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 Author Grismer, Lee Author Wood, Perry L. Author Quah, Evan S. H. Author Anuar, Shahrul Author Muin, Abdul Author Sumontha, Montri Author Ahmad, Norhayati Author Bauer, Aaron M. text Zootaxa 2012 3520 1 55 journal article 10.5281/zenodo.211847 9ac583a7-c77c-49a4-93a5-dedcd7789325 1175-5326 211847 Cyrtodactylus lekaguli sp. nov. Tuk-kai Boonsong, Bent-toed Gecko Figs. 7 , 8 Cyrtodactylus pulchellus Taylor 1963 :714 ; Manthey and Grossmann 1997 :226 ; Cox, van Dijk, Nabhitabhata and Thirakhupt 1998:86; Pauwels et al . 2000 :129–130, 2002:27. Holotype .—Adult male ( FMNH 215987) collected on 7 November 1979 by D. L. Damman from Khao Chong waterfall, Khao Chamao, Trang Province, Thailand at approximately 07° 35.28 N , 99° 50.25 E at 165 meters above sea level as estimated from Google Earth. Paratypes .—Adult females ( FMNH 215985–86) bear the same data as the holotype . The adult male FMNH 176885 was collected by E. H. Taylor from Khao Chao (date unknown). Adult female IRSNB 2678 collected on 21 July 1998 by O. S. G. Pauwels and Chucheep Chimsunchart from Tham Phung Chang (cave) Phang-nga City, Muang District, Phang-nga Province, Thailand . Diagnosis .—Adult males reaching 103.5 mm SVL, adult females reaching 97.3 mm SVL; 10–12 supralabials, 9–11 infralabials; tubercles of dorsum moderate, with no intervening smaller tubercles; no tubercles on ventral surfaces of forelimbs, gular region, or in ventrolateral body fold; 30–50 paravertebral tubercles; 20–24 longitudinal rows of dorsal tubercles; 31–43 rows of ventral scales; 20–25 subdigital lamellae on fourth toe; 30–36 femoroprecloacal pores in males; dorsum not bearing a scattered pattern of white tubercles; four or five dark body bands in adults lacking lightened centers and light colored tubercles; body band to interspace ratio 1.00–2.00; 12–14 dark caudal bands; and white caudal bands infused with dark coloration and the posterior portion of the tail in hatchlings and juveniles is white. These characters are scored across all species of the Cyrtodactylus pulchellus complex in Table 6 . Description of holotype .—Adult male SVL 92.2 mm ; head large, moderate in length (HL/SVL 0.28) and wide (HW/HL 0.76), somewhat flattened (HD/HL 0.41), distinct from neck, and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded anteriorly; snout elongate (ES/HL 0.43), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.25); ear opening elliptical, moderate in size (EL/HL 0.10), vertically 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 and medial postrostral (=internasal), bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large, anterior supranasal and smaller posterior supranasal, posteriorly by large single postnasal, ventrally by first supralabial; 10(R,L) square supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; second supralabial not larger than first; 10,9(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 largest; scales of occiput intermixed with small tubercles; large, boney frontal ridges bordering orbit confluent with boney, V-shaped, transverse, parietal ridge; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by left and right, rectangular postmentals which contact medially for 50% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to 5th infralabial; small, granular, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales. FIGURE 7. Cyrtodactylus lekaguli . Upper left: LSUDPC 6317; adult female (gravid) from La-ngu District, Satun Province, Thailand (photo by M. Sumontha). Right: LSUDPC 6325, Krabi, Krabi Province, Thailand (photo by N. Baker). Middle left: MS 519; adult male from Petch Phanomwung Cave, Kanchanadit District, Surat Thani Province, Thailand (photo by M. Sumontha). Lower left: LSUDPC 6327, Khamin Cave, Ban Nasan District, Surat Thani Province, Thailand (photo by M. Sumontha). Lower right: LSUDPC 6326, Krabi, Krabi Province, Thailand (photo by N. Baker). Body relatively short (AG/SVL 0.42) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles not separated by smaller intervening tubercles; tubercles extend from occipital region to caudal constriction and onto original portion of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 23 longitudinal rows of tubercles at midbody; 35 paravertebral tubercles; 41 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 surface of forelimbs raised, intermixed with larger tubercles; scales of ventral surface of forearm flat, rounded, 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 0.21), larger tubercles on dorsal surface of thigh separated by smaller granular scales, tubercles on dorsal surfaces of foreleg smaller; 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; 33 contiguous, pore-bearing femoro-precloacal scales forming an inverted T bearing a deep, precloacal groove in which eight pore-bearing scales are found (four 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 rounded; 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; 23,24(R,L) subdigital lamellae on 4th toe. Tail original, 125 mm in length, 9.0 mm in width at base, tapering to a point; dorsal scales small, square, weakly keeled, in transverse rows; subcaudal region bearing large, transverse scales for first half of tail, second half bearing smaller, irregularly shaped scales; base of tail bearing hemipenial swellings; four smaller, postcloacal tubercles on hemipenial swellings; postcloacal scales smooth, flat, large, imbricate. Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail brown; wide, dark brown, nuchal band extends from posterior margin of one eye to posterior margin of other eye; nuchal band edged with thin, white, lines; four similarly colored dorsal bands lacking lightened centers occur between limb insertions, first band terminates at shoulders, second and third bands terminate dorsal to ventrolateral fold, fourth band terminates at anterior margin of hind limb insertions; white edging on body bands generally restricted to the tubercles giving edging a spotted appearance; body band/interspace ratio 1.00; one additional dark brown band posterior to hind limbs; no band on posterior margin of thigh; 14 dark, caudal bands; 13 light caudal bands infused with dark pigment; scales of all ventral surfaces heavily stippled. Variation. The paratypes closely approach the holotype in coloration ( Fig. 8 ). FMNH 176885 (SVL 100.0 mm) and 215985 (SVL 80.5 mm ) have a slightly lighter ground color and in FMNH 215985, the ends of the second and third body bands contact one another on the left side of the body just dorsal to the ventrolateral fold. IRSNB 2678 has an overall more faded color pattern ( Fig. 8 ). Juveniles have white tail tips ( Fig. 7 ). Additional specimens examined. MS 441 and 442 from Khao Chong, Trang Province closely approximate the holotype in overall coloration other than being slightly darker ( Fig. 8 ). MS 442 has a completely regenerated tail and MS 441 has relatively narrow body bands. MS 519 from Petch Phanomwung Cave, Kanchanadit District, Surat Thani Province, MS 79 from Suwankuha Cave, Muang District, Phang-nga Province, and IRSNB 16558 from Nakon Si Thammarat Province, Thailand are much lighter in overall coloration. Meristic differences between individuals of the type series and additional specimens examined are presented in Table 8 . Taylor (1963) reported a male (No. 35747) from the Khao Chong Forest Station, Trang Province with a SVL of 102 mm and 38 femoroprecloacal pores. FIGURE 8. Type series and additional specimens examined of Cyrtodactylus lekaguli . Upper row: type series from left to right FMNH 215987, 215895, 215986, 176885. Additional specimens examined from type locality from left to right: MS 441, 442. Lower row: additional specimens examined from Satun Province, Thailand from left to right: MS 401, 445, 446 and from Surat Thani Province, Thailand MS 519 and Phang-nga Province, Thailand MS 79; paratype IRSNB 2678 from Phang-nga Privince, Thailand. Distribution. Cyrtodactylus lekaguli sp. nov. is known from the type locality at Khao Chong waterfall, Khao Chamao, Trang Province, from Petch Phanomwung Cave, Kanchanadit District, Surat Thani Province, Krabi, Krabi Province, and from Suwankuha Cave, Phung Chang Cave, Reusi Cave, Tham Phung Chang Cave, Phangnga City, Muang District, Phang-nga Province, Thailand ( Fig. 3 ). Natural history. Taylor (1963) reported specimens being found at the Khao Chong Forest Station, Trang Province on the trunks of trees in the forest near streams, in the crevices of trees, and in the rest house. These data would suggest this population may be less confined to rocky areas than are other members of the Cyrtodactylus pulchellus complex. Specimens from Krabi, Phang-nga, and Surat Thani Provinces were found in association with karst boulders and caves. Pauwels et al. (2000) reported that during a one-hour visit in late July in Phung Chang Cave, six individuals were observed, and the species was abundant in Reusi Cave as well. They reported a female (IRSNB 15143) collected in July that contained two eggs of about 11 mm diameter, and another one collected on the same day that contained an egg of about 9 mm diameter and another much less developed. They also reported on a hatchling that hatched one day after its egg was collected deep from within the Reusi Cave on January 11th . That egg had been deposited along with 15 others, of which 11 had recently hatched, in a small fissure filled with soft soil. The eggs were about 16 mm by 13 mm . The hatchling (MNHN 1999.7706) had a SVL of 38 mm and a total length of 79 mm at birth and a white tail tip. Etymology. The specific epithet lekaguli honors Dr. Boonsong Lekagul ( 1907–1992 ), biologist and conservationist of Thailand and the first biologist to suggest the Thai common name " Tuk-Kai " for geckos of the genus Cyrtodactylus ( Lekagul 1977 ) . TABLE 8. Morphological data of type series and additional specimens of Cyrtodactylus lekaguli from Thailand. m=male; f=female; 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. FMNH FMNH FMNH FMNH IRSNB MS MS MS MS MS MS MS IRSBN 176885 215985 215986 215987 2678 441 442 401 445 446 519 79 16558 Paratype Paratype Paratype Holotype Paratype

Khao Chong	Khao Chong	Khao Chong	Khao Chong	Phang- nga	Khao Chong	Khao Chong	La- ngu	La- ngu	La- ngu	Surat Thani	Phang -nga	Nakkon Si Thammarat
Sex	m	f	f	m	f	m	f	f	m	f	m	m	m
Supralabials	11	11	12	10	11	11	10	11	12	12	12	12	11
Infralabials	9	10	9	10	9	9	10	10	9	9	9	11	9
Tubercles on ventral surface of forelimbs	0	0	0	0	0	0	0	0	0	0	0	0	0
Tubercles in gular region	0	0	0	0	0	0	0	0	0	0	0	0	0
Ventrolateral fold tuberculate	0	0	0	0	0	0	0	0	0	0	0	0	0
No. of paravertebral tubercles	40	44	50	35	39	30	37	38	41	43	35	39	35
No. longitudinal rows of tubercles	22	22	22	23	22	21	22	20	20	20	24	22	22
No. of ventral scales	38	43	40	41	40	38	37	33	31	35	38	33	36
Proximal subdigital lamellae square	0	0	0	0	0	0	0	0	0	0	0	0	0
No. of subdigital lamellae on 4th toe	23	24	21	23	22	20	21	23	20	23	25	24	22
No. of femoro- precloacal pores	34	/	/	33	/	34	/	/	36	/	30	33	36
Bands on base of thigh	1	1	0	0	0	0	0	0	0	0	0	0	0
No. body bands	4	4	4	4	5	4	4	4	4	4	5	4	4
Band/interspace width	1.00	1.00	1.00	1.00	1.00	2.00	1.25	1.00	1.25	1.25	1.00	1.25	1.00
No. dark bands on original tail	/	/	12	14	/	/	/	/	/	/	/	/	/
Hatch./juv. with white tail tips	/	/	/	/	/	/	/	/	/	/	/	/	/
SVL	100.0	80.5	85.7	92.2	95.0	103.5	97.3	93.9	93.7	95.6	102.7	97.2	103.4
TL	82B	93R	115	125	97.8PR
TW	9.4	7.2	6.9	9	7.5
FL	17.3	13	13.6	14.8	15.4
TBL	20	15.8	15.9	19.5	19.7
AG	45	35.4	35.7	39	45.0
HL	28.2	23	24	25.4	26.5
HW	20	15.8	16.7	19.2	17.3
HD	12.1	9.7	9	10.3	10.7
ED	6.3	5	5.8	6.4	7.0
EE	8.5	7.3	6.9	7.9	8.3
ES	12.3	10	10.1	11	11.6
EN	9.7	7.3	7.6	8.1	9.0
IO	5.8	4.9	4.9	5.4	5.6
EL	1.9	1.8	2	2.1	1.5
IN	3.5	2.5	2.6	2.4	2.6

Comparisons. Cyrtodactylus lekaguli 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, in the ventrolateral body fold, and in having 31–43 as opposed to 17–28 ventral scales. Cyrtodactylus lekaguli sp. nov. has fewer (20–24) longitudinal rows of dorsal tubercles than C. trilatofasciatus sp. nov. (23–27). Cyrtodactylus lekaguli sp. nov. is separated from C. australotitiwangsaensis sp. nov. , C. bintangtinggi sp. nov. , C. bintangrendah sp. nov. , and C. trilatofasciatus sp. nov. in having less than less than 37 femoro-precloacal pores. From C. trilatofasciatus sp. nov. , C. lekaguli sp. nov. differs in having four or five narrow body bands with a band/ interspace ratio of 1.00–2.00 as opposed to having three wide body bands with a band/interspace ratio of 2.00–2.75 and it differs from C. langkawiensis sp. nov. in having wider body bands (1.00–2.00 vs. 0.75–1.00, respectively). It differs from C. astrum sp. nov. in lacking a scattered dorsal pattern of white tubercles. Having 12–14 dark caudal bands differentiates it from C. australotitiwangsaensis sp. nov. , C. bintangtinggi sp. nov. , C. macrotuberculatus , C. pulchellus and C. trilatofasciatus sp. nov. which have less than 11 dark caudal bands. Heavy dark mottling in the white caudal bands differentiate it from all other species except C. astrum sp. nov. , and C. langkawiensis sp. nov. The small maximum SVL ( 103.5 mm ) of C. lekaguli sp. nov. separates it from the larger species C. astrum sp. nov. , C. australotitiwangsaensis sp. nov. , C. bintangtinggi sp. nov. , C. bintangrendah sp. nov. , C. macrotuberculatus sp. nov. , C. pulchellus , and C. trilatofasciatus sp. nov. that reach over 108 mm SVL. Remarks. Cyrtodactylus lekaguli sp. nov. , C. langkawiensis sp. nov. and C. astrum sp. nov. , are the only species previously considered to be C. pulchellus that have white caudal bands infused with dark pigment. Like C. langkawiensis sp. nov. , C. lekaguli sp. nov. is a relatively small species (neither reaches 104 mm SVL; [ Taylor 1963 ]) and the hatchlings of all three species have white tail tips. Based on this, and the fact that these species are found collectively only in extreme northern Malaysia and southern Thailand , we hypothesize that C. lekaguli sp. nov. forms a close, but yet unresolved, phylogenetic relationship with C. astrum sp. nov. and C. langkawiensis sp. nov. Three specimens from La-ngu, Satun Province, Thailand (MS 401, 445–446) 75 km south of the type locality are only weakly diagnosable from the three individuals of the type series and other individuals from Khao Chong, Phnag-nga, and Nakon Si Thammarat provinces on the basis of having fewer ventral scales (31–35 vs. 37–43) and fewer longitudinal rows of dorsal tubercles (20 vs. 22 or 23; Table 8 ). Dorsal tuberculation is generally weaker and the overall color pattern is less contrasted ( Fig. 8 ). These morphological data suggest they may not be conspecific with C. lekaguli sp. nov. from the type locality and northward. However, until molecular evidence can be brought to bear on this hypothesis, we consider this population to be C. cf. lekaguli .