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Cnemaspis harimau sp. nov. Tiger Rock Gecko Figure 2 A, B Holotype. Adult male (ZRC 2.6894) collected on 17 March 2010 by Chan Kin Onn, Lee Grismer, Jesse Grismer, Anna Savage, Shahrul Anuar, Mohd. Abdul Muin, and Evan Quah at 2030 hrs at 601 m a.s.l. from Sungai Badak (=Badak river), Gunung Jerai, Kedah, Peninsular Malaysia (N 05° 48 ΄ 59 ʺ, E 100 ° 23 ΄ 53 ʺ). Paratypes. Collection locality and collector of the paratypes is the same as that of the holotype. The paratypes were collected between 2030 and 2130 hrs. ZRC 2.6897 (female; 16 March 2010); LSUHC 9669 (male; 17 March 2010); and ZRC 2.6895, 2.6896, LSUHC 9665, 9667 (female; 17 March 2010). Diagnosis. Cnemaspis harimau differs from all other Southeast Asian congeners except C. affinis, C. biocellata, C. kumpoli, C. mcguirei, C. pseudomcguirei, and C. shahruli in having a black shoulder patch with a white or yellow ocellus anteriorly. It can be differentiated from the above-mentioned species in the unique combination of adult males reaching 40.7 mm SVL, females 39.4 mm SVL; nine or 10 supralabials, 9–10 infralabials; three postmentals; ventral scales keeled; four precloacal pores in males, arranged in a chevron, medially separated by one non-pore bearing scale; 18–20 paravertebral tubercles; 25–30 subdigital lamellae under the fourth toe; keeled subcaudals; enlarged, spinose lateral caudal tubercles on anterior 1 / 3 of tail; caudal tubercles encircling the tail, and males having an orange-colored gular region in life. Description of holotype. Adult male; SVL 40.7 mm; head oblong in dorsal profile, moderate in size (HL/ SVL 0.26), somewhat narrow (HW/SVL 0.17), flat (HD/HL 0.39), distinct from neck; snout short (ES/HL 0.50), concave in lateral profile; postnasal region constricted medially; scales of rostrum low, rounded, juxtaposed, weakly keeled, larger than similarly shaped scales on occiput; weak, supraorbital ridges; frontonasal sulcus present, moderate; canthus rostralis smoothly rounded; eye large (ED/HL 0.20); extrabrillar fringe scales small in general but slightly larger anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 80 % divided by longitudinal groove; rostral bordered posteriorly by two supranasals, a smaller, azygous postrostral, and nostrils; bordered laterally by first supralabials; 10 (R,L) raised supralabials of similar size, but smallest posteriorly; 11 (R,L) infralabials, decreasing gradually in size posteriorly; nostrils small, oblong, oriented dorsolaterally; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, flat, extending to level of second infralabials, bordered postero-laterally by two enlarged, elongate postmentals of similar size which are separated by a single, small, azygous scale; gular scales smooth, rounded, juxtaposed; throat scales keeled. Body slender, elongate (AG/SVL 0.43); small, raised, keeled, dorsal scales generally equal in size throughout body, intermixed with numerous, large, multi-keeled, semi-longitudinally arranged spinose, tubercles; tubercles extend from occiput to base of tail and are smallest on the occiput; 20 paravertebral tubercles; pectoral and abdominal scales smooth, flat, imbricate; abdominal and pectoral scales keeled, imbricate, larger than dorsals; four pore-bearing precloacal scales arranged in a chevron, two left and two right ones medially separated by a single non pore-bearing scale; forelimbs moderately long, slender, dorsal scales raised, multicarinate; ventral scales of forearm smooth, slightly raised, juxtaposed to subimbricate; palmar scales smooth, raised, subimbricate; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; subdigital lamellae wide beneath the proximal and distal phalanges, bearing a larger scale at digital inflections; scales under intermediate phalanges smaller; interdigital webbing absent or extremely reduced; fingers increase in length from first to fourth with fifth same length as fourth; hind limbs longer and thicker than forelimbs; dorsal scales of thigh, keeled, raised, juxtaposed to subimbricate; dorsal scales of tibia multicarinate, raised, juxtaposed; ventral scales of hind limbs keeled, imbricate; plantar scales smooth, flat, subimbricate; no enlarged subtibials or submetatarsals; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide beneath the proximal and distal phalanges except at base and intermediate phalanges 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; 26 subdigital lamellae on fourth toe; dorsal, caudal scales arranged in segmented whorls, slightly larger than dorsal body scales; caudal scales low, keeled, juxtaposed anteriorly, raised and more keeled posteriorly; shallow, middorsal and lateral caudal furrow; subcaudal scales keeled, subimbricate anteriorly, degree of keeling and scale-overlap increase posteriorly; no median row of enlarged subcaudal scales; three longitudinal rows of tubercles on base of tail on either side of midline; single row of paravertebral tubercles throughout length of tail; single row of dorsolateral and lateral tubercles on anterior 25 % of tail; no tubercles within lateral, caudal furrow; three postcloacal tubercles; tail approximately 125 % of SVL. Color in life. Base color of dorsum grayish-brown; top of head yellowish with indistinct gray speckling on the occiput; a faint pre-orbital stripe lines the canthus rostralis; three black stripes radiate from the anterior region of the neck to the base of the occiput; light gray, irregular blotches along the vertebral column beginning from the nape to the base of the tail; broken, dark, paravertebral longitudinal streaks border each side of the midline, beginning with a paired streak on the pre-scapular region and alternating with the vertebral blotches towards the posterior, terminating at the base of the tail; black shoulder patch, bordered anteriorly by a short yellow band enclosing a whitish ocellus (corresponds to an enlarged, spinose, pigmented tubercle), and bordered posteriorly by a longer yellow band; a series of four yellow bands continue along the flanks, decreasing in length posteriorly with the last band terminating just anterior to the hindlimb insertion; dorsal surfaces of limbs irregularly speckled with dark, light and yellowish blotches; tail distinctly marked with dark gray and whitish bands which are not sharply demarcated; gular region and throat orange; ventral surfaces of limbs, remainder of body and tail base light gray; postcloacal tubercles yellow; ventral surfaces of tail with dark stippling, corresponding to the caudal bands. Color in preservative. Dorsal base color including head grayish-brown; faint, post-orbital stripes are visible in dorsal profile; vertebral blotches indistinct; yellow bands faded to light gray; dorsal surfaces of limbs with irregular light and dark blotches; entire ventral surface whitish gray, orange coloration on gular region and throat undetectable; ventral surfaces of tail with dark stippling, corresponding to the caudal bands. Variation and sexual dimorphism. Adult male, LSUHC 9669 has a darker base color and a distinct pre- and post-orbital stripe which connects to the occipital streaks with which it forms a continuous stripe; medial occipital spot present; paired paravertebral streaks are dark and more distinct than the holotype; precloacal pores were not assessable due to torn skin. Sexual dimorphism is distinct with females lacking precloacal pores, shoulder patches and orange coloration on the gular region and throat; other body markings are similar to males; the paratype females, ZRC 2.6896, 2.6897 and LSUHC 9667 have a darker base color; ZRC 2.6897 has a wide, distinct, continuous vertebral stripe extending from the base of the occiput to the tail base. It should be noted that base color of specimens, be it in life or in preservative, should not be taken into account when diagnosing this species because of their ability to substrate match. Other morphological variations are presented in Table 1. Values that were identical throughout the type series are not indicated. Distribution. Gunung Jerai, in the state of Kedah, Peninsular Malaysia. Natural history. Sungai Badak was mostly dry during our visit and was a rocky stream bed, lined with dense vegetation and medium sized (0.5–2 m diameter) granite boulders. Lizards were observed on the boulders and at the base of tree trunks near the boulders during the day and night, although many more were seen at night. They were wary and when provoked, fled downwards to seek refuge in porous matrices formed by the dried expansive soil at the base of boulders. Etymology. The specific epithet, harimau means tiger in the Malay language and is in reference to the yellow banding on the flanks of this species which are arbitrarily analogous to the banding on a tiger. Additionally, the year 2010 coincides with the Chinese zodiac year of the tiger. Comparisons. Cnemaspis harimau can be differentiated from all other Sundaland congeners except C. affinis, C. biocellata, C. kumpoli, C. mcguirei, C. pseudomcguirei, and C. shahruli in having a black shoulder patch with a white or yellow ocellus anteriorly. It differs from C. biocellata by having a single, shoulder ocellus instead of two ocelli; four precloacal pores vs. 8–12; precloacal pores separated vs. continuous; keeled vs. smooth subcaudals; and 25–30 vs. 29–37 subdigital lamellae under the fourth toe; from C. kumpoli by having a maximum SVL of 40.7 mm vs. 63.0 mm; keeled vs. smooth ventral scales; four vs. 7–8 precloacal pores; 18–20 vs. 29–35 paravertebral tubercles; keeled vs. smooth subcaudals; 25–30 vs. 34–41 subdigital lamellae under the fourth toe; from C. mcguirei in having a maximum SVL of 40.7 mm vs. 65.0 mm; four vs. 5–10 precloacal pores; 18–20 vs. 26–32 paravertebral tubercles; the absence vs. presence of tubercles within the lateral caudal furrow; and 25–30 vs. 30–35 subdigital lamellae under the fourth toe; from C. pseudomcguirei in having 18–20 vs. 26–32 paravertebral tubercles; mental bordered posteriorly by two enlarged lateral postmentals which are separated by a single, small azygous scale vs. six small postmentals; precloacal pores separated vs. continous; caudal tubercles encircle vs. do not encircle tail; the absence vs. presence of tubercles within the lateral, caudal furrow; and having vs. lacking yellow bands on the flanks; from C. shahruli in having a maximum SVL of 40.7 mm vs. 36.5 mm; four vs. zero precloacal pores; and caudal tubercles encircle vs. do not encircle tail. Cnemaspis harimau is most similar in appearance to its sister species C. affinis, but differs by having a maximum SVL of 40.7 mm vs. 50.8 mm; 18–20 vs. 20–28 paravertebral tubercles; lateral postmentals separated by one vs. more than one (usually three) smaller, azygous scales; caudal tubercles encircle vs. do not encircle tail; lateral caudal tubercles on anterior 25 % of tail highly spinose and protruding vs. slightly spinose; and an overall higher degree of scale keeling (most prominent on the tail). Differences between species within the shoulder patch group are summarized across Table 2. . sp nov harimau affinis biocellata kumpoli mcguirei pseudomcguirei shahruli Max. SVL (mm) 40.7 50.8 40.1 63 65 42.5 36.5 Supralabials 9,10 9– 13 6–10 7,8 7–9 9 10,11 Infralabials 9,10 8– 10 5–7 6,8 7,8 8, 9 8–10 No. of postmentals 3 5 3 3 3, rarely 5 6 3–6 Ventral scales keeled (1) or not (0) 1 1 1 0 1 1 1 No. of precloacal pores 4 5–6 8–12 7, 8 5–10 1–5 0 Preliminary molecular analysis show that Cnemaspis harimau (n= 3) and C. affinis (n= 5) are sister species with a pairwise sequence divergences of 2.9–3.6 % based on 749 base pairs on ND 2 (Grismer et al. in prep.). This suggests that a common ancestor once ranged throughout lowland areas from at least Penang Island to Gunung Jerai at a time when Penang Island was connected to Peninsular Malaysia (Inger & Voris, 2001). A subsequent rise in sea levels separated Penang Island from the mainland and for a brief time, rendered Gunung Jerai as an island as well. This is likely to have resulted in the allopatric speciation event that gave rise to each species. Despite the relatively recent separation, both species have diverged enough to attain reciprocal monophyly and unique morphological differences. Although C. harimau is known only from 601 m elevation, it may range much lower, following a microhabitat of rocky stream corridors down the mountain. Cnemaspis harimau is likely to be endemic to Gunung Jerai being that Jerai is bordered by the Indian Ocean in west and paddy fields to the east. Remarkably, Cnemaspis harimau is the first endemic reptile reported from this isolated mountain range. The ranid frog, Odorrana monjerai (Matsui & Jaafar 2006), which was previously thought to be endemic to Gunung Jerai, has also been found in Bukit Hijau, Kedah (Chan et al. 2010). This discovery adds to the growing number of upland endemics that have been found in the vastly unexplored mountain ranges of Peninsular Malaysia (Chan et al. 2009; Grismer 2007; Grismer et al. 2008 b, 2009; Wood et al. 2009). |
