Cyrtodactylus linnoensis, Grismer & Wood & Jr. & Thura & Zin & Quah & Murdoch & Grismer & Lin & Kyaw & Lwin, 2018

CYRTODACTYLUS LINNOENSIS View in CoL SP. NOV.

LINNO CAVE BENT-TOED GECKO

( FIG. 34; TABLE 19)

Holotype: Adult male LSUHC 12826 View Materials collected on 4 October 2016 at 2000 h by L. Lee Grismer, Perry L. Wood, Jr., Evan S. H. Quah, Matthew L. Murdoch, Marta S. Grismer, Myint Kyaw Thura, Thaw Zin and Htet Kyaw from the Linno Cave region 5 km south-west of Hpa-an, Hpa-an District, Kayin State, Myanmar (N16°50.551, E97°36.402; 25 m in elevation). GoogleMaps

Paratypes: Adult males BYU 52230–31 View Materials , 52323 View Materials , LSUHC 12829 View Materials , 12832–34 View Materials , juvenile male BYU 52223 View Materials and adult females LSUHC 12824–25 View Materials bear the same collection data as the holotype GoogleMaps .

total variation ( Fig. 12; Fig. S5) and load most heavily for numbers of paravertebral tubercles and ventral scales ( Table S4). Cyrtodactylus yathepyanensis sp. nov. is well-differentiated from C. linnoensis sp. nov. and C. sadanensis sp. nov. by having varying combinations of statistically different mean values of supralabial and infralabial scales, paravertebral tubercles, unmodified and total fourth toe lamellae, ventral scales, enlarged femoral scales, and longitudinal rows of body tubercles and precloacal pores ( Table 4). It differs further from C. sadanensis sp. nov. and some C. linnoensis sp. nov. in having continuous vs. discontinuous enlarged femoral and precloacal scales and pigmented vs. unpigmented thighs and brachia, and from C. sadanensis sp. nov. in having a smaller maximum SVL (72.8 mm vs. 78.0 mm; Table 8). Morphological and colour pattern differences from other species in the Indo-Chinese clade are listed in Table 8. Genetic distances among Diagnosis: Cyrtodactylus linnoensis sp. nov. differs from all congeners by having the unique combination of nine or ten supralabials; seven or eight infralabials; 13–15 rows of longitudinal body tubercles; 26 or 27 paravertebral tubercles; 35–38 ventral scales; relatively long digits with nine or ten expanded subdigital lamellae proximal to the digital inflection on the fourth toe, 13 or 14 unmodified, distal, subdigital lamellae, 22–24 total subdigital lamellae; raised, moderately to strongly keeled, dorsal, body tubercles extending beyond base of tail; 12–37 enlarged femoral scales, proximal scales one-half to one-third the size of distal scales; 12–14 femoral pores in males; 9–13 enlarged precloacal scales; 4–6 precloacal pores in males; three or four rows of enlarged post-precloacal scales; medial subcaudal scales three times as wide as long, extending onto lateral surface of tail; top of head bearing diffuse, dark mottling, no yellow reticulum; nuchal loop lacking an anterior, azygous notch, posterior border usually sinuous; five or six dark, jagged, dorsal bands lacking paravertebral elements, wider than interspaces, faintly lightened centres, edged with light-coloured tubercles; band on nape; dark markings but no light-coloured tubercles in dorsal interspaces; ventrolateral folds whitish; anterodorsal margins of thighs lack pigment; brachia at least faintly pigmented; 15 light-coloured caudal bands bearing dark markings, not encircling tail; 14 dark caudal bands wider than light caudal bands; and mature, regenerated tail not spotted.

Description of holotype: Adult male SVL 74.9 mm; head moderate in length (HL/SVL 0.29), wide (HW/HL

LSUHC 12834 View Materials paratype 6.7 2.9 2.3

LSUHC 12833 View Materials paratype 5.9 3.0 2.1

LSUHC 12832 View Materials paratype 6.9 2.7 2.3

BYU 52233 View Materials paratype 4.4 2.5 1.9

BYU 52232 View Materials paratype 6.4 2.9 2.2. broken

,

b

LSUHC 12829

paratype 6.3 3.0 2.2; regenerated

,

r

BYU 52231

paratype 6.6 3.1 2.3 applicable;

not

BYU 52230

paratype 6.4 3.1 2.2 or unobtainable

LSUHC 12825 View Materials paratype 6.6 3.1 2.5;, data / left

,

L

;

LSUHC 12824

paratype 6.5 3.7 2.0 R., right

Methods

LSUHC 12826 View Materials holotype 6.6 2.9 2.5 and

Material

Continued

in listed are. 19 Table IO EL IN Abbreviations

0.62), flat (HD/HL 0.41), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.47), rounded in dorsal profile, moderately flat in lateral profile; eye large (ED/HL 0.23); ear opening round, moderate in size (EL/HL 0.13); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally by inverted Y-shaped furrow, bordered posteriorly by left and right supranasals contacting on midline and minute azygous scale, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial; 9(R,L) square supralabials extending to below midpoint of eye; 8(R,L) infralabials tapering posteriorly to below orbit; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with small tubercles laterally; dorsal superciliaries not elongate or keeled; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting for 70% of their length posterior to mental; one row of slightly enlarged chinshields bordering all infralabials; and gular and throat scales small, flat, grading posteriorly into larger, subimbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.46) with well-defined ventrolateral folds; dorsal scales small, raised and interspersed with large, conical, semi-regularly arranged, moderately to strongly keeled tubercles extending from nape to beyond base of tail; tubercles on nape smaller than those on posterior portion of body, less sharply keeled; approximately 13 longitudinal rows of body tubercles; 26 paravertebral tubercles; 38 flat, subimbricate, ventral scales larger than dorsal scales; 13 enlarged precloacal scales; five precloacal pores; three rows of large, post-precloacal scales; no deep, precloacal groove or depression.

Forelimbs moderate in stature, relatively short (FL/ SVL 0.18); slightly raised scales of forearm larger than those on body, lacking tubercles; palmar scales low, rounded; digits well-developed, relatively long, inflected at basal, interphalangeal joints; digits much more narrow distal to inflections; widened, proximal, subdigital lamellae do not extend onto palm; claws well-developed, sheathed by a dorsal and ventral scale at base; hindlimbs more robust than forelimbs, moderate in length (TBL/SVL 0.22), covered dorsally by raised scales intermixed with large tubercles and bearing flat, slightly larger scales anteriorly; ventral scales of thigh flat, imbricate, larger than dorsal scales, one row 17(R,L) of enlarged femoral scales in contact with enlarged precloacal scales, proximal femoral scales one-half to one-third the size of distal femoral scales; 6(R,L) femoral bearing diffuse, dark, lineate markings; superciliary scales yellowish; dark-brown, nuchal loop bearing a sinuous posterior border connects with dark, nape band laterally; five jagged, body bands wider than interspaces, bearing faintly lightened centres, lacking paravertebral elements, edged with light tubercles; one nape band; one chevron-shaped postsacral band; interspaces bearing large dark markings, especially on flanks; line of yellowish spots on lower flanks; banding on limbs indistinct, mottled with yellowish markings; anterodorsal margins of thighs unpigmented; anterodorsal margins of brachia faintly to heavily pigmented; ventrolateral folds whitish; dark caudal bands not bearing lightened centres, wider than light caudal bands; light caudal bands bearing dark markings, not encircling tail; ventral surfaces pigmented, dusky in appearance; and subcaudal region dark with lighter mottling.

Variation ( Fig. S12): The paratypes reasonably approximate the holotype in aspects of colour pattern. The nuchal loop is divided in BYU 52230 View Materials , LSUHC 12824 View Materials , 12829 View Materials and 12832. In LSUHC 12825 View Materials , the dorsal bands are so wide and dark, they are difficult to discern. Although BYU 52233 View Materials is a juvenile, its colour pattern is not notably more contrasted or different from the range of variation observed in the adults, indicating that an ontogenetic change is absent. Meristic and mensural differences are presented in Table 19 .

pores; small, postfemoral scales form an abrupt union with larger, flat ventral scales of posteroventral margin of thigh; subtibial scales flat, imbricate; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 9(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that do not extend onto sole, 14(R,L) unmodified subdigital lamellae distal to inflection, 23 total subdigital lamellae; and claws well-developed, base of claw sheathed by a dorsal and ventral scale.

Tail original, moderate in proportions, 79.0 mm in length, 7.6 mm in width at base, tapering to a point; dorsal scales of tail flat; medial subcaudal scales three times as wide as long, extending onto lateral surface of tail; 2(R,L) enlarged, postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales flat.

Coloration in life: Dorsal ground colour of head body, and limbs grey, that of anterior portion of tail very dull-yellow; top of head bearing, diffuse, dark mottlings, yellow reticulum absent; rostrum Distribution: Cyrtodactylus linnoensis sp. nov. is known only from the Linno Cave region 5 km south-west of Hpa-an, Hpa-an District, Kayin State, Myanmar ( Fig. 20).

Etymology: The specific epithet, linnoensis , is a noun in apposition in reference to the type locality.

Natural history: The Linno Cave region is situated at the north-east end of a small karst hill approximately 340 m wide, 430 m long and 57 m high located on the west bank of Salween River. It is actually the north-west section of a much larger karst hill that has been bisected by the Salween River. The collecting area is a vertical karst wall approximately 250 m in length and 20 m tall that parallels the river and leads to the Linno Cave at its north-eastern terminus ( Fig. 35). The habitat along the wall has several deep cracks, holes, side chambers and deep incisions that serve as refuge sites for geckos during the day. Additionally, the wall is sheltered by vegetation. The last 50 m of the wall near the cave lack vegetation and here we saw only Hemidactylus brookii and Gekko gecko both day and night. At night, Cyrtodactylus linnoensis sp. nov. were abundant and occurred along the entire wall of statistically different mean values of supralabial and infralabial scales, paravertebral tubercles, expanded, unmodified and total fourth toe lamellae, ventral scales, enlarged femoral scales, longitudinal rows of body tubercles and precloacal pores ( Table 8). It differs further from C. sadanensis sp. nov. in having a larger maximum SVL (78.0 mm vs. 73.8 mm). Morphological differences from other species in the Indo-Chinese clade are listed in Table 8. Genetic distances among the species of this group range from 11.0 to 12.7% ( Table 10).