The phylogenetic relationships of three new species of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae) from poorly explored regions in northeastern Peninsular Malaysia Author Grismer, L. Lee Author Wood, Perry L. Author Anuar, Shahrul Author Quah, Evan S. H. Author Muin, Mohd Abdul Author Mohamed, Maketab Author Onn, Chan Kin Author Sumarli, Alexandra X. Author Loredo, Ariel I. Author Heinz, Heather M. text Zootaxa 2014 3786 3 359 381 journal article 46000 10.11646/zootaxa.3786.3.6 839c26e6-bb6f-4258-84f3-75174e7fabdb 1175-5326 250549 98B6A0EF-CAFC-4A78-B044-C5B9D6136CFB Cyrtodactylus timur sp. nov. Banjaran Timur Bent-toed Gecko Figs. 5 , 6 Holotype . Adult male LSUHC 11207 collected on 1 July 2013 by Shahrul Anuar at 2350 hrs from Punca Air, Gunung Tebu, Terengganu, Peninsular Malaysia (5° 36.11 N 102° 36.19 E ; at 650 m elevation). Paratypes . LSUHC 11183–85 have the same collection data as the holotype . LSUHC 11288 has the same collection data as the holotype except that it was collected on 2 July 2013 . Diagnosis. Cyrtodactylus timur sp. nov. can be differentiated from all other species of Cyrtodactylus by having 10–12 supralabials; 8–10 infralabials; weak body tuberculation; no tubercles on ventral surface of forelimbs, gular region, or in ventrolateral body folds; 38–43 paravertebral tubercles; 21–24 longitudinal tubercle rows; 31–40 ventral scales; 21–25 subdigital lamellae on fourth toe; 21 or 22 femoroprecloacal pores; deep precloacal groove; four dark dorsal body bands; body band/interspace ratio 1.00–1.25; no scattered white tubercles on dorsum; 8–10 dark caudal bands on original tail; white caudal bands nearly immaculate; and a maximum SVL of 120.5 mm . These characters are scored across all species of the C. pulchellus complex in Table 5. Description of holotype . Adult male SVL 117.7 mm ; head large, moderate in length (HL/SVL 0.28), wide (HW/HL 0.73), somewhat flattened (HD/HL 0.41), distinct from neck, triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded anteriorly; snout elongate (ES/HL 0.43), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.23); ear opening subelliptical, moderate in size (EL/HL 0.05), vertically oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by a small azygous scale; an inverted Y-shaped furrow, bordered posteriorly by left and right supranasals, and one medial postrostral (=internasal), bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large anterior supranasal and smaller posterior supranasal, posteriorly by one postnasal, ventrally by first supralabial; 12(R), 10(L) rectangular supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; first supralabial largest; 8(R,L) infralabials tapering in size posteriorly; scales of rostrum and lores flat, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales on posterior portion of occiput intermixed with small tubercles; no tubercles on top of head between eyes; large, boney frontal ridges bordering orbit confluent with boney, transverse, parietal ridge; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by left and right, rectangular postmentals which contact medially for 40% of their length; single row of slightly enlarged, elongate sublabials extending posteriorly to third infralabials; small, granular to flat, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales. FIGURE 5. Upper left: adult male holotype (LSUHC 11207) of Cyrtotdactylus timur sp. nov. from Punca Air, Gunung Tebu, Terengganu, Peninsular Malaysia. Upper right: female paratype LSUHC 11185. Middle left: uncataloged hatching LSUDPC 8249. Lower left: adult male paratype LSUHC 11183. Lower right: juvenile female paratype LSUHC 11184. Body relatively short (AG/SVL 0.46) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with larger, unicainate, regularly arranged, low tubercles; smaller intervening tubercles absent; tubercles extend from occiput to caudal constriction and onto base of tail where they fade out; caudal tubercles absent from posterior nine-tenths of tail; tubercles on occiput and nape small, those on body larger; approximately 21 longitudinal rows of tubercles at midbody; 38 paravertebral tubercles; 31 flat imbricate ventral scales between ventrolateral body folds, ventral scales larger than dorsal scales; precloacal scales large, smooth. Forelimbs moderate, relatively short (FL/SVL 0.18); scales on dorsal surfaces of forelimbs juxtaposed, intermixed with larger tubercles; scales of ventral surface of forearm flat, subimbricate, lacking tubercles; palmar scales weakly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae rectangular proximal to joint inflection, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.21), larger tubercles on dorsal surface of thigh separated by smaller granular scales, tubercles on dorsal surfaces of foreleg same size as those on thigh; ventral scales of thigh flat, smooth, imbricate, larger than dorsal granular scales; ventral, tibial scales flat, smooth, imbricate; single row of greatly enlarged, flat, rectangular, imbricate, femoral scales extend from knee to knee through precloacal region where they are continuous with enlarged, precloacal scales; 21 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which 12 pore-bearing scales are found (six on left and right); postfemoral scales immediately posterior to enlarged scale row smaller, forming an abrupt union on posteroventral margin of thigh; plantar scales low, slightly raised; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae proximal to joint inflection rectangular, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; 20(R) 21(L) subdigital lamellae on 4th toe. Tail original, 143.3 mm in length, 11.2 mm in width at base, tapering to a point; dorsal scales of tail flat, nearly square; tail weakly segmented with 8–10 transverse scale rows per segment; subcaudal region bearing a median row of large transverse scales; very shallow dorsal and lateral caudal furrows extend entire length of tail; base of tail bearing hemipenial swellings; three small, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate. Coloration in life ( Figs. 5 ). Dorsal ground color of head, body, and limbs tan, immaculate; wide, dark-brown nuchal loop edged anteriorly and posteriorly by thin, faint yellow line; four wide, brown, body bands between nuchal loop and hind limb insertions weakly edged anteriorly and posteriorly by thin, faint, yellowish lines; band/ interspace ratio 1.00; no markings on posterior margin of thigh; eight white caudal bands infused with brown pigment, one-half to one-third width of dark-brown caudal bands; ventral surface of head, body, and limbs beige, immaculate except for black stipples in each scale; dark-brown and white caudal bands encircle tail. Variation ( Figs. 5 , 6 ). Paratypes LSUHC 11183–85 resemble the holotype in patterning but are slightly darker in overall coloration. LSUHC 11184 is a juvenile with a more contrasting color pattern and lighter ground color. LSUHC 11288 is considerably darker than the remainder of the type series and the thin whitish lines edging the anterior and posterior margins of the body bands are more discontinuous, appearing more as a series of transverse spots. LSUHC 11183 has a completely regenerated, unicolor brown tail. A small juvenile (LSUHC 10886, SVL 60.3 mm ) not part of the type series has a very bold, contrasting banding pattern. Meristic differences are listed in Table 6. Distribution. This species may be endemic to Gunung Tebu, Terengganu, Peninsular Malaysia . It is expected that its distribution is more extensive both above and below the type locality at Punca Air. Etymology. The specific epithet “ timur ” is an invariable noun in apposition in reference to this species being found in the Banjaran Timur of northeastern Peninsular Malaysia . “ Timur ” means east in the Malay language and is in reference to the mountain range being on the east coast of Peninsular Malaysia . Natural History. All adults were collected at night in hill dipterocarp forest in the vicinity of a rocky stream ( Fig. 6 ) between 640 m and 700 m in elevation. Lizards were most common on the granite boulders along the stream but others were seen on the bases of trees and one individual was observed on an earthen bank. A small juvenile (LSUHC 10886, SVL 60.3 mm ) was found on a twig at the base of a large tree deep in the forest and not along the stream. Comparisons (Tables 5, 6). Cyrtodactylus timur sp. nov. is differentiated from all other members of the C. pulchellus complex by having a combination of moderately sized body tubercles and no tubercles in gular region, ventrolateral body fold, or on ventral surfaces of forelimbs; 38–43 paravertebral tubercles; 21–24 longitudinal rows of body tubercles; 31–40 ventral scales; 20–25 subdigital lamellae on the fourth toe; 21 or 22 femoroprecloacal pores; four dorsal body bands between nuchal loop and hind limb insertions; body band/interspace ratio 1.00–1.25; no scattered pattern of white dorsal tubercles; 8–10 dark caudal bands on original tail; white caudal bands infused with brown pigment in adults; and a maximum SVL of 120.5 mm . Within the C. pulchellus complex, C. timur sp. nov. is the sister species to C. jelawangensis ( Fig. 2 ). See comparisons between these two species above (Table 6).