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Zootaxa 3760 (1): 067 078 www.mapress.com/zootaxa/ Copyright 2014 Magnolia Press Article http://dx.doi.org/10.11646/zootaxa.3760.1.4 http://zoobank.org/urn:lsid:zoobank.org:pub:18b56f00-45b7-4f46-a9d5-a8420fca7eba ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from northwestern Thailand L. LEE GRISMER 1,2, PERRY L. WOOD, JR. 3 & MICHAEL COTA 4,5 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 Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: pwood@byu.edu 4 Natural History Museum, National Science Museum, Thailand, Technopolis, Khlong 5, Khlong Luang, Pathum Thani 12120 Thailand. E-mail: herpetologe@gmail.com 5 Faculty of Science and Technology, Suan Sundandha Rajabhat University, 1 U-thong Nok Rd, Dusit, Bangkok 10300 Abstract A new species of gekkonid, Hemiphyllodactylus chiangmaiensis sp. nov., from northwestern Thailand is separated from all other species of Hemiphyllodactylus by a set of features including: a maximum SVL of 41.2 mm; 8 12 chin scales extending transversely from unions of second and third infralabials and posterior margin of mental; lamellar formula on hand 3 3 3 3 or 3 4 3 3; lamellar formula on foot 3 3 3 3 or 3 4 4 4; continuous precloacal and femoral pores; a unique dorsal color pattern; and caecum and oviducts pigmented. These characters place this species in the speciose H. typus group. Hemiphyllodactylus chiangmaiensis sp. nov. fills a biogeographical hiatus in the distribution of this genus across northern Indochina. Key words: Gekkonidae, Hemiphyllodactylus, Hemiphyllodactylus chiangmaiensis sp. nov., Thailand, Chiang Mai, new species Introduction The gekkonid genus Hemiphyllodactylus Bleeker, 1860 currently composes 22 confirmed species (Grismer et al. 2013) that collectively extend from the Mascarene Islands in the western Indian Ocean, eastward through southern Asia and Indochina. From here the genus ranges southward through the Philippines and Sundaland, through the Indo-Australian Archipelago, and continues into much of Oceania to as far eastward as Hawaii. Many of these species are geographically restricted upland or insular populations ranging throughout mainland Asia or are restricted to islands in western Indonesia and the Philippines. Grismer et al. (2013) demonstrated that Hemiphyllodactylus was far more diverse than the most recent taxonomic revision based solely on morphology (see Zug 2010) indicated. Ten of the 22 species they identified were done so on the basis of genetic and or preliminary morphological evidence and their descriptions were deferred to subsequent, more in depth morphological analyses. One of these species from Chiang Mai, Chiang Mai Province in northwestern Thailand (Fig. 1), was originally identified as H. yunnanensis (Zug 2010) but is actually the sister species to a lineage containing H. longlingensis Zhou & Liu and an unnamed species from Mandalay Division, Myanmar (Grismer et al. 2013: Hemiphyllodactylus sp. nov. 8 in Figure 2). We were able to examine nine specimens from Chiang Mai and present here morphological data supporting the molecular phylogenetic analysis that initially indicated this population was a distinct species. It is described below. Accepted by A. Bauer: 4 Dec. 2013; published: 30 Jan. 2014 67

FIGURE 1. Map showing the type locality of Hemiphyllodactylus chiangmaiensis sp. nov. at Chiang Mai, Chiang Mai Province, Thailand. Materials and methods Color notes were taken using digital images of some specimens prior to preservation. For purposes of comparison the terminology and methodology involving the evaluation of mensural and meristic characters follows Grismer et al. (2013) which is generally based on Zug (2010). Mensural data 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 (TailL), taken from the vent to the tip of the tail, original or regenerated; trunk length (TrunkL), taken from the posterior margin of the forelimb at its 68 Zootaxa 3760 (1) 2014 Magnolia Press GRISMER ET AL.

FIGURE 2. Maximum Likelihood phylogram (-In L 22097.690183) of the genus Hemiphyllodactylus Bleeker, 1860 with Bayesian posterior probabilities and Maximum Likelihood bootstrap values, respectively after Grismer et al. (2013). A NEW SPECIES OF HEMIPHYLLODACTYLUS Zootaxa 3760 (1) 2014 Magnolia Press 69

insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HeadL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HeadW), measured at the angle of the jaws; eye diameter (EyeD), the greatest horizontal diameter of the eyeball; snout-eye length (SnEye), measured from anteriormost margin of the eyeball to the tip of snout; nareseye length (NarEye), measured from the anterior margin of the eye ball to the posterior margin of the external nares; and internarial width (SnW), measured between the nares across the rostrum. Meristic character states evaluated on the holotype and comparative material (see Appendix; Zug [2010]) were the number of scales contacting the nares (circumnasal scales); the number of scales between the supranasals (postrostrals); the numbers of supralabial and infralabial scales counted from the largest scale immediately posterior to the dorsal inflection of the posterior portion of the upper jaw to the rostral and mental scales, respectively; the number of longitudinal ventral scales at midbody contained within one eye diameter; the number of longitudinal dorsal scales at midbody contained within one eye diameter; the number of subdigital lamellae wider than long beneath the first finger and toe; lamellar formulae determined as the number of U-shaped subdigital lamellae on the digital pads on digits 2 5 of the hands and feet; the total number of precloacal and femoral pores (i.e. the contiguous or discontinuous rows of femoral and precloacal scales bearing pores); and the number of cloacal spurs. Color pattern characters evaluated were the presence or absence of dark pigmentation in the gonadal tracts and caecum; presence or absence of a dark postorbital stripe extending to at least the neck; and the presence or absence of a linear series of white postorbital spots above the dark postorbital stripe. Some of the information on character states and their distribution in other species was obtained from Zug (2010). THNHM refers to the Natural History Museum, National Science Museum, Thailand, Pathum Thani, Thailand; 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. Systematics Hemiphyllodactylus chiangmaiensis sp. nov. Chiang Mai Dwarf Gecko Ching-chok-khaosung-chiang-mai Figs. 3,4 Holotype. Adult male (THNHM 15194) collected in the vicinity of Chiang Mai, Chiang Mai Province, Thailand. Precise locality, collector, and date of collection unknown. Paratypes. THNHM 15192 93, 15195 200 bear the same data as the holotype. Diagnosis. Hemiphyllodactylus chiangmaiensis sp. nov. can be separated from all other species of Hemiphyllodactylus by having the unique combination of a maximum SVL of 41.2 mm; 8 12 chin scales extending transversely from unions of second and third infralabials and posterior margin of mental; enlarged postmental scales; three or four circumnasal scales; 1 3 scales between supranasals (=postrostrals); 9 11 supralabials; 9 11 infralabials; 11 21 longitudinally arranged dorsal scales at midbody contained within one eye diameter; 6 10 longitudinally arranged ventral scales at midbody contained within one eye diameter; lamellar formula on hand 3 3 3 3 or 3 4 3 3; lamellar formula on foot 3 3 3 3 or 3 4 4 4; continuous precloacal and femoral pores; dorsal body pattern consisting of dark, irregularly shaped, paravertebral blotches; postsacral mark cream-colored, bearing anteriorly projecting arms; and caecum and oviducts pigmented. These characters and potentially diagnostic morphometric characters are scored across all species in Table 1. The taxonomy of H. yunnanensis follows Zug (2010). Description of holotype. Adult male; head triangular in dorsal profile depressed, distinct from neck; lores and interorbital regions flat; rostrum relatively long (NarEye/ HeadL = 0.32); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening oval, small; eye to ear distance greater than diameter of eye; rostral wider than high, partially divided dorsally, bordered posteriorly by large supranasals; three internasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial 70 Zootaxa 3760 (1) 2014 Magnolia Press GRISMER ET AL.

(=circumnasals 3R,L); 9 (R,L) square supralabials tapering to below posterior margin of orbit; 10 (R,L) square infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest; dorsal superciliaries flat, rectangular, imbricate; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single sublabial; row of smaller scales extending transversely from juncture of second and third infralabials and contacting mental; gular scales triangular small, granular, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals. FIGURE 3. Holotype (upper) and type series (lower) of Hemiphyllodactylus chiangmaiensis sp. nov. Body somewhat elongate, dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 11 scales contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, seven scales contained within one eye diameter; no enlarged, precloacal scales; 25 pore-bearing scales extending from midway between the knee and hind limb insertion of one leg to the other; forelimbs short, robust in stature, covered with granular scales dorsally and with slightly larger, flat, subimbricate scales ventrally; palmar scales flat, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits II V 3 4 3 3 (R,L); four transversely expanded lamellae on digit I; claws on digits II V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with slightly pointed, juxtaposed scales dorsally and by A NEW SPECIES OF HEMIPHYLLODACTYLUS Zootaxa 3760 (1) 2014 Magnolia Press 71

larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits II V 3 3 3 3 (R,L); four transversely expanded lamellae on digit I; claws on digits II V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; posterior section of tail broken, round in cross-section; all caudal scales flat, imbricate, not forming distinct caudal segments. Morphometric data are presented in Table 2. Coloration in alcohol (Fig. 3). Top of head and body nearly unicolor dull brown with weak, darker mottling; faint, dark, diffuse preorbital stripe; slightly more prominent postorbital stripe extends to occipital region, becoming more faint and diffuse and wider as it continues posteriorly onto body; faint, dark, dorsal, paravertebral markings weakly counter-shaded with lighter coloration; light postsacral marking bearing anteriorly projecting arms and dark medial chevron; dorsal surface of limbs same color as body, nearly unicolor; original portion of tail bearing irregular shaped, alternating, diffuse light and dark bands. Variation. The paratypes closely match the holotype in coloration and pattern (Fig. 3). THNHM 15198 has a faint, wide, lateral stripe extending from the posterior margin of the eye to the groin. Coloration of this species in life is much more rich (Fig. 4). Variation in scale counts and morphometric data are presented in Table 2. FIGURE 4. Hemiphyllodactylus chiangmaiensis sp. nov. (LSUDPC 6668) from the type locality of Chiang Mai, Chiang Mai Province, Thailand. Distribution. Hemiphyllodactylus chiangmaiensis sp. nov. is known only from the type locality near the city of Chiang Mai in Chiang Mai Province (Fig. 1) but is expected to range more widely throughout the northwestern portion of Thailand and perhaps into Myanmar. Natural History. Hemiphyllodactylus chiangmaiensis sp. nov. occurs in rocky areas in the vicinity of 600 m in elevation, where it takes refuge between rocks when threatened at night and during the day. This species is also found on the vertical surfaces of man-made structures, which they occupy when the human commensals Hemidactylus and Gehyra are absent. Etymology. This species is named after the Thai Province of Chiang Mai where the type locality is located. 72 Zootaxa 3760 (1) 2014 Magnolia Press GRISMER ET AL.

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Comparisons. The taxonomy of Zug (2010) is used in the comparisons below for H. titiwangsaensis Zug, H. typus Bleeker and H. yunannensis (Boulenger) except for H. zugi Nguyen, Lehmann, Le, Duong, Bonkowski & Ziegler which has been removed from the latter species (Nguyen et al. 2013). Hemiphyllodactylus chiangmaiensis sp. nov. differs from H. ganoklonis Zug in having a maximum known SVL of 41.2 mm versus 34.2 mm and from H. margarethae Brongersma, H. titiwangsaensis, H. typus, and H. yunnanensis by having a maximum SVL less than 46.1 mm 49.3 mm. It differs from H. aurantiacus Beddome, H. ganoklonis, and H. insularis Taylor in having enlarged as opposed to small postmentals. Hemiphyllodactylus chiangmaiensis sp. nov. has three or four circumnasal scales that separates it from H. tehtarik which has five and is further separated by having 6 10 as opposed to 12 ventral scales. Hemiphyllodactylus chiangmaiensis sp. nov. has a lamellar hand formula of 3 3 3 3 or 3 4 3 3 which separates it from H. aurantiacus (2 2 2 2), H. ganoklonis (3 4 4 3), H. margarethae (4 4 4 4), H. titiwangsaensis and H. typus (3 4 4 4). From H. titiwangsaensis and H. typus, H. chiangmaiensis sp. nov. differs in having three or four transversely expanded subdigital lamellae beneath digit 1 on the hand as opposed to 5 8. It can be separated further from H. harterti (Werner) in having 17 25 continuous femoral and precloacal pores as opposed to 42 45. The caecum and gonadal tracts of H. chiangmaiensis sp. nov. are pigmented, further differentiating it from H. harterti, H. insularis, some H. margarethae, H. tehtarik, H. titiwangsaensis, and H. yunnanensis. Hemiphyllodactylus chiangmaiensis sp. nov. differs from H. zugi in having a smaller maximum SVL (41.2 versus 46.6 mm); 6 10 versus 15 or 16 ventral scales; having a 3 3 3 3 or 3 3 4 3 versus a 3 4 4 4 lamellar formula on the hand; having as opposed to lacking dark dorsal transverse blotches on the body; and a pigmented caecum and gonads. From H. larutensis Boulenger, H. chiangmaiensis is separated on the basis of having a maximin SVL of 41.2 versus 52.2 mm; 17 25 continuous femoral and precloacal pores as op[p[osed to 27 36; one as opposed to two or three cloacal spurs on each side; having a banded to blotched dorsal pattern as opposed to a unicolor dorsal pattern; and a pigmented caecum and gonads as opposed to these structures being unpigmented. Four morphometric ratios, HeadL/SVL, HeadW/SVL, HeadW/HeadL, and EyeD/HeadL of other species of Hemiphyllodactylus differ discretely from the corresponding ratios in H. chiangmaiensis sp. nov. (Table 1). Discussion All well-studied continental populations of Hemiphyllodactylus are generally upland species with restricted distributions (Grismer et al. 2013). Thus, the presence of an endemic Hemiphyllodactylus in the uplands of northern Thailand is not surprising given that this area has a number of other endemic species as well as geographic variants that may themselves represent distinct lineages (see Chan-ard 2003; Chan-ard et al. 2011; Cox et al. 1998; Das 2010; Manthey & Grossmann 1997; Matsui et al. 1998; Nabhitabhata et al. 2000; Nutphund 2001 Taylor 1962, 1963, 1965). Many of these, such as H. chiangmaiensis sp. nov., occupy niches in montane regions that are filled by related taxa elsewhere and fill a biogeographical gap across northern Indochina (Wood & Grismer in prep.). References Chan-ard, T. (2003) A Photographic Guide to Amphibians in Thailand. Darnsutha Press, Bangkok, Thailand, 176 pp. Chan-ard, T., Cota, C. & Makchai, S. (2011) Amphibians of Eastern Thailand and Checklist of Thailand. National Science Museum, Ministry of Science and Technology, Bangkok, Thailand, 160 pp. Chan-ard, T., Grossmann, W., Gumprecht, A. & Schulz, K.-D. (1997) Amphibians and Reptiles of Peninsular Malaysia and Thailand. Bushmaster Publications, Wuerselen, 240 pp. Cox, M.J., van Dijk, P.P., Nabhitabhata, J. & Thirakhupt, K. (1998) A Photographic Guide to Snakes and Other Reptiles of Peninsular Malaysia, Singapore and Thailand. New Holland Publishers (UK) Ltd., London, 144 pp. Das, I. (2010) A Field Guide to the Reptiles of South-East Asia. New Holland Publishers, United Kingdom, 376 pp. Grismer, L.L., Wood, P.L., Anuar, S., Mohd., M.A., Quah, E. H., McGuire, J.A., Brown, R.M., Ngo, V.T. & Hong, P. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169 (4), 849 880. http://dx.doi.org/10.1111/zoj.12064 Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Südostasiens. Natur und Tier Verlag, Münster, 512 pp. Matsui, M., Nabhitabahata, J. & Panha, S. (1998) A new Ansonia from northern Thailand (Anura: Bufonidae). Herpetologica, 54, 448 454. A NEW SPECIES OF HEMIPHYLLODACTYLUS Zootaxa 3760 (1) 2014 Magnolia Press 77

Nabhitabhata, J. & Chan-ard, T. & Chuaynkern, Y. (2000) Checklist of Amphibians and Reptiles of Thailand. Environmental Policy and Planning, Bangkok, Thailand, 152 pp. Nutphund, W. (2001) Amphibians of Thailand. Amarin Printing and Publishing Public Co. Ltd., Bangkok, Thailand, 191 pp. Taylor, E.H. (1962) The amphibians of Thailand. University of Kansas Science Bulletin, 43, 265 599. Taylor, E.H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 667 1077. Taylor, E.H. (1965) The serpents of Thailand and adjacent waters. University of Kansas Science Bulletin, 45, 609 1096. Zug, G.R. (2010) Speciation and dispersal in a low diversity taxon: the Slender geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, 631, 1 70. http://dx.doi.org/10.5479/si.00810282.631 78 Zootaxa 3760 (1) 2014 Magnolia Press GRISMER ET AL.