Cyrtodactylus trilatofasciatus, Grismer, Lee, Wood, Perry L., Quah, Evan S. H., Anuar, Shahrul, Muin, Abdul, Sumontha, Montri, Ahmad, Norhayati & Bauer, Aaron M., 2012

Grismer, Lee, Wood, Perry L., Quah, Evan S. H., Anuar, Shahrul, Muin, Abdul, Sumontha, Montri, Ahmad, Norhayati & Bauer, Aaron M., 2012, A phylogeny and taxonomy of the Thai-Malay Peninsula Bent-toed Geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species, Zootaxa 3520, pp. 1-55 : 37-40

Cyrtodactylus trilatofasciatus sp. nov.

Cameron Highlands Bent-toed Gecko Fig. 15 , 16

Cyrtodactylus pulchellus Lim et al. 2002:51 ; Grismer 2011:420 (in part).

Holotype.—Adult male ( ZRC 2.6976) collected on 29 March 2011 by L. Grismer and Chan, K. O. from Ringlet, Cameron Highlands, Pahang, Peninsular Malaysia (04° 24.516 N, 100° 22.591 E) at 1109 meters above sea level.

Paratypes.— Paratypes ZRC 2.6977–78, and LSUHC 0 461 (adult female, adult male and adult female, respectively) have the same collection data as the holotype.

Diagnosis.—Adult males reaching 122.2 mm SVL, adult females reaching 119.1 mm SVL; 10–12 supralabials, 8–11 infralabials; tubercles of dorsum small to moderate with no intervening smaller tubercles; no tubercles on ventral surfaces of forelimbs, gular region, or in ventrolateral body fold; 34–38 paravertebral tubercles; 23–27 longitudinal rows of dorsal tubercles; 33–36 rows of ventral scales; 22–27 subdigital lamellae on fourth toe; 41–46 femoro-precloacal pores in males; dorsum not bearing a scattered pattern of white tubercles; three body bands in adults lacking lightened centers and light colored tubercles; body band to interspace ratio 2.00–2.75; six or seven wide, dark, caudal bands on original tail; white caudal bands in adults not heavily infused with dark pigment; and posterior portion of tail in hatchlings and juveniles banded not white. These characters are scored across all species of the Cyrtodactylus pulchellus complex in Table 6 .

Description of holotype.—Adult male SVL 122.2 mm; head large, moderate in length (HL/SVL 0.26) and wide (HW/HL 0.89), somewhat flattened (HD/HL 0.38), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis rounded anteriorly; snout elongate (ES/HL 0.44), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.23); ear opening elliptical, moderate in size (EL/HL 0.07), obliquely oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by an inverted Y-shaped furrow, bordered posteriorly by left and right supranasals, a medial postrostral (=internasal), bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal and small posterior supranasal, posteriorly by two small postnasals, ventrally by first supralabial; 11(R,L) square supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; first supralabial largest; 11(R,L) infralabials tapering in size posteriorly; scales of rostrum and lores flat, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales of occiput and top of head between eyes intermixed with small tubercles; large, boney frontal ridges bordering orbit confluent with boney, V-shaped, transverse, parietal ridge; dorsal superciliaries elongate, smooth, largest anteriorly, damaged on right; mental triangular, bordered laterally by first infralabials and posteriorly by left and right, rectangular postmentals which contact medially for 50% of their length; single row of enlarged, elongate sublabials extending posteriorly to 3rd infralabial; small, granular to flat, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.47) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with larger, trihedral, regularly arranged, keeled tubercles, smaller intervening tubercles generally absent; tubercles extend from top of head from between eyes to caudal constriction and onto tail where they occur in transverse rows separated by six or seven small, flat scales; caudal tubercles largest dorsally, absent laterally and ventrally; caudal tubercles decrease in size posteriorly; tubercles on occiput and nape small, those on body largest; approximately 24 longitudinal rows of tubercles at midbody; 38 paravertebral tubercles; 34 flat imbricate ventral scales between ventrolateral body folds, ventral scales larger than dorsal scales; precloacal scales large, smooth; distinct precloacal groove.

Forelimbs moderate, relatively short (FL/SVL 0.16); scales on dorsal surfaces of forelimbs slightly raised, intermixed with larger tubercles; scales of ventral surface of forearm flat, subimbricate, lacking tubercles; palmar scales rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae rectangular proximal to joint inflection, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.18), larger tubercles on dorsal surface of thigh separated by smaller granular scales, tubercles on dorsal surface of foreleg same size as those on thigh; ventral scales of thigh flat, smooth, imbricate, larger than dorsal granular scales; ventral, tibial scales flat, smooth, imbricate; single row of greatly enlarged, flat, rectangular, imbricate, pore-bearing femoral scales extend from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 41 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which 10 pore-bearing scales are found (five on each side of groove); postfemoral scales immediately posterior to pore-bearing scale row small, forming an abrupt union with pore-bearing postfemoral scales on posteroventral margin of thigh; plantar scales low, slightly raised; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae proximal to joint inflection rectangular, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; 25(R) 24(L) subdigital lamellae on 4th toe.

Tail 150 mm in length, 11.1 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; tail segmented with six or seven transverse scale rows per segment; posterior margin of segments bordered by four larger tubercles dorsally in anterior one-third of tail, fewer posteriorly; subcaudal region bearing large, transverse scales; shallow dorsal and lateral caudal furrows extend entire length of tail; base of tail bearing hemipenial swellings; three small, postcloacal tubercles on left hemipenial swellings, two on right; postcloacal scales smooth, flat, large, imbricate.

Coloration in life ( Fig. 15 ). Dorsal ground color of head, body, limbs, and tail dark brown; no white chevron on rostrum; wide, very dark brown nearly black, nuchal band extends from posterior margin of one eye to posterior margin of other eye, edged with prominent thin, white, dotted lines of tubercles; three wide, similarly colored dorsal bands between limb insertions with immaculate, white tubercles forming a dotted line edging the bands; first band terminates at shoulders, second band terminates just dorsal to ventrolateral fold midway between limb insertions, third band terminates on anterior margin of hind limb, groin, and the area anterior to the groin; body band/interspace ratio 2.75; one additional dark brown band posterior to hind limbs; oblique band outlined with white tubercles on posterior margin of thigh; six wide, dark bands approximately four times the width of five white bands extend onto tail; all bands encircle tail, no bands on tail or body have lighten centers or encompass yellowish to white tubercles; ventral surfaces of head, limbs, and tail heavily smudged with brown; gular scales stippled; abdomen immaculate, beige except for slightly darker, anterolateral regions, and weakly stippled ventral scales.

Variation. The dorsal color pattern of the paratypes (LSUHC 10064–65, 10461 female, male and female, respectively) is generally more vivid overall, especially in LSUHC 10065 ( Fig. 16 ). Grismer (2011:420) illustrated a juvenile from Cameron Highlands with an anomalous dorsal pattern and seven black caudal bands. Additionally, the posterior one-half of the tail of LSUHC 10065 is regenerated. LSUDPC 6612–13 are newly born captive specimens with completely banded tails. Meristic differences in the type series and additional specimens examined are presented in Table 12.

Additional specimens examined. ZRC 2.1178 has a completely regenerated tail that lacks a pattern and its interspaces between the dorsal bands are slightly wider ( Fig. 16 ).

Distribution. Cyrtodactylus trilatofasciatus sp. nov. is known only from the type locality at Ringlet, Cameron Highlands, Pahang, Malaysia ( Fig. 3 ). This species most certainly occurs in other localities throughout the upland regions of Cameron Highlands and to the north where rocks are present.

Natural history. Cyrtodactylus trilatofasciatus sp. nov. occurs in hill dipterocarp and montane forests from approximately 1000–1500 meters in elevation. In these habitats lizards emerge at night and forage among rocks or on nearby vegetation ( Fig. 15 ). This species has not been found in forested areas where rocks are absent.

Etymology. The specific epithet trilatofasciatus comes from the Latin words ter meaning three, latus meaning wide, and fasciatus meaning banded and refers the diagnostic character state of having three wide bands.

Comparisons. Cyrtodactylus trilatofasciatus sp. nov. is separated from C. macrotuberculatus in lacking large tubercles on the dorsal surface of the head, body and limbs, on the underside of the forearms, in the gular region, and in the ventrolateral body fold. From C. astrum sp. nov. it differs in having 34–38 paravertebral tubercles as opposed to 40–57. It differs from C. lekaguli sp. nov. in having 23–27 longitudinal rows of dorsal tubercles as opposed to 20–24 rows of tubercles. It is separated from C. bintangtinggi sp. nov. and C. langkawiensis sp. nov. in having less than 37 ventral scales (overlapping with C. bintangtinggi sp. nov. by one scale at 36) and from C. macrotuberculatus by having more than 32 ventral scales. It differs from all species except C. australotitiwangsaensis sp. nov. and C. bintangrendah sp. nov. in having more than 40 femoro-precloacal pores (overlapping with C. bintangtinggi sp. nov. by one pore at 41). It differs from all species except C. australotitiwangsaensis sp. nov. by having three body bands as opposed to four or more (one specimen [LSUHC 9009] out of 14 C. bintangtinggi sp. nov. has three bands). It differs from all species by having body band/ interspace ratio of 2.00–2.75 as opposed to a ratio of 2.00 or less (one specimen [ZRC 2.6964] of 11 C. astrum sp. nov., two specimens [LSUHC 9107–08] of 13 C. australotitiwangsaensis sp. nov., and one specimen [MS 441] of 16 C. lekaguli sp. nov. have a ratio of 2.00). Cyrtodactylus trilatofasciatus sp. nov. is further separated from C.

astrum sp. nov. in lacking a scattered dorsal pattern of white tubercles. Cyrtodactylus trilatofasciatus sp. nov. differs from all species except C. australotitiwangsaensis sp. nov. and C. macrotuberculatus by having no more than seven dark caudal bands. Cyrtodactylus trilatofasciatus sp. nov. differs from all species except C. australotitiwangsaensis sp. nov. and C. macrotuberculatus in having a maximum SVL greater than 115 mm.

SVL=snout-vent length; TL=tail length; TW=tail width; FL=forelimb length; TBL=tibia length; AG=axilla-groin length;

HL=head length; HW=head width; HD=head depth; ED=eye diameter; EE=eye to ear distance; ES=eye to snout

distance; EN=eye to nostril distance; IO=interorbital distance; EL=ear length; and IN=internarial distance B = broken;

PR = partially regenerated; R = regenerated; /=data unavailable.

ZRC ZRC ZRC LSUHC ZRC ZRC Remarks. Despite the fact that Cyrtodactylus trilatofasciatus sp. nov. from Cameron Highlands, Pahang and C. australotitiwangsaensis sp. nov. from Fraser’s Hill and Genting Highlands, Pahang are sister species separated by only 76 km of continuous upland forest, the genetic distance between them ranges from 6.0–9.3%. This biogeographical pattern of well-differentiated sister lineages is consistent with other taxa from these areas such as Cnemaspis flavolineata (Grismer et al. in prep), Larutia miodactyla (Boulenger) ( Grismer 2011:592) , and Sphenomorphus bukitensis Grismer (Grismer 2011:631) as well as with the general high degree of herpetological endemism in Cameron Highlands (e.g. the megophryid Kalophrynus youngi Matsui ; the microhylid Leptolalax kecil Matsui, Belabut, Norhayati & Yong ; the agamid Pseudocalotes flavigula Smith ; the scincid Sphenomorphus cameronicus Smith ; the colubrids Smedley; Macrocalamus schulzi Vo g e l & D a v i d and M. tweediei Lim ; and the viperid Popeia nebularis Vogel, David & Pauwels ). Reasons for this high level of endemism are uncertain but it may have to do with the fact that the upland plateau of Cameron Highlands is extensive whereas the regions of Fraser’s Hill and Genting Highlands are much more restricted, being little more than montane peaks.

Figures

FIGURE 3. Distribution of the species of the Cyrtodactylus pulchellus complex. Closed circles represent localities from which specimens have been examined genetically and / or morphologically (see appendix and Table 1). Closed squares are localities from which specimens have been only examined morphologically or photographically (see appendix). Closed stars represent the type localities from where specimens have been examined genetically, morphologically and photographically. 1. Suwankuha Cave, Muang District, Phang-nga Province, Thailand. 2. Petch Phanomwung Cave, Kanchanadit District, Surat Thani Province, Thailand. 3. Phuket Island 4. Khao Chong waterfall, Khao Chamao, Khaochong District, Trang Province, Thailand. 5. Phattalung Province, Thailand. 6. Rattabhumi District, Songkhla, Thailand. 7. La-ngu District, Satun Province, Thailand (three squares). 8. Gua Wang Burma, Perlis State Park, Perlis, Malaysia (two circles). 9. Wang Kelian, Perlis, Malaysia. 10. Adang Island, Satun Province, Thailand. 11. Pulau Langkawi, Kedah, Malaysia (two stars). 12. Gua Kelam and Bukit Chabang, Malaysia (two circles). 13. Kuala Perlis, Perlis, Malaysia. 14. Chuping, Perlis, Malaysia. 15. Bukit Wang, Kedah, Malaysia. 16. Ulu Muda, Kedah, Malaysia. 17. Gunung Jerai and Hutan Lipur Sungai Tupah, Kedah, Malaysia (two circles). 18. Bukit Palong, Perak, Malaysia. 19. Ulu Paip, Kedah, Malaysia. 20. Bukit Mertajam, Penang, Malaysia. 21. Pulau Pinang, Penang, Malaysia. 22. Lengong Valley, Perak, Malaysia. 23. Bukit Panchor, Penang, Malaysia. 24. Bukit Larut, Perak, Malaysia. 25. Bukit Pondok, Perak, Malaysia. 26. Ringlet, Cameron Highlands, Pahang, Malaysia. 27. Gunung Bubu, Perak, Malaysia. 28. Fraser’s Hill, Pahang, Malaysia. 29. Genting Highlands, Pahang, Malaysia. 30. Gunung Angsi, Negeri Sembilan, Malaysia. P = Tham Phung Chang Cave, Phang-nga City, Muang District, Phang-nga Province, Thailand. U = Nakon Si Thammarat Province; specific locality unknown.

FIGURE 15. Cyrtodactylus trilatofasciatus from Ringlet, Cameron Highlands, Pahang, Malaysia. Upper left: LSUHC 10461 (paratype); adult female. Upper right: LSUDPC 5892; adult female. Lower left: ZRC 2.6976 (holotype); adult male. Lower right: Lower montane forest habitat of C. trilatofasciatus at Cameron Highlands, Pahang, Malaysia. All photos by L. Grismer.

FIGURE 16. Type series and additional specimens examined of Cyrtodactylus trilatofasciatus. Upper row: type series from left to right; holotype ZRC 2.6976, paratypes ZRC 2.6977 – 78, LSUHC 10461. Lower row: additional specimens examined from type locality from left to right: LSUHC ZRC 2.1178, 2.1179.