Cnemaspis lineatubercularis, Ampai & Jr & Stuart & Aowphol, 2020

Ampai, Natee, Jr, Perry L. Wood, Stuart, Bryan L. & Aowphol, Anchalee, 2020, Integrative taxonomy of the rock-dwelling gecko Cnemaspis siamensis complex (Squamata, Gekkonidae) reveals a new species from Nakhon Si Thammarat Province, southern Thailand, ZooKeys 932, pp. 129-159 : 129

Cnemaspis lineatubercularis sp. nov. Figures 3 , 4 , 5 , 6 , 7 , 8

Type material.

Holotype (Figs 3 - 5 ). ZMKU R 00828, adult male from Thailand, Nakhon Si Thammarat Province, Lan Saka District, Kam Lon Subdistrict, Wang Mai Pak Waterfall (8°26.807'N, 99°46.525'E; 96 m a.s.l.), collected on 25 January 2019 by Natee Ampai, Anchalee Aowphol, Attapol Rujirawan, Korkwan Termprayoon and Siriporn Yodthong.

Paratypes (Figs 6 - 8 ). Eighteen paratypes (adult males = 11, adult females = 7). ZMKU R 00821-00825 (five adult males), and ZMKU R 00826 (adult female), same data as holotype except that they were collected on 25 October 2016. ZMKU R 00827, ZMKU R 00829-00831 (four adult males), ZMKU R 00832-00835 (four adult females), THNHM 28694-28695 (two adult males) and THNHM 28696-28697 (two adult females), same data as holotype.

Diagnosis.

Cnemaspis lineatubercularis sp. nov. can be distinguished from all other Cnemaspis by having the following combination of characters: (1) maximum SVL of 40.6 mm (mean 38.8 ± SD 1.4, N = 12) in adult males and maximum SVL of 41.8 mm (mean 39.5 ± SD 1.9, N = 7) in adult females; (2) 8-9 supralabial and infralabial scales; (3) gular, pectoral, abdominal, and subcaudal scales keeled; (4) rostral, interorbitals, supercilium, palmar scales, and ventral scales of brachia smooth; (5) 5-6 small, subconical spine-like tubercles present on flanks (6) 19-21 paravertebral tubercles linearly arranged; (7) 27-29 subdigital lamellae under the 4th toe; (8) 4-7 pore-bearing precloacal scales, pores rounded, arranged in chevron shape and separated in males; (9) one postcloacal tubercle each side in males; (10) ventrolateral caudal tubercles anteriorly present; (11) caudal tubercles restricted to a single paravertebral row on each side; (12) single median row of subcaudal scales keeled and lacking enlarged median row; and (13) gular region, abdomen, limbs and subcaudal region yellowish only in males. These differences are summarized among geographically close congeners in the siamensis group (Table 5 ).

Description of holotype.

Adult male; SVL 40.1 mm; head moderate in size (HL/SVL 0.26), narrow (HW/SVL 0.16), flattened (HD/HL 0.41) and head distinct from neck; snout moderate (ES/HL 0.47), snout slightly concave in lateral view; postnasal region concave medially; scales of rostrum smooth, larger than conical scales on occiput; weak supraorbital ridges; gular marking absent; gular and throat scales granular, keeled and round; shallow frontorostral sulcus; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.23); pupil round; extral-brillar fringe scales largest anteriorly; scales on interorbitals and supercilium smooth; ear opening oval, taller than wide; rostral slightly concave; rostral bordered posteriorly by supranasals and laterally by first supralabials; 9, 9 (Right, Left) supralabials decreasing in size posteriorly; 9, 9 (Right, Left) infralabials decreasing in size posteriorly; nostril elliptical, oriented dorsoposteriorly, bordered posteriorly by small postnasal scales; mental scales large, triangular, concave, bordered posteriorly by three large postmentals.

Body slender, elongate (AG/SVL 0.43); small, keeled, dorsal scales equal in size throughout body intermixed with several large, keeled, multicarinate tubercles; 19 paravertebral tubercles linearly arranged; tubercles present on lower flanks; tubercles extend from occiput to tail; five small, subconical spine-like tubercles on flanks; dorsal scales raised and keeled; pectoral and abdominal scales keeled, round, flat to concave, slightly larger than dorsal and not larger posteriorly; ventral scales of brachia smooth, raised and juxtaposed; six separated pore-bearing precloacal scales with rounded pores; precloacal depression absent; femoral pores absent.

Fore and hind limbs moderately long, slender; scales beneath forearm slightly raised, smooth and sub-imbricate; subtibial scales keeled; palmar scales smooth and juxtaposed; digits elongate, slender, inflected joint and bearing slightly recurved claws; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular; lamellae of distal phalanges wide; lamellae beneath inflection large; interdigital webbing absent; enlarged submetatarsal scales on 1st toe absent; total subdigital lamellae on fingers: 17-21-25-28-26 (right manus), 17-16 (broken)-25-28-26 (left manus); fingers increase in length from first to fourth with fourth and fifth nearly equal in length; relative length of fingers IV>V>III>II>I; total subdigital lamellae on toes: 13-21-24-29-25 (right pes), 13-21-24-29-25 (left pes); toes increase in length from first to fifth with fourth and fifth nearly equal in length; relative length of toes IV>V>III>II>I.

The original tail cylindrical, swollen at the base and longer than head and body (TL/SVL 1.36); subcaudal scales keeled, juxtaposed, similar to dorsal scale of the tail size; shallow, middorsal furrow; deeper lateral caudal furrow present; enlarged, transverse caudal tubercles arranged in segmented whorls, not encircling tail; enlarged median subcaudal scale row absent; caudal tubercles absent from lateral furrow; tail length (TL) 54.7 mm; a single postcloacal tubercle on each side at lateral surface of hemipenial swellings at the base of tail.

Measurements of holotype

(in mm; Table 6 ). SVL 40.1; TL (original) 54.7; TW 3.9; FL 5.8; TBL 7.2; AG 17.4; HL 10.3; HW 6.3; HD 4.2; ED 2.4; EE 3.1; ES 4.8; EN 3.9; IO 2.9; EL 1.0; IN 1.0.

Coloration in life

(Fig. 4 ). Dorsal ground color of head light brown, top of head bearing small, diffuse, faint black and yellowish markings; dark postorbital stripes faint, extending to nape; large, round, whitish marking on nape; single light-yellowish prescapular crescent on the shoulder, located dorsoanteriorly of forelimb insertion; dorsal ground color of body, limbs and tail light brown with black irregular blotches; ground color of ventral surfaces grayish-white intermixed with yellowish blotches; ventral pattern sexually dimorphic, anterior gular, abdominal, and caudal regions yellowish in males; two dark blotches on nape form a bipartite pattern; light sage vertebral blotches extending from the nape to tail; flanks with irregular, incomplete brown to yellowish blotches becoming smaller posteriorly; tubercles on the whole body white or yellow; widely separated, white or yellow tubercles occur on flanks; subconical spine-like yellowish tubercles on flanks; limbs beige with dark brown mottling; tail faintly marked with dark brown.

Coloration in preservative

(Figs 5 , 7 , 8 ). Color pattern similar to that in life with some fading of markings. Dorsal ground color of head, body, limbs and tail darker brown than in life, with indistinct, irregular markings. Yellow coloration in gular, pectoral, abdominal regions, flanks, and tail faded to light-yellow and creamy-white.

Variation.

Most paratypes approximate the holotype in general aspects of morphology (Figs 6 - 8 ), with most differences found in the degree of vertebral blotches. All adult female paratypes lack the yellowish coloration in the gular, abdominal, and caudal regions. ZMKU R 00821-00825, ZMKU R 00829-00831, THNHM 28694 (nine adult males) and ZMKU R 00826, ZMKU R 00832 and ZMKU R 00833 (three adult females) have regenerated tails of uniform tan coloration. ZMKU R 00821, 00824, 00827 (three adult males) and ZMKU R 00832, 00835 (two adult females) have lighter dorsal markings that appear more as transverse bands than as paravertebral blotches. THNHM 28696 (adult female) has a broken tail. Differences in meristic and morphometric characters within the type series are presented in Table 6 .

Distribution and natural history.

Cnemaspis lineatubercularis sp. nov. is known only from Wang Mai Pak Waterfall (96 m a.s.l.), Kam Lon Subdistrict, Lan Saka District, Nakhon Si Thammarat Province, southern Thailand (Fig. 9 ). The type locality is surrounded by lowland evergreen forest along a river basin in the southern part of the Nakhon Si Thammarat mountain range. Specimens were found only along granitic rocky streams of Wang Mai Pak Waterfall. The rocky boulder microhabitats of this species are dry with cool surface temperatures (24.8-26.7 °C, 73.2-86.1% relative humidity). When disturbed, some individuals retreated deeper into rock crevices, cracks, more shaded areas or beneath rock boulders.

Seven specimens (ZMKU R 00822-00825, ZMKU R 00827, THNHM 28696-28697) were collected during the day (1650-1847 h) and 12 specimens (ZMKU R 00821, ZMKU 00826, ZMKU R 00828-00832, THNHM 28694-28695 and ZMKU R 00833-00835) were collected at night (1913-1951 h).

The male holotype was found during the night (1943 h) perched head down on a vertical surface in a crevice of a granitic rock boulder near a stream. A female paratype (ZMKU R 00832) was found with the male holotype, separated by only a distance of approximately 10 cm.

Paratypes that were found during the day were in shaded areas, crevices of boulders, rock walls and on boulder outcrops near streams. Paratypes found at night were in shaded surfaces of the boulders, within deep crevices, or perched on vegetation near a rocky stream. Three gravid females (ZMKU R 00832-00834) contained one or two eggs during January 2019. Some juveniles (SVL <30 mm; not collected) were found in rock cracks and perched on a rock near a stream on 25 January 2019.

Cnemaspis lineatubercularis sp. nov. appears to be a diurnal species in that observed specimens during daytime were active and fast-moving when disturbed, but those at night were inactive, slow-moving or asleep on dry granitic rocks and vegetations. At night, Cyrtodactylus lekaguli and Gehyra mutilata were found in syntopy with the new species on a rock wall and vegetation near a stream. A summary of ecological parameters of activity periods, elevation (lowland <600 m), microhabitat preference and presence or absence of ocelli (eyespots) of Cnemaspis in Thailand is shown in Table 7 .

Etymology.

The specific epithet lineatubercularis is taken from linea (Lat. for line) and tubercularis (Lat. for having tubercles), in reference to the new species having paravertebral tubercles linearly arranged.

Comparisons.

Cnemaspis lineatubercularis sp. nov. can be distinguished from other members of the siamensis group ( C. adangrawi , C. chanardi , C. huaseesom , C. kamolnorranathi , C. omari , C. phangngaensis , C. punctatonuchalis , C. roticanai , C. siamensis , C. thachanaensis , and C. vandeventeri ; Table 5 ) by having a smaller maximum SVL of 41.8 mm (vs. 44.9 mm in C. adangrawi , 43.5 mm in C. huaseesom , 42.0 mm in C. phangngaensis , 49.6 mm in C. punctatonuchalis , 47.0 mm in C. roticanai , 44.7 mm in C. vandeventeri ) and by having a larger maximum SVL 41.8 mm (vs. 40.9 mm in C. chanardi , 37.8 mm in C. kamolnorranathi , 41.3 mm in C. omari , 39.7 mm in C. siamensis , 39.0 mm in C. thachanaensis ).

Cnemaspis lineatubercularis sp. nov. is distinguished from C. adangrawi , C. phangngaensis , and C. thachanaensis by having fewer 8-9 supralabial scales (vs. 10 in C. adangrawi and C. phangngaensis , 10-11 in C. thachanaensis ). This species is distinguished from C. phangngaensis by having fewer 8-9 infralabial scales (vs. 10 in C. phangngaensis ). This species is distinguished from C. huaseesom and C. punctatonuchalis by having keeled ventral scales (vs. smooth ventral scales in C. huaseesom and C. punctatonuchalis ). This species is distinguished from C. punctatonuchalis , C. siamensis , and C. thachanaensis by presence of precloacal pores (vs. precloacal pores absent in C. punctatonuchalis , C. siamensis , C. thachanaensis ). This species is distinguished from C. huaseesom , C. kamolnorranathi , and C. phangngaensis by having a separated row of precloacal pores (vs. continuous in C. huaseesom , C. kamolnorranathi , C. phangngaensis ). This species is distinguished from C. kamolnorranathi by having rounded precloacal pores (vs. pores elongated in C. kamolnorranathi ).

Cnemaspis lineatubercularis sp. nov. is distinguished from C. adangrawi , C. chanardi , C. omari , C. phangngaensis , C. punctatonuchalis , C. roticanai , and C. vandeventeri by having fewer 19-21 paravertebral tubercles (vs. 23-25 in C. adangrawi , 22-25 in C. chanardi , 22-29 in C. omari , 22 in C. phangngaensis , 24-27 in C. punctatonuchalis , 25-27 in C. roticanai , 25-29 in C. vandeventeri ). This species is distinguished from C. adangrawi , C. chanardi , C. huaseesom , C. omari , C. roticanai , C. siamensis , and C. vandeventeri by having paravertebral tubercles linearly arranged (vs. randomly arranged in C. adangrawi , C. chanardi , C. huaseesom , C. omari , C. roticanai , C. siamensis , C. vandeventeri ). This species is distinguished from C. adangrawi and C. phangngaensis by having tubercles on lower flanks (vs. absent in C. adangrawi and C. phangngaensis ). This species is distinguished from C. siamensis and C. thachanaensis by having more 27-29 lamellae under 4th toe (vs. 24-26 in C. siamensis and 24 in C. thachanaensis ).

Cnemaspis lineatubercularis sp. nov. is distinguished from C. chanardi , C. huaseesom , C. kamolnorranathi , C. omari , C. roticanai , C. siamensis , and C. vandeventeri by the presence of ventrolateral caudal tubercles anteriorly (vs. lacking in C. chanardi , C. huaseesom , C. kamolnorranathi , C. omari , C. roticanai , C. siamensis , C. vandeventeri ). This species is distinguished from C. vandeventeri by having lateral caudal furrows (vs. lacking in C. vandeventeri ). This species is distinguished from C. huaseesom and C. punctatonuchalis by having keeled subcaudal scales (vs. lacking in C. huaseesom and C. punctatonuchalis ). This species is distinguished from C. chanardi , C. huaseesom , C. omari , C. punctatonuchalis , C. roticanai , and C. siamensis by having single median row of keeled subcaudals (vs. lacking in C. chanardi , C. huaseesom , C. omari , C. punctatonuchalis , C. roticanai , C. siamensis ). This species is distinguished from C. chanardi , C. punctatonuchalis , C. siamensis , and C. vandeventeri by lacking enlarged median subcaudal scales (vs. present in C. chanardi , C. punctatonuchalis , C. siamensis , C. vandeventeri ). This species is distinguished from C. adangrawi , C. chanardi , C. huaseesom , C. kamolnorranathi , C. omari , C. punctatonuchalis , C. roticanai , C. siamensis , and C. vandeventeri by having caudal tubercles restricted to a single paravertebral row on each side (vs. lacking in C. adangrawi , C. chanardi , C. huaseesom , C. kamolnorranathi , C. omari , C. punctatonuchalis , C. roticanai , C. siamensis , C. vandeventeri ).

Cnemaspis lineatubercularis sp. nov. is distinguished from C. thachanaensis by having one postcloacal tubercle in males (vs. lacking in C. thachanaensis ). This species is distinguished from C. huaseesom by having keeled subtibial scales (vs. smooth in C. huaseesom ). This species is distinguished from C. adangrawi , C. kamolnorranathi , C. punctatonuchalis , C. siamensis , C. thachanaensis , and C. vandeventeri by having yellow coloration in the subcaudal region (vs. lacking in C. adangrawi , C. kamolnorranathi , C. punctatonuchalis , C. siamensis , C. thachanaensis , C. vandeventeri ). This species is distinguished from C. huaseesom , C. phangngaensis , C. punctatonuchalis , and C. roticanai by lacking dorsal color pattern sexually dimorphic (vs. having in C. huaseesom , C. phangngaensis , C. punctatonuchalis , C. roticanai ).

Figures

Figure 3. Male holotype (ZMKU R 00828) of Cnemaspis lineatubercularis sp. nov. from Wang Mai Pak Waterfall, Lan Saka District, Nakhon Si Thammarat Province, Thailand.

Figure 4. Male holotype (ZMKU R 00828) of Cnemaspis lineatubercularis sp. nov. from Wang Mai Pak Waterfall, Lan Saka District, Nakhon Si Thammarat Province, Thailand, in life A dorsal view B ventral view C lateral view of the head D dorsal view of trunk E precloacal region showing distribution of pore-bearing scales (red arrows) F dorsal view of tail G ventral view of tail. Scale bar: 10 mm (in dorsal and ventral views).

Figure 5. Male holotype (ZMKU R 00828) of Cnemaspis lineatubercularis sp. nov. from Wang Mai Pak Waterfall, Lan Saka District, Nakhon Si Thammarat Province, Thailand, in preservative A lateral view of head B dorsal view of trunk C ventral view of chin D dorsal view E ventral view. Scale bar: 10 mm (D, E).

Figure 6. Coloration of Cnemaspis lineatubercularis sp. nov. in dorsal (above) and ventral (below) views of A male paratype ZMKU R 00830 and B female paratype ZMKU R 00835. Note yellowish ventral coloration that is present in males but absent in females.

Figure 7. Male paratypes of Cnemaspis lineatubercularis sp. nov. in preservative in A dorsal view B ventral view; from left to right, top panel: ZMKU R 00821, ZMKU R 00822, ZMKU R 00823, ZMKU R 00824 and ZMKU R 00825; bottom panel: ZMKU R 00827, ZMKU R 00829, ZMKU R 00830, ZMKU R 00831, THNHM 28694 and THNHM 28695.

Figure 8. Female paratypes of Cnemaspis lineatubercularis sp. nov. in preservative in A dorsal view B ventral view; from left to right, top panel: ZMKU R 00826, ZMKU R 00832, THNHM 28696 and THNHM 28697; bottom panel: ZMKU R 00833, ZMKU R 00834 and ZMKU R 00835.

Figure 9. Habitats of Cnemaspis lineatubercularis sp. nov A Wang Mai Pak Waterfall at type locality B microhabitat of holotype in granitic rocky stream (white arrow) C microhabitat of paratypes in granitic rocky outcrops (white arrows) at Wang Mai Pak Waterfall, Lan Saka District, Nakhon Si Thammarat Province, Thailand.