Cnemaspis siamensis Smith

Grismer, Lee, Sumontha, Montri, Cota, Michael, Grismer, Jesse L., Wood, Perry L., Pauwels, Olivier S. G. & Kunya, Kirati, 2010, A revision and redescription of the rock gecko Cnemaspis siamensis (Taylor 1925) (Squamata: Gekkonidae) from Peninsular Thailand with descriptions of seven new species, Zootaxa 2576, pp. 1-55 : 6-24

Cnemaspis siamensis Smith

Siamese Rock Gecko Djing Djok Niew Yaow Thai Figure 2

Gonatodes siamensis Smith, M. A., 1925:21 . IV.—Contributions to the herpetology of Borneo. Sarawak Museum Journal 8:15–34. Type locality: Maprit, near Patiyu (=Pathio), Peninsular Siam [ Thailand].

Gonatodes kendallii Smith, M. A. 1916:151 . non Boulenger fide Taylor, 1963:740.

Gonatodes siamensis Smith, 1930:16 .

Cnemaspis siamensis Smith, 1935:71 .

Cnemaspis (Cnemaspis) siamensis Rösler, 2000:63 .

Diagnosis. Adult males and females reaching 37.3 mm SVL; eight or nine supralabials; 6–8 infralabials; gular scales smooth; forearm, subtibials, ventrals, subcaudals, and dorsal tubercles keeled; 19–25 paravertebral tubercles; tubercles on flanks not linearly arranged; no ventrolateral caudal tubercles; caudal tubercles do not encircle tail; lateral, caudal tubercles may or may not occur with lateral, caudal furrow; scales of median, subcaudal scale row slightly enlarged and keeled; precloacal pores absent; one or two postcloacal tubercles; shield-like subtibials absent; metatarsal scales may be slightly enlarged; 22–26 subdigital lamellae on fourth toe; dark, longitudinal gular markings present; head not yellow in adult males; no dark shoulder or neck patch enclosing a white to yellow ocellus; no yellow to white, prescapular crescent or lightly colored, transverse bars on flanks. These differences are summarized across all species in TABLES 1 and 2 .

Description. Head oblong in dorsal profile, moderate in size, somewhat narrow, flat, distinct from neck; snout short, slightly concave in lateral profile; postnasal region constricted medially; raised scales of rostrum keeled, larger than similarly shaped scales on occiput; low, supraorbital ridges; shallow frontonasal sulcus; canthus rostralis smoothly rounded; eye large; extra-brillar fringe scales small in general but largest anteriorly; pupil round; ear opening oval, taller than wide; rostral slightly concave dorsally, dorsal 70% divided by longitudinal groove; rostral bordered posteriorly by two supranasals and one smaller azygous internasal, bordered laterally by first supralabials and nostrils; eight or nine slightly raised supralabials of similar size; 6–8 infralabials, decreasing gradually in size posteriorly; nostrils small, round, oriented posterolaterally, bordered posteriorly by small, granular, postnasal scales; mental large, triangular, medially concave, extending to level of second infralabials, bordered posteriorly by two or three (usually three) postmentals; gular scales slightly raised, smooth; throat scales smaller, weakly keeled.

Body slender, elongate; small, weakly keeled, dorsal scales equal in size throughout body, intermixed with numerous, large, multi-keeled, semi-longitudinally arranged tubercles; tubercles extend from top of head to base of tail and are smallest anteriorly; 19–25 paravertebral tubercles; pectoral and abdominal scales slightly raised, subimbricate, keeled; abdominal scales slightly larger than pectoral and dorsal scales; pore-bearing, precloacal scales and precloacal depression absent; forelimbs moderately long, slender; dorsal scales of forelimbs keeled; ventral scales of forearm smooth, juxtaposed to subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched, wide throughout length of digits, bearing a greatly enlarged, projecting scale at digital inflections; interdigital webbing absent; fingers increase in length from first to fourth with fifth same length as fourth; hind limbs longer and thicker than forelimbs; dorsal scales keeled, raised, juxtaposed; ventral scales of thigh smooth to weakly keeled; subtibials keeled, larger than dorsal tibials; plantar scales smooth, slightly raised, juxtaposed; slightly enlarged, submetatarsal scales beneath first metatarsal may be present; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout length of digits except at base where scales are more granular; greatly enlarged, projecting scale at digital inflections; interdigital webbing absent to weak; toes increase in length from first to fourth with fourth and fifth nearly equal in length; 22–26 subdigital lamellae on fourth toe; dorsal caudal scales arranged in segmented whorls; caudal scales flat anteriorly, weakly keeled, juxtaposed; very shallow, middorsal furrow; deep, single, lateral furrow; median row of slightly enlarged, keeled, subcaudal scales with three or four scales per caudal segment; other subcaudals keeled; paravertebral, dorsolateral, and lateral rows of keeled, caudal tubercles; paravertebral and lateral, caudal tubercles largest, spinose; caudal tubercles do not encircle tail, not present in lateral, caudal furrow; one or two enlarged, postcloacal tubercles on lateral surfaces of hemipenal swellings at base of tail; tail approximately 1.3% times SVL.

Coloration (in life, Fig. 2 ). Dorsal ground color of head, body, limbs and tail pale gray; top of head bearing numerous small, brown markings with dark lines radiating out from eyes; dark, irregular, postorbital stripe of varying intensities extends onto nape followed by paired, dark brown, offset, paravertebral markings countershaded by larger, lightly colored blotches extending to level of axillae; posterior to axillae markings become elongate and extend to base of tail; dark paravertebral markings on body often connect medially to form short, irregularly shaped bands; flanks bearing small, light brown and cream colored, irregularly shaped, randomly distributed blotches; limbs covered with distinct, light and dark markings, appearing weakly banded; dark bands on digits; original tail distinctly marked with dark brown and cream colored bands; regenerated tail uniform gray; gular region, throat, and anterior portion of pectoral region yellow; gular region in males bearing a dark brown, median stripe or longitudinally arranged spots flanked by longitudinally arranged dark brown spots; gular markings faint in females; ventral surfaces of limbs, remainder of body, and anterior one-half of tail, beige with dark stippling on limbs and lateral edges of belly; posterior one-half of original subcaudal region yellow with cream colored bands. There appears to be no sexual dimorphism with respect to color with the exception of the gular markings.

Distribution. Cnemaspis siamensis ranges throughout the lowland, hilly regions east of the Tenasserim and Phuket Mountains from Kaeng Krachan National Park, Phetchaburi Province in the north, southward to Khao Mod, Surat Thani Province on the east coast and at least Kaeng Krung National Park, Surat Thani Province in the west ( Fig. 1 ). We have examined close-up photographs (LSUDPC 5273–78) of an adult male and female (BM 1916.6.22.2–3) from Khlong Banlai, Phuket Island which have longitudinal gular markings and no precloacal pores indicating they are C. siamensis .

Natural history. The only natural history observations made on Cnemaspis siamensis were that of Pauwels et al. (2003) just west of Kaeng Krachan National Park, Phetchaburi Province. They reported finding a dead specimen 1.6 m above the ground beneath a detached piece of bark on the trunk of a tree that had been smashed by passing elephants. During October of 2009, we observed two specimens at the type locality in Pathio, Chumpon Province. One (LSUHC 9474; Fig. 2 ), a gravid female carrying two eggs, was found in disturbed forest near sea level at the foot of a hill composed of karst outcroppings ( Fig. 3 ) at 2300 hrs. The specimen was 35 m from the nearest karst formation and was running and hopping throught the leaf litter on the forest floor beneath a dead tree branch. An adult male (LSUDPC 5241; Fig. 2 ) was found at 2400 hrs on a forested hillside covered with scattered, small rocks (some of which had been cemented together to form walls) as it was emerging from a crack in a rock wall. Pauwels et al. (2000) reported finding what may be C. siamensis at Phang-Nga Wildlife Breeding Station beneath a decaying stump in an evergreen forest, but these observations may also apply to C. vandeventeri sp. nov. (see below). Collectively, these observations suggest that C. siamensis is not a saxicolous, microhabitat specialist as are the majority of other species of Cnemaspis but is a nocturnal, lowland, scansorial, forest dwelling species that opportunistically utilizes rocky microhabitats when available.

Remarks. Smith’s (1925) brief, nearly anecdotal description of Cnemaspis siamensis reported it on the Malay Peninsula from Phuket; Tasan and Pak Chan ( C. vandeventeri sp. nov.); Krabi ( C. chanardi sp. nov.); and Khao Sebab (=Khao Sa Bap), Chanthaburi ( C. chanthaburiensis ) in southeastern Thailand. In later treatments, Smith (1930, 1935) provided slightly expanded descriptions of C. siamensis but they were still based on multiple species from throughout Thailand and reported on additional material from the mountains of Nakhon Si Thammarat ( C. chanardi sp. nov.). Smith (1925) designated the type locality for C. siamensis as “Maprit, near Patiyu” which currently is Pathio, Chumpon Province and lizards from this locality bear the diagnostic character states of lacking precloacal pores; a lightly colored, prescapular crescent; lightly colored, transverse bars on the flanks; and having dark, longitudinal markings on the gular region ( Fig. 2 ). Smith (1925) noted in the series he examined that males had 0–8 precloacal pores which is a clear indication he was examining multiple species. He made no mention of the presence or absence of gular markings, markings on the flanks, or a prescapular crescent.

Although Taylor’s (1963) description of what he thought was Cnemaspis siamensis was far more complete than any of Smith’s (1925, 1930, 1935), it was based on a different species, C. chanardi sp. nov. from Khao Chong, Trang Province (see below), and thus continued to conflate undescribed taxa with C. siamensis . Taylor (1963) did note the prescapular, crescent-shaped markings in his material and expressed surprise that according to Smith (1925, 1930, 1935), some males had precloacal pores and others did not.

Dring (1979) was the first to begin to untangle the cumulative errors of Smith (1925, 1930, 1935) and Taylor (1963) by noting character differences between many of the Thai populations. Although he gave these populations no formal names, he flagged them for subsequent researchers. Referring to specimens from southeastern Thailand as species A, led Bauer & Das (1998) to describe Cnemaspis chanthaburiensis . Similarly, he was the first to indicate that populations of C. siamensis north of the Isthmus of Kra lacked precloacal pores and had dark, gular markings and those south of the Isthmus did not. This was emphasized in his key to the species of Cnemaspis showing that these two populations of C. siamensis emerged separately. Similar contributions by Dring (1979) led directly to the descriptions of C. dringi (Das & Bauer 1998) from Sarawak, Malaysian Borneo and C. flavigaster ( Chan & Grismer 2008) from Selangor in southern Peninsular Malaysia.

Material examined. Chumpon Province: Krom Luang District THNHM 0372; Pha To District, Ngao National Park THNHM 1086; Pathio LSUHC 9474, 9485, MCZ 39025; Kapoh Water Fall FMNH 215977. Phetchaburi Province: Muang District THNMH 1441–42, 1448–49. Prachuap Khiri Khan Province:Pa-La-U, Kaeng Krachan National Park, Hua Hin District, Prachuap Khiri Khan THNHM 1336–37; Thap Sakae District THNHM 2000. Surat Thani Province: Kanchanadit District MS16; Kaeng Krung National Park THNHM 1084.

affinis argus aurantiacopes baueri continued.

bayuensis biocellata boulengeri caudanivea

Maximum SVL 46.1 40.1 69.0 47.1 Supralabials 9,10 6– 10 8–9 8,9 Infralabials 8,9 5–7 6,7 7,8 Ventral scales keeled (1) or not (0) 1 1 0 0 No. of precloacal pores 5–9 8–12 0 0–3 Precloacal pores continuous (1) or separated (0) 0 1 / 0 Precloacal pores elongate (1) or round (0) 0 0 / 0 No. of paravertebral tubercles 23–30 21–23 32–38 20–27 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 1 0

continued next page bayuensis biocellata boulengeri caudanivea

Tubercles present (1) or absent (0) on flanks 1 1 0 weak Caudal tubercles in lateral furrow (1) or not (0) 0,0 / / 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 0 1 Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 0 0 1 0 Subcaudals keeled (1) or not (0) 1 0 0 0 Single median row of keeled subcaudals (1) or not (0) 0 0 0 0 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 1 1 0 No. of postcloacal tubercles 2 1 1 1,2 Enlarged femoral scales present (1) or absent (0) 0 0 1 0 Shield-like subtibial scales present (1) or absent (0) 0 0 1 1 Subtibial scales keeled (1) or not (0) 1 0 0 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 1 1 No. of 4th toe lamellae 27–30 29–37 29–32 23–30 continued.

chanthaburiensis dringi flavigaster flavolineata

Maximum SVL 42.1 46.4 50.1 46.7 Supralabials 8–10 11 9 ,10 8–10 Infralabials 7–9 9 8–10 8–10 Ventral scales keeled (1) or not (0) 0 1 0 1 No. of precloacal pores 7–9 5,6 7,8 7 Precloacal pores continuous (1) or separated (0) 1 0 1 1 Precloacal pores elongate (1) or round (0) 0 0 0 0 No. of paravertebral tubercles 23–25 25–27 21–24 22–24 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0 0 Tubercles present (1) or absent (0) on flanks 1 0 1 1 Caudal tubercles in lateral furrow (1) or not (0) 1 0 0 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 0 0 0 0 Subcaudals keeled (1) or not (0) 0 1 0 1 Single median row of keeled subcaudals (1) or not (0) 0 1 0 0 Caudal tubercles encircle tail (1) or not (0) 0 0 0 1 Enlarged median subcaudal scale row (1) or not (0) 0 1 0 0 No. of postcloacal tubercles 1–3 2 1,2 2,3

continued next page chanthaburiensis dringi flavigaster flavolineata

Enlarged femoral scales present (1) or absent (0) 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 Subtibial scales keeled (1) or not (0) 0 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 1 0 No. of 4th toe lamellae 25–29 32–35 29–34 23–26 continued.

harimau karsticola kendallii kumpoli limi Maximum SVL 40.7 48.1 58.0 63.0 88.2 Supralabials 9,10 7, 8 10–12 7, 8 8–11 Infralabials 9,10 6 ,7 7–9 6,8 7–9 Ventral scales keeled (1) or not (0) 1 1 1 0 1 No. of precloacal pores 0 7,8 0 7,8 0 Precloacal pores continuous (1) or separated (0) / 1 / 0 / Precloacal pores elongate (1) or round (0) / 0 / 0 / No. of paravertebral tubercles 18–20 17–19 20–26 29–35 25–33 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0 0 0 Tubercles present (1) or absent (0) on flanks 1 1 weak 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 1 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 0 1 Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 0 0 0 0 0 Subcaudals keeled (1) or not (0) 1 1 1 0 0 Single median row of keeled subcaudals (1) or not (0) 0 0 1 0 0 Caudal tubercles encircle tail (1) or not (0) 1 0 1 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 1 0 weakly No. of postcloacal tubercles 0 2,3 1,3 2,3 1,2 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 No. of 4th toe lamellae 26–30 27–30 23–36 34–41 30–35

continued.

mcguirei monachorum nigridia nuicamensis

Maximum SVL 65 32.9 75.5 47.5 Supralabials 7– 9 7, 8 10,11 7–9 Infralabials 7,8 5–7 9–11 6–7 Ventral scales keeled (1) or not (0) 1 0 1 0

continued next page mcguirei monachorum nigridia nuicamensis

No. of precloacal pores 5–10 3 12 –14 4–6 Precloacal pores continuous (1) or separated (0) 0 1 0 0 Precloacal pores elongate (1) or round (0) 0 0 0 0 No. of paravertebral tubercles 26– 32 11–19 39–43 20–22 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0 1 Tubercles present (1) or absent (0) on flanks 1 0 0 1 Caudal tubercles in lateral furrow (1) or not (0) 1 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 ant 1 1 Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 0 0 0 0 Subcaudals keeled (1) or not (0) 1 0 0 0 Single median row of keeled subcaudals (1) or not (0) 0 0 0 weak Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 1 1 1 No. of postcloacal tubercles 2,3 1,2 2–4 2–4 Enlarged femoral scales present (1) or absent (0) 0 0 0 weak Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 0 1 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 1 0 No. of 4th toe lamellae 30–35 25–27 28,29 29–33 continued.

paripari pemanggilensis perhentianensis pseudomcguirei

Maximum SVL 50.7 76.0 47 42.5 Supralabials 12 10– 13 8–10 9 Infralabials 10,11 10– 14 7–8 8,9 Ventral scales keeled (1) or not (0) 1 1 0,1 1 No. of precloacal pores 2–6 0 6–8 1–5 Precloacal pores continuous (1) or separated (0) 0 / 1 1 Precloacal pores elongate (1) or round (0) 0 / 0 0 No. of paravertebral tubercles 26–31 30–37 22–27 26–32 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0 0 Tubercles present (1) or absent (0) on flanks 0 1 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 0 0 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 1 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 ant

continued next page paripari pemanggilensis perhentianensis pseudomcguirei

Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 0 0 0 0 Subcaudals keeled (1) or not (0) 1 1 1 1 Single median row of keeled subcaudals (1) or not (0) 0 1 0 0 Caudal tubercles encircle tail (1) or not (0) 0 1 0 0 Enlarged median subcaudal scale row (1) or not (0) 1 1 0 0 No. of postcloacal tubercles 2 2 3–4 2,3 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 0 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 0 0 0,1 No. of 4th toe lamellae 26–31 27–31 28–31 23–25 continued.

psychedelica roticanai shahruli siamensis tucdupensis

Maximum SVL 75.3 47 36.5 39.7 51 Supralabials 7, 8 8, 9 10,11 8, 9 8–10 Infralabials 5–7 7, 8 8–10 6–8 7,8 Ventral scales keeled (1) or not (0) 0 1 1 1 0 No. of precloacal pores 0 3–6 0 0 0 Precloacal pores continuous (1) or separated (0) / 0 / / / Precloacal pores elongate (1) or round (0) / 0 / / / No. of paravertebral tubercles 34–48 25–27 19–22 19–25 18–21 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0 0 1 Tubercles present (1) or absent (0) on flanks 0 1 1 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 1 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 0 0 1 Lateral caudal tubercle row present (1) or absent (0) 0 0 1 1 0 Caudal tubercles restricted to a single paravertebral

row on each side (1) or not (0) 1 0 0 0 1 Subcaudals keeled (1) or not (0) 0 1 1 1 0 Single median row of keeled subcaudals (1) or not (0) 1 1 0 weak 1 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 1 weak 0 1 1 No. of postcloacal tubercles 1,2 1,2 2,3 1,2 0–3 Enlarged femoral scales present (1) or absent (0) 1 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 1 0 0 0 0 Subtibial scales keeled (1) or not (0) 0 1 1 1 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 0 0 0 1 No. of 4th toe lamellae 25–28 26–29 21–30 24–26 27–30 continued.

huaseesom punctatonuchalis niyomwanae

Maximum SVL 37.9 49.6 56.8 Supralabials 7–9 8 8–11 Infralabials 6–8 7,8 6–8 Ventral scales keeled (1) or not (0) 0 0 0 No. of precloacal pores 5,6 0 3 Precloacal pores continuous (1) or separated (0) 1 / 1,0 Precloacal pores elongate (1) or round (0) 0 / 0 No. of paravertebral tubercles 18–24 24–27 26–31 Tubercles of ventralmost row on flank linearly arranged

and in contact or nearly so (1) or tubercles on flank

widely spaced and more randomly distributed (0) 0 0 0

continued next page Holotype. Adult male ( THNHM 6983) collected on 22 June 2005 at Ban Chong, Chong, Nayong District, Trang Province, Thailand. Squares represent type localities between 400 and 600 m a.s.l. by Tanya Chan-ard.

Paratypes. FMNH 176863 was collected on 10 May 1959 by Edward H. Taylor (EHT 390) at the same locality as the holotype. CUMZ-R-2009,6,24-6 was collected on 16 February 2007 at the Dad Fa Waterfall in Tai Rom Yen National Park, Ban Nasan District, Surat Thani Province (8°51.87’N 99°31.13’E) at 362 m a.s.l. by Thanin Kaewmanee. THNHM 12434 was collected at Khao Nor Chuchi, Khlong Thom, Khlong Thom District, Krabi Province during 1996 by Tanya Chan-nard. THNHM 1334-35 were collected on 8 May 2003 at Khao Luang National Park, Lan Saka District, Nakhon Si Thammarat Province between 200 and 300 m a.s.l. by Tanya Chan-ard and Yodchaiy Chuaynkern. THNHM 14111 was collected on 6 December 2003 at Khao Luang National Park, Lan Saka District, Nakhon Si Thammarat Province between 200 and 300 m a.s.l. by Tanya Chan-nard and Y. Chuaynkern. THNHM 10705 collected on 6 December 2006 from Krung Nang Waterfall in Khao Nan National Park, Nopphitam Distrct, Nakhon Si Thammarat Province (08° 46.219 N, 99°35.29.6 E) at 129 m a.s.l. by T. Chan-nard, S. Mekchai, S. Laoteaw, and M. Cota. THNHM 10115 was collected at Khao Nan National Park, Nakhon Si Thammarat Province on 22 October 2006 at approximately 300 m a.s.l. by Tanya Chan-ard, Sunchai Makchai, Michael Cota, Suttinee Laoteow, and Huay Lak. THNHM 10135 was collected at Khao Nan National Park, Tha Sala District, Nakhon Si Thammarat Province on 23 October 2006 at 255 m a.s.l. by Tanya Chan-ard, Sunchai Makchai, Michael Cota, and Suttinee Laoteow. LSUHC 9564 was collected at the Phuphaphet Cave, Muang District, Satun Province on 6 October 2009 at approximately 220 m a.s.l. by Kirati Kunya. THNHM 10383 was collected on 14 December 2003 at Ban Yod Leong, Nopphitham District, Nakhon Si Thammarat Province at 355 m a.s.l. by T. Chan-ard and Y. Chuaynkern.

Diagnosis. Adult males reaching 40.1 mm SVL, adult females reaching 38.7 mm SVL; 7–10 supralabials; 6–8 infralabials; gulars smooth; forearm, subtibials, ventrals, subcaudals, and dorsal tubercles keeled; 20–30 paravertebral tubercles; tubercles on flanks not linearly arranged; ventrolateral caudal tubercles absent; caudal tubercles do not encircle tail; no lateral, caudal tubercles within lateral, caudal furrow; median row of subcaudals keeled, slightly enlarged; 6–8 precloacal, pore-bearing scales in adult males separated medially by non-pore-bearing scales; pores round; one postcloacal tubercle; shield-like subtibials and enlarged, submetatarsals absent; 25–30 subdigital lamellae on fourth toe; no dark, longitudinal gular markings or blotches; head not yellow in adult males; no dark patch on shoulder or neck enclosing a white to yellow ocellus; yellow to white, prescapular crescent present. These differences are summarized across all species in TABLES 1 and 2 .

Description of holotype. Adult male; SVL 37.7 mm; head oblong in dorsal profile, moderate in size (HL/ SVL 0.28), somewhat narrow (HW/SVL 0.17), flat (HD/HL 0.39), distinct from neck; snout short (ES/HL 0.48), concave in lateral profile; postnasal region constricted medially; raised scales of rostrum keeled, larger than similarly shaped scales on occiput; low, supraorbital ridges; no frontonasal sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.23); extra-brillar fringe scales small in general but largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 90% divided by longitudinal groove; rostral bordered posteriorly by two supranasals and two smaller azygous scales, bordered laterally by first supralabials and nostrils; 9R,10L raised supralabials of similar size; 7R,L infralabials, decreasing gradually in size posteriorly; nostrils small, oblong, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, subpentagonal, medially concave, extending to level of second infralabials, bordered posteriorly by three postmentals; gular scales raised, smooth, somewhat pointed; throat scales larger, taller, smooth, juxtaposed.

Body slender, elongate (AG/SVL 0.45); small, weakly keeled, dorsal scales equal in size throughout body, intermixed with numerous, large, multi-keeled, semi-longitudinally arranged tubercles; tubercles extend from top of head to base of tail and are smallest anteriorly; 24 paravertebral tubercles; pectoral and abdominal scales slightly raised, subimbricate, weakly keeled; abdominal scales slightly larger than pectoral scales, much larger than dorsals; seven contiguous, pore-bearing, precloacal scales arranged in a 3-4 chevron, followed posteriorly by three more contiguous, pore-bearing scales all in contact; pores round; forelimbs moderately long, slender, dorsal scales keeled; ventral scales of forearm smooth, juxtaposed to subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; subdigital lamellae wide throughout length of digits, bearing a larger scale at the digital inflections; interdigital webbing absent; fingers increase in length from first to fourth with fifth the same length as fourth; hind limbs longer and thicker than forelimbs; dorsal scales keeled, raised, juxtaposed; ventral scales of thigh, raised, weakly keeled; subtibials keeled, larger than dorsal tibials; plantar scales smooth, slightly raised, juxtaposed; slightly enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout length of digits except at base where scales are more granular; enlarged scale at digital inflections; interdigital webbing absent; toes increase in length from first to fourth with fourth and fifth nearly equal in length; 27 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal, caudal scales raised, keeled, subimbricate anteriorly; moderate, middorsal furrow; deep, single, lateral furrow; median row of slightly enlarged, keeled, subcaudals with four scales per caudal segment; other subcaudals keeled; paravertebral, dorsolateral, and lateral rows of large, keeled, nearly spinose, caudal tubercles; tubercles of paravertebral and lateral rows largest; ventrolateral, caudal tubercles absent; caudal tubercles do not encircle tail, absent from lateral, caudal furrow; 2R,1L postcloacal tubercles; tail approximately 1.1% times SVL; anterior 33.2 mm original, remainder regenerated.

Coloration (in alcohol). Dorsal ground color of head, body, limbs and tail pale brown; top of head bearing small, faint, brown markings and postorbital stripes; series of small, whitish, vertebral blotches extend from nape to level of hind limbs, becoming increasingly larger, separated, and irregular in shape posteriorly; from level of hind limbs to base of tail blotches become slightly offset and paravertebrally aligned, extending onto the tail to form lightly colored, caudal bands; three dark blotches on nape form a tripartite pattern, followed by paried dark, opposing blotches immediately anterior to shoulders which grade into small, diffuse, randomly arranged, darker markings on rest of body; thin, incomplete, light-colored, prescapular crescent preceded by whitish spot at base of head, followed by a series of thin, whitish to yellow, transverse bars on flanks; tubercles on anterior part of body tend to be white or black; posterior body tubercles tend to be yellow or black; enlarged, widely separated, white tubercles occur on flanks; limbs weakly mottled with diffuse, dark markings; dark bands on digits; tail faintly banded; all ventral surfaces cream colored to yellowish white with faint stippling being darkest on tail.

Variation. Differences in squamation and morphometrics are presented in TABLE 3. The paratypes closely approach the holotype in most aspects of coloration and pattern ( Fig. 4 ). The Surat Thani specimen (CUMZ-R-2009,6,24-6) has larger more spinose tubercles ( Fig. 5 ) and in life ( Fig. 6 ) a thin, dark, preorbital stripe and two postorbital stripes; the lower postorbital stripe extends onto upper portion of forelimb; the upper postorbital stripe is wider, incomplete, extending onto occiput where it is separated by opposing upper postorbital stripe by an elongate, medial, cream-colored marking; cream-colored marking bordered anteriorly by a dark, medial blotch producing a dark, tripartite, pattern on nape. The gular region is yellow, immaculate; throat, ventral portion of forelimbs, pectoral region beige, immaculate; belly, ventral portions of hind limbs, and subcaudal region yellow, immaculate ( Fig. 6 ). This species appears to not be sexually dimorphic for dorsal color pattern. Ventral coloration in life for females is light yellow except for the hind limbs and tail which are somewhat beige ( Fig. 6 ). The prescapular crescent is wider in THNHM 10135 but faint in THNMH 13838. Baring on the flanks is weak in THNHM 10135. The prescapular crescent is wider in THNHM 10115 and nearly absent in THNHM 1335. Baring on the flanks is weak in THNHM 10115. Figure 7 shows the color pattern variation between specimens from Satun and Nakhon Si Thammarat provinces.

FMHN THNHM THNHM THNHM THNHM THNHM CUMZ R

176863 6983 12434 1334 1335 10115 2009,6,24-9

Paratype Holotype Paratype Paratype Paratype Paratype Paratype SVL 40.1 37.7 35.1 33.5 34.6 38.6 37.2 Sex m m m m f f m Supralabials 10 9 9 8 8 9 8

Infralabials 8 7 8 7 7 7 8

Precloacal pores 7 4 6 3 0 0 6

Paravertebral tubercles 25 24 23 23 21 23 30 4 th toe lamellae 26 27 27 30 26 30 25 TL 49.5 43 20.8 40.5 45.5 53.3 40.6 TW 4.7 4.2 4.2 3.7 3.9 4.0 3.8 FL 5.7 5.8 5.7 5.7 5.5 6.2 5.3 TBL 8.0 7.1 7.1 6.9 6.4 7.4 6.8 AG 19.6 16.8 14.6 13.8 18.8 12.6 13.9 HL 10.7 10.5 4.8 9.0 9.6 10.6 10.3 HW 7.4 6.7 6.6 5.9 5.7 6.5 6.5 HD 5.0 4.1 4.2 4.1 4.0 4.6 4.4 ED 2.3 2.4 2.0 2.0 2.2 2.4 2.1 EE 3.3 2.9 3.1 2.5 2.6 3.0 3.0 ES 5.3 5.0 4.6 4 4.6 4.8 4.7 EN 4.0 3.9 3.4 3.1 3.4 3.6 3.7 IO 3.0 2.8 2.5 2.2 2.5 2.8 2.4 EL 1.3 0.8 1.2 0.9 1.0 1.2 1.2 IN 1.1 1.0 0.9 0.8 0.8 1.1 0.7 Continued.

LSUHC THNHM THNHM THNHM THNHM

9564 14111 10135 10705 13838

Paratype paratype paratype paratype paratype Distribution. Cnemaspis chanardi sp. nov. occurs in the foothills of the Nakhon Si Thammarat and Sankalakhiri Mountains and lowland areas to the west, extending from the southern terminus of the Isthmus of Kra in Donsak District, Surat Thani Province, southward along the western foothills through Khao Chong and Nayong district, Trang Province to at least the Phuphaphet Cave, Satun Province in the south. It probably continues approximately 45 km further south to the Banjaran Nakawan mountains that form the physiographic barrier between Thailand and Peninsular Malaysia. It extends westward from the foothills through the lowlands to at least Khlong Thom District, Krabi Province ( Fig. 1 ). It ranges from near sea level at Khlong Thom to just under 600 m at Khao Chong. Smith (1930) reported specimens from Khao Whip (=Khao Wang Hip), Nakhon Si Thammarat Province but the elevation is unknown. Currently, C. chanardi sp. nov. is not known to occur in the foothills or lowlands east of the crests of the Nakhon Si Thammarat and Sankalakhiri mountains ( Fig. 1 ).

Its presence on Ko Tao Island, Surat Thani Province, approximately 85 km off the coast from Muang District, Chumpon Province, as opposed to the presence of Cnemaspis siamensis which is geographically closer to Ko Tao Island ( Fig. 1 ), is consistent with the geological history of this part of Peninsular Thailand. The island chain consisting of Ko Tao and the intervening islands Ko Samui and Ko Phangan, are offshore extensions of the Nakhon Si Thammarat Mountains to which they were connected during the last glacial maximum ( Sathiamurthy & Voris 2006) and lie to the east of the Isthmus of Kra ( Fig. 1 ). It is likely that C. chanardi also occurs on Ko Samui and Ko Phangan.

Natural history. Taylor (1963) stated that at Khao Chong, Trang, Cnemaspis chanardi sp. nov. is primarily a rock dweller but can be found on the buttresses of trees and trunks in the general vicinity of rocks.

Taylor (1963) noted that lizards sought refuge in the crevices of boulders and moved across their rocky microhabitat with considerable speed. Taylor (1963) further noted that lizards were seen abroad during the morning hours, indicating this species is not strictly nocturnal. We made similar observations at the Phuphaphet Cave area in Satun ( Fig. 8 ). Here lizards were collected and observed during the day on the buttresses of trees and within tree holes between 1.5–2 m above the ground along a foot path in old, secondary forest. All the trees upon which the lizards were observed had holes into which the lizards would retreat upon provocation ( Fig. 8 ). Several rock outcrops were nearby but no lizards were observed on them. At the Dad Fa Waterfall, Surat Thani Province, lizards were found at 20:30 hrs beneath rocks 30 m from the nearest stream. Specimens at Khao Luang were found during the day on the ground beneath small rocks on forested hillsides between 200 and 300 m in elevation. There were no large outcrops of rock nearby. These observations indicate C. chanardi sp. nov. is a diurnal, scansorial species that utilizes large, open, above-ground substrates (i.e. tree trunks and boulders) and does not occupy habitats that do not contain both trees and rocks. A gravid female carrying two eggs was found near the Phuphaphet cave on 7 October 2009. THNHM 10115 from near Khao Nan National Park contains two eggs. USNM 76144 from Ko Tao Island and THNHM 13838 collected on 14 December from Bad Yod Leong also contain two eggs.

Etymology. The specific epithet chanardi , a masculine name in the genitive case, is in reference to Mr. Tanya Chan-ard of the Thailand Natural History Museum, National Science Museum, Bangkok for his extensive contributions to the herpetology of Thailand, his gracious assistance with this project, and for collecting much of the material used in this study.

Comparisons. Cnemaspis chanardi sp. nov. is most similar to C. kamolnorranathi sp. nov., C. roticanai Grismer & Chan 2010 , C. siamensis , and C. vandeventeri sp. nov. of Peninsular Thailand ( C. roticanai is biogeographically considered a Thai species owing to the geological history that Pulau Langkawi shares with the mountains of Peninsular Thailand despite the fact that it is politically a Malaysian island). In fact, C. chanardi sp. nov., C. siamensis , and C. vandeventeri sp. nov. were at one time considered conspecific ( Smith 1925, 1930, 1935; Taylor 1963). Cnemaspis chanardi sp. nov. can be differentiated from all Southeast Asian species of Cnemaspis ( TABLE 1 ) except C. roticanai and C. vandeventeri sp. nov. on its having a lightly colored, prescapular crescent ( Fig. 4 ). It is differentiated from C. roticanai and C. vandeventeri sp. nov. by being much smaller (maximum SVL 40.1 mm vs. 44.7 mm for C. vandeventeri and 47.0 mm for C. roticanai ) and from C. vandeventeri sp. nov. in having, as opposed to lacking, tubercles on the flanks ( Fig. 5 ). It differs further from C. siamensis in having, as opposed to lacking precloacal pores in males and from C. siamensis in lacking, as opposed to having, dark, longitudinal, gular markings. From C. kamolnorranathi sp. nov. it can be separated further in having keeled as opposed to smooth ventral and subtibial scales. Cnemaspis chanardi sp. nov. is easily diagnosed from all other species of Southeast Asian Cnemaspis on the basis of several aspects of squamation ( TABLE 1 ).

Remarks. The precloacal pores of juvenile and subadult male Cnemaspis chanardi sp. nov. are difficult to discern. THNHM 14111, a juvenile (SVL 23.5 mm), has one pigmentless, poorly developed, precloacal pore that appears to have just erupted through the surface of the enclosing scale. Contiguous, under-developed, pigmentless scales with glandular secretions that have not yet erupted through the scale surface, flank the pore-bearing scale. THNHM 1334, a subadult (SVL 33.5 mm), has three, nearly fully developed, precloacal pores flanked by pigmentless, contiguous, under-developed scales. The remaining adult males (SVL 35.1– 40.1 mm) have 4–8, small but fully developed precloacal pores that are not flanked by under-developed, pigmentless scales. Nonetheless, the precloacal pores in adults of this species are generally much smaller and harder to see than precloacal pores in most other species of Cnemaspis ( Fig. 13 ).

Cnemaspis chanardi sp. nov. is superficially similar to C. kamolnorranathi sp. nov. and the two species may actually be sympatric in Tai Rom Yen National Park where the former occurs at the Dad Fa Water Fall and the latter approximately 10 km to the northeast at the Petchphanomwat Water Fall. They can be differentiated on the basis of C. chanardi sp. nov. having well-developed, transversely elongate, precloacal pores that do not form a continuous row as opposed to having poorly developed precloacal pores that form a continuous row ( Fig. 13 ). Additionally, C. chanardi sp. nov. has large tubercles on its flanks ( Fig. 5 ) and C. kamolnorranathi sp. nov. does not. Cnemaspis chanardi sp. nov. utilizes both rocks and tree trunks whereas C. kamolnorranathi sp. nov. may be more restricted to rocks. A molecular analysis (Grismer et al., in prep.) will provide additional data to futher test the hypothesis that these nominal taxa are separate species as is suggested by their morphology, ecology, and distribution.

Additional material examined. Krabi Province: Khlong Thom THNHM 12439–40. Nakhon Si Thammarat Province: Khao Luang National Park THNHM 14111; Thum Panra District; Thum Thong Panra, PSUZC-RT 2010.53-54. Satun Province: Muang District, Phuphaphet Cave LSUHC 9565. Surat Thani Province: Donsak District, near Donsak Pier MS 395; Kanchanadit District, Petphanomwung Cave ZMKU Rep-000313, KZM 009; Ko Tao Island USNM 76143–44.

Figures

FIGURE 1. Distribution of Cnemaspis in Thailand. Squares represent type localities.

FIGURE 2. Cnemaspis siamensis from Pathio, Chumpon Province. Upper: male LSUHC 9485. Middle: female LSUHC 9474. Lower: ventral view of male LSUHC 9485 (photos by L. Grismer).

FIGURE 3. The urbanized type locality of Cnemaspis siamensis in the town of Pathio, Chumpon Province (photo by L. Grismer).

FIGURE 4. Type series of Cnemaspis chanardi sp. nov.

FIGURE 5. Upper: flank of Cnemaspis chanardi sp. nov. (CUZM R- 2009,6,24 - 9). Lower: flank of C. vandeventeri sp. nov. (THNHM 8261).

FIGURE 6. Left and lower right: Cnemaspis chanardi sp. nov. (male; CUZM R- 2009,6,24 - 9) from Dad Fa Waterfall in Tai Rom Yen National Park, Surat Thani Province. Upper right: C. chanardi sp. nov. (female; PSUZC-RT 2010.53) from Thum Thong Panra, Thum Panra District, Nakhon Si Thammarat (photos by M. Sumontha).

FIGURE 7. Upper: Cnemaspis chanardi sp. nov. (LSUHC 9565) from Phuphaphet Cave area, Satun Province (photo by M. Sumontha). Middle: C. chanardi sp. nov. (THNHM 1335; gravid female) from Khao Luang, Nakhon Si Thammarat Province (photo by M. Cota). Lower: uncataloged specimen of C. chanardi sp. nov. from Khao Nan Yai, Nakhon Si Thammarat Province (photo by T. Sarimanon).

FIGURE 8. General habitat (left) and tree hole microhabitat (right) of Cnemaspis chanardi sp. nov. at the Phuphaphet Cave area, Satun Province (photos by M. Sumontha).

FIGURE 13. Left: precloacal pores of Cnemaspis kamolnorranathi sp. nov. (holotype THNHM 15908). Right: precloacal pores of C. chanardi (CUZM R- 2009,6,24 - 9).

Tables

TABLE 1. Diagnostic characters of Southeast Asian species of Cnemaspis. New species in bold type. / = character not evaluated.

Maximum SVL	48.0	65.2	56.6	64.9
Supralabials	9–13	15	9–11	11–13
Infralabials	8–10	10–12	8–10	10–12
Ventral scales keeled (1) or not (0)	0	1	0	1
No. of precloacal pores	5–6	10	0	0
Precloacal pores continuous (1) or separated (0)	0	1	/	/
Precloacal pores elongate (1) or round (0)	0	0	/	/
No. of paravertebral tubercles	20–28	28–31	23–29	18–27
Tubercles of ventralmost row on flank linearly arranged
and in contact or nearly so (1) or tubercles on flank
widely spaced and more randomly distributed (0)	0	0	1	0
Tubercles present (1) or absent (0) on flanks	1	1	1	1
Caudal tubercles in lateral furrow (1) or not (0)	0	0	0	0
Ventrolateral caudal tubercles anteriorly (1) or not (0)	0	0	1	1
Lateral caudal tubercle row present (1) or absent (0)	1	1	1	1
Caudal tubercles restricted to a single paravertebral
row on each side (1) or not (0)	0	0	0	0
Subcaudals keeled (1) or not (0)	1	1	0	0
Single median row of keeled subcaudals (1) or not (0)	0	0	1	0
Caudal tubercles encircle tail (1) or not (0)	0	0	0	1
Enlarged median subcaudal scale row (1) or not (0)	0	0	1	0
No. of postcloacal tubercles	2	3,4	2	2
Enlarged femoral scales present (1) or absent (0)	0	0	0	0
Shield-like subtibial scales present (1) or absent (0)	0	0	0	0
Subtibial scales keeled (1) or not (0)	1	1	1	1
Enlarged submetatarsal scales on 1st toe (1) or not (0)	0	0	1	0
No. of 4th toe lamellae	17–20	37–39	27–30	26–27

TABLE 2. Diagnostic characteristics of color pattern of the Cnemaspis of Thailand. 1 = character state present; 0 = character state absent. / = character not evaluated.

biocellata Eye spots on occiput 1	chanardi 0	chanthaburiensis 0	kamolnorranathi 0	kumpoli 0	huaseesom 0	mcguirei 0	punctatonuchalis 0	narathiwatensis 0	siamensis 0	vandeventeri 0
Black shoulder patch enclosing ocellus in males 1	0	0	0	1	0	1	0	1	0	0
Wide, yellow, nuchal loop in males 1	0	0	0	0	0	0	0	0	0	0
Black neck patch enclosing an ocellus 0	0	0	0	0	0	0	1	0	0	0
Head yellow in adult males 0	0	0	0	0	1	0	0	0	0	0
Light colored, prescapular cresent 0	1	0	0	0	0	0	0	0	0	1
Light colored, postscapular bar 0	0	0	0	0	0	1	0	1	0	0
Dark, longitudinal, gular markings 0	0	0	0	0	0	0	0	0	1	0
Red bands on limbs in males 0	0	0	0	1	0	0	0	0	0	0
Light colored, transverse bars on flanks 0	1	0	0	1	0	1	0	0	0	0
Venter red-orange in males 0	0	1	0	0	0	0	0	0	0	1
Cnemaspis chanardi sp. nov. Chan-ard’s Rock Gecko Djing Djok Niew Yaow Tanya Figures 4,5,6,7,13
Gonatodes siamensis Smith, 1930:16 , Cnemaspis siamensis Smith, 1935:71 Cnemaspis siamensis (in part) Taylor, 1963:740