Cyrtodactylus sungaiupe, Termprayoon & Rujirawan & Grismer & Wood Jr & Aowphol, 2023

Termprayoon, Korkhwan, Rujirawan, Attapol, Grismer, L. Lee, Wood Jr, Perry L. & Aowphol, Anchalee, 2023, Two new karst-adapted species in the Cyrtodactylus pulchellus group (Reptilia, Gekkonidae) from southern Thailand, ZooKeys 1179, pp. 313-352 : 313

Cyrtodactylus sungaiupe sp. nov.

Figs 5 , 6 , 7 Thung Wa Bent-toed Gecko

Type material.

Holotype. Adult male (ZMKU R 01009, Figs 5 , 6 ) collected from Thailand, Satun Province, Thung Wa District, Thung Wa Subdistrict (7°05.865'N, 99°58.506'E; 58 m a.s.l.), on 29 April 2022 by Korkhwan Termprayoon, Akrachai Aksornneam, Attapol Rujirawan, and Siriporn Yodthong.

Paratypes. Two adult males (ZMKU R 01004-01005) and two adult females (ZMKU R 01007-01008), same data as holotype, except collected on 12 May 2019, by Korkhwan Termprayoon, Attapol Rujirawan, Natee Ampai, Piyawan Puanprapai and Siriporn Yodthong. One adult male (ZMKU R 01010) and two adult females (ZMKU R 01011-01012), same data as holotype.

Referred specimens.

ZMKU R 01006 (immature male), same data as holotype, except collected on 12 May 2019, by Korkhwan Termprayoon, Attapol Rujirawan, Natee Ampai, Piyawan Puanprapai and Siriporn Yodthong. ZMKU R 01013-01014 (two juveniles), same data as holotype.

Diagnosis.

Cyrtodactylus sungaiupe sp. nov. can be distinguished from all other species of the C. pulchellus group by a combination of the following characters: (1) SVL 89.7-102.7 mm in adult males (n = 4), 87.3-104.6 mm in adult females (n = 4); (2) 12-16 supralabial and 10-13 infralabial scales; (3) weak tuberculation on body; (4) no tubercles on ventral surfaces of forelimbs, gular region or in ventrolateral body folds; (5) 30-38 paravertebral tubercles; (6) 19-22 longitudinal rows of dorsal tubercles; (7) 32-40 rows of ventral scales; (8) 20-24 subdigital lamellae on the fourth toe; (9) 29-34 femoroprecloacal pores in adult males; (10) absence of precloacal pores in adult females; (11) deep precloacal groove in males; (12) absence of scattered pattern of white tubercles on dorsum; (13) four dark dorsal body bands; (14) nine or twelve dark caudal bands on original tail; (15) light caudal bands in adults infused with dark pigmentation; (16) caudal tubercles extended 1/8-1/10 of anterior portion of tail and (17) posterior portion of tail in hatchlings and juveniles white.

Description of holotype.

Adult male SVL; 89.7 mm; head moderate in length (HL/SVL 0.29) and wide (HW/HL 0.65), flattened (HD/HL 0.37), distinct from neck and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions concave; canthus rostralis rounded anteriorly; snout elongated (ES/HL 0.40), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.22); ear opening elliptical, moderate in size (EL/HL 0.07), obliquely orientated; eye to ear distance slightly greater than diameter of eye; rostral rectangular, divided dorsally by an inverted Y-shaped furrow, bordered posteriorly by left and right supranasals and internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal, posteriorly by two postnasals, ventrally by first supralabial; 9/10 (left/right) rectangular supralabials extending to below mid-point of eye, 13/15 to below the posterior margin of the eye-ball, decreasing abruptly just posterior to mid-point of eye; 8/7 infralabials extending to below mid-point of eye, 13/11 to upturn the labial margin, decreasing gradually in size posteriorly; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales on top of head and occiput intermixed with rounded, small tubercles; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, 2.5 mm in width, 3.3 mm in length, bordered laterally by first infralabials and posteriorly by left and right, trapezoidal postmentals which contact medially for approximately 50% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to the sixth (left) and seventh (right) infralabials; small, granular, 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, weakly-keeled tubercles, smaller intervening tubercles occasionally present; tubercles extend from occiput to base of tail, but not further than 1/10 of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 19 longitudinal rows of tubercles at mid-body; 36 paravertebral tubercles; 37 flat imbricate ventral scales between ventrolateral body folds; ventral scales larger than dorsal scales; precloacal scales large, smooth; deep precloacal groove.

Forelimbs moderately slender, relatively short (FL/SVL 0.17); dorsal scales on forelimbs raised, granular, larger than those on body; dorsal scales on forearm intermixed with enlarged, subconical and weakly-keeled tubercles, brachium without tubercles; scales of ventral surface of forearm flat, subimbricate, tubercles absent; palmar scales small, weakly rounded; digits well-developed, inflected at basal, interphalangeal joints; 19/19 (left/right) subdigital lamellae on the fourth finger, 6/6 proximal subdigital lamellae rectangular, broadly expanded proximal to joint inflection, 13/13 distal subdigital lamellae slightly expanded distal to inflection becoming gradually more expanded near the claw; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.20), enlarged, subconical, weakly-keeled tubercles on dorsal surface of legs separated by smaller juxtaposed scales; ventral scales of thigh flat, smooth, imbricate, larger than dorsal granular scales; ventral, tibial scales flat, smooth, imbricate; a single row of 35 enlarged femoroprecloacal scales extending nearly from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 33 contiguous pore-bearing femoroprecloacal scales, forming an inverted T bearing a deep, precloacal groove (Fig. 6D ); six pore-bearing scales bordering groove (three on each side); 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 weakly rounded to flat; 20/20 (left/right) subdigital lamellae on fourth toe, 7/7 proximal subdigital lamellae rectangular, broadly expanded proximal to joint inflection, 13/13 distal subdigital lamellae slightly expanded distal to inflection becoming gradually more expanded near the claw; claws well-developed, sheathed by a dorsal and ventral scale.

Original tail 122.5 mm in length, slightly longer than SVL (TL/SVL = 1.37), 6.4 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; original portion segmented, approximately 7-8 transverse scales rows per segment; one transverse row of four dorsal tubercles on posterior margin of 1st segment and one tubercle on 2nd segment; caudal tubercles extended 1/10 of anterior portion of original tail (Fig. 6E ); subcaudal region bearing large median row of transverse scales; shallow dorsal and lateral caudal furrow; base of tail bearing hemipenial swellings; two rows of (1+4)L/(2+3)R medium-sized postcloacal tubercles on each hemipenial swelling; postcloacal scales smooth, flat, large, imbricate.

Colouration in life

(Fig. 5 ). Ground colour of head, body, and limbs light-brown; superciliaries yellow anteriorly and posteriorly; supralabial and infralabial scales light-brown with off-white markings posteriorly; wide, dark-brown nuchal band edged anteriorly and posteriorly by thin, yellowish lines bearing tubercles extending from posterior margin of one eye to posterior margin of another eye; four similar dark-brown body bands between nuchal loop and hind limb insertions edged anteriorly and posteriorly by broken, thin, creamy-white to yellow lines bearing tubercles, first band terminating at shoulders, second and third bands terminating just dorsal to ventrolateral folds, the fourth band terminating at femurs; dark body bands slightly larger than light-coloured interspaces; one additional dark-brown band posterior to hind limbs (postsacral band); ventral surfaces of head, abdomen and limbs greyish-white; creamy pale yellow postcloacal tubercles; tail bearing nine dark bands separated by ten light-brown (anteriorly) to white (posteriorly) bands, white caudal band infused with dark pigmentation; subcaudal region off-white anteriorly, becoming darker posteriorly.

Colouration in preservative

(Fig. 6 ). The overall colour pattern of head, body, limbs and tail similar to that in life with some fading. Ground colour of head, body, limbs and dorsum tan; dark bands on dorsum and tail brown; yellow-coloured tuberculation on dorsum fading to off-white or white; light-beige coloured on the ventral surface.

Variation.

Meristic and morphometric data for the type series and referred specimens of Cyrtodactylus sungaiupe sp. nov. are given in Tables 6 , 7 and Suppl. material 3. All paratypes and referred specimens resemble the holotype in general aspects of morphology with variations in colouration and banding pattern. All specimens have continuous dark body bands, except one specimen (ZMKU R 01010) where the second to fourth dorsal bands do not connect on the mid-line and are offset (Fig. 7A ). Dorsal scale on brachium of two specimens (ZMKU R 01010 and ZMKU R 01012) intermixed with tubercles, whereas absent in the holotype and other specimens. ZMKU R 01005 has dark marking on left femur. Three male paratypes have discontinuous pore-bearing femoroprecloacal scales; ZMKU R 01004 has three poreless scales (one on the left and two on the right), ZMKU R 01005 has one enlarged and one small poreless scales on the left and ZMKU R 01010 has two poreless scales (one on each side). Last row of caudal tubercles of paratypes (ZMKU R 01006 and ZMKU R 01008) extending to 4th-5th segment of original portion (to 2nd light caudal band), approximately 1/8 of anterior portion of original tail. Juveniles (ZMKU R 01013-01014) have body pattern similar to adults, but less prominent tuberculation, light-yellow ground colour of body, edged anteriorly and posteriorly by yellowish lines bearing tubercles, the original tail has approximately six or ten dark caudal bands, the posterior portion of tail is white (Fig. 7B ).

Distribution.

Cyrtodactylus sungaiupe sp. nov. is currently known from an unnamed karst formation in Thung Wa Subdistrict, Thung Wa District, Satun Province, Thailand (Figs 1 , 8 ).

Natural history.

All individuals of Cyrtodactylus sungaiupe sp. nov. were collected from karst forest (Fig. 8 ) at the type locality which was surrounded by rubber plantations. Specimens were collected on October 2016 between 1900 and 2000 h with temperature 26.3 °C and relative humidity 90.3%, and on April 2022 between 1900 and 2030 h with temperature 28.3 °C and relative humidity 86.0%. The specimens were found on small rocks, karst boulders, adjacent vegetation and on the forest floor. The holotype (ZMKU R 01009) was collected from the branch of a shrub approximately 1.3 m above the ground. Four specimens (ZMKU R 01004, ZMKU R 01006, ZMKU R 01010, ZMKU R 01013) were found on karst boulders. Three female specimens (ZMKU R 01008, ZMKU R 01011-01012) were collected from tree trunks. One adult male (ZMKU R 01005) was collected off a fallen dry stick and one adult female (ZMKU R 01007) was found on the forest floor covered with leaf litter. A juvenile (ZMKU R 01014) was taken from a perch on a small rock near the ground.

A gravid female (ZMKU R 01008) carrying two eggs (externally visible) was found on a tree trunk in October 2016. Two juveniles (ZMKU R 01013-01014) were collected during April 2022. Other lizard species found in this area, include Gehyra mutilata (Wiegmann, 1834) and Gekko gecko (Linnaeus, 1758).

Etymology.

The specific epithet sungaiupe is derived from the old name of Thung Wa District (Sungai Upe District), the type locality of the new species.

Comparison.

Cyrtodactylus sungaiupe sp. nov. can be distinguished from other species in the C. pulchellus group by having a combination of weak tuberculation on the body; no tubercles on ventral surface of forelimbs, gular region or in ventrolateral body folds; 12-16 supralabial scales; 30-38 paravertebral tubercles; 19-22 longitudinal tubercle rows; 32-40 ventral scales; 29-34 femorprecloacal pores in males; deep precloacal groove in males; nine or twelve dark caudal bands on original tail; light caudal bands on original tail, infused with dark pigmentation in adults; caudal tubercles extended 1/8-1/10 of anterior portion of tail; and juveniles with white tail tip. Additional comparisons between Cyrtodactylus sungaiupe sp. nov. and other species in the C. pulchellus group are in Suppl. material 4.

Cyrtodactylus sungaiupe sp. nov. is a member of Clade A which comprises C. astrum , C. dayangbuntingensis , C. langkawiensis , C. lekaguli and C. stellatus . Cyrtodactylus sungaiupe sp. nov. differs from those five species by uncorrected pairwise distances of ND2 of 8.02-10.69% (Table 2 ). It differs from C. astrum by having smaller maximum SVL of 104.6 mm (vs. 108.3 mm); absence of scattered pattern of white tubercles on dorsum (vs. present); nine or twelve dark caudal bands on the original tail (vs. 13 or 14); caudal tubercles extending between 1/8 and 1/10 of anterior portion of tail (vs. ≥ 1/3 of the tail); and having statistically significant different mean values of mensural characters of FLadj, TBLadj, AGadj, HWadj, HDadj, EEadj, ESadj, ENadj and INadj (p <0.001-0.043; Table 5 ). It can be further separated from C. astrum in having statistically significant different mean values of meristic characters of SL, IL, PVT and LRT (p <0.001-0.011; Table 5 ).

Cyrtodactylus sungaiupe sp. nov. differs from C. dayangbuntingensis by having larger maximum SVL of 104.6 mm (vs. 99.0 mm); absence of scattered pattern of white tubercles on dorsum (vs. present); caudal tubercles extending between 1/8 and 1/10 of anterior portion of tail (vs. 1/5 of the tail); and one to two rows of postcloacal tubercles (vs. up to three rows). Additionally, PCA showed that Cyrtodactylus sungaiupe sp. nov. is clearly separated in morphospace from C. dayangbuntingensis (Fig. 3 ).

Cyrtodactylus sungaiupe sp. nov. morphologically differ from C. langkawiensis by having larger maximum SVL of 104.6 mm (vs. 99.8 mm); and caudal tubercles extending between 1/8 and 1/10 of anterior portion of tail (vs. ≥ 1/3 of the tail); and having statistically significant different mean values of mensural characters of AGadj, HLadj, HWadj and INadj (p <0.001-0.033; Table 5 ). It can be further separated from C. langkawiensis in having statistically significant different mean values of meristic characters of SL, IL, PVT, LRT and VS (p <0.001-0.046; Table 5 ).

Cyrtodactylus sungaiupe sp. nov. differ from C. lekaguli by having smaller maximum SVL of 104.6 mm (vs. 108.5 mm); and caudal tubercles extending between 1/8 and 1/10 of anterior portion of tail (vs. ≥ 1/3 of the tail); and having statistically significant different mean values of mensural characters of HWadj, HDadj, EEadj, IOadj and ELadj (p <0.001-0.008; Table 5 ). It can be further separated from C. lekaguli in having statistically significant different mean values of meristic characters of SL, IL and LRT (p <0.001-0.006; Table 5 ).

Cyrtodactylus sungaiupe sp. nov. differ from C. stellatus by having larger maximum SVL of 104.6 mm (vs. 96.1 mm); absence of scattered pattern of white tubercles on dorsum (vs. present); absence of precloacal in female (vs. present); and having statistically significant different mean values of mensural characters of SVL, FLadj, TBLadj, AGadj, HLadj, HWadj, HDadj, EEadj, ESadj and ENadj (p <0.001-0.035; Table 5 ). It can be further separated from C. stellatus in having statistically significant different mean values of meristic characters of PVT (p = 0.003; Table 5 ).

Figures

Figure 1. Map showing the type localities of Cyrtodactylus sungaiupe sp. nov. (red star) in Thung Wa District and Cyrtodactylus wangkhramensis sp. nov. (blue star) in La-ngu District, Satun Province, Thailand and the type localities of closely-related species, C. astrum, C. dayangbuntingensis, C. langkawiensis, C. lekaguli and C. stellatus. Circles represent additional localities of specimens used in molecular analyses.

Figure 3. Principal Component Analysis (PCA) of Cyrtodactylus sungaiupe sp. nov., Cyrtodactylus wangkhramensis sp. nov. and the closely-related species in Clade A of the C. pulchellus group. The letters in the scatter plots refer to holotype (= H), paratype (= P) and topotype (= T).

Figure 5. Adult male holotype of Cyrtodactylus sungaiupe sp. nov. (ZMKU R 01009) from the type locality in Thung Wa Subdistrict, Thung Wa District, Satun Province, Thailand A dorsolateral view of specimen in life B dorsal and ventral views immediately after euthanasia.

Figure 6. Adult male holotype of Cyrtodactylus sungaiupe sp. nov. (ZMKU R 01009) in preservation A dorsal and ventral views of body B head dimensions showing lateral, dorsal and ventral views C ventrolateral fold on the left side D precloacal depression and pore-bearing femoroprecloacal scales E dorsal view of tail showing last row of tubercles.

Figure 7. Variation in colouration and banding pattern of Cyrtodactylus sungaiupe sp. nov. immediately after euthanasia A male paratype (ZMKU R 01010) showing disconnected bands on dorsal B juvenile (ZMKU R 01013) having light-yellow on the body and bearing white tail tip.

Figure 8. Habitat of Cyrtodactylus sungaiupe sp. nov. at the type locality in Thung Wa Subdistrict, Thung Wa District, Satun Province, Thailand A landscape view of karst tower B vegetation structure and C karst boulders surrounding karst body D immature female (not collected) on karst boulder.