Cnemaspis puterisantubongae, Kurita & Nishikawa & Hossman & Mizuno & Sato & Gumal, 2024

Kurita, Takaki, Nishikawa, Kanto, Hossman, Mohamad Yazid, Mizuno, Takafumi, Sato, Hirotoshi & Gumal, Melvin, 2024, Description of a new sandstone-dwelling species of genus Cnemaspis (Sauria: Gekkonidae) from Gunung Santubong National Park, southwestern Sarawak, Malaysia, Zootaxa 5468 (2), pp. 361-378 : 365-375

Cnemaspis puterisantubongae sp. nov. Kurita & Nishikawa

English: Santubong rock gecko

Malay: Cicak puteri Santubong

Holotype. Adult male, SFC-GRB-00517 (field no.: KUHE Swk23-186) collected on September 24, 2023, by T. Kurita and K. Nishikawa from Gunung (= Mt.) Santubong, Kuching, Kuching District, Sarawak, Malaysia (1.75791°N, 110.32685°E, ca. 250 m a.s.l.).

Paratype. Five adult males and three adult or young females (SFC-GRB-00515, 00516, 00518, 00519, SFC-GRB-00528–00531; field nos. KUHE Swk23-184, 185, 187, 188, 197–200, respectively) collected on September 24–25, 2023, by T. Kurita and K. Nishikawa from the type locality .

Diagnosis. Cnemaspis puterisantubongae differs from all other species in the unique combination of the following characters:SVL up to 66.7 mm; 9–12 supralabial scales; 9–12 infralabials; 0–1 internasal; 4–7 postmentals; head, ventral, forearm, tibial, subcaudal, and dorsal tubercles keeled; 29–34 paravertebral tubercles between limb insertions, not linearly arranged; few or no tubercles on ventrolateral surface of trunk; chevron arranged 10–14 precloacal pore scales in adult males and 9–15 pore-bearing scales in adult females, separated to right and left at midline by 1–3 poreless scales; no femoral pores; 1–3 postcloacal tubercles on each side; enlarged submetatarsal scales beneath first toe; 27–33 subdigital lamellae on fourth toe; tubercles present within lateral caudal furrow, at least anterior part; ventrolateral caudal tubercles present anteriorly; single enlarged, smooth median row of subcaudals; yellow ocher head, trunk, and limb dorsum in both sexes; distinct paravertebral black blotches; faint yellow dotted transverse markings on flanks; black and white banding pattern on original tail.

Description of holotype ( Fig. 3 ). Adult male; SVL 60.9 mm; head moderate in size (head length/SVL = 25.9%), oblong in dorsal profile (head width/head length = 66.7%), flat (head depth/head length = 42.2%), distinct from neck (nape width/head width = 62.9%); snout short (snout length/head length = 43.0%), beak concave at middle between snout tip and anterior edges of eyes in lateral profile (beak depth/head depth = 73.7%), raised at middorsal region, dented posterior to nostril; head slightly concave middorsally and convex at orbital edges; canthus rostralis smoothly rounded; nostrils round, open posterodorsally; scales of rostrum juxtaposed, conical, weakly keeled; occipital area covered with conical scales smaller than those of rostrum, intermixed with larger conical scales posterior to the level of anterior edge of ear opening; eye moderate in size (eye diameter/head length = 16.5%); eyeball located about three granular scales inward from lateral margin of snout and not visible from the ventral side; pupil round; strongly prominent, pointed extrabrillar fringe scales from anterior to dorsal edge of eye, largest anterior; ear opening oval, oriented dorsoventrally; rostral rectangular, wider than high in dorsal and lateral views, high enough to cover entire snout tip, bordered by first supralabials, two large supranasals, and nostrils; nostrils bordered by rostral, first supralabials, supranasals, and six/seven postnasals; medial longitudinal groove of rostral reaches two thirds of height on dorsal edge; 11/12 supralabials and 10/10 infralabials, gradually decreasing in size posteriorly; mental large, fan-shaped with U-shaped posterior edge extending to the level of middle of third infralabials, bordered posteriorly by six postmentals; postmentals bordered by first infralabials elongate anteroposteriorly, larger than the others; gular scales juxtaposed, slightly keeled, raised, oval, gradually decreasing in size posteriorly; throat scales larger than gular scales, slightly imbricate, keeled, weakly raised, oval.

Body slender, elongate (axilla–groin length/SVL = 47.3%); small, smooth or weakly keeled granular scales throughout trunk dorsum, intermixed with keeled conical tubercles more or less randomly arranged, infilled with a few small amorphous scales between granular scales and tubercles; one long median keel on anterior part of dorsal tubercle, with an additional pair of dorsolateral short keels on larger tubercles; tubercles dense dorsally, decreasing in number laterally, absent on ventrum; 31/33 paravertebral tubercles; abdominal scales square, flat, keeled, slightly imbricate anteriorly, stretching longitudinally, slightly raised, strongly keeled, imbricate posteriorly; precloacal scales oval, raised, keeled, juxtaposed; six/seven precloacal pores or pore-bearing scales forming chevron arrangement, medially separated by two keeled, poreless scales; precloacal depression absent; no femoral pores; postcloacal scales oval, raised, keeled, slightly imbricate; five/five enlarged, a pair of postcloacal tubercles on lateral surfaces of hemipenal swellings at base of tail.

Brachium and antebrachium long (brachial length/SVL = 13.6%, antebrachial length/SVL = 17.0%); forearms covered with conical, keeled, slightly imbricate dorsal scales and low-conical, keeled, juxtaposed ventral scales; palmar scales conical, weakly keeled, slightly enlarged on wrist; fingers long with an inflected joint, increasing in length from first to fourth, with fifth slightly shorter than fourth; interdigital webbing weak; subdigital lamellae smooth, almost entirely wide and saddle-shaped, unnotched; lamellae on joint between second and third distal phalanges (base of digit in first finger) distinctively enlarged; subdigital lamellae from first to fifth fingers 15, 20, 26, 27, 20/16, 20, 26, 27, 20; claws recurved, bordered by a smooth dorsal scale and distalmost subdigital lamella; hind limbs longer (femur length/SVL = 21.6%, crus length/SVL = 20.0%), more robust than forelimbs; hind limbs covered with conical, keeled, juxtaposed dorsal scales; ventral scales on hind limbs weakly raised, keeled, slightly imbricate proximally, gradually increasing in size, degree of keel distally; plantar scales round, raised, smooth, slightly imbricate; slightly-enlarged flattened smooth metatarsal scales beneath first metatarsal; toes long with an inflected joint, increasing in length from first to fourth with fifth slightly shorter than fourth; subdigital lamellae similar to fingers; toe lamellae from first to fifth digit 13, 19, 26, 29, 24/12, 20, 25, 27, 24; claws recurved, bordered by a smooth dorsal scale and distalmost lamella.

Tail original, long (tail length/SVL = 121.7%), segmented; shallow contiguous middorsal caudal furrow from first to ninth segments; lateral caudal furrow continues from first to 14th segments; caudal scales arranged in segmented whorls; dorsocaudal scales round, weakly raised, smooth or keeled, slightly imbricate; ventrocaudal scales oval, strongly keeled, slightly imbricate, larger and more flattened than dorsocaudals; single median row of enlarged, smooth, imbricate, subcaudal scales, with 4–5 scales per caudal segment in anterior part of tail; conical enlarged caudal tubercles with strong keel arranged on paravertebral, dorsolateral, lateral (in lateral furrow), and ventrolateral rows, gradually decreasing in size posteriorly; 1–2 ventrolateral tubercles per caudal segment on basal, one per segment on distal; lateral tubercles in lateral caudal furrows on basal segments; paravertebral tubercles largest, decreasing in size towards venter; tubercles not encircling tail.

In life, head yellow ocher, with scattered faint yellow and black spots posterior to the level of anterior eye edge; black longitudinal stripes extending from nostrils to middorsal forehead; eight black lines radially extending from eye, of which two on anterior eye edge reach supralabials, three on dorsal edge short, other three on posterior edge stretching to nape, anterior part of lateral trunk, and anterior edge of forelimb insertion; supralabials yellowish gray with ocher blob; infralabials yellowish gray; iris brownish yellow and black; nape yellow ocher dorsally, gray laterally with faint yellow spots; trunk dorsum yellow ocher, with yellowish gray region around forelimb insertions; vertebral region of trunk faint yellowish gray; eight faint yellow blotches along vertebrae from nape to base of tail; eight clear black dorsal blotches from nape to base of tail, of which first is on middorsal and others are separated to right and left along faint vertebral region; two black dorsal blotches on shoulder region separated paravertebrally and dorsolaterally; dorsal surface of forelimbs, hind limbs, and digits yellow ocher with black and faint blown transversally elongated blotches; four faint yellow dotted lines on lateral sides of trunk; abdominal surface of head, trunk, and limbs gray, with slightly darker stain on head ventrum; yellow spots on marginal regions of head, trunk, and limbs in abdominal view; seven poorly-demarcated black and seven maculate white bands encircling or forming saddles on tail; tail with black tip; black bands approximately twice as long as adjacent white bands; enlarged median subcaudal scales same colouration as caudal bands on 10 proximal caudal segments, immaculate white on subsequent segments.

Variation. Male paratypes are highly similar to the holotype in colouration and pattern. Compared with males, female paratypes have pore-bearing precloacal scales, obviously convex at centre (versus precloacal pores), clear black and white bands encircling original tail (versus white midventral and adjacent scales on posterior part of tail), and darker body colouration ( Fig. 4 ). Tails regenerated from tip are dorsally black and ventrally white in males, uniformly dark gray in females; those regenerated from the middle are dorsally black and ventrally yellow in males, unknown in females. Scales on regenerated tail are small, raised, keeled, slightly imbricate, and equal in size. Scalation and other meristic characters are described in Table 2 .

Comparisons ( Table 3 ). Cnemaspis puterisantubongae sp. nov. is differentiated from all other Southeast Asian congeners except C. leucura , C. matahari , and probably C. sirehensis in having an ocher yellow dorsal surface of the head, trunk, and limbs and in black and white banding/saddling of the posterior part of original tail; the regenerated tail has a yellow subcaudal portion in males. Cnemaspis puterisantubongae is distinct from known Bornean species in the following characters: moderate size, with SVL up to 66.7 mm (versus 76.9 mm in C. nigridia and <60 mm in C. paripari , C. matahari , C. sirehensis , C. dringi , C. lagang , and C. kendallii ); 4–7 postmental scales (versus 10 in C. dringi and 8–10 in C. lagang ); 10–14 precloacal pores in males (versus 2–5 in C. paripari , 7–8 in C. matahari , 4–9 in C. leucura , 5–6 in C. dringi , 4–9 in C. lagang , and absent in C. kendallii ); 9–14 putatively precloacal pore-bearing convex scales in females (versus no obvious convex precloacal scales in C. paripari , C. matahari , C. leucura , C. lagang , and C. kendallii ); 29–34 paravertebral tubercles (versus indistinct dorsal tubercles in C. lagang and 12–18 in C. kendallii ); at least a few caudal tubercles present in a lateral furrow (versus absent in C. sirehensis and C. dringi ); single median row of smooth, flat subcaudal scales (versus smooth but folded scales in C. paripari , C. matahari , and possibly C. sirehensis , and keeled scales in C. dringi , C. lagang , and C. kendallii ); 1–3 postcloacal tubercles (versus absent in C. dringi ); sharp tetragonal dorsal black marks (versus diffuse thin marks in C. paripari , C. matahari , C. sirehensis , and C. leucura ; sharp thin marks in C. lagang ; and sharp round marks in C. kendallii ); and black–gray dorsal banding and white median subcaudal of original tail (versus black–dark yellow banding in C. nigridia , immaculate white in C. paripari and C. lagang , and black-blobbed dorsal and immaculate ventral in C. leucura ).

Etymology. The generic name is feminine in gender. The specific epithet puterisantubongae is derived from Puteri (= princess) Santubong, who is one of two princesses appearing in a local myth. These princesses are said to have descended from the celestial realm to a human village; their remarkable beauty and weaving skills are represented in the species by yellow ocher body colouration, distinct bold black marks, and pale/faint yellow spots in both sexes. Santubong is also the name of the promontory for which the Santubong National Park is named.

Distribution. Cnemaspis puterisantubongae is known from Santubong National Park, Kuching District, Sarawak, East Malaysia ( Fig. 1 ).

Natural History. The species has been found exclusively on the surfaces of sandstone boulders or outcrops containing pebble conglomerate, sometimes along small streams in lowland mixed dipterocarp forest ( Fig. 5 ). Individuals have been observed and/or collected at about 01:00, 09:00, 16:00, and 20:00. From morning to dusk, geckos positioned themselves on the edges and inside of large rock crevices. Geckos tended to avoid humans and scattered when approached, particularly at distances of several metres. However, at around midnight, they strayed farther from crevices and remained exposed even when approached by humans, suggesting that this species is diurnal.

Cnemaspis puterisantubongae shares the same habitat with the sympatric species C. kendallii , Gekko monarchus (Schlegel, 1836 in Duméril & Bibron, 1836), and Tropidophorus brookei ( Gray, 1845) . The two sympatric Cnemaspis species appear to use different microhabitats. Compared with C. kendallii , which uses trees, logs, and the upper and central parts of large boulders above human height, C. puterisantubongae has never been observed on such open surfaces but remains within closed or complex interspaces ( Fig. 5 ). Typically, one Cnemaspis individual is found on a single boulder; on a larger boulder two or three individuals can be observed, sometimes including both C. puterisantubongae and C. kendallii , while at other times only one of the two species is present. This behaviour may indicate a competitive relationship among these individuals. However, although both species have been observed in partially open spaces around midnight, they were slow to react, and interspecific interactions were not observed during surveys. Whether microhabitat selection by either species is affected by the occurrence of the other remains unknown because the range of C. puterisantubongae is within that of C. kendallii .

One female C. puterisantubongae individual had two large eggs and others did not ( Fig. 4 ), suggesting that the reproductive season of these species includes September. We observed more than 20 individuals, but none exhibited tail waving when they were threatened or startled as opposed to C. kendallii , which frequently waves or wags its tail when threatened.

Figures

FIGURE 1. The type localities of the nigridia group of the Bornean Cnemaspis (thus not shown for C. lagang and C. kendallii) and phylogenetic relationship of its members. The map is tilted 45° counterclockwise. Igneous and sedimentary bases within Sarawak were shown in light and dark gray, respectively. Support of the internal branches of the maximum likelihood tree was evaluated by ultrafast bootstrap/SH-aLRT values, and asterisks indicate those higher than 95/95. The black, brown, and gray bands under the tips of the tree indicate granite, sandstone, and limestone dwelling species, respectively.

FIGURE 3. The holotype (SFC-GRB-00517; temporal field number KUHE Swk23-186) of Cnemaspis puterisantubongae sp. nov. The lateral (upper), dorsal (lower left), and ventral views (lower right) were provided before fixation, with close-up pictures of some discriminant characters after fixation. The label KUHE Swk23-186 is a temporal field number.

FIGURE 4. The dorsal (upper) and ventral views (middle) of an adult female paratype (SFC-GRB-00516; temporal field number KUHE Swk23-185) of Cnemaspis puterisantubongae sp. nov. The magnified view of precloacal pore-bearing scales on the left side of body (bottom) are indicated by black arrows.

FIGURE 5. The habitat (sandstone boulders) of Cnemaspis puterisantubongae sp. nov. in Santubong National Park (upper). The microhabitat preferences of C. puterisantubongae (lower left; inner surface of boulders) and co-occurring C. kendallii (lower right; both of inner and outer surfaces) are shown, with arrows pointing the positions of geckos.