Hemidactylus siva, Srinivasulu & Srinivasulu & Kumar, 2018

Srinivasulu, Chelmala, Srinivasulu, Aditya & Kumar, Gandla Chethan, 2018, A new cryptic rock-dwelling Hemidactylus Oken, 1817 (Squamata: Gekkonidae) from northern Karnataka, India, Zootaxa 4444 (1), pp. 25-42 : 28-38

Hemidactylus siva sp. nov.

FigS. 1–3 .

Holotype. NHM.OU. REP.H77-2017, adult female, Underground ŚiVa temple (15.317° N, 76.464° E; 442 m a.S.l.), Hampi , Karnataka, India; collected on 3rd February 2017 by Gandla Chethan Kumar & Tariq A. Shah.

Paratypes. NHM.OU.REP.H79-2017, Subadult male, NHM.OU.REP.H80-2017, Subadult male, NHM.OU. REP.H78-2017, adult female, collection detailS are Same aS the holotype. NHM.OU.REP.H35-2017, adult female, NHM.OU.REP.H36-2017, adult female, locality detailS Same aS the holotype; collected on 4th February 2017 by Tariq A. Shah & C. SriniVaSulu.

Diagnosis. A large Sized Hemidactylus, Snout-Vent aVeraging 99.8 ± 5.3 mm (n=4, adult femaleS) up to 104.7 mm. DorSal pholidoSiS heterogeneouS, With 16 irregularly arranged longitudinal roWS of enlarged, moderately to feebly keeled tubercleS at midbody. FirSt Supralabial iS in contact With noStril. TWo, rarely three, Well-deVeloped pairS of poStmentalS, the inner pair Slightly larger than the outer pair, and in contact With the outer pair. The primary pair of poStmentalS are broadly in contact With each other. Ventrolateral foldS indiStinct, 27–30 Scale roWS acroSS Venter. Enlarged ScanSorS on all digitS; 10–13 (manuS) and 9–11 (peS) diVided ScanSorS beneath firSt digit, and 13– 14 (manuS) and 13–15 (peS) beneath fourth digit; 17–18 femoral poreS on each Side of the thigh, Separated by 5 poreleSS ScaleS in maleS. Original tail depreSSed, oVal in tranSVerSe Section Without a median dorSal furroW; ScaleS on the tail Slightly larger than dorSalS of body, imbricate, With a longitudinal SerieS of tWo enlarged, Weakly keeled, Striated, and flattened tubercleS on either Side of the median furroW, only on the firSt tWo tail SegmentS.

The large Size (to 104.7 mm SVL) of Hemidactylus siva sp. nov. eaSily diStinguiSheS it from many other Indian and Sri Lankan congenerS: H. albofasciatus GrandiSon & Soman , H. aquilonius McMahan & Zug , H. brookii Gray , H. frenatus Duméril & Bibron , H. garnotii Duméril & Bibron, H. cf. gleadowi Murray, H. gracilis Blanford , H. gujaratensis Giri, Bauer , VyaS & Patil, H. imbricatus Bauer, Giri, Greenbaum, Jackman, Dharne & Shouche , H. kushmorensis Murray , H. parvimaculatus Deraniyagala , H. persicus AnderSon , H. platyurus Schneider , H. reticulatus Beddome , H. robustus Heyden , H. sataraensis Giri & Bauer , H. scabriceps Annandale , and H. treutleri Mahony , all of Which are Significantly Smaller, reaching approximately 70 mm.

Other congenerS can be diStinguiShed on the baSiS of a Suite of characterS (differing or non-oVerlapping characterS indicated parenthetically): dorSum With conical, granular, Striated ScaleS intermixed With enlarged, irregularly arranged, longitudinal roWS of 16 Sub-trihedral, feebly to moderately keeled, Striated tubercleS (enlarged dorSal tubercleS uSually feW, SometimeS abSent in H. leschenaultii Duméril & Bibron ; dorSum With Very feW enlarged tubercleS, more often abSent altogether in H. flaviviridis Rüppell ; no enlarged dorSal tubercleS in H. giganteus Stoliczka ; dorSum With 18–20 roWS of irregularly arranged, enlarged, rounded and feebly keeled tubercleS in H. aaronbaueri Giri ; heterogeneouS Scalation With 10–12 irregularly arranged longitudinal roWS of enlarged, rounded tubercleS in H. yajurvedi Murthy, Bauer, Lajmi, AgarWal & Giri ); 17–18 femoral poreS on each Side of the thigh in maleS (6–14 femoral poreS on each Side in H. lankae Deraniyagala and H. triedrus Daudin , and 20 femoral poreS on each Side in H. sykesii Günther , a junior Synonym of H. maculatus ; vide Mirza & Sanap 2014); a combination of 13–15 ScanSorS on IV digit of peS and femoral poreS Separated by 5 poreleSS ScaleS (10–11 ScanSorS on IV digit of peS and femoral poreS Separated by 2–4 poreleSS ScaleS in H. depressus Gray ; vide BatuWita & Pethiyagoda 2012) and a combination of 14–15 SupralabialS and femoral poreS Separated by 5 poreleSS ScaleS (11–12 SupralabialS and femoral poreS Separated by 1–3 poreleSS ScaleS in H. pieresii Kelaart ; vide BatuWita & Pethiyagoda 2012).

BaSed on the dorSal pholidoSiS and general appearance, H. siva sp. nov. iS moSt Similar to Hemidactylus maculatus Duméril & Bibron , H. hunae Deraniyagala , H. graniticolus , and H. acanthopholis ; hoWeVer, it differS from theSe by haVing a dorSal pholidoSiS With conical, granular ScaleS intermixed With enlarged, irregularly arranged longitudinal roWS of 16 Sub-trihedral, moderately to feebly keeled, Striated tubercleS of unequal Size that are comparatiVely larger toWardS the diStal midbody (versus back With conical, granular, Striated ScaleS intermixed With enlarged, fairly regularly arranged longitudinal roWS of 16–18 Sub-trihedral, Weakly keeled, Striated tubercleS in H. graniticolus and H. hunae ; dorSal Scalation on trunk granular, intermixed With enlarged fairy regularly arranged longitudinal roWS of 18–20 trihedral, moderately keeled, Striated tubercleS in H. acanthopholis ; dorSum With Small juxtapoSed, conical, granular ScaleS intermixed With large trihedral tubercleS arranged in 20 fairly regular roWS in H. maculatus ; back With Small granular ScaleS intermixed With much larger ScaleS); femoral poreS in maleS 17–18 on each Side of the thigh Separated by 5 poreleSS ScaleS (versus 22–24 femoral poreS on each Side With a gap of 3–6 ScaleS in H. hunae ; 23–28 femoral poreS on each Side Separated by 1–3 poreleSS ScaleS in H. graniticolus ; 19–21 femoral poreS on each Side Separated by 13–14 poreleSS ScaleS in H. acanthopholis ). H. siva sp. nov. differS from H. sushilduttai Giri, Bauer, Mohapatra , SriniVaSulu & AgarWal in haVing 17–18 femoral poreS on each Side Separated by 5 poreleSS ScaleS (versus 21–24 on each Side Separated by 4 poreleSS ScaleS); tWo roWS of Subtrihedral tubercleS on the firSt and Second caudal SegmentS (versus 6–8 conical tubercleS on the dorSum of firSt caudal Segment).

Description. The holotype iS generally in good condition With minor preSerVation artefactS ( Fig. 1A ). Body Straight, 4th and 5th fingerS of left hand are Slightly upturned, tail Slightly curVed, Venter flattened.

Head Short (HL/SVL ratio 0.30), Slightly elongate (HW/HL ratio 0.68), not Strongly depreSSed (HH/HL ratio 0.42), diStinct from neck ( Fig. 1B ). Loreal region Slightly inflated, canthuS roStraliS not prominent. Snout Short (SE/ HL ratio 0.41); Slightly longer than eye diameter (OD/SE ratio 0.50); ScaleS on Snout, canthuS roStraliS, forehead and interorbital region homogenouS, juxtapoSed, and Weakly pointed; ScaleS on Snout, canthuS roStraliS and forehead tWice the Size of thoSe on the occipital and interorbital region, canthuS roStraliS With Slightly enlarged patch of ScaleS ( Fig. 1D ). Eye Small (OD/HL ratio 0.20); pupil Vertical With crenulated marginS; SupraciliarieS Small, pointed, thoSe at the anterior end of orbit Slightly larger. Ear opening oVal (greateSt diameter 2.25 mm); eye to ear diStance Slightly greater than diameter of eye (EE/OD ratio 1.17). RoStral Wider (4.2 mm) than deep (2.1 mm), diVided to half by moderately deVeloped roStral grooVe; three internaSalS, tWo internaSalS contacting the naSal enlarged, enlarged, medial naSal ScaleS (2) Smaller in Size (leSS than half of the adjacent internaSalS) arranged longitudinally aS a SerieS in betWeen, to all oVer the length of enlarged internaSalS, proViding no contact, one SupranaSal on each Side Which iS Smaller than internaSal, tWice the Size bigger to the medial naSal Scale, roStral in contact With noStril, Supralabial I, and internaSal; noStrilS large (0.85 mm), circular, each Surrounded by SupranaSal, internaSal, roStral, Supralabial I and poStnaSal; 2–4 roWS of ScaleS Separate orbit from SupralabialS. Mental triangular, mental Wider (4.3 mm) than deep (3.9 mm), tWo Well-deVeloped poStmentalS, the inner pair Slightly longer (4.2 mm) than mental (3.9 mm), and in broadly contact With each other (2.4 mm) behind mental, outer pair about half the Size of the inner pair, Separated from each other by inner pair ( Fig. 1C ). Inner poStmental bordered by mental, infralabial I, outer poStmental and tWo to three gular ScaleS; outer poStmental bordered by infralabialS I, II, III, inner poStmental, and 5–8 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, 2–8 roWS of ScaleS beloW infralabialS III to VIII are enlarged and Weakly imbricate. SupralabialS (to midorbital poSition) 10 (right)–10 (left); SupralabialS (to angle of jaW) 15 (right)–14 (left); infralabialS (to angle of jaW) 12 (right)–11 (left).

Body relatiVely Stout, not elongate (TRL/SVL ratio 0.43), With Ventrolateral foldS Without denticulate ScaleS; Skin Very delicate. DorSal pholidoSiS heterogeneouS, compoSed of conical, granular, Striated ScaleS intermixed With enlarged, irregularly arranged, longitudinal roWS of 16 Subtrihedral, moderately to feebly keeled, Striated tubercleS at midbody, extending from occipital region to tail, more prominent toWardS the diStal midbody, each enlarged tubercle roughly tWo to three timeS longer than adjacent granuleS, Surrounded by roSette of 12–14 Small granuleS, 3–5 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 on flank SmalleSt after medial roWS; enlarged tubercleS are feebly to moderately 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, pointed ( Fig. 1E ). Ventral ScaleS larger than dorSal, Smooth, imbricate, Slightly larger on precloacal and femoral region than on cheSt and abdominal region; midbody Scale roWS acroSS belly 29; gular region With Still Smaller, Sub-imbricate ScaleS, thoSe on lateral aSpect of neck granular, anterior gular ScaleS Slightly larger than the reSt.

Femoral and precloacal poreS abSent. ScaleS on the palm and Sole Smooth, imbricate, not much rounded; ScaleS on dorSal aSpect of upper arm larger than granuleS 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, Subtrihedral tubercleS, Which are numerouS on Shank compared to anterior aSpect of thigh; poSterior aSpect of thigh lackS enlarged tubercleS.

Fore- and hindlimbS relatiVely Short, Stout; forearm Short (FL/SVL ratio 0.15); Slightly leaner in compariSon With hindlimbS (CL/SVL ratio 0.17); digitS moderately Short, Strongly claWed; all digitS of manuS and digitS I–IV of peS indiStinctly Webbed; 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 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: 11-6*- 13-13-12 (left manuS) 11-14-13-13 -12 (right manuS; Fig. 2A ) 9-13-13-13 -12 (left peS) 9-13-13-13 -13 (right peS; Fig. 2B ). RelatiVe length of digitS (meaSurementS in mm in parentheSeS): III (8.26)> IV (8.02)> II (7.8)> V (7.1)> I (5.9) (right manuS); V (8.9)> III (8.0)> II (7.9)> IV (7.8)> I (5.5) (right peS).

Regenerated tail depreSSed, flat beneath, Verticillate, With Well-defined median furroW; oVal in croSS Section, longer than Snout-Vent length (TL/SVL ratio 1.12); tail coVered aboVe With Small (Slightly larger than thoSe on the dorSal granuleS), poSteriorly-pointed, imbricate, Striated ScaleS and a SerieS of tWo enlarged, moderately keeled and Weakly Striated, poSteriorly pointed and flattened tubercleS on either Side of the median furroW (arranged With a diStinct median Space) on firSt and Second caudal SegmentS ( Fig. 2C ); Ventral ScaleS larger, imbricate, median roW (Subcaudal plateS) Slightly broader, about tWice aS broad aS adjacent ScaleS, not extending acroSS Width of the tail proximally, but diStally they extend almoSt acroSS the Width of the tail ( Fig. 2D ).

Coloration in life. DorSal markingS are much more eVident in vivo. Vertebral Stripe faintly ViSible, mildly darker in colour to the adjacent dorSal coloration, extendS from behind the pectoralS to the tail. DorSum light yelloWiSh-beige With fiVe faint White-bordered broWn bandS: one acroSS the occiput, one acroSS the ShoulderS, tWo acroSS the region betWeen fore- and hindlimbS, and one acroSS the pelVic region ( Fig. 3A ). The broWn colouration of the bandS iS only Slightly darker than the reSt of the dorSum. The band on the ShoulderS iS horSeShoe-Shaped. A diStinct White-bordered broWn lateral line, continuouS With the band acroSS the Shoulder, iS preSent poSterior to the eyeS. Regenerated tail faintly banded; alternating light and dark bandS Similar to dorSum in colour; darker bandS SomeWhat Wider than pale bandS in the proximal Section. The diStal portion of the tail poSSeSS equal to Subequal band portionS of darker and paler bandS along the tapering tail. The bandS become more diStinct aS the tail taperS. ForelimbS and hindlimbS With dull mottling and faint croSS-banding extended to the manuS and peS. IriS broWn.

Coloration in preservative. The general colouration of the dorSum iS light broWn, to a deeper broWn on the SideS of the trunk. The bandS on the dorSum and tail are more prominent after preSerVation. BandS on the forelimbS and hindlimbS completely faded, remain aS a darkiSh broWn colour extending to the manuS and peS. The anterior dorSal Surface of the head haS a Slight orange-broWn colouration, poSSibly aS an artefact of collection or preSerVation. Mid-dorSal line diStinct from occiput to the tail, lighter on the dorSum than on the tail. Tail diStinctly lighter than the body, faded to a light broWn-beige colour.

Etymology. The Specific epithet iS a patronym, noun in appoSition, honouring the deity ŚiVa, referring to the underground ŚiVa temple from Where the SpecieS WaS firSt collected.

Suggested common name. Hampi Rock Gecko.

Variation. MenSural data for the type SerieS and additional material iS giVen in Table 3. A total of four femaleS ranging in Size from 91.6 mm to 104.7 mm and tWo Subadult maleS, ranging in Size from 74.5 mm and 77.5 mm. MaleS haVe a SerieS of 17–18 femoral poreS Separated medially by 5 poreleSS ScaleS ( Fig. 4 ). All paratypeS reSemble the holotype in moSt of the morphological characterS except aS folloWS: range of SupralabialS iS from 12–15 (9–10 beloW eye) and infralabialS from 10–14. The ScaleS acroSS belly range from 27–30 in the paratypeS.

Colouration in Subadult male paratypeS iS much more prominent than in the femaleS ( Fig. 3B ). Vertebral Stripe ViSible but faint. In life, dorSum light broWn With fiVe White- to yelloW-bordered dark broWn WaVy bandS: one acroSS the occiput, one acroSS the ShoulderS, tWo acroSS the region betWeen fore- and hindlimbS, and one acroSS the pelVic region. The broWn colouration of the bandS iS darker than in the adult holotype. The band on the ShoulderS iS horSeShoe-Shaped With a Slight W-Shaped curVe on the concaVe Side. A diStinct yelloW-bordered dark broWn lateral line, continuouS With the band acroSS the occiput, iS preSent poSterior to the eyeS. SupraciliarieS light broWniSh yelloW. TubercleS on flankS pale. Tail diStinctly banded With alternating WhitiSh and dark broWn bandS; darker bandS Wider than White bandS in the proximal Section, becoming equal to Subequal toWardS the tip. The White colour of the lighter bandS becomeS brighter toWardS the tip. A total of Six dark bandS and SeVen light bandS on the tail. Tail tip black. ForelimbS and hindlimbS paler than dorSum, dull pinkiSh-broWn extending to the manuS and peS. IriS bronze-broWn. Adult maleS Similar to adult femaleS in colouration and patternS ( Fig. 3C ).

Phylogenetic relationships. Phylogenetic analySiS baSed on the concatenated nuclear (RAG-1 and PDC) and mitochondrial (Cytb) gene SequenceS ShoWed H. siva sp. nov. to be placed baSally to the entire H. prashadi clade ( Fig. 5 & 6 ). The poSition of H. siva sp. nov. WaS highly Supported (>0.95 poSterior probability) by both analySeS, but the phylogenetic poSitionS of Some other SpecieS Were not. We belieVe that the relationShip of theSe SpecieS With each other—and With H. siva sp. nov. —could be Subject to further reSolution, and muSt be corroborated With analySeS of additional loci. Furthermore, pairWiSe diStance analySiS ShoWS a genetic diStance of 11.3–18.8% from memberS of the H. prashadi clade ( Table 4). Our phylogenetic analySiS SupportS the diStinctiVeneSS of H. siva sp. nov. from the reSt of itS congeneric SpecieS of the genuS Hemidactylus in India.

Distribution and Natural History. Hemidactylus siva sp. nov. iS only knoWn from the temple ruinS of Hampi, a UNESCO World Heritage Site in Bellary diStrict, Karnataka, India ( Fig. 7 ). It iS relatiVely uncommon at the locality and WaS obSerVed to be inhabiting dark areaS in the leSS croWded abandoned temple complexeS ( Fig. 8 ). All the SpecimenS Were found on the rock WallS of the old temple ruinS during diurnal SearcheS. The habitat around the type locality compriSeS boulderS and hillockS adjacent to the Southern bank of RiVer Tungabhadra, dominated by open Scrub foreStS. The typeS Were found Sympatrically With Hemidactylus flaviviridis , H. giganteus , H. leschenaultii , Cnemaspis adii , and Psammophilus blanfordanus Stoliczka.

Figures

FIGURE 1. Hemidactylus siva sp. nov. (hOlOtype, NHM.OU.REP.H77.2017). A) DORsAl vIew Of the Adult feMAle, B) DORsAl vIew Of heAd, C) LAteRAl vIew Of heAd, D) VentRAl vIew Of heAd, E) DetAIls Of dORsAl phOlIdOsIs. PhOtO by AdItyA SRInIvAsulu & C. SRInIvAsulu. ScAle bAR= 10 MM

FIGURE 2. Hemidactylus siva sp. nov. (hOlOtype, NHM.OU.REP.H77.2017). A) VentRAl vIew Of RIght MAnus, B) VentRAl vIew Of RIght pes, C) DORsAl vIew Of tAIl, And D) VentRAl vIew Of tAIl. PhOtO by AdItyA SRInIvAsulu & C. SRInIvAsulu. ScAle bAR= 10 MM

FIGURE 3. PhOtOgRAphs Of Hemidactylus siva sp. nov. In lIfe. A) hOlOtype (NHM.OU.REP–H77.2017, Adult feMAle), B) pARAtype (NHM.OU.REP.H79–2017, subAdult MAle), C) Adult feMAle, nOt cOllected. PhOtOs A & B by G. ChethAn KuMAR, And C by TARIq A. ShAh & BhARgAvI SRInIvAsulu.

FIGURE 4. PReclOAcAl RegIOn Of pARAtype (NHM.OU.REP.H79.2017) Of Hemidactylus siva sp. nov. shOwIng A seRIes Of 18 feMORAl pORes On left And RIght sIde sepARAted MedIAlly by fIve pOReless scAles. PhOtO by C. SRInIvAsulu & AdItyA SRInIvAsulu. ScAle bAR=10 MM.

FIGURE 5. MAxIMuM clAde cRedIbIlIty tOpOlOgy estIMAted fROM BAyesIAn InfeRence (BI) Of the cOMbIned RAG-1 And PDC gene dAtAset. VAlues At the nOdes ARe pOsteRIOR pRObAbIlIty vAlues.

FIGURE 6. MAxIMuM clAde cRedIbIlIty tOpOlOgy estIMAted fROM BAyesIAn InfeRence (BI) Of Cytb gene dAtAset. VAlues At the nOdes ARe pOsteRIOR pRObAbIlIty vAlues.

FIGURE 7. MAp Of IndIA depIctIng the type lOcAlIty (Red sOlId cIRcle) Of Hemidactylus siva sp. nov. In BellARy DIstRIct, KARnAtAkA, IndIA; And dIstRIbutIOn Of lARge-bOdIed tubeRculAted Hemidactylus specIes: H. maculatus (lIght blue sOlId cIRcles), H. graniticolus (dARk blue sOlId cIRcles), H. acanthopholis (ORAnge sOlId cIRcles), H. paaragowli (yellOw sOlId cIRcles), H. sushilduttai (pInk sOlId cIRcles), H. vanam (gRey sOlId cIRcles) And H. kangerensis (gReen sOlId cIRcles). SIte RecORds thAt need cOnfIRMAtIOn Of H. cf. maculatus RepORted In eARlIeR lIteRAtuRe ARe depIcted As blAck sOlId cIRcles.

FIGURE 8. UndeRgROund ŚIvA teMple, HAMpI In BellARy DIstRIct, KARnAtAkA, IndIA wheRe the hOlOtype Of Hemidactylus siva sp. nov. wAs cOllected, (Inset, StOne RelIef In the sAMe teMple depIctIng A lIZARd neAR ŚIvA LIṅgA - An AbstRAct RepResentAtIOn Of ŚIvA). PhOtO by C. SRInIvAsulu.