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 WGKK Grismer, Chan, Nurolhuda & Sumontha, 2008 Grismer, Chan, Nurolhuda & Sumontha 2008 [151,993,1124,1150] Reptilia Gekkonidae Cnemaspis Animalia Squamata 33 34 Chordata species biocellata   Figs. 11, 12     Holotype. ZRC 2.66693. Type locality: “ Kuala Perlis, Perlis, Peninsular Malaysia( 06°24.437N, 100°08.564E) at  37 min elevation.   Diagnosis.Maximum SVL 40.1 mm; 6–10 supralabials; 5–9 infralabials; ventral scales smooth; 6–12 contiguous, pore-bearing precloacal scales with round pores; 21–27 paravertebral tubercles; body tubercles randomly arranged, present on flanks, absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a median row of enlarged scales; one postcloacal tubercle on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 29–37 subdigital fourth toe lamellae; paired ocelli on occiput in males; wide, white to yellow nuchal loop in males; single ocellus in shoulder region in males; original tail yellow and regenerated tail orange in males; and throat, pectoral region, ventral surface of hind limbs, abdomen, and subcaudal region orange or yellow in males (Tables 6,7).  Color pattern in life (Figs. 11,12).Adult males: ground color of dorsal surface of head, body, limbs, and tail dull yellow; rostrum grayish with faint, light markings highlighting bright yellow, anterior, extra brillar fringe; interorbital region yellow with two, distinct, white, immaculate, well-defined occipital ocelli; ocelli accentuated by wide, black, occipital band forming anterior border of a series of closely spaced, large, white to yellow spots forming a nuchal band extending from posterior margin of one eye to posterior margin of other eye; small, black, shoulder patch enclosing a single, white to yellow ocellus; body overlain by five yellow, butterfly-shaped, vertebral blotches extending from shoulder region to base of tail; small, faint yellow blotches on flanks and limbs; blotches tend to form caudal bands; ventral surfaces yellow or orange, nearly immaculate with only a faint amount of subcaudal mottling. Adult females: ground color of dorsal surfaces light brown lacking black occipital and shoulder markings of males; occiput bears faint, straw colored, ocelli homologous to those in males; poorly defined, straw colored, butterfly-shaped, vertebral markings extend from nape to base of tail continuing posteriorly to form poorly defined, caudal bands; small, irregularly shaped, faint spots on flanks and limbs; and Ventral surfaces beige, nearly immaculate with only faint, subcaudal mottling.   Distribution.  Cnemaspis biocellataextends through the karst system of the Banjaran Nakawan from Thale Ban National Park, Satun Province, Thailandsouthward through Wang Kelian and Perlis StatePark to Gua Kelam, Tasik Meranti, Kampung Bukit Cabang, and Kuala Perlis, Perlisin northern Peninsular Malaysia( Grismer 2011a; Fig. 4).  Natural history. Grismer (2011a)noted that  Cnemaspis biocellataoccurs in lowland areas from near sea level to approximately 200 meters in elevation and appears to be a karst-substrate specialist. Lizards have been observed on trees and cement walls ( Fig. 11) but only where they were adjacent to a karst formation.  Cnemaspis biocellatais often observed abroad during the day on the shaded, vertical surfaces of karst walls and boulders as well as within crevice microhabitats and beneath small limestone rocks piled on the ground. During the day, lizards are wary and difficult to approach and will retreat into nearby crevice microhabitats at the slightest provocation. At night, lizards move farther away from these retreats and are more approachable. At GuaKelam, Perlisthe habitat is highly disturbed and continually frequented by visitors to the neighborhood park yet  C. biocellataare abundant and easily observed both day and night. At Tasik Meranti, Perlis, lizards occur on large karst rocks in lowland dipterocarp forest but restrict their daytime activities to shaded areas. At Wang Kelian, they are abroad only during the day, perhaps due to the presence of the much larger  Cyrtodactylus astrumGrismer, Wood, Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer & Pauwelsand  Gekko geckoLinnaeusat night which may even prey on them. At Kampung Bukit Cabang, lizards have been found at night within caves. At Kuala Perlis, lizards are abundant during the day and night on karst formations in the vicinity of parks, parking lots, and housing communities in areas with no immediate native vegetation. These observations suggest that the most important environmental component for this species is the karst substrate and the microhabitats it offers, regardless of the condition of the surrounding forest. This small species is amazingly quick and agile and effortlessly moves from one inclined surface to another. Upon capture, lizards release large sections of their skin in much the same manner as the Stump-tailed Gecko,  Gehyra mutilata(Wiegmann).  Cnemaspis biocellatais the second smallest species of  Cnemaspisand females carrying one or two eggshave been observed only during March.  Relationships.  Cnemaspis biocellatais a member of the Pattaniclade and most closely realeted to the sister species  C. niyomwanaeand  C. kumpoli( Fig. 2).   Material examined.  Malaysia: Perlis, Kuala Perlis ZRC 2.6693– 98( typeseries). Specimens examined since Grismer et al.(2008a):  Malaysia: Perlis, Gua Kelam LSUHC 8787–92, 8802, 8804–05, 9682.  Kampung Bukit Chabang LSUHC 9683–84. 3407812429 ZRC Malaysia 37 6.4072833 Kuala 1 100.14273 Peninsular 33 34 ZRC 2.66693 1 Perlis holotype 3407812330 ZRC Malaysia Kuala 35 36 ZRC 2.6693, 98 1 Perlis 3407812407 [428,1239,548,573] LSUHC Malaysia Gua Kelam 35 36 LSUHC 8787-92, 8802, 8804-05, 9682 1 Perlis 3407812312 LSUHC Malaysia Kampung Bukit Chabang 35 36 LSUHC 9683-84 1 Perlis