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

Grismer, L. Lee, Wood, Perry L., Jr., Thura, Myint Kyaw, Zin, Thaw, Quah, Evan S. H., Murdoch, Matthew L., Grismer, Marta S., Lin, Aung, Kyaw, Htet & Lwin, Ngwe, 2018, Twelve new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from isolated limestone habitats in east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism, Zoological Journal of the Linnean Society 182, pp. 862-959 : 929-935

CYRTODACTYLUS WELPYANENSIS SP. NOV.

WEL PYAN CAVE BENT-TOED GECKO

( FIG. 30; TABLE 17)

Holotype: Adult male LSUHC 12874 collected on 3 October 2016 at 1500 h by Myint Kyaw Thura, L. Lee Grismer, Perry L. Wood, Jr., Matthew L. Murdoch, Evan S. H. Quah, Thaw Zin, Htet Kyaw and Marta S. Grismer from Wel Pyan Cave 35 km north of Hpa-an, Hpa-an District, Kayin State, Myanmar (N17°12.188, E97°37.066; 21 m in elevation).

Paratypes: Adult female LSUHC 12785 and adult male 12786 bear the same collection data as the holotype .

Abbreviations are listed in the Material and Methods. R, right; L, left; /, data unobtainable or not applicable; r, regenerated; b, broken.

Diagnosis: Cyrtodactylus welpyanensis sp. nov. differs from all congeners by having the unique combination of eight or nine supralabials; seven infralabials; 16 longitudinal rows of body tubercles; 30–33 paravertebral tubercles; 28–30 ventral scales; relatively long digits with eight expanded subdigital lamellae proximal to the digital inflection on the fourth toe, 11–13 unmodified, distal, subdigital lamellae, 19–21 total subdigital lamellae; raised, moderate to strongly keeled, dorsal body tubercles not extending beyond base of tail; enlarged femoral and precloacal scales continuous; 30–31 nearly equally sized, enlarged, femoral scales; 16 femoral pores in males; 11–13 enlarged precloacal scales; 7–8 precloacal pores in males; three rows of enlarged post-precloacal scales; subcaudal scales twice as wide as long, not extending onto lateral surface of tail; top of head darkly mottled, no yellow reticulum; nuchal loop not divided medially, lacking an anterior, azygous notch, posterior border jagged; six dark dorsal bands lacking paravertebral elements, wider than interspaces, lacking lightened centres, not edged with white tubercles; nape band present; dark markings in dorsal interspaces; ventrolateral folds whitish; anterodorsal margins of thighs and brachia darkly pigmented; nine light caudal bands bearing dark markings, not encircling tail; ten dark caudal bands wider than light caudal bands; and mature regenerated tail unicolour.

Description of holotype: Adult male SVL 69.2 mm; head moderate in length (HL/SVL 0.30), wide (HW/ HL 0.62), flat (HD/HL 0.37), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region shallowly concave, canthus rostralis rounded; snout elongate (ES/HL 0.42), rounded in dorsal profile, not flat in lateral profile; eye large (ED/HL 0.28); ear opening round, moderate in size (EL/HL 0.08); 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 and one azygous internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals and ventrally by first supralabial; 8(R)10(L) square supralabials extending to below midpoint of eye; 7(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 45% 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, subconical, semi-regularly arranged, moderate to strongly keeled tubercles; tubercles extend from nape to base of tail but no farther; tubercles on nape smaller than those on posterior portion of body and less sharply keeled; approximately 16 longitudinal rows of body tubercles; 30 paravertebral tubercles; 30 flat, subimbricate, ventral scales larger than dorsal scales; 11 enlarged precloacal scales; seven precloacal pores; three rows of large post-precloacal scales; and no deep, precloacal groove or depression.

Forelimbs moderate in stature, relatively short (FL/ SVL 0.17); flat scales of forearm larger than those on body, not interspersed with small tubercles; palmar scales flat; 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 0.20), covered dorsally by raised scales intermixed with large tubercles; flat, slightly larger scales anteriorly; ventral scales of thigh flat, imbricate, larger than dorsal scales, one row of 14(R)16(L) nearly equally sized, enlarged, femoral scales in contact with enlarged precloacal scales; 7(R)9(L) femoral pores; subtibial scales flat, imbricate; small, postfemoral scales form abrupt union with larger, flat ventral scales of posteroventral margin of thigh; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 8(R,L) expanded subdigital lamellae on fourth toe proximal to joint inflection, 11(R,L) unmodified subdigital lamellae distal to inflection, 19 total subdigital lamellae; and claws well-developed, base of claw sheathed by a dorsal and ventral scale.

Tail moderate in proportions, 86.0 mm in length, last 50.0 mm regenerated, 6.8 mm in width at base, tapering to a point; dorsal scales of tail flat; medial subcaudal scales twice as wide as long, not extending onto lateral surface of tail; 4(R,L) enlarged postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales flat.

Coloration in life ( Fig. 30): Dorsal ground colour of head body, and limbs faintly magenta, that of anterior portion of tail dull-white; top of head and rostrum bearing a network of dark, diffuse mottling, no yellow reticulum; superciliary scales whitish; dark-brown, jagged, nuchal loop, unnotched anteromedially, deeply incised posteromedially; short, jagged incomplete band on nape; six jagged, dark, body bands not bearing lightened centres, wider than interspaces, not bearing paravertebral elements; one sacral band; interspaces bearing faint, dark markings, most extensive on flanks; irregularly shaped, faint bands on thighs; thighs and brachia darkly pigmented; ventrolateral body folds whitish; dark caudal bands not bearing lightened centres, wider than light caudal bands; light caudal bands with dark markings, not encircling tail; and ventral surfaces pigmented, dusky in appearance, more so beneath limbs and tail, less so in gular region.

Variation ( Fig. S 10): The paratypes closely approximate holotype in aspects of dorsal colour pattern. The dorsal band in the shoulder region in LSUHC 12786 is not bifurcated and it has a completely, nearly unicolour tan, regenerated tail. LSUHC 12785 is much darker overall. Meristic and mensural differences are presented in Table 17 .

Distribution: Cyrtodactylus welpyanensis sp. nov. is known only from Wel Pyan Cave 35 km north of Hpa-an, Hpa-an District, Kayin State, Myanmar ( Fig. 29).

Etymology: The specific epithet, welpyanensis (pronounced way-pee-an-ensis), is a noun in apposition in reference to the type locality of Wel Pyan Cave.

Natural history: Wel Pyan Cave is located along the west side of the Salween River and situated on the eastern flank of a small karst hill approximately 0.5 km wide, 1.3 km long and 256 m high that is surrounded by paddy fields. The opening of the cave is narrow (~10 × 10 m) and situated approximately 25 m above the base of the hill. The interior of the cave is complex and filled with blind caverns, stalactites and a maze of small, underground passageways through which water courses and through which a flatulent monk guided us ( Fig. 31). We found three specimens of C. welpyanensis sp. nov. at approximately 1500 h 2–6 m up on clean, dry, smooth surfaces of the cave walls all within 10 m of each other and all within 50 m of the cave entrance where light still penetrated. Outside the cave, the base of the karst hill was jagged, deeply incised with small, steep ravines and riddled with boulders that had broken away from the hillside. We do not believe that C. welpyanensis sp. nov. is restricted to the interior of the cave and if we had not been unexpectedly rushed away due to safety concerns, we believe many more specimens would have been found outside the cave after dark. The only other gekkonid seen was H. frenatus .

Comparisons: Cyrtodactylus welpyanensis sp. nov. is part of the sinyineensis group. The PCA and DAPC analyses indicate that the species of this group are completely separated in morphospace where the first two principal components account for 63% of the total variation ( Fig. 12; Fig. S5) and load most heavily for numbers of infralabials, longitudinal rows of body tubercles, expanded subdigital lamellae on the fourth toe and post-precloacal scale rows ( Table S3). Cyrtodactylus wlepyanensis sp. nov. is well-differentiated from C. aequalis , C. sinyineensis sp. nov. and C. dammathetensis sp. nov. by having varying combinations of statistically different mean values of supralabial scales, infralabial scales, ventral scales, longitudinal rows of body tubercles, precloacal scales and body bands ( Table 3). It differs further from the other species in the sinyineensis group in that the body tubercles do not extend past the base of the tail and from C. aequalis and C. sinyineensis sp. nov. in that the dark body bands are not edged with light tubercles ( Table 8). Morphological and colour pattern 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 16.5% ( Table 10).

The oldhami group

The monophyletic oldhami group is composed of Cyrtodactylus cf. peguensis zebraicus, a polyphyletic C. oldhami , C. thirakhupti , C. payarhtanensis and C. lenya from the Thai-Malay Peninsula just north of the Isthmus of Kra and C. saiyok from southwestern Thailand ( Figs 2, 9). This group undoubtedly contains more Thai species but until researchers describing new species from this region realize that taking tissue samples in an age of integrative taxonomy is imperative, we will never know. The oldhami group is highly variable and defined by the following characters from Smith (1935), Taylor (1963), Pauwels et al. (2004), Grismer & Norhayati (2008), Panitvong et al. (2014) and in part by Connette et al. (2017): 9–13 supralabials; 9–12 infralabials; 14–20 longitudinal rows of body tubercles; 23–40 ventral scales; 7–20 subdigital lamellae; femoral pores absent; 0–8 precloacal pores in males; post-preclocal scales large; wide, transverse, medial, subcaudal scales; no anterior azygous notch in nuchal loop; no band on nape; 4–7 variably shaped body bands with lightened centres or dorsal pattern composed of light lines or pairs of dark paravertebral blotches; anterodorsal margins of thighs and brachia and ventrolateral folds pigmented; and maximum SVL 61.0–83.0 mm. Connette et al. (2017) recovered C. payarhtanensis as the sister species of a genetic sample of C. oldhami from the pet trade that has no voucher or locality. We sequenced two vouchered southern Thai specimens (MS 460, 585) from Ranong and Phang-nga Provinces, respectively, that form a monophyletic group but exclude the pet trade specimen, indicating C. oldhami may be polyphyletic. Our data further indicate that C. thirakhupti is the sister species of C. oldhami (MS 460, 585) and that C. payarhtanensis is sister to that group ( Fig. 9).

The yathepyanensis group

The monophyletic yathepyanensis group is composed of Cyrtodactylus yathepyanensis sp. nov. and the sister

Abbreviations are listed in the Material and Methods. R, right; L, left; /, data unobtainable or not applicable.

species C. linnoensis sp. nov. and C. sadanensis sp. nov. from the lowland flood plain of the Salween River Basin of Kayin and Mon and states ( Figs 9, 20). The yathepyanensis group is defined by the flowing range of characters: 9–11 supralabials; 7–9 infralabials; dorsal body tubercles raised, moderately to strongly keeled, extending beyond base of tail; 13–19 longitudinal rows of body tubercles; 26–33 paravertebral tubercles; 30–38 ventral scales; 21–24 subdigital lamellae; 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; 2–6 precloacal pores in males; three or four post-preclocal scale rows; transverse caudal scales two or three times as wide as long, extending onto lateral surface of tail; top of head bearing dark, mottled pattern; no anterior, azygous notch in nuchal loop; band on nape; 4–6 regularly shaped body bands lacking or with only faint, lightened centres, edged with light tubercles; anterodorsal margins of thighs darkly pigment; ventrolateral folds not whitish; 13–16 light caudal bands not encircling tail; 13–17 dark caudal bands; and maximum SVL 72.3–78.0 mm ( Table 8). The description and diagnosis of each species follows.