Cyrtodactylus santana, Chan & Grismer & Santana & Pinto & Loke & Conaboy, 2023

Chan, Kin Onn, Grismer, L. Lee, Santana, Fernando, Pinto, Pedro, Loke, Frances W. & Conaboy, Nathan, 2023, Scratching the surface: a new species of Bent-toed gecko (Squamata, Gekkonidae, Cyrtodactylus) from Timor-Leste of the darmandvillei group marks the potential for future discoveries, ZooKeys 1139, pp. 107-126 : 107

Cyrtodactylus santana sp. nov.

Figs 4 , 5 (Nino Konis Santana Bent-toed Gecko)

Material examined.

Holotype. ZRC 2.7672 (Fig. 4 ), adult male collected by Chan Kin Onn, Iffah Iesa, Fernando Santana, and Pedro Pinto on 30 August 2022 at 2230 hrs from Napana Wei cave (8.411758°S, 127.293321°E; 152 m a.s.l.) in the northeastern sector of NKS . Paratypes. ZRC 2.7673-77 (adult males) and ZRC 2.7678-81 (adult females) with the same collection information as the holotype.

Diagnosis.

The new species is a distinct evolutionary lineage that is closely related to C. batucolus , C. seribuatensis , C. petani , and C. sadleiri . It can be differentiated from other congeners by the following combination of characters: strong dorsal tuberculation present, 23-27 paravertebral tubercles, 15-19 subdigital lamellae on 4th toe, 42-48 ventral scales across midbody, deep precloacal groove absent, enlarged femoral and precloacal scales present, distinct blotches on top of the head absent, dorsal bands faint, whitish, lightly counter-shaded with dark brown.

Description of holotype.

Adult male SVL 68.6 mm; head moderate in length (HL/SVL 0.30), wide (HW/HL 0.65), somewhat flattened (HD/HL 0.40), distinct from neck, triangular in dorsal profile; lores weakly inflated, prefrontal region concave, canthus rostralis smoothly rounded; snout elongate (ES/HL 0.43) rounded in dorsal profile; eye large (ED/HL 0.23); ear opening elliptical, moderate in size (EL/HL 0.11), obliquely oriented; eye to ear distance greater than diameter of eye; rostral wider than high, concave, partially divided dorsally, bordered posteriorly by left and right supranasals and smaller 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 postnasals, ventrally by first supralabial; 10 (R) 10 (L) squarish supralabials extending to just beyond dorsal inflection of labial margins tapering in size abruptly below midpoint of eye, first supralabial largest; nine (R) and eight (L) infralabials tapering smoothly posteriorly slightly beyond last supralabial posteriorly; scales of rostrum and lores raised, larger than granular scales on top of head and occiput; scales of occiput intermixed with slightly enlarged tubercles; dorsal superciliaries elongate, keeled; mental triangular, bordered laterally by first infralabials and posteriorly by left and right rectangular postmentals which contact medially; one row of slightly enlarged, elongate sublabials extending posteriorly to 6th infralabial; gular scales small, granular, grading posteriorly into slightly larger, flatter, throat scales which grade into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.43) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with moderately sized, conical, semi-regularly arranged, keeled tubercles; tubercles extend from occiput to anterior one-third of tail; tubercles on occiput and nape relatively small, increases in size and density posteriorly; tubercles on pelvic region and hindlimbs largest and densest; approximately 15 longitudinal rows of tubercles at midbody; 27 paravertebral tubercles on body; 44 flat, imbricate, ventral scales between ventrolateral body folds, ventral scales much larger than dorsal scales; precloacal scales large, seven scales across base of precloacal region; precloacal depression weak (Fig. 4E ).

Forelimbs moderate in stature, relatively short (ForL/SVL 0.17); granular scales of forearm slightly larger than those of body, interspersed with large, keeled tubercles; palmar scales slightly raised; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded throughout its length; digits slightly more narrow distal to inflection; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TibL/SVL 0.18), covered dorsally by granular scales interspersed with larger, keeled tubercles and covered anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals; ventral tibial scales flat, imbricate; two rows of enlarged, flat, imbricate, femoral scales extend from knee to knee through the precloacal region where they are continuous with enlarged, precloacal scales; posterior row of enlarged femoral scales contains 41 contiguous pore-bearing scales extending from knee to knee forming a V-shape bordering the precloacal depression; postfemoral scales immediately posterior to the row of pore-bearing scales nearly one-half their size, forming an abrupt union on posteroventral margin of thigh; plantar scales low, slightly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae transversely expanded throughout length of digit; digits more narrow distal to joints; 17 subdigital lamellae on right 4th toe, 16 on left; claws well-developed, sheathed by a dorsal and ventral scale.

Tail robust, original, tip broken; dorsal scales at base of tail granular becoming flatter posteriorly; no median row of transversely enlarged, subcaudal scales; subcaudal scales much larger than dorsal caudal scales; one pair of paravertebral and dorsolateral tubercle rows on either side of midline; distance between paravertebral tubercle rows much greater than distance between paravertebral and adjacent dorsolateral rows; caudal tubercles decrease in size posteriorly, extending approximately 40% length of tail; four enlarged, postcloacal tubercles at base of tail on hemipenial swelling; all postcloacal scales flat, imbricate.

Colouration in life.

Dorsal ground colour of head yellowish; neck, trunk, limbs, and tail brown; no distinct markings on top of head; pale loreal stripe extend from nostril to eye and continuing as a postorbital stripe that forms a faint forked pattern on occiput; area dorsal and ventral to the loreal and postorbital stripe counter-shaded with dark brown; six pale, faint, thin, irregular bands from nape to base of tail faintly counter-shaded anteriorly and posteriorly with dark brown; dark speckling and faint, cream-coloured blotches on limbs; pale body banding extend onto tail but not encircling tail (Fig. 5 ). Ventral surfaces of head, body and limbs lightly stippled with grey; subcaudal region darkened with fine mottling; iris greenish brown.

Variation.

ZRC 2.7674-76, ZRC 2.7678, and ZRC 2.7680-81 have broken tails. Some specimens have more distinct dorsal markings than others (Fig. 6 ). The level of yellowness of the head also varies and does not appear to be a sexually dimorphic character. Meristic differences are listed in Table 4 .

Comparisons.

Due to the large number of Cyrtodactylus species, we restrict our comparison to species within the Cyrtodactylus darmandvillei group. The new species differs from C. batucolus by having fewer paravertebral tubercles (23-27 vs. 30-35), more ventral scales (42-48 vs. 38-42), lacking distinct blotches on top of a yellowish head, lacking dark paravertebral dorsal blotches on the body and tail, and having less distinct but well-defined pale-coloured dorsal bands (Fig. 5 ). From C. darmandvillei Weber, 1890, it differs by having more paravertebral tubercles (23-27 vs. 17-20), more ventral scales (42-48 vs. 36-40), and lacking enlarged median subcaudals. From C. jellesmae Boulenger, 1897 it differs by being larger in size (max SVL 74 mm vs. 63 mm) and having as opposed to lacking enlarged femoral and precloacal scales. From C. kimberleyensis Bauer & Doughty, 2012, it differs by being larger in size (max SVL 74 mm vs. 45 mm), having moderate to strong dorsal tubercles (vs. weak to absent), more paravertebral tubercles (23-27 vs. 16-18), more ventral scales (42-48 vs. 36), and having as opposed to lacking enlarged femoral and precloacal scales. From C. petani , it differs by being larger in size (max SVL 74 mm vs. 57.2 mm), having more ventral scales (42-48 vs. 30-35), and more precloaco-femoral pores in males (43-45 vs. 31-35). From C. sadleiri , it differs by being smaller in size (max SVL 74 mm vs. 88 mm), having less subdigital lamellae on 4th toe (15-19 vs. 19-24), more ventral scales (42-48 vs. 34-42), and lacking a deep precloacal groove. From C. seribuatensis , it differs by having fewer subdigital lamellae on 4th toe (15-19 vs. 19-22), more ventral scales (42-48 vs. 32-39), and lacking distinct blotches on top of a yellowish head, lacking dark paravertebral dorsal blotches on the body and tail, and having less distinct but well-defined pale-coloured dorsal bands (Fig. 5 ).

Distribution.

Cyrtodactylus santana sp. nov. occurs in Lene Hara and Napana Wei caves within NKS. The nearest village is Tutuala in the municipality of Lautém. The larger distribution of this species is not yet known but it likely occurs in other limestone caves within NKS. There is a report of a similar-looking and unidentified Cyrtodactylus on Ataúro island ( Kaiser et al. 2013: fig. 3B). However, the specific identity of the Ataúro Cyrtodactylus cannot be ascertained at this point due to the lack of comparative material. As such, we consider the distribution of Cyrtodactylus santana sp. nov. to be restricted to NKS until new data suggest otherwise (see Discussion for more information about the Cyrtodactylus from Ataúro).

Natural history.

Lizards were considerably more abundant in Napana Wei cave compared to Lene Hara cave (Fig. 7 ) even though the two caves are adjacent to each other and are less than 500 m apart but not connected by contiguous limestone. This disparity could be associated with the differences in the geomorphology of both caves. Lene Hara cave is cavernous and dome-like with a high ceiling (Fig. 7 ), whereas Napana Wei is low and narrow. In Lene Hara, a small number of lizards were observed under small rocks and on columnar formations but in Napana Wei, lizards were found in abundance on the underside and exterior surface of the cave wall. No lizards were observed on surrounding vegetation, suggesting that they could be limestone specialists. The caves are located less than 1 km from the coast. Cyrtodactylus santana sp. nov. is nocturnal and is found in sympatry with Gehyra sp.

Etymology.

Nino Konis Santana was a freedom fighter who led the Falintil militia against the Indonesian occupation of Timor-Leste. He was not only a fearless leader of the armed wing of the Resistance but also played a key role in peace initiatives, earning him a reputation as a peacemaker, diplomat, and statesman. The Nino Konis Santana National Park was named in honor of this national hero who was born in the suco (village in Tetum) of Tutuala, located within the boundaries of the park. The specific epithet Cyrtodactylus santana is used as a noun in apposition referring to Nino Konis Santana National Park, which is the type locality of the new species.

Figures

Figure 4. Holotype of Cyrtodactylus santana sp. nov. (ZRC 2.7672) A dorsal view of entire specimen B closeup of side of head C closeup of underside of head D ventral view of entire specimen E ventral view of pelvic region F ventral view of right foot G ventral view of right hand.

Figure 5. Live images of Cyrtodactylus santana sp. nov. paratype female (ZRC 2.7679) (top), C. batucolus from Pulau Besar, Malacca, Peninsular Malaysia (middle), and C. seribuatensis from Pulau Mentigi, Johor, Peninsular Malaysia (lower).

Figure 6. Paratypes of Cyrtodactylus santana sp. nov. and their corresponding voucher numbers.

Figure 7. Type locality of Cyrtodactylus santana sp. nov. at Lene Hara cava as viewed from the exterior (left) and interior (right). Photographs by Tan Heok Hui.