Cnemaspis paripari Grismer & Chan, 2009

Grismer, Lee, Wood, Perry L., Anuar, Shahrul, Riyanto, Awal, Ahmad, Norhayati, Muin, Mohd A., Sumontha, Montri, Grismer, Jesse L., Onn, Chan Kin, Quah, Evan S. H. & Pauwels, Olivier S. A., 2014, Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia, Zootaxa 3880 (1), pp. 1-147 : 100-103

Cnemaspis paripari Grismer & Chan, 2009

Fairy Rock Gecko

Figs. 52 , 53

Holotype. ZRC 2.6812 . Type locality: “ Gua Pari-pari , Bau District, Sarawak, [East] Malaysia (01°22.867 N, 110°07.164 E)” at approximately 30 m inelevation.

Diagnosis. Maximum SVL 50.7 mm; 12 or 13 supralabials; 10 or 11 infralabials; ventral scales keeled; 2–6, discontinuous, pore-bearing, precloacal scales with round pores; 26–31 paravertebral tubercles; body tubercles randomly arranged, absent on flanks and from lateral caudal furrows; ventrolateral caudal tubercles absent; lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals keeled but bearing an enlarged median row of smooth scales; two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe enlarged; subtibials keeled; 26–31 subdigital fourth toe lamellae; head, limbs, and regenerated tail yellow in males; posterior one-half of original tail white in males (Tables 6,7).

Color pattern in life ( Figs. 52 , 53 ). Males: dorsal ground color of body yellowish brown; limbs yellow; head (especially snout) bright yellow bearing small, irregularly shaped, brownish, occipital flecks and a faint, brownish, postorbital stripe; ground color of nape and shoulder region gray bearing paravertebral, irregularly shaped, black blotches; incomplete, transverse, yellow bands between forelimb and hind limb insertions; smaller, scattered, dark spots between bands; limbs generally immaculate; anterior one-half of tail gray bearing faint, dark bands; posterior one-half of tail immaculate, white; all ventral surfaces gray except for posterior half of tail which is white and beige; regenerated tail bright yellow. Females: adult females have an overall brown ground color, lack a yellow head and a yellow or white tail; yellow banding on body faint; dark blotching pattern on nape; tail brown at base, gradually turning to gray posteriorly; weakly banded.

Distribution. Cnemaspis paripari is known only from the karst formations that extend approximately 4.2 km from Gua Angin to Gua Pari-pari within the Bau Limestone Area, Sarawak, Malaysia ( Fig. 4 ).

Natural History. According to Grismer and Chan (2009), Cnemaspis paripari is a diurnal, lowland, saxicolous species that appears to be restricted to the karst outcroppings extending from Gua Angin to Gua Paripari that are surrounded by lowland dipterocarp forest ( Fig. 53 ). They reported seeing several specimens around and slightly within the openings of caves where light could still penetrate as well as on rocks along the periphery of the outcroppings. No specimens were observed deep within the caves. Most lizards were observed on vertical surfaces in shaded areas and would retreat into nearby cracks at the slightest provocation. Males would often curl their tail up over their back and wave the bright yellow (regenerated) or white (original) posterior section from side to side.

Relationships. Cnemaspis paripari and its sister species of C. nigridia from the nigridia species group ( Fig. 2 ).

Material examined. East Malaysia: Sarawak, Gua Pari-pari ZRC 2.6812 ; Gua Angin LSUHC 9185 , ZRC 2.6813 – 14 (type series) .

Kendallii group. The kendallii group is a well-supported lineage containing a morphologically diverse group of endemic, insular species from the Seribuat ( Cnemaspis baueri and C. pemanggilensis ) and Natuna ( C. mumpuniae sp. nov. and C. sundainsula sp. nov.) archipelagos along with C. bidongensis from Pulau Bidong from Peninsular Malaysia; C. peninsularis sp. nov. from Peninsular Malaysia and Singapore, and C. kendallii from East Malaysia and Indonesia ( Figs. 2 , 4 ). As noted above, the monophyly of this group is further supported in that these seven species lack precloacal pores ( Fig. 5 ). The relationships within the kendallii group clearly indicates that C. kendallii is polyphyletic, being that it has at least six separate, independent origins ( Fig. 2 and see below): C. kendallii from Borneo; C. kendallii from Peninsular Malaysia (= C. peninsularis sp. nov.), and C. kendallii from the Natuna Archipelago, Indonesia ( Gonatodes kendallii fide De Rooij [1915] = C. mumpuniae sp. nov. and Gonatodes kendallii fide Günther [1895] = C. sundainsula sp. nov.). The precise phylogenetic placement of C. kendallii ( Gonatodes kendallii fide Smedley [1928] = C. sundagekko sp. nov.) from Pulau Siantan of the Anambas Archipelago, Indonesia is not yet known but will likely add an additional independent origin (see below). The polyphyletic nature of C. kendallii was first noted by Grismer et al. (2008b) and Grismer (2011a:334). Leong et al. (2003) indicated that populations on a number of Indonesian Islands were also likely to be different species. In their revision of C. kendallii, Das & Bauer (1998) considered the population from the upland region of Bukit Larut in Peninsular Malaysia, all Peninsular Malaysian and Seribuat Archipelago populations, and the Natuna Besar and Anambas Island populations to compose C. kendallii which in fact is a composite of six species ( Grismer et al. 2008b and herein), some of which occur in different clades. The phylogeny also indicates that the Peninsular Malaysian and Seribuat Archipelago populations (i.e. C. peninsularis sp. nov.) and the Natuna Besar Island populations are not closely related to Bornean C. kendallii , which which is the provenance of the holotype ( Gray 1845). Additionally, examination of the population from Pulau Siantan from the Anambas Archipelago reveals that it too is not conspecific with Bornean C. kendallii but likely related to a clade containing C. baueri , C. pemanggilensis , C. mumpuniae sp. nov., C. bidongensis , and C. peninsularis sp. nov. being that it has the derived character state of caudal tubercles encircling the tail. Therefore, C. kendallii sensu lato is reclassified and the new species are described below.

In the molecular analysis, Cnemaspis sundainsula sp. nov. is the basal species in a well-supported monophyletic group comprised of it and C. pemanggilensis , C. kendallii sensu stricto, C. baueri , C. mumpuniae sp. nov., C. bidongensis , and C. peninsularis sp. nov. The monophyly of these latter species is supported in that they are the only species of Cnemaspis that have caudal tubercles encircling the tail and the monophyly of C. baueri , C. mumpuniae sp. nov., C. bidongensis , and C. peninsularis sp. nov. is supported in that they are the only Cnemaspis in which the posterior portion of the original tail is black.

The kendallii group is diagnosed by having a maximum SVL 58.1–84.5 mm; 9–13 supralabials; 7–12 infralabials; keeled ventral scales; 0–6, contiguous, round, pore-bearing, no precloacal pores; body tuberculation moderate; 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; and 25–38 subdigital fourth toe lamellae.

Figures

FIGURE 2. Maximum-likelihood phylogram (–InL 73957.608688) of the species of the genus Cnemaspis with Bayesian posterior probabilities (BPP) and maximum-likelihood (ML) bootstrap values, respectively based on the concatenated 3 gene dataset. Black circles are nodes supported by BBP and ML values greater than 0.95 and 70, respectively. Gray circles are nodes supported only by ML values greater than 70. White circles are nodes supported only by BPP values greater than 0.95. The distribution maps on the right delimit the ranges of the four major clades.

FIGURE 4. Distribution of the species in the Pattani and Southern Sunda clades. Stars indicate type localities.

FIGURE 5. Ecological parameters of activity period, elevation (upland species are> 600 m in elevation), substrate preference (microhabitat), and the presence or absence of ocelli (eyespots) mapped onto the molecular phylogeny (Fig. 2) of the species of Cnemaspis. Nodes lacking significant statistical support in Figure 2 are collapsed here. Morphological and color pattern synapomorphies are indicated at the appropriate nodes and considered as additional support of the molecular phylogeny. These character states were polarized following the methodology of Maddison et al. (1984) and were not simply mapped onto the tree. Branch lengths do not represent genetic distance.

FIGURE 52. Adult male Cnemaspis paripari (LSUDPC 4920) from Gua Pari-pari, Sarawak, East Malaysia in the dark color pattern phase. Photograph by LLG.

FIGURE 53. Cnemaspis paripari from Gua Angin, Sarawak, East Malaysia. Upper left: adult male (LSUDPC 4924) in the dark color pattern phase. Upper right: gravid female (LSUDPC 4923) in the dark color pattern phase. Lower: karst microhabitat at Gua Angin. Photographs by LLG.