Cyrtodactylus gunungsenyumensis, Grismer, L. Lee, Wood, P. L., Anuar, Shahrul, Davis, H. R., Cobos, A. J. & Murdoch, M. L., 2016

Grismer, L. Lee, Wood, P. L., Anuar, Shahrul, Davis, H. R., Cobos, A. J. & Murdoch, M. L., 2016, A new species of karst forest Bent-toed Gecko (genus Cyrtodactylus Gray) not yet threatened by foreign cement companies and a summary of Peninsular Malaysia’s endemic karst forest herpetofauna and the need for its conservation, Zootaxa 4061 (1) : 6-13

Cyrtodactylus gunungsenyumensis sp. nov.

Gunung Senyum Bent-toed Gecko Cicak Gunung Senyum

Figs 3 , 4

Holotype. Adult female (LSUHC 12209) from Gunung Senyum, Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia (3°41.530’N, 102°26.005’E; 75 m) collected by L. Lee Grismer, Perry L. Wood, Jr., Brandon T. Burch, Anthony J. Cobos, Hayden R. Davis, Shahrul Anuar, and Mathew L. Murdoch on 21 March 2015.

Paratypes. All paratypes (LSUHC 12199, 12201, 12204–06, 12220) bear the same collection data as the holotype.

Diagnosis. Cyrtodactylus gunungsenyumensis sp. nov. can be distinguished from all other Sundaland species by having the following suite of character states: adult SVL 65.1–74.7 mm; low, rounded, weakly keeled, body tubercles; tubercles absent from occiput but present on nape and limbs, and extending posteriorly only onto anteriormost base of tail; 34–40 paravertebral tubercles; weak ventrolateral body fold lacking tubercles; 38–41 ventral scales; no transversely enlarged, median, subcaudal scales; 20–23 subdigital lamellae on fourth toe; abrupt transition between posterior and ventral femoral scales; 31–39 enlarged femoroprecloacal scales; no femoral or precloacal pores; precloacal groove absent; wide, dark postorbital stripes extending from each eye and contacting on nape; no dark, postoccipital chevron; body bearing four or five wide, bold, dark, well-defined bands; light caudal bands wide. The meristic characters are scored across the sworderi complex in Table 5 and can be compared with all other Sundaland species in Grismer et al. (2012: Table 6 ).

Description. Adult female SVL 74.7 mm; head large, moderate in length (HL/SVL 0.27) and width (HW/HL 0.68), somewhat depressed (HD/HL 0.40), distinct from neck, triangular in dorsal profile; lores weakly inflated, prefrontal region concave; canthus rostralis rounded; snout elongate (ES/HL 0.43) and rounded in dorsal profile; eye large (ED/HL 0.23); ear opening triangular and moderate in size (EL/HL 0.1.0); eye-to-ear distance greater than diameter of eye; rostral subrectangular with a deep dorsomedial furrow, bordered posteriorly by large left and right supranasals, one medial postrostral (=internasal); external nares bordered anteriorly by rostral, dorsally by two supranasals (anterior one largest), posteriorly by five postnasals, and ventrally by first supralabial; nine, 10 (R, L) square to rectangular supralabials extending to and tapering gradually below posterior margin of orbit; eight (R, L) infralabials tapering gradually posteriorly to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small and granular; scales on top of occiput not intermixed with tubercles; dorsal and ventral superciliaries rectangular; mental triangular, bordered laterally by first infralabials and posteriorly by left and right rectangular postmentals contacting medially for approximately 30% of their length posterior to mental; one enlarged row of sublabials extending posteriorly to 4th infralabial; gular scales small and raised, grading posteriorly into slightly larger, flatter, throat scales, and thence into large, flat, imbricate pectoral and ventral scales.

Body relatively short (AG/SVL 0.48) with weak, non-tuberculate ventrolateral folds; dorsal scales small and granular, interspersed with larger, low, rounded, semi-regularly arranged, weakly keeled tubercles; tubercles extending from posterior margin of occiput to anteriormost section of tail; tubercles on posterior section of body largest; 35 paravertebral tubercles; 39 flat, imbricate ventrals, ventrals much larger than dorsals; patch of enlarged precloacal scales lacking pores; precloacal groove absent.

Forelimbs moderate in stature, relatively long (FL/SVL 0.16); granular scales of forearm larger than those of body, not interspersed with tubercles; palmar scales flat; digits well-developed, inflected at basal interphalangeal joints; subdigital lamellae slightly enlarged proximal to joint inflections, digits narrower distal to joint inflections; claws well-developed, sheathed by a dorsal and ventral scale.

Hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.19), covered dorsally by granular scales interspersed with larger tubercles and anteriorly by flat, subimbricate scales; ventral scales of femora flat and larger than dorsals; ventral tibial scales flat and imbricate; continuous, single row of flat, enlarged femoral scales extend medially from distal region of one femur to distal region of other femur and are confluent with a series of large, flat, precloacal scales totaling 39 scales; femoral pores absent; dorsal and ventral femoral scales meet abruptly on posteroventral margin of thigh; plantar scales low and slightly rounded; digits well-developed, inflected at basal interphalangeal joints; subdigital lamellae enlarged proximal to inflected joints, digits narrower distal to inflected joints; 22 (R, L) subdigital lamellae on 4th toe; claws well-developed, sheathed by a dorsal and ventral scale.

Tail widest at base, tapering to a point, approximately last four-fifths of tail regenerated; dorsal scales on original and regenerated portion of tail flat; subcaudals flat, larger than dorsals; no median row of transversely enlarged subcaudal scales; caudal scales arranged in semi-whorls; tubercles on base of tail only; base of tail not bearing lateral, bulbous swellings; no enlarged, lateral, postcloacal scales.

Coloration in life ( Fig. 3 ). Ground color of head, body, limbs and anterior portion of tail tan; snout and top of head brown bearing darker, faint mottling; wide, dark-brown, postorbital stripes from each eye extend posteriorly across nape where they merge; faint, dark-brown, transverse line on occiput; four wide, dark-brown, well-defined, body bands with slightly lightened centers extend from forelimb insertion points on body to sacral region and separated by tan interspaces of equal width; two dark caudal bands present on original anterior portion of tail, regenerated portion of tail lacks bands; dorsal surface of limbs bearing slightly darker mottling; ventral surface of head and body immaculate off-white except for fine black stippling on gular and lateral ventral scales; ventral surfaces of limbs slightly darker with denser stippling on scales; subcaudal region slightly darker than ventral surfaces of limbs and faintly mottled.

Variation. The paratypes closely resemble the holotype in overall coloration and pattern ( Fig. 4 ). The banding pattern of the juvenile LSUHC 12204 (SVL 43.7 mm) is much more bold than that of the adults (SVL> 65.1 mm). A n transition to the more faded adult pattern is observable in the subadult LSUHC 12201 (SVL 51.6 mm). LSUHC 121999 and 12204 have five dorsal body bands whereas all the other specimens have four. Specimens LSUHC 12200–01 are the only specimens with original tails which are distinctly banded. The color pattern of the adult LSUHC 12206 is considerably more faded than that of the other adults. LSUHC 12199 and 12204 lack tails. Meristic variation is presented in Table 4 .

Distribution. Currently, Cyrtodactylus gunungsenyumensis sp. nov. is known only from the type locality of Gunung Senyum, Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia ( Fig. 1 ). Exploration of the adjacent karst towers of Gunung Jebak Puyoh and Bukit Terus ( Fig. 5 ) to determine if this species occurs there as well is necessary.

Natural history. Hutan Lipur Gunung Senyum is a recreational area situated around three limestone hills: Gunung Senyum, Gunung Jebak Puyoh, and Bukit Terus ( Fig. 5 ). The northernmost hill, Gunung Senyum, is the largest, reaching 525 m in elevation and covering 1.75 km 2. Several large, open chambers and cave systems deeply incise and sculpt the periphery of this limestone hill, which is surrounded by karst forest vegetation. All specimens of the type series and two additional specimens not collected were active at night on the limestone walls or the adjacent karst vegetation ( Fig. 3 ). Lizards were observed at night between 0 1030 and 2400 hrs on the vertical trunks of small trees as well as on thin, horizontal branches of low-growing shrubs. Lizards only occurred in areas where karst rock-rubble had accumulated at the edge of the cliff faces and where the cliff faces were eroded and exfoliated, providing cracks and holes into which lizards were able to take refuge. This species is very wary and lizards would often flee during our approach. When captured, most quickly dropped their tails and only two specimens of the nine lizards observed had complete tails. This suggests this population may be under heavy predation. No gravid females or hatchlings were observed, indicating that the breeding season is not in March.

Etymology. The specific epithet gunungsenyumensis refers to the type locality of Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia. It is hoped this name will underscore the importance of this Hutan Lipur as a sanctuary for this endemic species and keep it safe from the quarrying interests of foreign cement companies.

Comparisons. Within the Cyrtodactylus sworderi complex, C. gunungsenyumensis sp. nov. most closely resembles the other karst forest species C. guakanthanensis ( Fig. 6 ) even though they are not each others closest relatives ( Fig. 2 ). Cyrtodactylus gunungsenyumensis is differentiated from C. guakanthanensis by having 34–40 as opposed to 32–34 paravertebral tubercles and rounded as opposed to conical dorsal tubercles on the body. These two species are very similar in dorsal banding and color pattern—the only consistent difference being that there is a dark, chevron-shaped nuchal blotch just posterior to the occiput in C. guakanthanensis that is lacking in C. gunungsenyumensis sp. nov. This similarity in pattern—which is quite divergent from the other species in the sworderi complex ( Fig. 6 )—is likely to have evolved in parallel ( Fig. 2 ) and is an adaptation to a karst forestdwelling life style. Similar banding patterns are seen in all other unrelated karst dwelling Cyrtodcatylus in Peninsular Malaysia ( Fig. 6 ) and many of the other 35 karst dwelling species throughout Indochina ( Table 6 ).

From its closest relative C. tebuensis , C. gunungsenyumensis sp. nov. differs in having low, rounded and weakly keeled dorsal tubercles as opposed to tubercles that are large, conical, and strongly keeled. Additionally, C. gunungsenyumensis sp. nov. differs in having 38–41 as opposed to 43–51 ventral scales; 20–23 as opposed to 17– 21 subdigital lamellae on the 4th toe; wide as opposed to narrow, light-colored caudal bands; and a banded as opposed to a striped or spotted dorsal pattern ( Fig. 6 ). Differences between C. gunungsenyumensis sp. nov. and C. sworderi and C. quadrivirgatus are highlighted in Table 5 .

Figures

FIGURE 1. Distribution of Cyrtodactylus gunungsenyumensis sp. nov., C. tebuensis, C. guakanthanensis, and C. sworderi of the sworderi complex. Stars indicate type localities.

FIGURE 2. Inferred phylogenetic relationships within the sworderi complex. The phylogram is a Maximum Likelihood topology with Bayesian posterior probabilities and Maximum Likelihood bootstrap values, respectively. The green circles indicate instances of independent evolution of karst ecosystem ecology.

FIGURE 3. Left: Cyrtodactylus gunungsenyumensis sp. nov. Upper left: LSUHC 12200 subadult female; middle left: 12201 juvenile; lower left; LSUHC 12209 adult female. Right: Microhabitat of C. gunungsenyumensis sp. nov. at Gunung Senyum, Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia.

FIGURE 4. Type series of Cyrtodactylus gunungsenyumensis sp. nov. from Gunung Senyum, Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia.

FIGURE 5. Google Earth photograph of Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia.

FIGURE 6. Upper left: Cyrtodactylus quadrivirgatus (LSUDPC 2509) from Fraser’s Hill, Pahang, Peninsular Malaysia. Lower left: C. guakanthanensis (LSUDPC 8072) from Gua Kanthan, Perak, Peninsular Malaysia. Upper right: C. tebuensis (LSUDPC 7990) from Gunung Tebu, Terengganu, Peninsular Malaysia. Lower right: C. sworderi (LSUDPC 5875) from Gunung Panti, Johor, Peninsular Malaysia.

Tables

TABLE 5. Diagnostic characters (in bold) separating Cyrtodactylus gunungsenymensis sp. nov. from all other species in the sworderi complex.

	gunungsenyumensis sp. guakanthanensis tebuensis quadrivirgatus nov.	sworderi
supralabials	9,10 9,10 11–13 8–11	10–13
infralabials	7–9 7,8 8–10 8–11	10
tubercles in occiput	no yes yes yes	yes
body tubercles paravertebral tubercles	low, rounded, weakly conical, weakly keeled large, conical, large, conical, keeled keeled keeled 34–40 32–34 32–39 32–39	large, conical, keeled 36–38
ventral scales	38–41 37–44 43–51 28–40	40–49
Postfemoral/ventral scale contact abrupt	yes yes yes no	no
4th toe lamellae	20–23 19–21 17–21 18–23	16–20
precloacal pores	absent absent absent 0–12	5–9
precloacal pores in females	absent absent absent absent	y
enlarged precloacal scales	yes yes yes yes	few
no. enlarged femorocloacal scales	31–39 36–41 31–37 31–44	0
wide, dark postorbital stripes contacing on nape	yes yes no no	no
dark, postoccipital cheveron	no yes no no	no
light caudal band width	wide wide narrow narrow	wide
dorsal pattern	banded banded striped/spotted variable spotted	spotted
maximum SVL	74.7 82.2 79.1 71	80.1

TABLE 4. Scale counts, color pattern, and measurements of the type series of Cyrtodactylus unungsenyumensis sp. nov. from Hutan Lipur Gunung Senyum, Pahang, Malaysia

	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC	LSUHC
	12209	12199	12200	12201	12204	12205	12206
	holotype	paratype	paratype	paratype	paratype	paratype	paratype
sex	female	male	female	female	/	male	male
supralabials	9	10	9	10	9	10	10
infralabials	8	9	7	8	7	7	8
paravertebral tubercles	35	34	38	40	37	36	39
ventral scales	39	39	39	41	41	38	39
postfemoral scale contact abrupt	yes	yes	yes	yes	yes	yes	yes
4th toe lamellae	22	20	22	20	23	21	20
precloacal pores	absent	absent	absent	absent	absent	absent	absent
precloacal pores in females	/	/	/	/	/	/	/
enlarged precloacal scales	y	y	y	y	y	y	y
no. enlarged femorocloacal scales	39	35	31	35	34	34	39
light caudal bands narrow (1) or wide (0)	0	/	0	0	/	0	0
dorsal pattern	banded	banded	banded	banded	banded	banded	banded
SVL	74.7	72.8	65.1	51.6	43.7	74.7	74.3
TL		/
TW	6.1	6.4	5.1	3.8	2.1	5.6	5.6
FL	11.7	11.7	11.1	8.5	5.2	11.5	10.1
TBL	14.2	13.6	12.4	10.0	7.2	13.4	13.1
AG	36.1	32.8	29	22.6	19.8	32.4	32.4
HL	20.1	19.8	18.5	15.3	11.6	19.3	19.9
HW	13.7	12.8	11.7	9.3	7.2	12.1	11.7
HD	8.0	7.3	7.0	5.6	3.7	7.0	7.1
ED	4.7	4.7	4.4	3.7	2.0	3.9	3.7
EE	6.3	5.7	5.4	4.1	2.4	4.3	4.3
ES	8.6	8.8	7.9	6.9	4.4	8.0	8.1
EN	7.1	6.6	5.9	5.0	3.0	5.8	5.9
IO	3.6	3.5	3.4	3.1	1.6	2.5	2.7
EL	2.1	2.2	2.0	1.2	0.1	0.4	0.4
IN	2.2	2.2	2.0	1.7	0.2	0.9	0.8