Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia Grismer, Lee Wood, Perry L. Anuar, Shahrul Riyanto, Awal Ahmad, Norhayati Muin, Mohd A. Sumontha, Montri Grismer, Jesse L. Onn, Chan Kin Quah, Evan S. H. Pauwels, Olivier S. A. Zootaxa 2014 2014-10-31 3880 1 1 147 WGPF Grismer & Wood & Anuar & Riyanto & Ahmad & Muin & Sumontha & Grismer & Onn & Quah & Pauwels, 2014 Grismer & Wood & Anuar & Riyanto & Ahmad & Muin & Sumontha & Grismer & Onn & Quah & Pauwels 2014 [151,415,151,177] Reptilia Gekkonidae Cnemaspis Animalia Squamata 92 93 Chordata species stongensis sp. nov.  Gunung Stong Rock Gecko   Fig. 49     Holotype. Adultmale LSUHC 11089collected on  26 June 2013by Chan Kin Onn, L.Lee Grismer and Jacob A. Chanat 1030 hrs at  10 mfrom Kem Baha, Gunung Stong, Kelantan, Peninsular Malaysia( 5°20.465 N, 101°58.001 E) at  461 melevation.    Paratypes.Adult males LSUHC 11091, 11093–94, 11100, 11139and adult female LSUHC 11092has the same collection data as the holotype.   Diagnosis.  Cnemaspis stongensis sp. nov.differs from all other Southeast Asia species of  Cnemaspisin having the unique combination of adult males reaching 49.3 mmSVL, adult females reaching 48.4 mmSVL; 8–11 supralabials; 8–10 infralabials; ventrals keeled; 5–8, continguous, pore-bearing precloacal scales with round pores; moderately prominent dorsal tubercles; 26–33 paravertebral tubercles; dorsal body tubercles generally randomly arranged; tubercles present on flanks; caudal tubercles not encircling tail; lateral caudal tubercles usually within lateral caudal furrows anteriorly only; ventrolateral caudal tubercles may or may not be present anteriorly only; lateral row of caudal tubercles present anteriorly only; subcaudals keeled; no enlarged, median subcaudal scale row; two or three postcloacal tubercles; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; and 28–32 subdigital lamellae on fourth toe. These differences are summarized across all Southeast Asian species in Table 6.  Cnemaspis stongensis sp. nov.lacks diagnostic color pattern characteristics.   Description of holotype.Adult male; SVL 49.3 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.25), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.43), distinct from neck; snout short (ES/HL 0.49), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum keeled, raised, slightly larger than similarly shaped scales on occiput; low, supraorbital ridges; moderate frontorostral sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.23); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave, dorsal 90% divided by longitudinal groove; rostral bordered posteriorly by two small supranasals and two large scales between the supranasals and laterally by first supralabials; 9R,L raised supralabials of similar size; 8R, 9L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, bordered posteriorly by four small postmentals of similar size; gular scales raised, keeled; throat scales same size, raised, keeled. Body slender, elongate (AG/SVL 0.43); small, keeled, dorsal scales equal in size throughout body, with intermixed larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles on flanks sparse, moderate in size; 32 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales slightly larger than dorsals; eight pore-bearing precloacal scales with round pores in a chevron pattern separated on the left side of the chevron by a single nonpore-bearing scale; forelimbs moderately long, slender (FL/SVL 0.18); dorsal scales of brachium raised, keeled; dorsal scales of forearm keeled, raised; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm raised, juxtaposed; palmar scales raised, smooth, juxtaposed; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wider; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.21); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales raised, smooth, juxtaposed; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally but wider distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wider; interdigital webbing absent; toes increase in length from first to fourth with fourth being slightly longer than fifth; 32 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; caudal scales flat anteriorly, weakly keeled, juxtaposed; middorsal and lateral furrows; no enlarged, median subcaudal scales; subcaudal scales keeled; caudal tubercles present in lateral furrow anteriorly; two enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail.   FIGURE 49.  Cnemaspis stongensis sp. nov.from Kem Baha, Gunung Stong, Peninsular Malaysia. Upper left: adult male holotype (LSUHC 11091) in dark color pattern phase. Upper right: ventral view of the holotype and a gravid female paratype (LSUHC 11091 upper and 11100 lower, respectively). Middle left: adult female paratype (LSUHC 11100) in dark color pattern phase. Middle right: granite boulder microhabitat at Kem Baha, Gunung Stong, Peninsular Malaysia. Lower: type series of  C. stongensis sp. nov.Photographs by LLG.  Color pattern in life ( Fig. 49).Dorsal ground color of head, limbs, and body grey; black, irregular striping on snout; black, transverse, azygous marking on top of head; black, postorbital stripe extending onto nape; black, medial, teardrop-shaped marking on nape; a series of seven, black, paravertebral markings extending from shoulder region to base of tail; similar black markings on flanks; dull-white, butterfly-shaped, paravertebral markings alternating with black, paravertebral markings and becoming completely separated posteriorly; dull-white, transverse bars on flanks; light-colored, caudal bands, anterior two yellowish and posterior seven white; light caudal bands alternate with black caudal bands; limbs bearing irregularly shaped, whitish bands; ventral surfaces of head, throat, pectoral, and abdominal regions, and limbs grey, gular region and abdomen slightly darker; caudal bands encircle tail but are less vivid in subcaudal region.  Variation.The paratypesclosely resemble the holotypein coloration and pattern ( Fig. 49). The dorsal pattern of LSUHC 11094 and 11139 appears slightly more speckled overall and LSUHC 11100 has a faint, wide, lightcolored, vertebral stripe. Morphometric variation and variation in scalation are presented in Table 11.   Distribution.  Cnemaspis stongensis sp. nov.is known only from the typelocality and surrounding areas on Gunung Stong, Kelantan, Peninsular Malaysia( Fig. 3).   Natural History. Lizards were observed to be active only at night on large granite boulders ( Fig. 49) ranging in elevation from 50 mat the base of Gunung Stong at Hutan Lipur Jelawang in lowland dipterocarp forest up to the typelocality of Kem Baha at 461 min hill dipterocarp forest. The majority of specimens were found in the vicinity of water but were not necessarily restricted to these areas. Within their microhabitat, lizards were most commonly seen on the undersides or on the lower sections of boulders ( Fig. 49) making escape much easier. We expect this species extends to higher elevations on Gunung Stong above Kem Baha.   Etymology.The specific epithet  stongensisis an adjective in reference to Gunung (mountain) Stong on which the typelocality of Kem Baha is located.   Comparisons.  Cnemaspis stongensis sp. nov.is a member of the  affinisgroup within which it is the sister species of  C. bayuensis. Cnemaspis stongensis sp. nov.differs from  C. bayuensisin having caudal tubercles anteriorly in the lateral caudal furrow as opposed to lacking them; having a lateral row of caudal tubercles present anteriorly as opposed to throughout the length of the tail; and lacking distinct black and white caudal bands as opposed to having them. However, these two species bear only a 2.2% sequence divergence from one another ( Table 4).  Cnemaspis stongensis sp. nov.differs from  C. pseudomcguirei, C. harimau,and  C. shahruliby having a greater maximum SVL ( 49.3 mmversus 36.5–43.2) and being considerably smaller than  C. mcguirei(maximum SVL 65.0 mm). It differs from  C. shahruliand  C. hangus sp. nov.in having as opposed to lacking pre-cloacal pores. From  C. harimau, C. shahruli, C. flavolineata, C. temiah sp. nov.and  C. hangus sp. nov.it differs in having more paravertebral tubercles (26–33 versus 18–27).  Cnemaspis stongensis sp. nov.can be further separated from all species of the  affinisgroup except  C. affinis, C. hangus sp. nov.,  C. selamatkanmerapoh,and  C. bayuensisby generally lacking tubercles in the lateral caudal furrows. It is seaparated from  C. harimauand  C. temiah sp. nov.by not having caudal tubercles that encircle the tail. Having 28–32 subdigital lamellae on the fourth toe separates it from  C. pseudomcguirei, C. flavolineata,and  C. temiah sp. nov.  Comments.The relatively close morphological and genetic similarity between  Cnemaspis stongensis sp. nov.and  C. bayuensisis surprising in that these sister species are separated by at least 85 kmof uninhabital terrane and the former is a granite boulder specialist and the latter is a karst specialist.  Relationships.  Cnemaspis stongensis sp. nov.is the sister species of  C. bayuensis( Fig. 2).   Additional material examined.  Malaysia: Kelantan; Gunung Stong, Kem Baha LSUHC 11090, 11095, 11138. 3407812452 2013-06-26 LSUHC Adult & Chan Kin Onn, L. & Jacob A. Chan Malaysia 10 5.3410835 Kelantan 1 101.96668 92 93 LSUHC 11089 1 Kelantan holotype 3407812400 2013-06-26 LSUHC Adult & Chan Kin Onn, L. & Jacob A. Chan Malaysia 10 5.3410835 Kelantan 1 101.96668 92 93 LSUHC 11091, 11093-94, 11100, 11139, LSUHC 11092 1 Kelantan paratype 3407812450 LSUHC Malaysia Kem Baha Gunung Stong 95 96 LSUHC 11090, 11095, 11138 1 Kelantan