Cnemaspis perhentianensis, Grismer, Lee & Onn, Chan Kin, 2008

Grismer, Lee & Onn, Chan Kin, 2008, A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia, Zootaxa 1771, pp. 1-15 : 3-13

Cnemaspis perhentianensis sp. nov.

Figures 2 , 3

Holotype. Adult male ( ZRC 2.6675) collected on 19 October 2007 by L. Lee Grismer at 1000 h at 40 m a.s.l. on Pulau Perhentian Besar, Trengganu, West Malaysia (05° 54.054N, 102° 44.343E).

Paratypes. All adult females collected at the same locality in 2007 by L. Lee Grismer and Peter Caron. ZRC 2.6676 was collected on 18 October, ZRC 2.6677–79 were collected on 20 October.

Diagnosis. Cnemaspis perhentianensis differs from all other Southeast Asian Cnemaspis in having the unique combination of a maximum SVL of 47.2 mm; 8–10 supralabials; seven or eight infralabials; scales of anterior portion of forearm keeled; ventral scales weakly keeled; no femoral pores; eight preanal pores in the single male; no linearly arranged row of tubercles on the flanks; paravertebral longitudinal rows of caudal tubercles and lateral caudal rows anteriorly; keeled subcaudals lacking an enlarged median row; two or three cloacal tubercles; no large, shield-like subtibial or submetatarsal scales; 28–31 subdigital lamellae on fourth toe; distinct, large, dark spots on neck and back alternating with transverse, white markings; white markings on flanks alternating with dark blotches; dark bands encircling tail; and posterior 25% of tail not white. These differences are summarized across all Southeast Asian species in Table 1 .

Description of holotype. Adult male; snout-vent length 46.1 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.24), somewhat narrow (HW/SVL 0.15), flattened (HD/HL 0.42), distinct from neck; snout short (ES/HL 0.43), slightly concave in lateral profile, longer than eye diameter; postnasal region constricted medially, concave; scales of rostrum weakly keeled, raised, larger than those on occiput; low, rounded, supraorbital ridges; frontorostral sulcus absent; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.28); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; eye to ear distance greater than diameter of eye; rostral slightly concave, dorsal 75% divided by deep longitudinal groove; rostral bordered dorsoposteriorly by two small supranasals and one small azygous scale and posterolaterally by one small, lower postnasal and first supralabial; supralabials eight (R and L), decreasing in size posteriorly; eight (R) seven (L) infralabials, decreasing in size posteriorly, first and second infralabials nearly same size; nostrils elliptical, oriented dorsoposteriorly; three small, granular, postnasal scales (R and L) undifferentiated from lateral scales of rostrum; mental triangular, slightly deeper than wide, concave medially, bordered posteriorly by three postmentals; postmentals bordered posteriorly by nine smaller, rounded scales; enlarged sublabials absent; gular scales granular, slightly raised; throat scales weakly keeled, flat, oriented vertically.

Tubercles of ventralmost row on flank linearly arranged 0 0 1 0 and in contact or nearly so (1) or tubercles on flank widely

spaced and more randomly distributed (0)

Caudal tubercles present (1) or absent (0) 0 1 1 1 Lateral caudal tubercles present (1) or not (0) 1 1 1 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 1 1 1 Caudal tubercles restricted to a single paravertebral row on 0 0 0

each side (1) or not (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 Postcloacal spurs present (1) or absent (0) 2 3–4 2 1 Shield-like subtial 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 subdigital (1) or 0 0 1 0 not (0)

Tubercles of ventralmost row on flank linearly arranged 1 0 0 0 and in contact or nearly so (1) or tubercles on

flank widely spaced and more randomly distributed (0)

Caudal tubercles present (1) or absent (0) 1 1 1 1 Lateral caudal tubercles present (1) or not (0) 0 1 1? Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 0? Caudal tubercles restricted to a single paravertebral row on 1 0 0 0 each side (1) or not (0)

Subcaudals keeled (1) or not (0) 0 0 0 1 Single median row of keeled subcaudals (1) or not (0) 0 0 0 1 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 1 0 0 1 Postcloacal spurs present (1) or absent (0) 1 1–2 2 1 Shield-like subtial scales present (1) or absent (0) 1 1 0 0 Subtibial scales keeled (1) or not (0) 0 0 0 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 1 0 0 Tubercles of ventralmost row on flank linearly arranged 0 0 0 0 and in contact or nearly so (1) or tubercles on flank widely

spaced and more randomly distributed (0)

Caudal tubercles present (1) or absent (0) 1 1 1 1 Lateral caudal tubercles present (1) or not (0)? 1? 1 Ventrolateral caudal tubercles anteriorly (1) or not (0)? 1? 1 Caudal tubercles restricted to a single paravertebral row on? 0 0 0 each side (1) or not (0)

Subcaudals keeled (1) or not (0) 1 1 1 1 Single median row of keeled subcaudals (1) or not (0)? 0 1 1 Caudal tubercles encircle tail (1) or not (0)? 1 0 1 Enlarged median subcaudal scale row (1) or not (0) / 0 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 0 0 0 Tubercles of ventralmost row on flank linearly arranged 0 0 0 0 and in contact or nearly so (1) or tubercles on flank widely

spaced and more randomly distributed (0)

Caudal tubercles present (1) or absent (0) 1 1 1 1 Lateral caudal tubercles present (1) or not (0) 1 1 1 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 1 0 1 Caudal tubercles restricted to a single paravertebral row on 0 0 0 0 each side (1) or not (0)

Subcaudals keeled (1) or not (0) 0 0 1 0 Single median row of keeled subcaudals (1) or not (0) 0 0 0 0 Preanal pores present (1) or absent (0) 0 0 0 No. of preanal pores / / / Tubercles of ventralmost row on flank linearly arranged 1 0 0 and in contact or nearly so (1) or tubercles on flank widely

spaced and more randomly distributed (0)

Caudal tubercles present (1) or absent (0) 1 1 1 Lateral caudal tubercles present (1) or not (0) 0? 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0? 1 Caudal tubercles restricted to a single paravertebral row on 1 0 0 each side (1) or not (0)

Subcaudals keeled (1) or not (0) 0 0 1 Single median row of keeled subcaudals (1) or not (0) 1 0 0 Caudal tubercles encircle tail (1) or not (0) 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 1 1 0 Postcloacal spurs present (1) or absent (0) 2–3 0 3–4 Shield-like subtial scales present (1) or absent (0) 0 0 0 Subtibial scales keeled (1) or not (0) 0? 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 0 0 Body slender, elongate; dorsal scales equal in size throughout body; dorsal tubercles more or less randomly distributed from occiput to level of hind limb insertions; dorsal tubercles smooth, lacking keels or ridges; pectoral and abdominal scales weakly keeled, flat, slightly elongate, imbricate, equal in size throughout; ventral scales slightly larger than dorsal scales; eight precloacal pores arranged in a chevron, not separated by intervening scales lacking pores; precloacal depression absent; femoral pores absent.

Forelimbs moderately long, slender; dorsal scales of brachium slightly raised, weakly keeled; scales of forearm same size as brachials, imbricate, those on anterior margin keeled; ventral scales of brachium smooth, rounded, juxtaposed; scales beneath forearm, smooth, flat, slightly raised; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws very slightly recurved; subdigital lamellae unnotched; lamellae beneath first phalanges slightly widened; slight interdigital webbing; fingers increase in length from first to fourth with fourth and fifth equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales of thigh smooth, slightly raised, juxtaposed to subimbricate; scales of anterior margin of thigh weakly keeled; ventral scales of thigh flat, imbricate; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws slightly recurved; subdigital lamellae unnotched; lamellae beneath first phalanx not widened; interdigital webbing weak; toes increase in length from first to fourth with fourth and fifth equal in length; 31 subdigital lamellae on fourth toe (R, L).

Tail 1.2 times SVL, swollen at base; caudal scales arranged in segmented whorls, first segment at base of tail six scales wide midventrally; anteriorly caudal scales raised, juxtaposed, weakly keeled becoming flatter, imbricate, and keeled posteriorly; shallow middorsal furrow; deeper, single lateral furrow; no enlarged, median subcaudal scales; subcaudal scales keeled; a paravertebral and lateral longitudinal row of large, flattened tubercles on either side of midline; ventrolateral row of tubercles anteriorly, fading posteriorly; transverse tubercle rows do not completely encircle tail; three (R) four (L) enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail.

Coloration (in life). Dorsal ground color grey to brown, overlain by irregularly shaped white markings on top of head and snout; paired white markings on occiput; no postorbital stripes; squarish, medial white marking on neck; six distinct, irregularly shaped, paravertebral, white markings on dorsum extend from shoulders to base of tail alternating with transversely elongate, distinct, white markings on flanks; three, small, elongate dark blotches at base of occiput; seven pairs of dark, poorly defined, paravertebral blotches extending from nape to anterior portion of tail, alternating with the light paravertebral markings; dark blotches on flanks alternating with white, transverse markings on flanks; black and dull white bands nearly encircle tail; irregularly shaped dark and light markings on limbs; dark and light bands encircling digits; ventral surfaces of neck, body, and limbs dull beige, immaculate; gular region smudged with dark stippling. This coloration has not been significantly altered by preservation.

Variation. The female paratypes closely approach the holotype in all aspects of coloration (Figs. 2,3) except that the white dorsal markings and the dark dorsal blotches are larger and even more distinctive in ZRC 2.6676 and ZRC 2.6678–79. The females lack preanal pores and their postcloacal tubercles are smaller. Morphometric variation and variation in scalation is presented in Table 2.

Distribution. Cnemaspis perhentianensis is known only from Pulau Perhentian Besar and Pulau Perhentian Kecil, just 0.5 km to the north. Three specimens on Pulau Perhentian Kecil were observed on 11 March 2008 but were not collected.

Natural history. The Perhentian Archipelago is composed of 11 relatively small islands lying 21 km off the east coast of the state of Terengganu ( Fig. 1 ). The largest of these islands, Pulau Perhentian Besar (ca. 857 hectares) is a rugged, hilly island reaching 249 m a.s.l. The majority of the island is covered in primary lowland dipterocarp forest and its granite bedrock is the source of extensive boulder outcrops that adds a significant degree of habitat and microhabitat heterogeneity to the island’s ecosystem, which in turn, supports TABLE 2. Descriptive measurements and characteristics of the type series of Cnemaspis perhentianensis . H = holotype; p = paratype

ZRC ZRC ZRC ZRC ZRC 2.6675 2.6676 2.6677 2.6678 2.6679 various saxicolous species ( Tamblyn 2005). One of these species, Cnemaspis perhentianensis , is solely restricted to outcroppings of granite boulders. Specimens collected or observed during the day were found on the shaded surfaces of both large and small rocks and would retreat into crevice microhabitats when threatened. Upon retreat, some would roll their tail over their back and wag the tip from side to side. At night, lizards could be found on all surfaces of the rock at greater distances from their crevice microhabitats. Egg scars were commonly found in the rock cracks and no specimens were seen on trees trunks or other vegetation.The above observations were made during or immediately following rain showers from 18-20 October 2007. During the second visit from 11-12 March 2008, no specimens were observed on Pulau Perhentian Kecil during the day or night. Following an afternoon rain shower, however, three specimens were observed during the evening of 12 March on Pulau Perhentian Kecil. This suggests that this species activity may be closely tied to precipitation.

Etymology. The specific epithet perhentianensis is in reference to the Perhentian Besar Island. The suffix ensis is a derivation meaning “from” or “inhabiting” and renders the specific epithet an adjective that must be in grammatical accord with the gender of Cnemaspis .

Comparisons to other species. Cnemaspis perhentianensis differs from C. argus , C. baueri , C. boulengeri , C. kendallii , C. kumpoli , C. limi , C. nigridius , C. pemanggilensis , and C. aurantiacopes in having a maximum SVL of less than 56.6 mm. It differs from C. dezwaani , C. jacobseni , C. modiglianiii , C. phuketensis , and C. siamensis in being larger, having a maximun SVL greater than 39.7 mm. Cnemaspis perhentianensis differs from C. argus , C. baueri , C. dringi , C. limi , and C. nigridius in having less than 11 supralabials and from C. dezwaani , C. jacobseni , C. modiglianii , C. phuketensis , and C. whittenorum in having more than seven supralabials. Cnemaspis perhentianensis has less than nine supralabials which differentiates it from C. affinis , C. flavolineatus , C. kendallii , C. kumpoli , C. limi , C. nigridius , C. pemanggilensis , C. siamensis , C. aurantiacopes , and C. tucdupensis and more than six infralabials differentiates it from C. whittenorum . Cnemaspis perhentianensis resembles all other species of Cnemaspis except C. affinis , C. argus , C. baueri , C. boulengeri , C. dringi , C. nigridius , C. pemanggilensis , and C. siamensis in having keeled scales on the forearm. All species of Southeast Asian Cnemaspis lack femoral pores except for C. affinis , C. dezwaani , and C. modiglianiii . Preanal pores occur in male C. perhentianensis but are lacking in male C. argus , C. baueri , C. boulengeri , C. jacobseni , C. kendallii , C. limi , C. pemanggilensis , C. phuketensis , C. whittenorum , C. aurantiacopes , and C. tucdupensis . At this point, C. perhentianensis (N=1) is differentiated from C. affinis , C. dringi , C. modiglianiii , C. caudanivea , and C. nuicamensis in having more than six preanal pores. It is differentiated from C. nigridius in having less than 16 preanal pores. Cnemaspis perhentianensis is differentiated from C. boulengeri , C. aurantiacopes , and C. tucdupensis in that the ventralmost row of body tubercles is not composed of tubercles that are linearly arranged. All Southeast Asian species of Cnemaspis except C. affinis and C. phuketensis have caudal tubercles and only C. boulengeri , C. nuicamensis , and C. tucdupensis lack a lateral row (this condition is unkown for C. dezwaani , C. dringi , C. jacobseni , and C. whittenorum ) and only C. affinis , C. boulengeri , C. chanthaburiensis , C. kumpoli , C. modiglianiii , C. caudanivea , and C. tucdupensis lack a ventrolateral row (this condition is unknown for C. dezwaani , C. dringi , C. jacobseni , C. siamensis , and C. whittenorum ). Only in C. boulengeri and C. tucdupensis are the caudal tubercles are restricted to a single paravertebral row on either side of the midline. Cnemaspis perhentianesis can be differentiated from C. baueri , C. boulengeri , C. chanthaburiensis , C. kumpoli , C. limi , C. nigridius , C. phuketensis , C. whittenorum , C. caudanivea , C. aurantiacopes , C. nuicamensis , and C. tucdupensis in having as opposed to lacking, keeled subcaudals although it lacks a single, median row of keeled subcaudals unlike C. dezwaani , C. jacobseni , C. kendallii , C. pemanggilensis , C. phuketensis , C. aurantiacopes , C. nuicamensis , and C. tucdupensis where this row is present (the condition in C. dringi is unknown). Cnemaspis perhentianensis is differentiated from all other Southeast Asian species except C. baueri , C. flavolineata , C. kendallii , C. pemanggilensis , and C. phuketensis in that the caudal tubercles do not encircle the tail (the condition in C. dringi is unknown). Unlike C. boulengeri , C. dezwaani , C. jacobseni , C. kendallii , C. limi , C. modigianii , C. nigridius , C. pemanggilensis , C. phuketensis , C. siamensis , C. whittenorum , C. aurantiacopes , C. nuicamensis , and C. tucdupensis , C. perhentianesis lacks an enlarged, median subcaudal row of scales. Cnemaspis perhentianensis differs from C. dringi and C. whittenorum in having enlarged postcloacal tubercles (this condition is not known for C. siamensis ) and differs from C. affinis , C. baueri , C. boulengeri , C. chanthaburiensis , C. dezwaani , C. flavolineata , C. jacobseni , C. kendallii , C. limi , C. modiglianiii , C. nigridius , C. pemanggilensis , C. phuketensis , C. caudanivea , C. aurantiacopes , and C. nuicamensis in having more than two postcloacal tubercles. Cnemaspis perhentianensis differs from C. boulengeri and C. caudanivea in lacking shield-like subtibial scales. Cnemaspis perhentianensis has keeled subtial scales unlike C. argus , C. boulengeri , C. chanthaburiensis , C. modiglianiii , C. pemanggilensis , C. caudanivea , C. nuicamensis , and C. tucdupensis which have smooth subtibial scales (the condition in C. nigridius , C. phuketensis , C. siamensis , and C. whittenorum is unknown). Cnemaspis perhentianensis differs from C. boulengeri , C. dringi , C. caudanivea , C. aurantiacopes , and C. tucdupensis in lacking enlarged submetatarsal scales on the first toe. Cnemaspis perhentianensis differs from all other Southeast Asian Cnemaspis except C. flavolineata , C. kumpoli , C. pemanggilensis , C. caudanivea , C. aurantiacopes , C. nuicamensis , and C. tucdupensis in having more than 27 subdigital lamellae on the fourth toe. Cnemaspis perhentianensis differs from all other Southeast Asian species of Cnemaspis except C. dringi , C. flavolineatus , C. kumpoli , and C. pemanggilensis in having as opposed to lacking, distinctive white markings on the flanks. Cnemaspis perhentianensis differs from all other Southeast Asian species of Cnemaspis except C. boulengeri and C. baueri in having as opposed to lacking, distinct large, dark, markings on the neck. Cnemaspis perhentianensis differs from C. baueri , C. boulengeri , C. kendallii , C. limi , C. nigridius , C.

phuketensis , C. aurantiacopes , and C. nuicamensis in having as opposed to lacking, black bands on the tail. Cnemaspis perhentianensis differs from C. caudanivea in lacking a white tail tip. These character states are summarized in Table 1 .

Discussion. The need for a phylogeny of Southeast Asian Cnemaspis can not be understated. With the presence of nine species of this genus in Peninsular Malaysia and 31 total, Cnemaspis comprises a significant portion of the lizard fauna ranging across much of Southeast Asia and until these relationships are resolved, little else can be said about the evolutionary biology of this large, widespread group. One of the most striking features of C. perhentianensis is its color pattern of alternating large, dark and light markings. This pattern is reminiscent of that found in C. argus , a relatively large montane endemic known from Gunung Lawit, Trengganu ( Dring 1979) just opposite the Perhentian Archipelago on the Malay Peninsula, and to a lesser extent C. kumpoli , also known from the adjacent peninsula. Although these putative relationship scenarios make reasonable biogeographic sense, their testing must await a phylogenetic analysis (Bauer et al. in prep.).

The presence of an endemic species in the Perhentian Archipelago is not surprising and still other species from this archipelago are in various stages of description (Grismer, in preparation). The archipelagos associated with Peninsular Malaysia are emerging as significant centers of endemism ( Grismer et al. 2006a, b; Grismer 2008) as so poignantly illustrated by the islands in the Seribuat Archipelago off the southeast coast of the Malay Peninsula that harbor at least 16 endemic species ( Grismer 2006). Such is the case for the islands of the Langkawi Archipelago off the northwestern coast of Peninsular Malaysia where endemic species are now beginning to be described ( Grismer 2008; Chan et al. in prep.) as research there continues. Additional field research in all these archipelagos including the Redang Archipelago, approximately 27 km southwest of the Perhentian Archipelago, will undoubtedly result in the discovery of additional new taxa. These island groups will ultimately prove to be significant centers of herpetological diversity for Peninsular Malaysia.

Figures

FIGURE 1. Distribution of Cnemaspis perhentianensis on Pulau Perhentian Besar, Terengganu, Peninsular Malaysia.

FIGURE 2. Type series of Cnemaspis perhentianensis. Left to right; Holotype ZRC 2.6675, paratypes ZRC 2.6677, 2.6676, 2.6679 and 2.6678.

FIGURE 3. Paratypes of Cnemaspis perhentianensis. Upper; ZRC 2.2279. Lower; ZRC 2.2278.

Tables

TABLE 1. Diagnostic characters of Southeast Asian species of Cnemaspis. / = character not applicable;? = character could not be evaluated from specimens or literature. New species in bold type.

Maximun SVL	affinis 48.0	argus 65.2	aurantiacopes baueri 56.6 64.9
Supralabials Infralabials	9–13 8–10	15 10–12	9–11 11–13 8–10 10–12
Forearm scales keeled (1) or not (0) Ventral scales keeled (1) or not (0)	0 0	0 1	1 0 0 1
Femoral pores present (1) or absent (0)	1	0	0 0
No. of femoral pores Preanal pores present (1) or absent (0)	/ 1	/ 0	/ / 0 0
No. of preanal pores	5–6	/	/ /