Zootaxa 3755 (5): 447 456 www.mapress.com/zootaxa/ Copyright 2014 Magnolia Press Article http://dx.doi.org/10.11646/zootaxa.3755.5.4 http://zoobank.org/urn:lsid:zoobank.org:pub:6de7daea-16aa-48f8-bee2-d485d0578151 ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) A new species of insular Rock Gecko (Genus Cnemaspis Strauch, 1887) from the Bidong Archipelago, Terengganu, Peninsular Malaysia L. LEE GRISMER 1, PERRY L. WOOD, JR. 2, AMIRRUDIN B. AHMAD 3, ALEXANDRA S.-I. SUMARLI 1, JESSIKA J. VAZQUEZ 1, LUKMAN H. B. ISMAIL 3, RONALD NANCE 1, MUHAMMAD AFIF B. MOHD-AMIN 3, MOHAMAD N. A. B. OTHMAN 3, SYED A. RIZAIJESSIKA 3, MARIA KUSS 1, MATTHEW MURDOCH 1 & ANTHONY COBOS 1 1 Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515 USA. E-mail: lgrismer@lasierra.edu 2 Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: pwood@byu.edu 3 Fakulti Sains dan Teknologi, Universiti Malaysia Terengganu, Kualau Terengganu, Malaysia E-mail: amirrundin@umt.edi.my Abstract A new insular species Cnemaspis bidongensis sp. nov. (Squamata: Gekkonidae), is described from Pulau Bidong, Terengganu, Peninsular Malaysia and bears a unique suite of morphological and color pattern characters that differentiate it from all other congeners. Cnemaspis bidongensis sp. nov. is the sister species to C. kendallii (Gray) and represents the fifth insular endemic species of Cnemaspis on archipelagos along the east coast of Peninsular Malaysia. This species survived massive deforestation of the small island of Bidong (260 ha) from the mid 1970s to the early 1990s when the island served as a Vietnamese refugee camp and harbored as many as 40,000 people at one time. We hypothesize that this species generalized lifestyle contributed to its survival, allowing it to seek refuge in rocky microhabitats. Key words: new species, Cnemaspis, endemic, Bidong Island, conservation, biodiversity, Terengganu, Peninsular Malaysia Introduction The archipelagos of the South China Sea are proving to be some of the most herpetologically diverse islands in Southeast Asia (Grismer et al. 2011; Leong et al. 2003). Even though those along the southern coast of Vietnam and the east coast of Peninsular Malaysia have been well-studied (Grismer 2011a; Grismer et al. 2006, 2011), the increase in their diversity and endemism shows no signs of abating (i.e., Chan & Norhayati 2010; Chan et al. 2011; Das & Grismer 2003; Das & Jim 2000; Diaz et al. 2004; J. Grismer et al. 2003; Grismer 2005, 2006; Grismer & Chan 2008; Grismer & Das 2006; Grismer & Ngo 2007; Grismer et al. 2003, 2004a,b, 2008, 2009, 2010a; Ngo 2008; Ngo & Grismer 2012; Ngo et al. 2008, 2010; Youmans & Grismer 2006). One of the island systems along Peninsular Malaysia just recently surveyed (Vazquez et al. in prep.) is the Bidong Archipelago, lying 35 km northeast of Kuala Terengganu, Terengganu (Fig. 1) that is composed of six wellvegetated, low-lying islands. Given that the Perhentian Archipelago to the north and the Tenggol Archipelago to the south harbor endemic species of lizards (Chan & Norhayati 2010; Grismer & Chan 2008; Grismer et al. 2009), it is not surprising that a recent survey of the Bidong Archipelago (Vazquez et al. in prep.) resulted in the discovery of a new, presumably endemic gekkonid. The six specimens collected have broad, flattened heads; large, somewhat forward and upwardly directed eyes bearing round pupils; a flattened body; and long, widely splayed limbs bearing long, inflected digits which place them in the genus Cnemaspis (Grismer et al. 2010b). However, they bear a suite of unique character states and a large, uncorrected percent sequence divergence from closely related species that indicates they belong to a new species. As such the population is described herein. Accepted by A. Bauer: 16 Dec. 2013; published: 24 Jan. 2014 447
FIGURE 1. Distribution of the endemic insular species of Cnemaspis on archipelagos off the east coast of Peninsular Malaysia. Material and methods Color characters were taken from digital images of living specimens cataloged in the La Sierra University Digital Photo Collection (LSUDPC) and in some cases, from living specimens. The following measurements on the type series were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate: snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90 to the inflection of the flexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90 to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eye-ball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eye-ball; eye to snout distance (ES), measured from anteriormost margin of the eye-ball to the tip of snout; eye to nostril distance (EN), measured from the anterior margin of the eye-ball to the posterior margin of the external nares; inner orbital distance (IO), measured between the anterior edges of the orbit; ear length (EL), the greatest horizontal distance of the ear opening; and internarial distance (IN), measured between the nares across the rostrum. Additional character states evaluated were numbers of supralabial and infralabial scales counted from below the middle of the orbit to the rostral and mental scales, respectively; size and number of postmental scales contacting the mental; the texture of the scales on the anterior margin of the forearm; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column (where applicable); the presence or absence of a row of enlarged, widely spaced tubercles along the ventrolateral edge of the body between the limb insertions; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the total number of precloacal pores, their orientation, shape, 448 Zootaxa 3755 (5) 2014 Magnolia Press GRISMER ET AL.
and degree of separation; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales, subtibial scales, and submetatarsal scales beneath the first metatarsal; and the number of postcloacal tubercles on each side of the tail base. Longitudinal rows of caudal tubercles on the nonregenerated portion of the tail are quite variable between species and useful in differentiating several taxa. Up to five pairs of the following rows may be present in varying combinations: paravertebral row the dorsal row adjacent to the middorsal, caudal furrow; dorsolateral row the row between the paravertebral row and the lateral, caudal furrow on the dorsolateral margin of the tail; lateral row the row immediately below the lateral, caudal furrow; and ventrolateral row the row below the lateral row on the ventrolateral margin of the tail. When present, this row is usually restricted to the anterior 25% (or less) of the tail. Rarely there may be a row of tubercles within the lateral, caudal furrow. Various color pattern characteristics were also evaluated (see description). A 1335 aligned base pair fragment of the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and its flanking t-rnas (t-rna-trp, t-rna-ala, t-rna-asn, and t-rna-cys) was amplified using the following primers L4437b 5 -AAGCAGTTGGGCCCATACC-3 and L5002 5 -AACCAAACCCAACTACGAAAAAT-3 (Macey & Schulte 1999) and sequenced for nearly all named species of Cnemaspis (Grismer et al. in prep.). Uncorrected pairwise sequence divergences were calculated in MEGA v5.2.1 (Tamura et al. 2011) for the undescribed Pulau Bidong species (two specimens: 11445, 11447) and its sister taxon C. kendallii (Gray) (27 specimens: 3773, 3797, 3820, 4707, 4756, 4958, 5056, 5073, 5184 85, 5523, 5703, 5731, 6380 81, 8122, 8126, 8910, 8965 67, 9376 77, 9380, 9382, 10238 39) from throughout its distribution in Peninsular Malaysia (see Grismer 2011b). Specimens examined as comparative material are listed in the appendices of Grismer et al. (2010b, 2013) and Wood et al. 2013). refers to the La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA. Results The molecular analysis places the Pulau Bidong population within the Cnemaspis kendallii group (see Comparisons section below) and within this group, it is the sister species to C. kendallii. Additionally, this population and C. kendallii bear a 6.1 9.7% sequence divergence. The morphological analysis shows that the Pulau Bidong population differs from all members of the C. kendallii group having relatively smaller maximum SVL; nine or ten supralabials; an enlarged, elongate mental scale; 21 26 paravertebral tubercles; no tubercles on the ventral portion of the flanks; no single, median row of keeled subcaudal scales; a median row of enlarged subcaudal scales; sexually dimorphic dorsal color pattern; the posterior two-thirds of the tail black in adult males; and the subcaudal region yellow in adult males (Table 1). Given that the Pulau Bidong population is not embedded within any other species in the molecular analysis; has a 6.1 9.7% sequence divergence from its sister species Cnemaspis kendallii; and the morphological analysis separates it from all other congeners we describe it below as: Cnemaspis bidongensis sp. nov. Pulau Bidong Rock Gecko Cicak Batu Pulau Bidong Figs. 1, 2 Holotype. Adult female ( 11455) collected on 26 August 2013 by Jacob A. Chan at 2200 hrs from Pulau Bidong, Terengganu, Peninsular Malaysia (5 37.201 N 103 03.244 E; 49 m elevation). Paratype. Adult males ( 11447, 11452 54) and adult female 11451 bear the same data as the holotype except they were collected between 1800 and 2400 hrs. Diagnosis. Cnemaspis bidongensis sp. nov. differs from all other Southeast Asian species of Cnemaspis in having the unique combination of adult males reaching at least 56.1 mm SVL, adult females reaching 58.1 mm SVL; nine or 10 supralabials; 7 9 infralabials; enlarged, elongate mental scale; keeled ventrals; no precloacal pores; moderately prominent, randomly arranged, dorsal tubercles; 21 26 paravertebral tubercles; no tubercles on A NEW INSULAR CNEMASPIS FROM MALAYSIA Zootaxa 3755 (5) 2014 Magnolia Press 449
lower flanks caudal tubercles encircling tail; no tubercles in lateral caudal furrows; ventrolateral caudal tubercles present anteriorly; subcaudals keeled; a median row of enlarged, keeled subcaudals; one or two postcloacal tubercles on each side; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; 26 30 subdigital lamellae on fourth toe; dark and light caudal bands distinct in both sexes; and no dark mottling on lateral portions of belly. These differences are summarized across all species of the C. kendallii species group in Table 1 and all across Southeast Asian species in Wood et al. (2013:Table 4). FIGURE 2. Upper left: adult female holotype ( 11455) of Cnemaspis bidongensis sp. nov. from Pulau Bidong, Terengganu, Peninsular Malaysia. Upper right: adult male paratype 11447. Lower left: ventral views adult male paratype 11452 (upper) and adult female holotype 11455 (lower). Lower right: adult male C. kendallii (LSUDPC 4426) from Gunung Ledang, Johor, Peninsular Malaysia. Description of holotype. Adult female; SVL 58.1 mm; head oblong in dorsal profile, moderate in size (HL/ SVL 0.26), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.44), distinct from neck; snout short (ES/HL 0.52), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum weakly keeled, raised, slightly larger than more rounded scales on occiput; low, supraorbital ridges; moderate frontal sulcus; canthus rostralis weak; eye large (ED/HL 0.21); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave, dorsal 95% divided by longitudinal groove; rostral bordered posteriorly by supranasals and laterally by first supralabials; nine R,L raised supralabials tapering in size posteriorly; nine R,L infralabials, decreasing in size posteriorly; nostrils elliptical, oriented posterolaterally, bordered posteriorly by small, granular, postnasal scales; mental large, triangular, greatly elongate, extending posteriorly to level of 4th infralabials, bordered posterolaterally by six scales (postmentals), two of which are elongate and contact the majority of the mental; gular scales raised, keeled; throat scales larger, raised, keeled and grading into larger, keeled pectoral scales. Body stature moderate (AG/SVL 0.44); small, keeled, dorsal scales equal in size throughout body, intermixed with a sparse array of slightly larger, randomly arranged multicarinate tubercles; tubercles extend from occiput to base of tail; no tubercles on lower flanks; 21 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, similar size throughout; abdominal scales slightly larger than dorsals; no precloacal pores; forelimbs 450 Zootaxa 3755 (5) 2014 Magnolia Press GRISMER ET AL.
moderately long, slender (FL/SVL 0.18); dorsal scales of brachium raised, weakly keeled; dorsal scales of forearm raised, keeled; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm weakly, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges not granular, more widened distally; lamellae beneath phalanx immediately following inflection not granular but wide like lamellae of distal phalanges; no interdigital webbing at base of digits; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs (HL/SVL 0.22); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior surface of thigh keeled; ventral scales of thigh raised, weakly keeled; subtibial scales keeled, flat, not subimbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges narrow but not granular proximally, wider distally; lamellae beneath phalanx immediately following inflection granular, wide like lamellae of distal phalanges; weak interdigital webbing at base of digits; toes increase in length from first to fourth with fourth being slightly longer than fifth; 29 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales raised anteriorly, weakly keeled, juxtaposed; no middorsal furrow; lateral furrows shallow; a median row of enlarged, keeled subcaudals; caudal tubercles encircle tail; caudal tubercles absent from lateral furrow; one enlarged postcloacal tubercle on lateral surface of hemipenial swellings at base of tail. TABLE 1. Diagnostic characters (bold) differentiating Cnemaspis bidongensis sp. nov. from other species in the C. kendallii group. bidongensis sp. nov. baueri kendallii pemanggilensis Maximum SVL (mm) 58.1 67.4 76.0 60.0 Supralabials 9,10 11 13 7 12 10,11 Enlarged, elongate mental scale Present Absent Absent Absent Paravertebral tubercles 21 26 18 27 17 26 30 37 Tubercles on ventral portions of flanks No No Yes No Single, median row of keeled subcaudal scales No No No Yes Enlarged median subcaudal scale row Yes Yes No Yes Sexually dimorphic dorsal pattern Yes No No No Dorsal pattern with elements of dark and light blotches Yes No Yes Yes Posterior two-thirds of tail black in males Yes Yes Yes No Subcaudal region yellow in adult males No No Yes No Color pattern in life (Fig. 2). Dorsal ground color of head, body, limbs and tail brown, and the posterior portion of tail black; a series of diffuse, yellowish lines on rostrum extend posteriorly onto frontal region; dorsal pattern on occiput consisting of a series of dull white spots surrounding a dark brown, tear-drop shaped, vertebral marking; three diffuse, dark brown, postorbital stripes radiate from eyes; dorsal pattern of neck and body consists of a vertebral series of six dull-white blotches extending to base of tail paralleled by similar blotches on flanks; a series of seven diffuse dark brown blotches extend from side of neck along flanks to base of tail on each side of body; intervening area between all body blotches consists of a network of dark and light mottling that extends onto the limbs; nine white caudal bands infused with faint black speckling encircle tail; inter-band areas bear black mottling; ventral surfaces of head, body and limbs beige to dull yellow with weak black stippling in each scale; anterior gular region yellow; subcaudal region bearing white, irregularly shaped bands; small, dark, elongate blotch on mental and medial postmental. Variation. The female paratype ( 11451) closely resembles the holotype in all aspects of coloration except that the overall pattern is slightly less bold and the posterior one-third of the tail is regenerated and bears an irregular lineate pattern (Fig. 2). The male paratypes ( 11447, 11452 54) depart significantly from the holotype in that sexual dimorphism in this species is marked (Fig. 2). Males have an overall yellowish to dullorange dorsal pattern on the head, body and tail and lack the dull-white blotching seen in females yet retain the darker blotching pattern. The caudal pattern is banded but the light bands are not white as in the females, the dark A NEW INSULAR CNEMASPIS FROM MALAYSIA Zootaxa 3755 (5) 2014 Magnolia Press 451
bands are dark brown as opposed to black, and the posterior portion of the tail is not black. The ventral pattern of males is similar to that of females except that the lateral margins of the abdomen tend to form a dark network enclosing small, lighter spots. Also, subcaudal banding is faint. Distribution. Cnemaspis bidongensis sp. nov. is presumed to be endemic to Pulau Bidong (Fig. 1) being that it has not been found in any other archipelago or any other island in the Bidong Archipelago (Vazquez et al. in prep.) Natural History. Cnemaspis bidongensis sp. nov. occurs in secondary, coastal forest and is widespread throughout the island. During the Vietnamese refugee period from May 1975 to October 1991, the island s primary forest was severely degraded by cutting. During this time, as many as 250,000 people fleeing the communist take over of southern Vietnam spent time on Pulau Bidong and in June 1979 it was considered the most heavily populated place on earth (http://en.wikipedia.org/wiki/bidong_island). Although this had a catastrophic effect on the native forest, C. bidongensis sp. nov. was able to survive because it is not a microhabitat specialist as are many other species of Cnemaspis (Grismer 2011a; Grismer & Chan 2009; Grismer & Ngo 2007; Grismer et al. 2010a,b; 2013; Wood et al. 2013). During the course of our fieldwork, lizards were observed day and night on both granite rocks and vegetation (Fig. 3). All lizards observed were wary, swift, agile and would seek shelter at the slightest provocation. During the day, lizards would often jump from rocks to nearby trees and escape by ascending 3 5 meters up the trunk a behavior we have not observed and do not know of being reported in any other species of Cnemaspis. Lizards would also avoid capture by retreating into rock cracks. During the evening, lizards were commonly seen on rocks, branches, and leaves where they appeared to be sleeping. When aroused, many would drop to the forest floor from as high as 1.5 meters and escape into the leaf litter a behavior also not known for species of this genus. Hatchlings and juveniles were not observed but the presence of gravid females carrying two eggs suggests that July is the beginning of the reproductive season. Etymology. The specific epithet bidongensis is an adjective in reference to the type locality Pulau Bidong, Terengganu, Peninsular Malaysia. FIGURE 3. Habitat structure of Cnemaspis bidongensis sp. nov. from Pulau Bidong, Terengganu, Peninsular Malaysia. 452 Zootaxa 3755 (5) 2014 Magnolia Press GRISMER ET AL.
TABLE 2. Meristic and mensural character states of the type series of Cnemaspis bidongensis sp. nov. Abbreviations are listed in Materials and Methods. 11455 11447 11451 11452 11453 11454 holotype paratype paratype paratype paratype paratype Sex F m f m m m Supralabials 9 9 10 9 9 9 Infralabials 9 8 9 8 8 7 Ventral scales keeled keeled keeled keeled keeled keeled No. of precloacal pores 0 0 0 0 0 0 No. of paravertebral tubercles 21 24 24 22 26 23 Body tubercles arrangement random random random random random random Tubercles on lower flanks 0 0 0 0 0 0 Caudal tubercles in lateral furrow 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly 1 1 1 1 1 1 Lateral caudal tubercle row 1 1 1 1 1 1 Only paravertebral tubercles on tail no no no no no no Subcaudals keeled keeled keeled keeled keeled keeled Single median row of keeled subcaudals no no no no no no Caudal tubercles encircle tail yes yes yes yes yes yes Enlarged median subcaudal row yes yes yes yes yes yes No. of postcloacal tubercles 1 1 1 2 2 1 Enlarged femoral scalas no no no no no no Shield-like subtibial scales no no no no no no Subtibial scales keeled keeled keeled keeled keeled keeled Enlarged submetatarsal scales on 1st toe no no no no no no No. 4th toe lamellae 29 30 27 26 29 28 SVL 58.1 54.8 57.1 53.4 51.3 56.1 TL 69 61.1 65.5 67.2 65.6 65.3 TW 6.2 5.8 6.1 6.5 5.3 6.4 FL 10.5 9.9 10.3 10.4 9.9 9.6 TBL 12.5 12.7 13.1 12.2 11.9 13.1 AG 25.8 25.7 25.3 24.6 24.7 25.3 HL 14.9 14.5 14.4 14.3 13.2 14.3 HW 9.7 9.4 10.2 9 8.6 9.2 HD 6.6 5.9 6.6 6 5.6 6.2 ED 3.1 3.2 3.1 3.1 2.9 2.8 EE 4.2 4.2 4.2 4.1 3.8 4.1 ES 7.7 7.1 7.1 7.1 6.4 7.1 EN 5.9 5.3 5.4 5.3 4.9 4.9 IO 3.4 3.2 3.2 2.8 2.8 3.3 EL 1.6 1.4 1.5 1.2 1.2 1.5 IN 1.6 1.4 1.5 1.5 1.3 1.4 A NEW INSULAR CNEMASPIS FROM MALAYSIA Zootaxa 3755 (5) 2014 Magnolia Press 453
Comparisons. Cnemaspis bidongensis sp. nov. is a member of a monophyletic lineage referred to as the kendallii group (Grismer et al. in prep.) which contains C. kendallii; C. baueri Das & Grismer and C. pemanggilensis Grismer & Das and is diagnosed by having a maximum SVL 50.5 81.0 mm; 7 13 supralabials; 7 12 infralabials; keeled ventral scales; no precloacal pores; 17 37 paravertebral tubercles; caudal tubercles not restricted to a single paravertebral row; lateral row of caudal tubercles present; 1 4 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe not enlarged; subtibials keeled; and 26 38 subdigital fourth toe lamellae. Cnemaspis bidongensis differs from all members of the kendallii group in having an enlarged, elongate mental scale extending posteriorly to the level of the fourth infralabials bordered posterolaterally by an enlarged postmental and a sexually dimorphic dorsal body pattern. Within this group C. bidongensis is most closely related to C. kendallii from which it differs by lacking as opposed to having tubercles on the ventral portions of the flanks; having as opposed to lacking an enlarged, median, subcaudal scale row; having as opposed to lacking a sexually dimorphic dorsal body pattern. From C. baueri, C. bidongensis sp. nov. differs by having a smaller maximum SVL (58.1 mm versus 67.4 mm); nine or 10 versus 11 13 supralabials; and a dorsal body pattern that contains elements of dark and light blotching (Fig. 1). Cnemaspis bidongensis sp. nov. can be differentiated from C. pemanggilensis in having 21 26 versus 30 37 paravertebral tubercles. These differences are represented in Table 1. Discussion Like mountain tops, the islands of Peninsular Malaysia have always been areas of relatively high endemism (Grismer 2011a; Grismer & Pan 2008; Grismer et al. 2010c, 2011) despite the fact they have received much less attention than continental regions. Nonetheless, field work on islands off the coasts of Peninsular Malaysia continues to uncover a surprising amount of herpetological diversity bearing an ever growing component of endemism. Thus, it is not surprising that a survey of the five islands of the previously herpetologically unexplored Bidong Archipelago resulted in the discovery of a new, endemic species of Cnemaspis. Endemic species of this genus are well-represented on Peninsular Malaysian islands with C. roticanai Grismer & Chan from the Langkawi Archipelago in the northwest; C. perhentianensis Grismer & Chan from the Perhentian Archipelago in the northeast and C. limi Das & Grismer, C. pemanggilensis Grismer & Das, and C. baueri Das & Grismer from the Seribuat Archipelago in the southeast (Grismer 2011a; Fig. 1). We expect that additional endemic species of reptiles will be found on the large island of Redang and from some of the small satellite islands in the Tenggol Archipelago that have yet to be thoroughly explored. We expect much the same pattern on islands of the Sembilan Archipelago along the west coast. Acknowledgements We wish to thank Mr. Baharim B. Mustapa, Head of Bidong Research Unit for facilitating the trips and Universiti Malaysia Terengganu for permission to conduct research on Bidong Island. This research was supported in part by grants to LLG from the College of Arts and Sciences, La Sierra University, Riverside, California. We would like to thank Jack W. Sites Jr. for funding the molecular work conducted in this study. References Chan, K.O., Grismer, L.L. & Grismer, J.L. (2011) A new insular endemic frog of the genus Kalophrynus Tschudi, 1838 (Anura: Microhylidae) from Tioman Island, Pahang, Peninsular Malaysia. Zootaxa, 3123, 60 68. Chan, K.O. & Norhayati, A. (2010) A new species of Cyrtodactylus (Squamata: Gekkonidae) from northeastern Peninsular Malaysia, Malaysia. Zootaxa, 2389, 47 56. Das, I. & Grismer, L.L. (2003) Two new species of Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Seribuat Archipelago, Pahang and Johor States, West Malaysia. Herpetologica, 59, 544 552. http://dx.doi.org/10.1655/02-22 Das, I. & Jim, L.L. (2000) A new species of Cyrtodactylus (Sauria: Gekkonidae) from Pulau Tioman, Malaysia. The Raffles Bulletin of Zoology, 48, 223 231. 454 Zootaxa 3755 (5) 2014 Magnolia Press GRISMER ET AL.
Diaz, R.E., Leong, T.M., Grismer, L.L. & Yaakob, N.S. (2004) A new species of Dibamus (Squamata: Dibamidae) from West Malaysia. Asiatic Herpetological Research, 10, 1 7. Grismer, J.L., Leong, T.M. & Norsham, S.Y. (2003) Two new Southeast Asian skinks of the genus Larutia and intrageneric phylogenetic relationships. Herpetologica, 59, 554 566. http://dx.doi.org/10.1655/02-56 Grismer, L.L. (2005) A new species of Bent-Toed Gecko (Cyrtodactylus Gray, 1827) from Pulau Aur, Johor, West Malaysia. Journal of Herpetology, 39, 424 432. http://dx.doi.org/10.1670/3-05a.1 Grismer, L.L. (2006) Two new species of skinks (Genus Sphenomorphus Fitzinger 1843) from the Seribuat Archipelago, West Malaysia. Herpetological Natural History, 9, 151 162. Grismer, L.L. (2011a) Field Guide to the Amphibians and Reptiles of the Seribuat Archipelago, Peninsular Malaysia. Edition Chimaira, Frankfurt am Main, 258 pp. Grismer, L.L. (2011b) Lizards of Peninsular Malaysia, Singapore and Their Adjacent Archipelagos. Edition Chimaira, Frankfurt am Main, 728 pp. Grismer, L.L. & Chan, K.O. (2008) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Zootaxa 1771, 1 15. Grismer, L.L. & Chan, K.O. (2009) A new species of karst dwelling Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Sarawak, Borneo. Zootaxa, 2246, 21 31. Grismer, L.L., Chan, K.O., Grismer, J.L., Wood, P.L. Jr. & Norhayati, A. (2010c) A checklist of the herpetofauna of the Banjaran Bintang, Peninsular Malaysia. Russian Journal of Herpetology, 17, 147 160. Grismer, L.L. & Das, I. (2006) A new species of gekkonid lizard of the genus Cnemaspis Strauch 1887 from Pulau Pemanggil, Johor, West Malaysia. Herpetological Natural History, 10, 1 7. Grismer, L.L., Das, I. & Leong, T.M. (2003) A new species of Gongylosoma (Squamata: Colubridae) from Pulau Tioman, West Malaysia. Herpetologica, 59, 567 574. http://dx.doi.org/10.1655/02-38 Grismer, L.L. Grismer, J.L. & Youmans, T.M. (2004b) A new species of Leptolalax (Anura: Megophryidae) from Pulau Tioman, West Malaysia. Asiatic Herpetological Research, 10, 8 11. Grismer, L.L., Grismer, J.L., Wood, P.L. Jr., Ngo, V.T., Neang, T. & Chan, K.O. (2011) Herpetology on the fringes of the Sunda Shelf: a discussion of discovery, taxonomy, and biogeography. Bonner Zoologische Monographien, 57, 57 97. Grismer, L.L. Kaiser, H. & Yaakob, N.S. (2004a) A new species of the Reed Snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad, 28, 1 6. Grismer, L.L. & Ngo, V.T. (2007) Four new species of the gekkonid genus Cnemaspis Strauch 1887 (Reptilia: Squamata) from southern Vietnam. Herpetologica, 63, 482 500. http://dx.doi.org/10.1655/0018-0831(2007)63[482:fnsotg]2.0.co;2 Grismer, L.L., Ngo, V.T. & Grismer, J.L. (2008) A new species of insular pitviper of the genus Cryptelytrops (Squamata: Viperidae) from southern Vietnam. Zootaxa, 1715, 57 68. Grismer, L.L., Ngo, V.T. & Grismer, J.L. (2010a) A colorful new species of insular rock gecko (Cnemaspis Strauch 1887) from southern Vietnam. Zootaxa 2352, 46 58. Grismer, L.L. & Pan, K.A. (2008) Diversity, endemism, and conservation of the amphibians and reptiles of southern Peninsular Malaysia and its offshore islands. Herpetological Review, 3, 270 281. Grismer, L.L., Sumontha, M., Cota, M., Grismer, J.L., Wood, P.L. Jr., Pauwels, O.S.G. & Kunya, K. (2010b) 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, 1 55. Grismer, L.L., Wood, P.L. Jr. & Grismer, J.L. (2009) A new insular skink of the genus Sphenomorphus Strauch 1887 (Squamata: Scincidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Tropical Life Sciences Research, 20, 51 69. Grismer, L.L., Wood, P.L. Jr., Maketab, M., Chan, K.O., Heinz, H.M., Sumaril, A.S.-I., Chan, J.A. & Loredo, A.I. (2013) A new species of karst-adapted Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from a threatened karst region in Pahang, Peninsular Malaysia. Zootaxa, 3746 (3), 463 472. http://dx.doi.org/10.11646/zootaxa.3746.3.5 Grismer, L.L., Youmans, T.M., Wood, P.L. Jr. & Grismer, J.L. (2006) Checklist of the herpetofauna of the Seribuat Archipelago, West Malaysia with comments on biogeography, natural history and adaptive types. Raffles Bulletin of Zoology, 54, 157 180. Leong, T.-M., Grismer, L.L. & Mumpuni (2003) Preliminary checklists of the Anambas and Natuna Islands (South China Sea). Hamadryad, 27, 65 174. Macey, J. & Schulte, J. (1999) Molecular phylogenetics, trna evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution, 12, 250 272. http://dx.doi.org/10.1006/mpev.1999.0615 Ngo, V.T. (2008) Two new cave-dwelling species of Cyrtodactylus Gray (Squamata: Gekkonidae) from southwestern Vietnam. Zootaxa, 1909, 37 51. Ngo, V.T. & Grismer, L.L. (2012) A new endemic species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu A NEW INSULAR CNEMASPIS FROM MALAYSIA Zootaxa 3755 (5) 2014 Magnolia Press 455
Island, southwestern Vietnam. Zootaxa, 3228, 48 60. Ngo, V.T., Grismer, L.L. & Grismer, J.L. (2008) A new endemic cave dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Kien Giang Biosphere Reserve, southwestern Vietnam. Zootaxa, 1967, 53 62. Ngo, V.T., Grismer, L.L. & Grismer, J.L. (2010) A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Phu Quoc National Park, Kien Giang Biosphere Reserve, southwestern Vietnam. Zootaxa, 2604, 37 51. Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) Mega5: Molecular Evolutionary Genetics Analysis Using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and Evolution, 28, 2731 2739. http://dx.doi.org/10.1093/molbev/msr121 Wood, P.L. Jr., Quah, E.S.H., Anuar, S. & Muin, M.A. (2013) A new species of lowland karst dwelling Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from northwestern Peninsular Malaysia. Zootaxa, 3691, 538 558. http://dx.doi.org/10.11646/zootaxa.3691.5.2 Youmans, T.M. & Grismer, L.L. (2006) A new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from the Seribuat Archipelago, West Malaysia. Herpetological Natural History, 10, 61 70. 456 Zootaxa 3755 (5) 2014 Magnolia Press GRISMER ET AL.