Article.

Copyright 2013 Magnolia Press TERMS OF USE Zootaxa 3616 (3): 239 252 www.mapress.com/zootaxa/ Article http://dx.doi.org/10.11646/zootaxa.3616.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:3063286b-b12e-4a49-b1eb-482d5509c292 ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) Phylogenetic relationships and description of a new upland species of Bent-toed Gecko (Cyrtodactylus Gray, 1827) of the C. sworderi complex from northeastern Peninsular Malaysia L. LEE GRISMER 1,2, SHAHRUL ANUAR 3, MOHD ABDUL MUIN 4, EVAN S. H. QUAH 3 & PERRY L. WOOD, JR. 5 1 Department of Biology, La Sierra University, Riverside, California, USA. E-mail: lgrismer@lasierra.edu 2 Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan, Malaysia 3 School of Biological Sciences, Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Penang, Malaysia. E-mail: evanquah@yahoo.com; shahrulanuar@gmail.com 4 Centre for Drug Research, Universiti Sains Malaysia, 11800 Minden, Pulau Pinang, Penang, Malaysia. E-mail: mamuin@gmail.com 5 Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: pwood@byu.edu Abstract Molecular and morphological analyses indicate that a new upland species of the Cyrtodactylus sworderi complex, C. tebuensis sp. nov. from Gunung Tebu, Terengganu, Malaysia is most closely related to C. sworderi and together they form the sister lineage to C. quadrivirgatus. Cyrtodactylus tebuensis sp. nov. is differentiated from all other species of Sundaland Cyrtodactylus on the basis of having the unique combination of large, conical, keeled body tubercles; tubercles present on top of head, occiput, nape, and limbs, and extending posteriorly beyond base of tail; 43 51 ventral scales; no transversely enlarged, median subcaudal scales; proximal, subdigital lamellae transversely expanded; 17 21 subdigital lamellae on fourth toe; an abrupt transition between posterior and ventral femoral scales; enlarged femoral scales; no femoral or precloacal pores; no precloacal groove; body bearing four wide, bold, dark brown stripes (lateral stripe on each flank and a pair of paravertebral stripes); and a pairwise sequence divergence of 13.0% from its closest relative C. sworderi based on the mitochondrial gene ND2. Cyrtodactylus tebuensis sp. nov. is the first endemic upland species of gekkonid from northeastern Peninsular Malaysia and underscores the necessity for additional field work in all upland systems. Key words: Gekkonidae, Cyrtodactylus, Cyrtodactylus tebuensis sp. nov., Malaysia, Terengganu, new species Introduction The gekkonid genus Cyrtodactylus Gray, 1827 is a morphologically diverse lineage of geckos. Although most species are terrestrial to scansorial forest-dwelling forms characterized by large lidless eyes with vertical pupils and having slender, inflected digits expanded only at their bases, the remarkable number of adaptive types (ranging from leaf and bark mimics to obligate cave-dwellers) within Cyrtodactylus is commensurate with the rate at which new species are being described. The genus currently contains 159 species (see Uetz 2012; Grismer et al. 2012) with tens of new species being added annually (e.g., David et al. 2011; Grismer et al. 2012a; Iskandar et al. 2011; Ngo 2011; Oliver et al. 2011; Schneider et al. 2011; Shea et al. 2011), and a number of these have been from Peninsular Malaysia. Of the 23 known species from Peninsular Malaysia and its associated islands, 17 of these have been added in only the last seven years (Chan & Norhayati 2010; Grismer 2005; Grimser & Leong 2005; Grismer & Norhayati 2008; Grismer et al. 2008a; Grismer et al. 2010; Grismer et al. 2012; Johnson et al. 2012; Rösler & Glaw 2008; Youmans & Grismer 2006). The discovery of some of these new species was the result of expeditions into poorly explored regions and islands (Chan & Norhayati 2010; Grismer 2005; Grismer & Leong 2005; Grismer et al. 2008a, 2010a; Youmans & Grismer 2006) whereas others stem from molecular alayses of existing species indicating they were actually species complexes (Grismer et al. 2012a; Johnson et al. 2012). Accepted by A.M. Bauer: 28 Dec. 2012; published: 20 Feb. 2013 239

FIGURE 1. Location of Cyrtodactylus tebuensis sp. nov. at Gunung Tebu, Terengganu and distribution of C. sworderi (Grismer 2011; Grismer et al. 2007). Despite the recent rapid uptick in herpetological studies and fieldwork in Peninsular Malaysia (see Grismer 2011 for a review), studies in the northeast, principally in the states of Kelantan and Terengganu, remained sparse and the only major peer-reviewed publication from this region was Dring s (1979) report on the herpetofauna of Gunung Lawit, Terengganu. To date, northeastern Peninsular Malaysia remains virtually unexplored from a 240 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

herpetological standpoint with peer-reviewed literature from this region only appearing for selected taxa (i.e., Grismer et al. 2008b). Joint expeditions between the School of Biological Sciences of Universiti Sains Malaysia and the Department of Biology at La Sierra University have been ongoing in various parts of the region since 2007. A summary report of these expeditions is in progress (Quah et al. in prep.) and here we report on some of the findings of our most recent expedition in August of 2012 to Gunung Tebu, Terengganu (Fig. 1). Here we collected five specimens of gekkonid lizards having the diagnostic characters of the genus Cyrtodactylus (see above). Although these specimens showed character state affinities to the sister species (Grismer et al. 2012b) C. quadrivirgatus Taylor (Johnson et al. 2012) and C. sworderi (Smith) (Grismer et al. 2007) they could not be ascribed to either species or to any other known species of Cyrtodactylus. Additionally, they had genetic distances from C. sworderi and C. quadrivirgatus of 13.0 % and 18.6%, respectively, based on the examination 1,502 base pairs of the mitochodrial gene NADH dehydrogenease subunit 2 (ND2) and its flanking trnas. We therefore describe them below as a new species. Materials and methods Morphological analysis. Color notes were taken using digital images of living specimens prior to preservation. The following measurements on the type series were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate: snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90º to the inflection of the dorsiflexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90º to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eyeball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eye ball; eye to snout distance (ES), measured from anteriormost margin of the eyeball to the tip of snout; eye to nostril distance (EN), measured from the anterior margin of the eye ball to the posterior margin of the external nares; interorbital distance (IO), measured between the anterior edges of the orbit; ear length (EL), the greatest horizontal distance of the ear opening; and internarial distance (IN), measured between the nares across the rostrum. Additional character states evaluated on the type series and comparative material (Appendix) were numbers of supralabial and infralabial scales counted from the largest scale immediately anterior to the dorsal inflection of the posterior portion of the upper jaw to the rostral and mental scales, respectively; the presence or absence of tubercles on the anterior margin of the forearm; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column starting at the midpoint between the forelimb insertions and ending at the midpoint between the hind limb insertions; the number of longitudinal rows of body tubercles counted transversely across the center of the dorsum from one ventrolateral fold to the other; the number of longitudinal rows of ventral scales counted transversely across the center of the abdomen from one ventrolateral fold to the other; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the total number of precloacal and femoral pores (i.e. the contiguous rows of femoral and precloacal scales bearing pores are combined as a single meristic); the presence or absence of a precloacal depression or groove; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales; the presence or absence of a white network of lines forming a reticulum on the top of the head; color pattern on body and nape (i.e., striped, banded, or blotched); degree of striping on the flanks and their degree of contact with a postorbital stripe; and the presence or absence of wide, dark bands on an original tail. Some of the information on character states and their distribution in other species was obtained from Chan & Norhayati (2010), Das & Lim (2000), De Rooij (1915), Dring (1979), Grismer et al. (2008a); Hikida (1990), Inger & King (1961), Manthey & Grossman (1997), Rösler & Glaw (2008), Smith (1930), and Taylor (1963). Additional specimens examined are listed in the appendix. Institutional abbreviations follow Sabaj-Pérez (2010), except we retain ZRC (Zoological Refernece Collection, Raffles Museum) for USDZ, following conventional usage. DWNP NEW CYRTODACTYLUS FROM MALAYSIA Zootaxa 3616 (3) 2013 Magnolia Press 241

refers to the Department of Wildlife and National Parks Collection, Krau, Pahang, Malaysia; LSUHC refers to the La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA; and LSUDPC refers to the La Sierra University Digital Photo Collection. Phylogenetic analysis. We obtained sequence data from a 1502 bp fragment of the mitochondrial NADH dehydrogenase subunit 2 (ND2) including the flanking trna s (Trp, Ala, Asn, Cys, Tyr) gene from six ingroup samples and 10 outgroup taxa based on Wood et al. (2012) (Table 1). We obtained 25 additional ingroup taxa from Genbank based on Johnson et al. (2012), Grismer et al. 2012b, and outgroup samples based on Wood et al. 2012 (see Tables 1). All new sequences are deposited in GenBank (Table 1) Total genomic DNA was isolated from liver or skeletal muscle specimens stored in 95% ethanol using the Qiagen DNeasy TM tissue kit (Valencia, CA, USA). ND2 was amplified using a double-stranded Polymerase Chain Reaction (PCR) under the following conditions: 1.0 µl genomic DNA, 1.0 µl light strand primer 1.0 µl heavy strand primer, 1.0 µl dinucleotide pairs, 2.0 µl 5x buffer, MgCl 10x buffer, 0.1 µl Taq polymerase, and 7.56 µl ultra-pure H 2 O. PCR reactions were executed on an Eppendorf Mastercycler gradient theromocycler under the following conditions: initial denaturation at 95 C for 2 min, followed by a second denaturation at 95 C for 35 s, annealing at 47 C for 35 s, followed by a cycle extension at 72 C for 35 s, for 31 cycles. All PCR pruducts were visulalized on a 10 % agarose gel electrophresis. Successful PCR products were vacuum purified using MANU 30 PCR plates (Millipore) and purified products were resuspended in ultra-pure water. Purified PCR products were sequence using the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler. Cycle sequencing reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequence on an ABI 3730xl DNA Analyzer are the BYU DNA Sequencing center. Primers used for amplification and sequencing are presented in Table 2. TABLE 1. A list of the samples used in the molecular analyses with Genbank accesion numbers. Abbreviated voucher numbers are as follows: LSUHC, La Sierra University Herpetological Collection, FMNH, Field Museum of Natural History, ZRC, Zoological Rederence Collection, Raffles Museum of Biodiversity Research. Voucher Species Locality ND2 GenBank Acession Numbers LSUHC 8933 Cyrtodactylus West Malaysia, Melaka, Pulau Besar JQ889178 batucolus LSUHC 8934 C. batucolus West Malaysia, Melaka, Pulau Besar JQ889179 LSUHC6471 C. elok West Malaysia, Pahang, Fraser's Hill, the Gap JQ889180 FMNH 255454 C. interdigitalis Lao PDR, Khammouan Province, Nakai District JQ889181 FMNH 265812 C. intermedius Thailand, Sa Kaeo, Muang Sa Kaeo JQ889182 LSUHC 9513 C. intermedius Thailand, Chantaburi Province JQ889183 LSUHC 9514 C. intermedius Thailand, Chantaburi Province JQ889184 ZRC 2.6951 C. majulah Nee Soon Swamp, Singapore JX988529 ZRC 2.6952 C. majulah Nee Soon Swamp, Singapore JX988530 LSUHC 8906 C. pantiensis West Malaysia, Johor, Gunung Panti FR, Bunker Trail JQ889185 LSUHC 8905 C. pantiensis West Malaysia, Johor, Gunung Panti FR, Bunker Trail JQ889186 LSUHC 10070 C. payacola West Malaysia, Penang, Bukit Panchor JQ889190 LSUHC 10071 C. payacola West Malaysia, Penang, Bukit Panchor JQ889191 LSUHC 9982 C. payacola West Malaysia, Penang, Bukit Panchor JQ889192 LSUHC 10853 C. quadrivirgatus West Malaysia, Terengganu, Gunung Tebu JX988523 LSUHC 9058 C. quadrivirgatus West Malaysia, Terengganu, Pulau Perhentian Besar JQ889260 LSUHC 5633 C. quadrivirgatus West Malaysia, Perak, Temengor, PITC Logging Camp JQ889204 LSUHC 5640 C. quadrivirgatus West Malaysia, Perak, Temengor, PITC Logging Camp JQ889206 LSUHC 7723 C. quadrivirgatus West Malaysia, Johor, Endau-Rompin, Peta, Sungai Semawak JQ889235...continued on the next page 242 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

TABLE 1. (Continued) Voucher Species Locality ND2 GenBank Acession Numbers LSUHC 8127 C. quadrivirgatus West Malaysia, Johor, Selai, Lubuk Tapah Trail JQ889236 LSUHC 9191 C. quadrivirgatus West Malaysia, Perak, Pulau Pangkor JQ889261 LSUHC 8859 C. quadrivirgatus West Malaysia, Perak, Bukit Larut JQ889241 LSUHC 8971 C. quadrivirgatus West Malaysia, Johor, Gunung Ledan NP JQ889257 LSUHC 8970 C. quadrivirgatus West Malaysia, Johor, Gunung Ledan NP JQ889239 LSUHC 6863 C. quadrivirgatus West Malaysia, Kedah, Pulau Langkawi, Gunung Raya JQ889227 LSUHC 6737 C. quadrivirgatus West Malaysia, Penang, Pulau Pinang, Empangan Air Hitam JQ889222 LSUHC 9620 C. quadrivirgatus West Malaysia, Kedah, Sungai Sedim JQ889262 LSUHC 10072 C. quadrivirgatus West Malaysia, Penang, Bukit Panchor JQ889193 LSUHC 10073 C. quadrivirgatus West Malaysia, Penang, Bukit Panchor JQ889194 LSUHC 6461 C. quadrivirgatus West Malaysia, Pahang, Fraser's Hill JQ889207 LSUHC 5022 C. quadrivirgatus West Malaysia, Pahang, Pulau Tioman, Sungai Mentawak JQ889199 LSUHC 5017 C. quadrivirgatus West Malaysia, Pahang, Sungai Lembing Logging Camp JQ889198 LSUHC 4980 C. quadrivirgatus West Malaysia, Pahang, Sungai Lembing Logging Camp JQ889197 LSUHC 6607 C. quadrivirgatus West Malaysia, Pahang, Genting Highlands JQ889218 LSUHC 6503 C. quadrivirgatus West Malaysia, Selangor, Templer's Park JQ889217 LSUHC 4823 C. quadrivirgatus West Malaysia, Selangor, Kepong, FRIM JQ889196 LSUHC 4018 C. quadrivirgatus West Malaysia, Selangor, Kepong, FRIM JQ889195 LSUHC 8900 C. semenanjungensis West Malaysia, Johor, Gunung Panti FR, Bunker Trail JQ889177 LSUHC 6349 C. seribuatebrisis West Malaysia, Johor, Pulau Nangka Kecil JQ889187 FMNH 265806 C. sp. Thailand, Loei, Phu Rua JQ889188 LSUHC 7685 C. sworderi West Malaysia, Johor, Endau-Rompin, Peta, Sungai Kawal JQ889189 LSUHC 10902 C. tebuensis sp. nov. West Malaysia, Terengganu, Gunung Tebu JX988527 LSUHC 10852 C. tebuensis sp. nov. West Malaysia, Terengganu, Gunung Tebu JX988525 LSUHC 10851 C. tebuensis sp. nov. West Malaysia, Terengganu, Gunung Tebu JX988524 LSUHC 10903 C. tebuensis sp. nov. West Malaysia, Terengganu, Gunung Tebu JX988528 LSUHC 10901 C. tebuensis sp. nov. West Malaysia, Terengganu, Gunung Tebu JX988526 TABLE 2. Primer sequences used in this study for the ND2 gene. Primer name Primer reference Sequence L4437b (Macey & Schulte 1999) External 5 -AAGCAGTTGGGCCCATACC-3 CyrtintF1 (Siler et al. 2010) Internal 5 -TAGCCYTCTCYTCYATYGCCC-3 CyrtintR1 (Siler et al. 2010) Internal 5 -ATTGTKAGDGTRGCYAGGSTKGG-3 H5934 (Macey et al. 1997) External 5 - AGRGTGCCAATGTCTTTGTGRTT-3 For the phylogenetic analyses we applied a pluralistic approach using the character based method of Maximum Parsimony (MP) and two model-based methods, Maximum Likelihood (ML) and Bayesian Inference (BI). The Akakine Information Criterion (AIC) as implemented in ModelTest v3.7 (Posada & Crandall 1998), was used to calculate the best-fit model of evolution for each codon position (Table 3). Maximum Parsimony (MP) criteria and bootstrap estimates for nodal support was employed in PAUP* v4.0 (Swofford 2002). One thousand bootstrap replicates for each heuristic search was run with ten random additional sequence replicates using tree bisection and reconnection (TBR) branch swapping. The 1000 bootstrap replicates were summarized as a 50% majority rule consensus tree. Maximum Likelihood analysis was performed using RAxML HPC v7.2.3 (Stamatakis et al. 2008), NEW CYRTODACTYLUS FROM MALAYSIA Zootaxa 3616 (3) 2013 Magnolia Press 243

1000 bootstrap pseudoreplicates via the rapid hill-climbing algorithm (Stamatakis et al. 2008). The Bayesian analysis was carried out in MrBayes v3.1 (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003) using default priors. Two simultaneous runs were performed with eight chains per run, seven hot and one cold following default priors. The analysis was run for 2,000,000 generations and sampled every 200 generations from the Markov Chain Monte Carlo (MCMC). The analysis was halted after the average standard deviation split frequency was below 0.01. The program Are We There Yet? (AWTY) (Nylander et al. 2008) was employed to plot the log likelihood scores against the number of generations to assess convergence and to determine the appropriate number of burnin trees. We conservatively discarded the first 25% of the trees as burnin. A consensus tree was then computed from the two parallel runs using TreeAnnotator v1.6.1 (Drummond & Rambaut 2007). Nodes that had posterior probabilities above 0.95 were considered significantly supported. TABLE 3. Selected models of evolution estimated by the AIC inmplemented in Model test v3.7 (Posada & Crandall 1998). Selcted models were applied when applicable and the next complex model was used if the selected model could not be applied due to computer programing limitations. Gene Model selected Model applied ND2 1 st pos GTR+I+Γ GTR+I+Γ 2 nd pos GTR+I GTR+I 3 rd pos GTR+Γ GTR+Γ trnas HKY+Γ HKY+Γ Results The molecular phylogeny (Fig. 2) supports the morphological data (Table 4) and indicates that the Gunung Tebu population is neither Cyrtodactylus sworderi or C. quadrivirgatus and indicates it is the sister species of the former. The Gunung Tebu population has an uncorrected pairwise sequence divergence of 13.0% from C. sworderi and 18.6% from C. quadrivirgatus. This level of sequence divergence far exceeds that found between multiple species pairs in the C. pulchellus complex and C. majulah clade (Grismer et al. 2012a,b). Based on this and the morphological data, we describe the Gunung Tebu population below as a new species and consider it, C. sworderi, and C. quadrivirgatus to comprise the C. sworderi complex. TABLE 4. Diagnostic characters separating Cyrtodactylus tebuensis sp. nov., C. sworderi, and C. quadrivirgatus. n=no; y=yes; /=data unavailable. tebuensis sp. nov. quadrivirgatus sworderi supralabials 11 13 8 11 10 13 infralabials 8 10 8 11 10 paravertebral tubercles 32 39 32 39 36 38 ventral scales 43 51 28 40 40 49 postfemoral scale contact abrupt y n n 4th toe lamellae 17 21 18 23 16 20 precloacal pores 0 0 12 5 9 precloacal pores in females n n y enlarged precloacal scales y y few enlarged femoral scales y y n femorocloacal scales 31 37 31 44 / white caudal bands narrow (1) or wide (0) 0 0 1 Max. SVL 79.1 71 80.1 244 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

FIGURE 2. Inferred phylogenetic relationships of the Cyrtodactylys sworderi complex based on 1502 bp of the mitochondrial ND2 gene. The tree is a Maximum Likelihood topology (lnl 12919.286678) with Bayesian posterior probabilities (BPP), Maximum Likelihood bootstrap support values (ML), and Maximum Parsimony bootstrap support values (MP), respectively, at the nodes. Cyrtodactylus tebuensis sp. nov. Tebu Mountain Bent-toed Gecko (Figs. 2,3,4) Holotype. Adult male (ZRC 2.6997) from Gunung Tebu, Terengganu, Malaysia (05 36.11 N 102 36.19 E; 650 m) collected by Shahrul Anuar and M. Yusof on 2 September 2012. Paratypes. All paratypes are from the same locality as the holotype. Subadult males ZRC 2.6998-6999 were collected on 31 August 2012 by Shahrul Anuar, Mohd. A. Muin, E. Quah, L. Grismer, B. Beltran, A. Cobos, A. Alonso, C. Thompson, and C. Ogle. Adult male LSUHC 10902 and adult female LSUHC 10903 were collected on the same date as the holotype by E. Quah, L. Grismer, B. Beltran, A. Cobos, A. Alonso, C. Thompson, and C. Ogle. Diagnosis. Cyrtodactylus tebuensis sp. nov. can be distinguished from all other Sundaland species by having the following suite of character states: 73.1 84.1 mm adult SVL; large, conical, keeled body tubercles; tubercles present on top of head, occiput, nape, and limbs and extend posteriorly beyond base of tail; 43 51 ventral scales; no transversely enlrarged, median subcaudal scales; proximal subdigital lamellae transversely expanded; 17 21 NEW CYRTODACTYLUS FROM MALAYSIA Zootaxa 3616 (3) 2013 Magnolia Press 245

subdigital lamellae on fourth toe; abrupt transition between posterior and ventral femoral scales; enlarged femoral scales; no femoral or precloacal pores; precloacal groove absent; and body bearing four wide, bold, dark brown stripes (lateral stripe on each flank and a pair of paravertebral stripes). The meristic characters are scored against C. sworderi and C. quadrivirgatus in Table 4 and against all other Sundaland species in Grismer et al. (2012b:Table 4). Description. Adult male SVL 79.1; head large moderate in length (HL/SVL 0.27) and width (HW/HL 0.63), somewhat depressed (HD/HL 0.41), distinct from neck, and triangular in dorsal profile; lores weakly inflated, prefrontal region slightly concave; canthus rostralis smoothly rounded; snout elongate (ES/HL 0.43) and rounded in dorsal profile; eye large (ED/HL 0.25); ear opening elliptical and small (EL/HL 0.06); eye-to-ear distance greater than diameter of eye; rostral subrectangular with a deep dorsomedial furrow containing a postrostral; rostral partially divided dorsally, bordered posteriorly by large left and right supranasals and one medial postrostrals (=internasal); external nares bordered anteriorly by rostral, dorsally by two supranasals (anterior one largest), posteriorly by five postnasals and ventrally by first supralabial; 11 (R, L) square supralabial scales extending to and tapering smoothly below posterior margin of orbit; 10,9 (R, L) infralabial scales tapering smoothly posteriorly to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small and granular; scales on top of head and occiput intermixed with slightly enlarged tubercles; dorsal and ventral superciliaries rectangular; mental triangular, bordered laterally by first infralabial and posteriorly by left and right rectangular postmentals contacting medially for approximately 50% of their length posterior to mental; one enlarged row of sublabials extending posteriorly to 4 6th infralabial; gular scales small and raised, grading posteriorly into slightly larger, flatter, throat scales, and thence into large, flat, imbricate pectoral and ventral scales. Body relatively short (AG/SVL 0.45) with weak, tuberculate ventrolateral folds; dorsal scales small and granular, interspersed with large, conical, regularly arranged, keeled tubercles; tubercles extending from top of head between eyes to anterior one-fifth of tail; tubercles on top of head, occiput, and nape relatively small, those on body largest; approximately 22 longitudinal rows of dorsal tubercles and 34 paravertebral tubercles; 44 flat, imbricate ventrals, ventrals much larger than dorsals; slightly enlarged patch of precloacal scales lacking pores; precloacal groove absent (Fig. 2). FIGURE 3. Holotype (ZRC 2.6997) of Cyrtodactylus tebuensis sp. nov. 246 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

FIGURE 4. Upper left: subadult male Cyrtodactylus tebuensis sp. nov. (paratype ZRC 2.6998). Upper right: adult female C. tebuensis sp. nov. (paratype LSUHC 10903). Lower left: subadult male C. tebuensis sp. nov. (paratype ZRC 2.6999). Lower right: adult female C. quadrivirgatus (LSUHC 10853) found syntopically with C. tebuensis sp. nov. Forelimbs moderate in stature, relatively short (FL/SVL 0.15); granular scales of forearm larger than those of body and interspersed with tubercles; palmar scales rounded; digits well-developed, inflected at basal interphalangeal joints; subdigital lamellae slightly enlarged proximal to joint inflections, digits narrower distal to joints; claws well-developed, sheathed by a dorsal and ventral scale. Hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.18), covered dorsally by granular scales interspersed with larger tubercles and anteriorly by granular scales; ventral scales of femora flat and larger than dorsals; ventral tibial scales flat and imbricate; a single row of enlarged femoral scales extend from just proximal to the knee medially making contact with the large precloacal scales; femoral pores absent; dorsal and ventral femoral scales meeting abruptly on posteroventral margin of thigh; plantar scales low and slightly rounded; digits well-developed, inflected at basal interphalangeal joints; subdigital lamellae enlarged proximal to inflected joints, digits narrower distal to joints; 17 subdigital lamellae on right 4th toe; claws well-developed, sheathed by a dorsal and ventral scale. Original tail widest at base, tapering to a point; dorsal scales on base of tail granular, becoming flatter posteriorly; no median row of transversely enlarged subcaudal scales; caudal scales arranged in semi-whorls; two enlarged tubercles at base of tail; base of tail with lateral, bulbous swellings in male; all postcloacal scales moderately sized, flat, and imbricate. Coloration in life (Fig. 3). Ground color of head, body, limbs and anterior portion of tail pale yellow; snout, top of head and eyes bearing light brown mottling; body bearing a wide, dark brown lateral stripe on each flank and a pair of dark brown paravertebral stripes beginning at the posterior margin of the eyes and extending posteriorly to the base of the tail; a longitudinal series of yellow spots occurs ventral to the dark lateral stripes; paired, dark brown triangular markings present on the nape followed by a dark brown, azygous irregularly shaped marking; brown irregularly shaped markings on limbs; 12 wide, dark brown to black caudal bands, anterior three bands irregularly shaped; five yellowish caudal bands anteriorly and seven white caudal bands posteriorly; all caudal bands encircle tail; anterior four yellowish bands contain dark blotching; all ventral surfaces beige except for soles NEW CYRTODACTYLUS FROM MALAYSIA Zootaxa 3616 (3) 2013 Magnolia Press 247

of hands and feet and subcaudal region which are slightly darker; a small, single dark spot occurs in all ventral scales except those of tail. Variation. The paratypes very closely resemble the holotype in all aspects of coloration (Fig. 4). ZRC 2.6999 has more evenly edged lateral stripes than the holotype but the nape markings are more irregularly shaped as opposed to being triangular. The dark stripes in LSUHC 10903 are considerably more crenulated and the paravertebral stripes even make contact at various places. Meristc variation is presented in Table 5. TABLE 5. Morphological characters of the type series of the Cyrtodactylus tebuensis sp. nov.. m=male; f=female; n=no; y=yes; /=data unavailable. ZRC ZRC ZRC LSUHC LSUHC 2.6999 2.6998 2.6997 10902 10903 paratype paratype holotype paratype paratype sex m m m m f supralabials 12 13 11 12 11 infralabials 9 9 9 8 10 paravertebral tubercles 36 35 34 39 32 ventral scales 49 43 44 48 51 postfemoral scale contact y y y y y abrupt 4th toe lamellae 19 20 17 20 21 precloacal pores 0 0 0 0 / precloacal pores in females / / / / n enlarged precloacal scales y y y y y enlarged femoral scales y y y y y no. femorocloacal scales 34 34 31 37 / white caudal bands narrow 0 0 0 0 0 (1) or wide (0) max. SVL 53.4 59.8 79.1 70.2 84.1 TL 57.1 65.1 83.4 82 93.2 TW 4.8 5.1 6.3 5.8 4.9 FL 8.1 9.1 11.5 11.5 9.6 TBL 9.5 9.9 13.9 12.3 10.6 AG 22.5 26.5 35.3 30.2 28.2 HL 14.8 16.4 21.2 18.8 17.2 HW 9.6 10.6 13.3 11.9 11.1 HD 5.8 6.8 8.7 8.1 6.5 ED 3.4 3.3 5.2 4.3 3.8 EE 4.5 5.3 6.3 5.6 4.9 ES 6.3 6.7 9.1 7.7 7.1 EN 4.8 5.5 6.9 6.1 5.1 IO 2.8 3.3 4.1 3.8 3.1 EL 1.2 1.2 1.3 1.4 1.2 IN 1.8 1.7 2.5 2.1 1.9 Distribution. Currently Cyrtodactylus tebuensis sp. nov. is known only from the type locality of Gunung Tebu, Terengganu, Malaysia (Fig. 1). It is expected to range more widely throughout the range of mountains of which Gunung Tebu is a part. 248 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

Natural history. All specimens were found active at night in primary hill dipterocarp forest (Fig. 5) on vegetation from 1.5 2 m above the ground including thin horizontal branches and the trunks of trees with their head facing towards as has been reported for their closest relative C. sworderi (Grismer et al. 2007). Syntopic with this species is C. quadrivirgatus. LSUHC 10903 was a gravid female carrying two eggs. Etymology. The specific epithet tebuensis refers to Gunung Tebu (Cane Mountain), in which the type locality is situated. FIGURE 5. Left: microhabitat in hill dipterocarp forest near the type locality of Cytrodactylus tebuensis sp. nov. Right: Trail along which the type series of C. tebuensis sp. nov. and C. quadrivirgatus were collected. Discussion As constituted here, the Cyrtodactylus sworderi complex comprises C. sworderi, C. tebuensis sp. nov., and C. quadrivirgatus. All are relatively robust, tuberculate, scansorial forest dwellers that collectively range from the Isthmus of Kra, Thailand southward through Peninsular Malaysia to Sumatra (Grismer 2011) but do not occur in Singapore (Grismer et al. 2012b). This complex is characterized by species having the unique combination of 8 13 supralabials, 8 10 infralabials, 32 39 paravertebral tubercles, 28 51 ventral scales, 16 23 subdigital lamellae on the fourth toe, 0 15 precloacal pores, and a maximun SVL ranging from 71.0 80.1. The unexplored areas of northeastern Peninsular Malaysia will undoubtedly prove to be a warehouse of a significant portion of Malaysia s herpetological diversity and, based on studies from other upland areas (i.e. Dring 1979; Grandison 1972; Grismer et al. 2010b and references therein) these mountain systems (collectively known as the Banjaran Timur) will harbor an untold number of yet to be discovered endemics. Current species of amphibians and reptiles known to range only in these systems are Kalophrynus robinsoni Smith, Psedocalotes dringi Hallermann & Böhme, Cnemaspis argus Dring, Sphenomorphus cophias Boulenger, Macrocalamus vogeli David NEW CYRTODACTYLUS FROM MALAYSIA Zootaxa 3616 (3) 2013 Magnolia Press 249

& Pauwels and to this list we add Cyrtodactylus tebuensis sp. nov. Additionally from Gunung Tebu, we collected a new species of the bufonid Ansonia Stoliczka, the megophryid Leptolalax Dubois, and the colubrid Pseudorhabdion Leviton & Brown whose descriptions are in progress. Dring (1979) reported on an extensive herpetofaunal survey from Gunung Lawit, Terengganu approximately 10 km due south of Gunung Tebu in the same mountain range. Curiously, his team found no Cyrtodactylus tebuensis sp. nov. but found C. quadrivirgatus (identification confirmed here LSUDPC 6715 24) to be relatively common from 43 790 m in elevation. This is similar to our findings in that we found C. quadrivirgatus on Gunung Tebu from 30 m to nearly 700 m in elevation. This could indicate that C. tebuensis sp. nov. is more localized than hypothesized above. In any event all these data underscore the unexplored nature of these upland systems and need for additional field work. Acknowledgments For field assistance we thank A. Alonso, A. Cobos, B. Beltran, C. Ogle, and C. Thompson, and M. Yusof. For the loan of specimens we are indebted to Kelvin K. P. Lim (ZRC). We thank the Terengganu State Forestry Department for their permission to conduct research in the Gunung Tebu Forest reserve. A research pass (40/200/19 SJ.1105) was issued to LLG by the Economic Planning Unit, Prime Minister s Department, Malaysia. This research was supported in part by grants to LLG from the College of Arts and Sciences, La Sierra University, Riverside, California and by a Universiti Sains Malaysia grant 815075 to Professor Shahrul Anuar. DNA sequencing was supported by Dr. Jack W. Sites Jr. and the department of biology at Brigham Young University. Field work for EQSH in Malaysia was partially supported by the USM Fellowship Scheme. References Chan, K.O. & Norhayati, A. (2010) A new species of Cyrtodactylus (Squamata: Gekkonidae) from northeastern Peninsular Malaysia, Malaysia. Zootaxa, 2389, 47 56. Das, I. & Lim, L.J. (2000) A new species of Cyrtodactylus (Sauria: Gekkonidae) from Pulau Tioman, Malaysia. The Raffles Bulletin of Zoology, 48, 223 231. David, P., Ngyuen, T.Q., Schneider, N. & Ziegler, T. (2011) A new species of the genus Cyrtodactylus Gray, 1827 from central Laos (Squamata: Gekkonidae). Zootaxa, 2833, 29 40. De Rooij, N. (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill Ltd., Leiden, 384 pp. Dring, J.C.M. (1979) Amphibians and reptiles from northern Trengganau, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum (Natural History), 34, 181 241. Drummond, A. & Rambaut, A. (2007) Beast: Bayesian Evolutionary Analysis by Sampling Trees. BMC Evolutionary Biology, 7, 214. http://dx.doi.org/10.1186/1471-2148-7-214 Grandison, A.G.C. (1972) The Gunong Benom Expedition 1967. 5. Reptiles and amphibians of Gunong Benom with a description of a new species of Macrocalamus. Bulletin of the British Museum of Natural History (Zoology), 23, 45 101. Grismer, L.L. (2005) New species of Bent-Toed Gecko (Cyrtodactylus Gray 1827) from Pulau Aur, Johor, West Malaysia. Herpetologica, 39, 424 432. http://dx.doi.org/10.1670/3-05a.1 Grismer, L.L. (2011) Lizards of Peninsular Malaysia, Singapore and Their Adjacent Archipelagos. Edition Chimiara, Frankfürt am Main, 728 pp. Grismer, L.L. & Leong, T.Z. (2005) New species of Cyrtodactylus (Squamata: Gekkonidae) from southern Peninsular Malaysia. Journal of Herpetology, 39, 584 591. http://dx.doi.org/10.1670/43-05a.1 Grismer, L.L., Chan, K.O., Grismer, J.L., Wood, P.L. Jr. & Belabut, D. (2008a) Three new species of Cyrtodactylus (Squmata: Gekkonidae) from Peninsular Malaysia. Zootaxa, 1921, 1 23. Grismer, L.L., Chan, K.O., Grismer, J.L., Wood, P.L., Jr. & Norhayati, A. (2010b) A checklist of the herpetofauna of the Banjaran Bintang, Peninsular Malaysia. Russian Journal of Herpetology, 17, 147 160. Grismer, L.L., Grismer, J.L., Wood, P.L.Jr., & Chan, K.O. (2008b) The distribution, taxonomy, and redescription of the geckos Cnemaspis affinis (Stoliczka 1887) and C. flavolineata (Nicholls 1949) with descriptions of a new montane species and two new lowland, karst-dwelling species from Peninsular Malaysia. Zootaxa, 1931, 1 24. Grismer, L.L. & Norhayati, A. (2008) A new insular species of Cyrtodactylus (Squamata: Gekkoidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia. Zootaxa, 1924, 53 68. Grismer, L.L., Shahrul, A.M. Quah, E.S., Muin, M.A., Chan, K.O., Grismer, J.L. & Norhayti, A. (2010a) A new spiny, prehensiletailed species of Cyrtodactylus (Squamata: Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa, 2625, 40 52. 250 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.

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APPENDIX Cyrtodctylus astrum Grismer et al. Peninsular Malaysia, Perlis, Gua Kelam: LSUHC 8806 10; Peninsular Malaysia, Perlis, Kuala Perlis: LSUHC 8815 16. Cyrtodactylus aurensis Grismer Peninsular Malaysia, Johor, Pulau Aur: LSUHC 3937, 7013, 7022 23, 7285 86, 7300, 8040. Cyrtodactylus baluensis (Mocquard) Malaysian Borneo, Sabah, Kinabalu: ZRC 2.5744, 2.5747; Malaysian Borneo, Sabah: ZRC 2.1116 21; Brunei Darussalam, Temburong, Kuala Belalong: ZRC 2.4729. Cyrtodactylus batucolus Grismer et al. Peninsular Malaysia, Melaka, Pulau Besar, Pantai Putera: LSUHC 8933, 8935 36, 8939, 8941 43 (type series). Cyrtodactylus brevipalmatus (Smith) Thailand (no data): LSUHC 1899. Cyrtodactylus cavernicolus Inger & King - Malaysian Borneo, Sarawak, Niah Cave: LSUHC 4055 56, ZRC 2.5327, 2.5775. Cyrtodactylus consobrinus (Smith) Peninsular Malaysia, Selangor, Kepong, Forest Research Institute Malaysia: LSUHC 4019, 4389, 4820 21; Peninsular Malaysia, Pahang, Sungai Lembing logging camp: LSUHC 4912, 4941 12; Malaysian Borneo, Sarawak, near Niah Cave: LSUHC 4062. Cyrtodactylus elok Dring Peninsular Malaysia, Pahang, Raub, Lakum Forest Reserve: ZRC 2.5649, 2.5668. Cyrtodactylus fumosus (Müller) Indonesia, north Sulawesi, Lembeh Island: FMNH 14249 50. Cyrtodactylus ingeri Hikida Malaysian Borneo, Sabah, Turtle Island: LSUHC 6167 76; Malaysian Borneo, Sabah, Poring: ZRC 2.5957. Cyrtodactylus jarakensis Grismer et al. Peninsular Malaysia, Perak, Pulau Jarak: LSUHC 9111 (holotype). Cyrtodactylus majulah Grismer et al. Singapore, Nee Soon Swamp-forest: ZRC 2.1133, 2.3190, 2.3205, 2.2361, 2.3283 84, 2.4576, 2.4862, 2.552, 2.5643, 2.6796, 2.6555, and 2.6950 53; LSUHC 10458. Cyrtodactylus malayanus (de Rooij) Malaysian Borneo, Sarawak, Kapit District, Nanga Tekalit Camp along Mengiong River: FMNH 149154, 149231. Cyrtodactylus marmoratus (Kuhl) Indonesia, west Java: ZRC 2.1134. Cyrtodactylus matsuii Hikida Malaysian Borneo, Sabah, Crocker Range: ZRC 2.5761. Cyrtodactylus oldhami (Theobald) Thailand, Phuket, Kathu: ZRC 2.5213, 2.5226, 2.5230. Cyrtotactylus pantiensis Grismer et al. Peninsular Malaysia, Johor, Gunung Panti Forest Reserve: LSUHC 8904 07 (type series). Cyrtodactylus payacola Johnson et al. LSUHC 10070 71, 10074, 10076 Bukit Panchor State Park, Penang, Malaysia. Cyrtodactylus pubisulcus Inger Malaysian Borneo, Sarawak, near Niah Cave: LSUHC 4069. Cyrtodactylus pulchellus Gray Peninsular Malaysia, Pahang, Genting Highlands: LSUHC 6637; Peninsular Malaysia, Perak, Bukit Larut: ZRC 2.5656 60, 2.5666; Peninsular Malaysia, Pulau Pinang, Ampangan Air Hitam: LSUHC 6668, 6725 29, 6785, 6829; Cyrtodactylus quadrivirgatus Taylor Peninsular Malaysia, Selangor, Kepong, Forest Research Institute Malaysia: LSUHC 4018, 4823; Peninsular Malaysia, Perak, Bukit Larut: ZRC 2.4865 66; Peninsular Malaysia, Pahang, Pulau Tioman: LSUHC 4813, 5022, 5101, 5173, 5517, 5562, 5582, 6136, 6146; Peninsular Malaysia, Pahang, Sungai Lembing logging camp: LSUHC 4980, 5017; Peninsular Malaysia, Perak, Temengor, PITC logging camp: LSUHC 5633 34, 5640; Peninsular Malaysia, Pahang, Pekan: LSUHC 6069, 6986, 6106. Cyrtodactylus sadleiri Wells & Wellington Australia, Indian Ocean, Christmas Island: ZRC 2.1158, 2.1154 55, 2.1164 65. Cyrtodactylus semenanjungensis Grismer & Leong Peninsular Malaysia, Johor, Gunung Panti foothills: ZRC 2.5603, 2.5627; Peninsular Malaysia, Johor, near Jemaluang: ZRC 2.5077 78. Cyrtodactylus seribuatensis Youmans & Grismer Peninsular Malaysia, Pahang, Pulau Seribuat: LSUHC 5191 52, 5218 31, 5243, 5578; Peninsular Malaysia, Pahang, Pulau Sembilang: LSUHC 5522; Peninsular Malaysia, Johor, Pulau Sibu: LSUHC 5604, 5778 80, 5784 89; Peninsular Malaysia, Johor, Pulau Sibu Tengah: LSUHC 5812 13. Cyrtodactylus tiomanensis Das & Lim Peninsular Malaysia, Pahang, Pulau Tioman: LSUHC 3771 72, 3793 94, 3847, 4581 84, 4587, 4590, 4597, 5027 30, 5044, 5411, 5479, 5512, 5519 20, 6250 51, 6268. Cyrtodactylus yoshii Hikida Malaysian Borneo, Sabah, Poring: ZRC 2.4851. 252 Zootaxa 3616 (3) 2013 Magnolia Press GRISMER ET AL.