6th infralabial of right and 6 scales ends on 6th infralabial on left. 11 left and 12 right supralabial having light to dark brown speculation, infralabials 10. Body slightly, somewhat elongate (AGL/SVL = 36.62%); ventrolateral fold present; dorsum covered with uniform flattened granules with irregularly arranged flattened to weakly conical slightly backwardly directed tubercles starts from poste-

Fig. 6. Chin scalation of the holotype of Hemidactylus hemchandrai sp. nov.

rior interorbital and temporal regions to tail; each tubercle surrounded by 9 – 10 scales and 4 – 5 scales in two adjacent enlarged tubercles on the dorsum, tubercles are

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Fig. 9. Ventral view of right pes of holotype of Hemidactylus hemchandrai sp. nov.

Fig. 7. Dorsal view of head of holotype of Hemidactylus hemchandrai sp. nov.

Fig. 8. Dorsal view of showing rostral of holotype of Hemidactylus hemchandrai sp. nov.

more compactly arranged on neck region; 12 to 15 rows of tubercles at midbody; ventral scales larger than dorsal,

Fig. 10. Femoral pores of the holotype male of Hemidactylus hemchandrai sp. nov.

subimbricate; scales on guller region are smaller than ventral; 42 to 46 rows of scales from midbody to the ventrolateral fold; Scales on palm are smooth and bit larger than radius, no tubercles on radius; scales on the femur are granular and intermixed with 3 rows of tubercles; preanal grove absent; 10 pores on right and 11 on left thigh, separated by 5 non pore bearing scales; tibia medium in length (TBL/SVL = 18.06%); digits short have curve claw on the tips; every digits with mostly divided scansors; lamellae numbering on digits of left manus (total: undivided) I (10: 4 basal, 1 distal), II (12: 2 basal 1 distal), III (12: 1 distal), IV (12: 1 basal 1 distal), V (13: 2 basal 1 Distal) and of left pes I (9:3 basal 1 distal), II (13: 2 basal 1 distal), III (13: 1 basal 1 distal), IV (14: 2 basal 1 distal). V (14: 3 basal 1 distal). Original portion of tail (9.2 mm) slightly depressed, flat beneath, verticillate; regenerated portion (58.4 mm) tapering in a point; length of tail is less to that of snout-vent length

A New Species of the Genus Hemidactylus Oken, 1817 from Central India

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TABLE 1. Meristic Data for Type Series Hemidactylus hemchandrai sp. nov. Dorsal tubercles

Scale rows across the ventre

12 to 15

40

12 to 15

40 to 41

12 to 15

40 to 41

the left manus

the right manus

the left pes

10, 12, 12, 12, 13 10, 12, 12, 12, 13 10, 12, 12, 13, 13

10, 12, 12, 12, 13 10, 12, 12, 12, 13 10, 12, 12, 13, 13

9, 13, 13, 14, 14 9, 12, 13, 13, 13 9, 13, 13, 14, 14

Femoral pores

the right pes

9, 13, 13, 10 on right and 14, 14 11 on left femor 9, 12, 12, 10 each side 13, 13 9, 13, 13, absent 14, 14

(TL/SVL 78.3%); original part of tail covered above with scales equal to those on dorsum), uniformly granular scales and a series of 6 to 7 somewhat enlarged tubercles, continuing from first segment of the tail up to the original part ends; caudal are broad and starting from second segment of the tail; regenerated portion of tail covered above with small scales, below with enlarged subcaudal plates; 40 scales across the belly. Coloration in life. In day time dorsum was dark brown in color, tubercles were more darker than that of forebody. In night dorsum was lighter brown with some pal bars on the dorsum. Eyes were in golden color where elliptical pupil was dark black. Venter was pal color. (Figs. 2 and 3) Comparison. H. hemchandrai may be distinguished from H. gracilis Blanford, H. reticulatus Beddome, H. albofasciatus Grandison et Soman, H. imbricatus Bauer et al., H. sataraensis Giri and Bauer, and H. anamallensis Günther, on the basis of its divided subdigital lamellae (vs. lamellae undivided or only distal lamellae divided or notched), from H. scabriceps Annandale by its granular (vs. imbricate) dorsal scalation; in presence of 12 to 15 rows of tubercles on the dorsum usually few in number; with fewer enlarged tubercles, more often absent altogether 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 et Bibron; large trihedral tubercles arranged in 20 fairly regular longitudinal rows in H. maculatus, conical, keeled, or subtrihedral tubercles arranged in from 16 – 20 more of less regular longitudinal series in H. brookii; 16 – 18 longitudinal rows of fairly regularly arranged, subtrihedral, weakly keeled, striated tubercles at midbody in H. granticolus; femoral pores in male 10 to 11 on each side separated by 5 to 6 scales (5 – 7 in H. flaviviridis; 19 – 25 femoral pores on each side in H. maculates; from 7 – 12 (16) preano-femoral pores on each side, usually interrupted mesially in H. brookii (Smith, 1935; Giri et al., 2003); 7 – 16 in H. parvimaculatus, Deraniyagala; 6 – 14 femoral pores on each side in H. lankae Deraniyagala; H. subtriedrus Jerdon and H. triedrus

Supralabial

Infralabial

left

right

left

right

6 to 7

11

12

10

10

6 to 7

11

11

9

8

6 to 7

11

11

10

10

Tubercle rows on the first segment of the tail

Lamellae under

(Daudin); 14 femoral pores in H. treutleri Mahony; it differs from H. karenorum and H. bowringii by having more number of lamellae, 10 under thumb and 12 under fourth finger of manus and 9 under the first and 14 under the fourth toe of pes vs. 5 under the thumb, 9 under the fourth finger, 5 or 6 under the first toe, and 10 to 12 under the fourth toe and 5 under the thumb, 7 or 8 under the fourth finger, 5 or 6 under the first toe, and 9 or 10 under the fourth toe, respectively, from H. robustus, H. persicus, H. turcicus, and H. porbandarensis by its lower number of precloacal femoral pores in males (10 – 11 on each thigh separated by 5 to 6 scales vs. 13 or fewer precloacal pores in a continuous series and no femoral pores). H. hemchandrai is looks closely similar to H. leschenaulti, H. giganteus, H. aaronbaueri, and H. gujaratensis can be distinguished by 9 lamellae under 1st and 14 lamellae under 4th toe of pes vs. 6 to 7 under 1st and 9 to 11 under 4th lamellae of pes, 11 supralabials and 8 to 10 infralabials vs. 10 to 12 supralabials and 8 to 9 infralabials, 12 TABLE 2. Measurements (mm) for Type Series of Hemidactylus hemchandrai sp. nov. Specimen SVL TL BW HL HW HD ED EL EE EN ES IN IO FAL TBL AG

Holotype male

Paratype male

Paratype female

86.4 67.6 15.5 18.3 17.4 10.2 5.7 2.9 7.5 8.3 12 2.9 8.3 13.5 15.6 31.6

67.3 69.6 15.4 15.8 13.7 8.2 5 2.4 5 6.6 9.3 2.3 7.3 10.7 13.2 24.6

62.9 71.7 13.4 15.2 12.6 7.9 4.8 2 4.6 6 8.7 2.2 6.5 8.8 11.4 24.5

Note. For abbreviations see Material and Methods.

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Parag H. Dandge and Ashish D. Tiple

Fig. 11. One of the locality (Pratapgad fort) of Hemidactylus hemchandrai sp. nov. exhibiting typical habitat.

to 15 rows of tubercles vs. absent or few number of tubercle on the dorsum mostly 6 scale rows present in the lateral side of the body in central Indian population, 10 to 11 femoral pores in males vs. 12 to 16 femoral pores present in males of H. leschenaulti. 11 to 12 lamellae under 1st and 13 to 15 under 4th toe of pes vs. 9 lamellae under 1st and 14 under 4th toe of pes, 11 to 12 supralabials and 8 to 10 infralabials vs. 12 to 15 supralabials and 11 to 13 infralabials, dorsum with 12 to 15 rows of enlarge tubercles vs. no enlarge tubercles on dorsum, 10 to 11 femoral pores on each thigh in males vs. 19 to 22 femoral pores present in males in H. giganteus. Medium size upto 86 mm vs. large size hemidactylus upto 128 mm, 12 to 15 rows of enlarge tubercles on the dorsum vs. 18 to 20 rows of enlarge tubercles on dorsum, 9 lamellae under 1st and 14 under 4th toe of pes vs. 9 to 12 under 1st lamellae and 11 to 13 under 4th toe of pes, 10 to 11 femoral pores in males separated by 5 scales vs. 15 to 19 femoral pores in males separated by 6 scales, two postnasal scales vs. 4 postnasal, pair of postmental are in touch with 1st and 2nd infralabials vs. postmental are in touch with 1st infralabial in H. aaronbaueri. 12 to 15 rows of tubercles vs. 12 to 16 rows of tubercles, 9 lamellae under 1st and 14 under 4th

toe of pes vs. 7 to 9 under 1st and 11 under 4th toe of pes, 10 to 11 femoral pores in males vs. 12 to 14 femoral pores in males, 40 t0 41 scale rows across the venter vs. 28 to 30 scale rows across the venter, incompletely or undeveloped rostral groove vs. strongly developed rostral groove in H. gujaratensis. All specimens studied and compare with collection available at Zoological Survey of India, Kolkata (Appendix 1). Variations and additional information in types series (Tables 1 and 2, Fig. 3). Mensural data for the types and additional material is given in Table 1 and 2, respectively paratypes are one male and one female ranging in size 63 mm to 67 mm. Both paratypes resembles to the holotype except as follows: supralabial ranges from 11 to 12 and infralabials from 8 to 10. Range of lamellae is 10 under the first and 12 to 13 under the fourth digit of the manus, 9 under first and 13 to 14 under the fourth digit of the pes. Male have a series of 10 to 11 femoral pores separated by 5 to 6 scales. Female without femoral pores. Etymology. The species name hemchandrai derived from the first Author (PHD) late father name. Distribution and habitat. H. hemchandrai is distributed almost throughout the Navegaon Bandh National

A New Species of the Genus Hemidactylus Oken, 1817 from Central India

239

Fig. 12. Rocks and boulders scattered on the Pratapgad fort.

park and peripheral region on the rocky hillock. It prefers the rock boulders for hiding. We observe the good population of H. hemchandrai at pratapgad fort (Figs. 11 and 12) which is at the Pratapgad range of the Navegaon Bandh National Park. Navegaon Bandh National Park is situated at Bhandara district of Maharashtra. Natural history. H. hemchandrai seem to be rupicolous gecko, but it come down on the ground during foraging. During the day time it hides under the rock crevices and with the last ray of sun come out for foraging. H. hemchandrai is a fast dwelling gecko which is active in night, in day time also its very fast dweller. As like most of the Hemidactylus, H. hemchandrai also feed mostly on insects, we observe Apis dorsata is the most preferable food (Fig. 13). H. hemchandrai is particular in habitat selection, it needs big rocky patches for hiding and foraging too. We collected the types from rocks behind the wireless station of Navegon bandh National park. H. hemchandrai is sympatric with H. cf. brookii, H. cf. leschenaulti at the type locality. Remarks. The new species from Navegaon Bandh National Park is compared with all Indian congeneric species, currently considered valid, differs from species

by the following combination of characters; SVL 63 to 86 mm, 12 to 15 rows of irregularly arranged flattened to weekly conical tubercles on the dorsum, 6 to 7 rows of tubercles on the first segment of the tail, 9 lamellae on first digit of pes, 14 lamellae on fourth digit of pes, 10 to 11 femoral pores in male separated by 5 to 6 scales, 11 to 12 supralabials and 8 to 10 infralabials. Total 122 species of genus Hemidactylus reported, the genus Hemidactylus is the second most speciose gecko genus (after Cyrtodactylus) and ranks among the top ten species-rich genera of reptiles. Hemidactylus geckos are widely distributed across all tropical and subtropical continental landmasses and hundreds of intervening continental and oceanic islands, from Southeast Asia westwards over Africa to the New World, Smid et al. (2013) H. hemchandrai is looks closely allied to recently described species H. aaronbaueri and H. gujaratensis and it represents the proper central Indian population. Discovery of this new species Hemidactylus from relatively unexplored area of India highlights the importance of intensive field surveys in revealing hitherto unknown species, the tally of Hemidactylus species in India stands at 27. Study of the reptile from the northwestern

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Parag H. Dandge and Ashish D. Tiple APPENDIX 1. Specimen studied in Zoological survey of India, Kolkata Species Name

Zoological survey of India registration No.

Hemidactylus brookii Hemidactylus frenatus Hemidactylus bowringii Hemidactylus flaviviridis Hemidactylus bowringii Hemidactylus giganteus Hemidactylus platyurus Hemidactylus prashadi Hemidactylus leschenaulti Fig. 13. Adult male (not collected) from type locality, feeding on Apis dorsata.

region of India resulted in a discovery of several new species (Giri, 2008; Giri and Bauer, 2008; Giri et al., 2003) and from south-eastern Andhra Pradesh and south Indian Karnataka state (Mahony, 2009 and Agarwal et al., 2011). Moreover, distribution and biology of genus Hemidactylus in Central India is poorly known. A new species from Vidarbha, Central India highlights the need for an intensive survey of reptile fauna of the region. Hence it is important to carry out detailed field studies to document biology and ecology of such species. Acknowledgments. We thank to State Forest Department of Maharashtra for permitting the specimen collection from locality and the Director, Zoological Survey of India, Kolkata and Officer-in-Charge and Dr. S. S. Talmale ZSI, CZRC, Jabalpur for facilities. We also thank to Mr. Kishor Mishrikotkar for valuable guidance, Mr. Vivek Sharma much needed help and Dr. S. Gomase, for their assistance during the field survey.

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19361, 19554 – 19557, 20583 – 20586 19507, 19508, 19568 – 19577, 19579 19547 – 19552 19558 – 19566, 24877, 24878 19997 – 20011 19567, 20272, 20273 19553, 19578, 19580 – 19587, 19910 20122 – 20129 20163, 20180, 20200, 20201, 20588, 20589 Hemidactylus reticulatus 20286 Hemidactylus maculatus 24959, 24772 – 24774

Bauer A. M., Jackman T. R., Greenbaum E., de Silva A., Giri V. B., and Das I. (2010b), “Molecular evidence for the taxonomic status of Hemidactylus brookii group taxa (Squamata: Gekkonidae),” Herpetol. J., 20, 129 – 138. Boulenger G. A. (1885), Catalogue of the Lizards in the British Museum (Natural History). Vol. I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Trustees of the British Museum, London. Giri V. B. and Bauer A. M. (2008), “A new ground-dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, with a key to the Hemidactylus of India,” Zootaxa, 1700, 21 – 34. Giri V., Bauer A. M., and Chaturvedi N. (2003), “Notes on the distribution, natural history and variation of Hemidactylus giganteus Stoliczka, 1871,” Hamadryad, 27, 217 – 221. Giri V. B, Bauer A. M., Vyas R., and Patil S. (2009), “New species of rock-dwelling Hemidactylus (Squamata: Gekkonidae) from Gujarat, India,” J. Herpetol., 43, 385 – 393. Giri V. B. (2008), “A new rock dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, India. Hamadryad,” 32, 25 – 33. Mahony S. (2009), “ A new species of gecko of the genus Hemidactylus (Reptilia: Gekkonidae) from Andhra Pradesh,” Russ. J. Herpetol., 16(1), 27 – 34. Smid J., Carranza S., Kratochvýl L., Gvozdýk V., Nasher A. K., and Moravec J. (2013),” Out of Arabia: A Complex Biogeographic History of Multiple Vicariance and Dispersal Events in the Gecko Genus Hemidactylus (Reptilia: Gekkonidae),” PLoS One, 8(5), e64018. Zug G. R., Vindum J. V., and Koo M. S. (2007), Burmese Hemidactylus (Reptilia, Squamata, Gekkonidae): taxonomic notes on tropical Asian Hemidactylus,” Proc. Calif. Acad. Sci. Fourth Ser., 58, 387 – 405.

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