Two new Bent-toed Geckos of the Cyrtodactylus pulchellus complex from Peninsular Malaysia and multiple instances of convergent adaptation to limestone forest ecosystems Author Grismer, L. Lee Author Wood, Perry L. Author Anuar, Shahrul Author Grismer, Marta S. Author Quah, Evan S. H. Author Murdoch, Matthew L. Author Muin, Mohd Abdul Author Davis, Hayden R. Author Aguilar, César Author Klabacka, Randy Author Cobos, Anthony J. Author Aowphol, Anchalee Author Sites, Jack W. text Zootaxa 2016 4105 5 401 429 journal article 39163 10.11646/zootaxa.4105.5.1 b0c40bc1-9e22-4422-bc18-6e5ddf1fd0ab 1175-5326 264750 3CBCC1EB-48BE-4730-A845-F64CFB196661 Cyrtodactylus hidupselamanya sp. nov. English: Chiku Bent-toed Gecko Malay: Cicak Jari-bengkok Chiku Fig. 3 , 4 Holotype . Adult male, LSUHC 12163 collected on 19 March 2015 at 2030 hrs by Shahrul Anuar from Felda Chiku 7, Kelantan, Peninsular Malaysia ( 5° 03.318” N 102° 08.573” E ; 110 m elevation). Paratypes . Paratypes LSUHC 12158–62, 12164–65, 12173–75 bear the same collection data as the holotype . Diagnosis. Cyrtodactylus hidupselamanya sp. nov. can be differentiated from all other species of Cyrtodactylus by having the combination of the following characters: maximum SVL of approximately 199 mm ; 8–12 supralabials; 9–12 infralabials; weak tuberculation on body; no tubercles on ventral surface of forelimbs, gular region, in ventrolateral body folds, or anterior one-third of tail; 39–48 paravertebral tubercles; 19–23 longitudinal tubercle rows; 26–33 ventral scales; 19–24 subdigital lamellae on fourth toe; no femoral pores; 17–22 precloacal pores; deep precloacal groove in males; four dark dorsal body bands; body bands as wide or slightly wider than interspaces; no rostral chevron; body bands and nuchal loop edged with a thin white, tubercle bearing lines; no scattered white tubercles on dorsum; no banding on base of thigh; 8–10 dark caudal bands on original tail; white caudal bands generally immaculate; and hatchlings and juveniles bearing white tail tips. These characters are scored across all species of the C. pulchellus complex in Table 5 . TABLE 4. Uncorrected p-distances among the species of the Cyrtodactylus pulchellu s complex based on 1473 bp of ND2 and its flanking tRNAs calculated in MEGA v6.06 (Tamura et al . 2013). Numbers in bold are within group distances.

C. astrum	0.014
C. australotitiwangsaensis	0.151	0.006
C. bintangrendah	0.156	0.095	0.011
C. bintangtinggi	0.149	0.100	0.086	0.004
C. hidupselamanya sp. nov.	0.166	0.152	0.160	0.146	0.000
C. jelawangensis	0.159	0.160	0.161	0.155	0.091	0.000
C. langkawiensis	0.095	0.146	0.147	0.142	0.174	0.164	0.001
C. lenggongensis sp. nov.	0.159	0.093	0.020	0.090	0.160	0.165	0.151	0.001
C. lekaguli	0.095	0.135	0.145	0.142	0.158	0.151	0.092	0.143	0.000
C. macrotuberculatus	0.167	0.115	0.119	0.113	0.171	0.161	0.177	0.119	0.174	0.034
C. pulchellus	0.167	0.113	0.131	0.127	0.171	0.153	0.174	0.135	0.164	0.095	0.010
C. sharkari	0.148	0.058	0.098	0.104	0.160	0.155	0.148	0.101	0.140	0.108	0.108	–
C. timur	0.171	0.146	0.153	0.156	0.056	0.099	0.163	0.158	0.155	0.183	0.181	0.159	0.004
C. trilatofasciatus	0.148	0.064	0.104	0.106	0.163	0.155	0.141	0.107	0.137	0.118	0.115	0.017	0.162	0.009

TABLE 5 . Diagnostic characters differentiating the 15 species of the Cyrtodactylus pulchellus complex. W=weak; P=prominent; /=data unavailable.

Supralabials	10–12	9–12	9–13	8–12	9–12	10–12	9–12
Infralabials	9–12	9–13	8–11	8 or 9	8–10	9–11	7–10
Tuberculation	W	P	P	P	W	W	P
Tubercles on ventral surface of forelimbs	No	No	No	No	No	No	Yes
Tubercles in gular region	No	No	No	No	No	No	Yes
Ventrolateral fold tuberculate	No	No	No	Yes	No	No	Yes
Paravertebral tubercles	45–57	37–45	31–42	36–44	38–44	30–50	34–44
Longitudinal rows of tubercles	22–29	23–30	21–26	22–25	22–25	20–24	19–27
Ventral scales	31–46	32–40	36–40	31–39	31–36	31–43	17–28
4th toe lamellae	20–24	21–27	21–25	21–24	19–21	20–25	20–24
Femoroprecloacal pores	31–38	39–45	37–41	41–46	30	33–36	28–41
Precloacal groove in males	Deep	Deep	Deep	Deep	Deep	Deep	Deep
Body bands	4	3(1) or 4	3(1) or 4	4	4 or 5	4 or 5	4
Body bands two times or more than the width of the interspace	No	No	No	No	No	No	No
Dorsum bearing scattered pattern of white tubercles	Yes	No	No	No	No	No	No
Hatchlings/juveniles with white tail tip	Yes	No	No	No	Yes	/	No
Adult posterior caudal region white	No	No	No	No	No	No	No
Dark caudal bands on original tail	12–14	7–8	8–10	8 or 9	11–16	12–14	7–10
White caudal bands in adults immaculate	No	Yes	Yes	Yes	No	No	No
Maximum SVL	108.1	119.8	111.4	114.4	99.1	103.5	118
Sample size	11	13	13	10	10	11	29

TABLE 5. (Continued)

Supralabials	11–13	9–11	10–12	11	9–12	10–12	9–12	10 or 11
Infralabials	9,10	8–10	8–11	10	9–11	8–10	8–11	8–10
Tuberculation	P	P	P	W	P	P	W	W
Tubercles on ventral surface of forelimbs	Yes	No	No	No	Yes	No	No	No
Tubercles in gular region	Yes	No	No	No	No	No	No	No
Ventrolateral fold tuberculate	Yes	No	No	No	No	No	No	No
Paravertebral tubercles	41–43	33–43	34–38	31	36–42	38–43	39–48	36–41
Longitudinal rows of tubercles	23,24	22–26	23–27	24	23–25	21–24	19–23	22–25
Ventral scales	22–24	29–34	33–36	41	31–36	31–40	26–33	32 or 33
4th toe lamellae	19	21–26	22–27	24	21–24	21–25	19–24	20–23
Femoroprecloacal pores	33–36	34–39	41–46	46	36	21 or 22	17–22	39–41
Precloacal groove in males	Shallow	Deep	Deep	Shallow	Deep	Deep	Deep	Deep
Body bands	3	4	3	4	4	4	4	4 or 5
Body band/interspace ratio	No	No	Yes	No	No	No	No	No
Dorsum bearing scattered pattern of white tubercles	No	No	No	No	No	No	No	No
Hatchlings/juveniles with white tail tip	No	No	/	/	Yes	No	Yes	/
Adult posterior caudal region white	No	No	No	No	No	No	Yes	No
Dark caudal bands on original tail	8,9	8–10	6	7	10	8–10	8–10	14
White caudal bands in adults immaculate	Usually	Yes	Yes	Yes	No	Yes	Yes	Yes
Maximum SVL	114.7	114.6	122.2	100.1	119.8	120.5	102.7	103.1
Sample size	3	13	6	1	4	5	14	4

Description of holotype . Adult male, 97.5 mm SVL; head large, moderate in length (HL/SVL 0.30), wide (HW/HL 0.67), somewhat flattened (HD/HL 0.36), distinct from neck, triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis rounded anteriorly; snout elongate (ES/HL 0.45), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.26); 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, and one medial postrostrals (=internasals), bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal and smaller postrostral, posteriorly by two postnasals, ventrally by first supralabial; 11(R, L) rectangular supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; first supralabial largest; 7(R) 8(L) infralabials tapering in size posteriorly; scales of rostrum and lores weakly raised, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales on occiput intermixed with small tubercles; posterior interorbital region tuberculate; large, boney frontal ridges bordering orbit confluent with boney, 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 70% of their length; single row of slightly enlarged, elongate chinshields extending posteriorly to fourth infralabials; 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 low, regularly arranged, keeled tubercles, smaller intervening tubercles occasionally present; tubercles extend from top of head to caudal constriction and onto anterior one-fifth of tail; tubercles on occiput and nape small, those on body largest; approximately 22 longitudinal rows of tubercles at midbody; 44 paravertebral tubercles; 27 flat imbricate ventral scales between ventrolateral body folds; ventral scales larger than dorsal scales; precloacal scales large, smooth; deep precloacal groove. Forelimbs moderate in stature, relatively short (FL/SVL 0.18); scales on dorsal surfaces of forelimbs, small, juxtaposed, intermixed with large tubercles in near contact with one another; scales of ventral surface of forearm flat, subimbricate, tubercles absent; palmar scales weakly 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 0.20), larger tubercles on dorsal surface of legs separated by smaller subimbricate scales; 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, femoroprecloacal scales extend nearly from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 19 contiguous, pore-bearing precloacal scales forming an inverted T bearing a deep, precloacal groove in which six pore-bearing scales are found (three on left, three on right); postfemoral scales immediately posterior to enlarged scale row small, nearly granular, forming an abrupt union with postfemoral scales on posteroventral margin of thigh; plantar scales low, slightly raised; digits welldeveloped, 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(R) 22(L) subdigital lamellae on 4th toe. Tail original 128.0 mm in length, 9.3 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; subcaudal region bearing large median row of transverse scales; no caudal furrows; base of tail bearing hemipenial swellings; three small, postcloacal tubercles on each hemipenial swelling; postcloacal scales smooth, flat, large, imbricate. Coloration in life. Dorsal ground color of head, body, limbs, and tail light-brown, immaculate; no V-shaped line on rostrum; wide, dark-brown nuchal loop edged anteriorly and posteriorly by thin, white line bearing tubercles; four dark-brown body bands between nuchal loop and hind limb insertions edged anteriorly and posteriorly by thin white lines bearing tubercles; body bands as wide as interspaces; no markings on posterior margin of thigh; ventral surface of head, body, and limbs beige, immaculate except for black stipples in each scale; tail bearing eight dark bands separated by seven, narrower, beige (anteriorly) to white (posteriorly) bands; subcaudal region tan. FIGURE 3. Cyrtodactylus hidupselamanya sp. nov. from Felda Chiku 7, Kelantan, Peninsular Malaysia. Upper left: male paratype LSUHC12175. Upper right: female paratype LSUHC 12173. Lower left: hatchling paratype LSUHC 12177. Lower right: juvenile paratype LSUHC 12176. Variation ( Figs. 3 , 4 ). The paratypes are remarkably similar to the holotype in overall coloration and pattern, likely due to substrate matching. The posterior section of the tail in the hatchling (LSUHC 12177) and the juvenile (LSUHC 12176) is whitish with faint bands—a condition found in the vast majority of cave-adapted Cyrtodactylus throughout Southeast Asia and all other limestone ecosystem adapted species in the C. pulchellus complex (Grismer et al. 2012, 2014a). However, unlike all other of the latter, the white coloration remains into adulthood when unstressed ( Fig. 4 ). Like other species in the C. pulchellus complex, hatchlings of C. hidupselamanya sp. nov. bear bright-yellow dorsal interspaces ( Fig. 3 ) that fade to brown in adulthood. Meristic and additional color pattern differences are listed in Table 6 . Distribution. Cyrtodactylus hidupselamanya sp. nov is known only from the type locality at Felda Chiku 7, Kelantan, Peninsular Malaysia ( Fig. 1 ). Another karst formation approximately 1 km to the south was not surveyed. There are other isolated karst formations 5–10 km away that were also not surveyed. Etymology. The specific epithet hidupselamanya is a modification of the Malay words “ hidup selamanya ” which, loosely translated means “live forever” and is in reference this species precarious future being that its limestone habitat is targeted to be completely quarried. Natural history. All specimens of Cyrtodactylus hidupselamanya sp. nov. were collected at night between 2000 and 2400 hrs inside a complex network of caves and caverns permeating and coursing through an isolated karst formation ( Fig. 5 ). A very narrow swath of undisturbed limestone forest closely surrounds the karst formation but the forest for several kilometers beyond this has been cleared for oil palm plantations ( Fig. 5 ). Some lizards were collected in open areas on the limestone walls near cavern entrances while others were found on walls much deeper within the cave systems. All specimens were found between 1–4 m above the cave floor. One specimen observed during the day at 1730 hrs inside the cave was taking refuge deep within a crack approximately 25 m from the cave entrance. It is likely that C. hidupselamanya sp. nov. ventures outside the caves at night to forage on the exterior walls of the karst formation although none were found. One juvenile (LSUHC 12176), however, was found on the limestone vegetation next to an exterior wall. Of the four adult females sampled (LSUHC 12159, 12162, 12173–74), none were gravid although one hatchling was collected (LSUHC 12177). This would suggest that C. hidupselamanya sp. nov. does not breed year-round and that April is at the end of its reproductive season. TABLE 6. Characters and measurements of the type series of Cyrtodactylus hidupselamanya sp. nov. from Felda Chiku 7, Kelantan, Peninsular Malaysia.M=male, F=female, /=data unavailable, R=regenerated, B=broken. Other abbreviations are listed in the Materials and Methods.

LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC
12173	12160	12158	12215	12161	12162	12163
paratype	paratype	paratype	paratype	paratype	paratype	holotype
sex	F	M	M	M	M	F	M
supralabials	11	11	11	12	11	9	11
infralabials	8	9	9	11	9	9	8
tuberculation weak, moderate, strong	Weak	Weak	Weak	Weak	Weak	Weak	Weak
tubercles on ventral surface of forelimbs tubercles in gular region	No No	No No	No No	No No	No No	No No	No No
ventrolateral fold tuberculate	No	No	No	No	No	No	No
no. of paravertebral tubercles	48	41	39	42	40	47	44
no. longitudinal rows of tubercles	22	21	24	20	21	21	22
tubercles on at least anterior 1/3 of tail	No	No	No	No	No	no	no
no. of ventral scales	27	28	33	26	29	29	27
no. of subdigital lamellae on 4th toe	23	22	24	22	23	22	22
no. of femoropreloacal pores	/	20	21	22	17	/	19
deep precloacal groove in males body bands	/ 4	Yes 4	Yes 4	Yes 4	Yes 4	/ 4	Yes 4
body band/interspace width ratio	1.25	1.25	1.25	1.25	1.00	1.25	1.00
scattered white dorsal tubercles	No	No	No	No	No	No	No
no. bands on original tail	8	/	/	9	9	9	8
posterior portion of tail white in adults immaculate white caudal bands in adults	Yes Yes	Yes Yes	Yes Yes	Yes Yes	Yes Yes	Yes Yes	Yes Yes
SVL	96	92.8	102.0	102.7	83.3	100.1	97.5
TL	122	117	80R	/	112	127	128.0
TW	7.6	8.3	9.5	8.5	7.1	7.9	9.3
FL	16.8	17.1	17.9	17.8	15.6	17.2	17.2
TBL	20.8	19.4	21.4	20.5	18.0	19.7	19.4
AG	50.1	45.1	48.7	46.6	39.9	47.8	46.3
HL	29.4	26.7	29.2	28.3	24.3	27.4	26.9
HW	19.1	17.9	20.2	18.5	15.7	19.6	18.1
HD	10.2	9.6	10.7	10.7	8.7	10.2	9.6
ED	7.0	6.9	7.5	7.3	5.8	6.3	7.0
EE	8.4	6.7	8.0	7.6	6.3	7.3	7.4
ES	12.2	12.1	13.3	12.8	10.8	12.0	12.1
EN	9.4	9.4	9.6	9.6	7.9	8.7	9.0
IO	7.5	7.8	7.3	7.3	6.7	6.4	6.4
EL	2.2	2.2	2.0	1.8	2.1	2.3	1.9
IN	3.0	3.0	3.1	3.2	3.0	2.8	2.8

……continued on the next page TABLE 6. (Continued)

LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC
12174	12165	12175	12159	12164	12176	12177
paratype	paratype	paratype	paratype	paratype	paratype	paratype
sex	F	M	M	F	M	F	M
supralabials	10	12	12	11	10	12	10
infralabials	9	8	9	9	10	8	9
tuberculation weak, moderate, strong	Weak	Weak	Weak	Weak	Weak	Weak	Weak
tubercles on ventral surface of forelimbs tubercles in gular region	No No	No No	No No	No No	No No	No No	No No
ventrolateral fold tuberculate	No	No	No	No	No	No	No
no. of paravertebral tubercles	43	40	39	46	48	43	39
no. longitudinal rows of tubercles	23	23	22	21	21	23	19
tubercles on at least anterior 1/3 of tail	no	no	no	no	no	no	no
no. of ventral scales	28	27	28	27	28	27	27
no. of subdigital lamellae on 4th toe	22	22	23	21	22	19	22
no. of femero-precloacal pores	/	21	19	/	19	/	/
deep precloacal groove in males	/	Yes	Yes	/	Yes	/	Yes
body bands	4	4	4	4	4	4	4
body band/interspace width ratio scattered white dorsal tubercles	1.25 No	1.25 No	1.25 No	1.00 No	1.25 No	1.25 No	1.25 No
no. bands on original tail	10	10	8	N/A	9	8	10
posterior portion of tail white in adults	Yes	Yes	Yes	Yes	Yes	Yes	Yes
immaculate white caudal bands in adults	Yes	Yes	Yes	Yes	Yes	Yes	Yes
SVL	87.3	84.9	94.9	86.6	95.0	67.5	43.4
TL	117	115	109	19B	121	88	115
TW	6.7	6.4	8.3	6.6	7.5	5.3	3.1
FL	15.3	15.1	17.6	15.4	16.7	12.1	7.6
TBL	17.9	17.3	19.7	17.9	19.2	13.4	8.7
AG	42.9	39.7	47.9	40.0	45.2	32.3	19.3
HL	24.8	25.0	26.7	24.5	27.1	19.4	13.5
HW	16.2	16.2	16.8	16.1	17.2	12.4	8.4
HD	8.9	8.7	10.8	8.9	10.3	7.3	5.3
ED	5.5	5.9	5.7	5.5	6.3	4.7	2.9
EE	6.5	6.3	7.1	6.2	7.1	4.8	2.9
ES	11.3	10.6	11.5	10.4	11.8	8.2	5.4
EN	8.5	8.2	8.2	8.1	9.2	6.2	4.0
IO	5.5	5.5	5.4	5.5	5.7	5.2	3.1
EL	2.4	1.7	1.1	1.7	1.7	1.3	1.1
IN	2.6	2.7	2.9	2.3	3.0	2.2	1.3

FIGURE 4. Type series of Cyrtodactylus hidupselamanya sp. nov. from Felda Chiku 7, Kelantan, Peninsular Malaysia. Voucher numbers are in the LSUHC series. LSUHC 12163 is the holotype. Lizards were quite common, generally unwary of our approach, and usually did not try to escape capture. We also noted that only two of 14 specimens (14%) collected had regenerated tails. This would suggest that predation within the cave systems is not high. This is in contrast to other karst ecosystem species that utilize the surrounding limestone vegetation such C. gunungsenyumensis and C. metropolis . Both species are far less numerous, extremely wary and difficult to catch, and have a high frequency of regenerated tails—nearly 100% in the former ( Grismer et al. 2014c ; 2016 a). Comparisons. Cyrtodactylus hidupselamanya sp. nov. is differentiated from all other species of the C. pulchellus complex being that it is the only species with the exception of C. jelawangensis in which the posterior caudal region is whitish in adults. It is further differentiated from all other species by having a combination of weak tuberculation on body; no tubercles on the ventral surfaces of the forelimbs, on the gular region, or in the ventrolateral body folds; 19–23 longitudinal rows of dorsal tubercles; 39–48 paravertebral tubercles; 26–33 ventral scales; 19–24 subdigital lamellae on the fourth toes; no femoral pores but 17–22 precloacal pores; a deep precloacal groove; four body bands; body bands as wide or slightly wider than interspaces; body bands and nuchal loop edged with a thin white line bearing tubercles; no scattered white tubercles on the dorsum; 8–10 dark caudal bands on the original tail separated by immaculate (posteriorly) white caudal bands ( Table 5 ). Within the C. pulchellus complex, C. hidupselamanya sp. nov. is the sister species of C. jelawangensis but separated from it on the basis of having low and rounded as opposed to prominent tubercles; lacking as opposed to having tubercles on the ventral surfaces of the forelimbs; and having fewer rows of longitudinal dorsal tubercles (19–23 versus 23–25) ( Table 5 ). FIGURE 5. Left: limestone forest habitat and karst tower of the type locality of Cyrtodactylus hidupselamanya sp. nov. from Felda Chiku 7, Kelantan, Peninsular Malaysia. Right: Three dimensional Google Earth satellite image view of the karst formation encompassing the type locality illustrating its island-like nature being that it is surrounded by oil palm agriculture. White circles denote approximate collecting sites of the type series. Bottom: edge of the karst tower at the type locality. Remarks. The karst formation in which Cyrtodactylus hidupselamanya sp. nov. occurs is scheduled to be completely quarried for its limestone and other raw materials by ASN Cement Sdn Bhd. From what we currently know about this species, this will result in its extinction.