Cyrtodactylus payacola, Johnson, Chelséa B., Quah, Evan, Anuar, Shahrul, Wood, Perry L., Grismer, Jesse L., Greer, Lee F., Onn, Chan Kin & Ahmad, Norhayati, 2012

Johnson, Chelséa B., Quah, Evan, Anuar, Shahrul, Wood, Perry L., Grismer, Jesse L., Greer, Lee F., Onn, Chan Kin & Ahmad, Norhayati, 2012, Phylogeography, geographic variation, and taxonomy of the Bent-toed Gecko Cyrtodactylus quadrivirgatus Taylor, 1962 from Peninsular Malaysia with the description of a new swamp dwelling species, Zootaxa 3406, pp. 39-58 : 53-57

Cyrtodactylus payacola sp. nov.

Figs. 6 , 7

Cyrtodactylus cf. quadrivirgatus Grismer 2011:429 .

Holotype. Adult male ( LSUHC 10074) collected by E. Quah S. H. on 4 September 2011 from Bukit Panchor State Park, Penang, West Malaysia 05º09.465' N, 100º32.885' E at an elevation of 47 m a.s.l.

Paratypes. All paratopotypes were collected in various combinations by M. A. Muin, E. Quah S.H., S. Anuar, C. K. Onn, and L. L. Grismer from the same locality as the holotype. LSUHC 9982 was collected on 6 March 2010; LSUHC 10070 and 10071 were collected on 29 June 2011.

Additional specimens examined. LSUHC 10076 was collected from Shah Alam, Selangor (3°05.262’N, 101°31.466”E; 15 m a.s.l.) by Daicus Belabut. ZRC 2.1127 was collected from the Sungai Chikus Forest Reserve, Perak (4° 09.426N, 101°00.562’E; 7 m a.s.l.) during April 1925.

Diagnosis. Cyrtodactylus payacola sp. nov. is distinguished from all other Sundaland species by having a maximum SVL of 67.7 mm; moderately sized, conical, keeled body tubercles; tubercles occurring on the occiput, forelimbs, hind limbs, and beyond base of tail; 44–51 ventral scales; no transversely enlarged, median subcaudal scales; 18–21 subdigital lamellae on fourth toe; abrupt transition between postfemoral and ventral femoral scales; no femoral pores; 11 or 12 contiguous, pore-bearing precloacal scales; shallow, longitudinal, precloacal groove; a pair of posteromedially elongate, triangular to semilunar-shaped, paravertebral blotches on nape prominently outlined in light yellow; no wide, dark, ventrolateral stripes on flanks; no white reticulum on top of head; paired, semitransversely arranged, dark blotches on body.

Description of holotype. Adult male SVL 67.7 mm; head moderate in length (HL/SVL 0.26), wide (HW/HL 0.59), flat (HD/HL 0.38), distinct from neck, triangular in dorsal profile, lores weakly inflated, prefrontal region deeply concave, canthus rostralis smoothly rounded; snout elongate (ES/HL 0.40), rounded in dorsal profile; eye large (ED/HL 0.23); ear opening elliptical, obliquely oriented, moderate in size (EL/HL 0.06), eye to eye distance greater than diameter of eye; rostral square, punctuated with pores, partially divided dorsally by three postrostral granular scales (=internasals), bordered posteriorly by large left and right supranasals, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal and small posterior supranasal, posteriorly by two large postnasals, and ventrally by first supralabial; 10 (R, L) rectangular supralabials extending to just beyond dorsal inflection of labial margins, tapering abruptly below midpoint of eye; nine (R, L) infralabials tapering smoothly posteriorly to below orbit; scales of rostrum and lores raised, slightly larger size than granular scales on top of head and occiput; scales of occiput intermixed with slightly enlarged tubercles; supraorbitals smooth; dorsal superciliaries elongate, smooth; mental triangular, bordered laterally by first infralabials, posteriorly by left and right rectangular postmentals contacting medially for 40% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to fifth infralabial; gular scales small, granular, grading posteriorly into slightly larger, flatter, throat scales which grade into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.44) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with moderately sized, conical, semi-regularly arranged, keeled tubercles being most dense on flanks; tubercles extend from occiput to anterior one-sixth of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 21 longitudinal rows of tubercles at midbody, 37 paravertebral tubercles on body; 51 flat, imbricate, ventral scales, ventral scales larger than dorsal scales; precloacal scales not large; 11 contiguous, pore-bearing precloacal scales forming a “V” bordering a longitudinal, shallow precloacal groove.

Forelimbs moderate in stature, relatively short (FL/SVL 0.13); granular scales of forearm slightly larger than those on body, interspersed with large, keeled tubercles; palmar scales rounded, flat; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded proximal to joint inflection, more granular distal to inflection; digits slightly more narrow distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.14), covered dorsally by granular scales interspersed with large, conical tubercles, covered anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals; ventral tibial scales flat; femoral scales imbricate; small postfemoral scales form an abrupt union with large ventral scales on posteroventral margin of thigh; plantar scales low, flat; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded proximal to inflections, more granular distal to inflections, digits more narrow distal to inflections; 19R, 18L subdigital lamellae on fourth toe; claws well-developed, sheathed by a dorsal and ventral scale.

Tail moderate, 90.7 mm in length, tapering to a point, 5.3 mm in width at base; dorsal scales of base of tail granular becoming flatter posteriorly; no median row of transversely enlarged, subcaudal scales; subcaudal scales larger than dorsal caudal scales; one pair of paravertebral and dorsolateral tubercle rows on either side of midline; paravertebral rows not widely separated; caudal tubercles decrease in size posteriorly, extending approximately one-sixth length of tail; one enlarged, postcloacal tubercle on left and right base of tail on hemipenial swelling; all postcloacal scales flat.

Coloration in life. Dorsal: ground color of head, neck, body, limbs, and tail light straw-yellow; light brown mottling on top of head and rostrum; diffused, light brown, postorbital patch; paired, symmetrical, triangular to semilunar-shaped, dark blotches on entire portion of nape prominently outline in ground color; seven dark brown rectangular-shaped bands extending from below nape to base of tail; bands counter shaded with lighter color; wide, dark, elongate markings above shoulders; irregularly shaped, dark blotches on flanks; body darkly speckled overall; brown blotching on limbs, obscure banding on hind limbs; dark body bands extend onto tail to form brown bands alternating with cream-colored bands neither of which encircle tail; anterior caudal bands transform into black and white, respectively, on posterior two-third of tail. Ventral: surfaces of head, body and limbs lightly stippled in gray; subcaudal region darkened by fine mottling; iris reddish. Lizards lighten considerably at night ( Fig. 7 ).

Variation. The paratypotypes ( Fig. 6 ) and the additional specimens examined approximate the holotype in all aspects of coloration. The caudal bands in LSUHC 9982 encircle only the posterior one-half of the tail. The additional specimens examined (LSUHC 10076 from Shah Alam, Selangor and ZRC 2.1127 from the Chikus Forest Reserve, Perak) share the species-level diagnostic characters of the holotype and paratopotypes and approximate the holotype in general aspects of coloration although the dark, dorsal markings form semi-transversely arranged bands as opposed to forming more paired markings. Furthermore, LSUHC 10076 has paired, non-symmetrical, semilunar-shaped, dark blotches only on the upper portion of nape as opposed to extending along the entire nape of the neck and the caudal bands encircle the tail along its entire length. Meristic differences are shown in Table 6.

Distribution. Cyrtodactylus payacola sp. nov. is known from the Bukit Panchor State Park, Penang; the Sungai Chikus Forest Reserve, Perak; and Shah Alam, Selangor, Peninsular Malaysia ( Fig. 1 ) and most likely ranges throughout all lowland coastal areas west of the Banjaran Bintang and Titiwangsa to as far south perhaps as Melaka, Melaka where its sister species C. pantienesis has also been reported to occur (Grismer et al. 2008). To the north, it may not range beyond the Thai –Malaysian border owing to the potential dispersal barrier of the Banjaran Nakawan.

Natural history. The three localities from which Cyrtodactylus payacola sp. nov. is known are all wet, swampy, lowland forests no higher than 15 m in elevation ( Fig. 8 ). The lizards from Bukit Panchor and Shah Alam were collected at night from the surface of leaves up to 2 m above the forest floor. During this period this species’ coloration lightens considerably ( Fig. 7 ) making them easily confused with C. quadrivirgatus with which it is narrowly sympatric at Bukit Panchor and Shah Alam. At Bukit Panchor, C. quadrivirgatus appears to occur only up to the edge of the swampy areas, remaining in the more elevated drier regions (LSUHC 10072–73). LSUHC 10070, a gravid female carrying two eggs, was collected on 29 June indicating that breeding takes place at least during this month.

Etymology. The specific epithet payacola is derived from the word paya meaning “swamp” in the Malay language and the Latin suffix colo, which means to inhabit or dwell in and is in reference to microhabitat in which this species is found.

Comparisons. Cyrtodactylus payacola sp. nov. is readily differentiated from all other Sundaland species of Cyrtodactylus on the basis of color pattern and morphology (see Chan & Norhayati 2010: Table 1). It is separated from C. quadrivirgatus , a species to which it was provisionally ascribed ( Grismer 2011), based on having 11 or 12 contiguous, pore-bearing precloacal scales along a shallow, longitudinal precloacal groove as opposed to having 0–5 pore-bearing scales and no groove and a pair of posteromedially elongate, triangular to semilunar-shaped, paravertebral blotches on the nape that are prominently outlined in light yellow as opposed to lacking such markings. It differs from its sister species, C. pantiensis , in having more precloacal pores (11 or 12 versus eight or nine); having fewer subdigital lamellae on the fourth toes (18–21 versus 22 or 23); and having a generally immaculate ground color as opposed to the dense, dark brown speckling seen in C. pantiensis .

The Shah Alam specimen (LSUHC 10076) is more similar to C. payacola sp. nov in all aspects of coloration and morphology than it is to any other species of Cyrtodactylus except for having paired dark, dorsal markings and paired, non-symmetrical, semilunar-shaped, dark blotches on only the upper portion of nape. Although it is a female, there are 12 large, non-pore-bearing scales homologous to those pore-bearing scales in males. Therefore, we tentatively consider this specimen C. payacola sp. nov. and await the acquisition of additional material.

Figures

FIGURE 1. Closed red circles are known localities of Cyrtodactylus payacola sp. nov. Closed white circles are localities from which specimens were sampled for the molecular and morphological analysis of Cyrtodactylus quadrivirgatus. Closed black circle (Shah Alam, Selangor) was not sampled for the molecular analysis and closed black squares are the remaining known localities for C. quadrivirgatus (fide Grismer 2011) from which no specimens were examined. Numbers correspond to the following localities: 1. Pulau Langkawi, Kedah. 2. Pulau Tuba, Kedah. 3. Ulu Muda, Kedah. 4. Gunung Jerai, Kedah. 5. Sungai Sedim, Kedah. 6. Pulau Pinang, Penang. 7. Bukit Larut, Perak. 8. Pulau Pangkor, Perak. 9. Temengor Forest Reserve, Perak. 10. Sungai Pelus, Perak. 11. Pulau Perhentian Besar, Terengganu. 12. Gunung Lawit, Terengganu. 13. Sungai Tembak, Terengganu. 14. Kuala Aring, Terengganu. 15. Kuala Tahan, Pahang. 16. Sungai Lembing, Pahang. 17. Gunung Benom, Pahang. 18. Fraser’s Hill, Pahang. 19. Genting Highlands, Pahang. 20. Ulu Kali, Pahang. 21. Ulu Gombak, Selangor. 22. Lakum Forest Reserve, Pahang. 23. Bukit Rengit, Pahang. 24. Bukit Bau, Pahang. 25. Pasoh, Negri Sembilan. 26. Gunung Ledang, Johor. 27. Pulau Tulai, Johor. 28. Pulau Tioman, Pahang. 29. Endau – Rompin, Johor. 30. Gunung Panti, Johor. 31. Singapore. 32. Shah Alam, Selangor. 33. Ulu Temiang Forest Reserve, Kelantan. 34. Serasa Forest Reserve, Kelantan. 35. Empangan Tembat, Terengganu. 36. Templer’s Park, Selangor. 37. Bukit Panchor, Penang. 38. Sungai Chikus Forest Reserve, Perak.

FIGURE 6. Type series of Cyrtodactylus payacola sp. nov. Holotype LSUHC 10074.

FIGURE 7. Cyrtodactylus payacola sp. nov. showing dark (upper; LSUHC 10074) and light (lower; LSUHC 10074) color phases.

FIGURE 8. Swampy microhabitat of Cyrtodactylus payacola sp. nov. at Bukit Panchor, Penang.