Cyrtodactylus sanpelensis, 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 : 919-923

CYRTODACTYLUS SANPELENSIS SP. NOV.

SANPEL CAVE BENT-TOED GECKO

( FIG. 24; TABLE 14)

Holotype: Adult male LSUHC 12877 collected on 8 October 2016 between 1600 and 2000 h by Myint Kyaw Thura, L. Lee Grismer, Marta S. Grismer, Evan S. H. Quah, Perry L. Wood, Jr., Matthew L. Murdoch, Thaw Zin and Htet Kyaw from Sanpel Cave 21.3 km south-east of Mawlamyine, Mawlamyine District, Mon State, Myanmar (N16°22.427, E97°46.388; 44 m in elevation).

Paratypes: Adult males BYU 52223 , LSUHC 12875 , 12881 , 12883 , 12887 , 12889–90 , adult females BYU 52221–22 , 52224 , LSUHC 12878–80 , 12886 and juvenile LSUHC 12885 bear the same collection data as the holotype .

Diagnosis: Cyrtodactylus sanpelensis sp. nov. differs from all congeners by having the unique combination of 9–11 supralabials; seven or eight infralabials; 12–14 rows of longitudinal body tubercles; 27–31 paravertebral tubercles; 35–41 ventral scales; relatively long digits with eight or nine expanded subdigital lamellae proximal to the digital inflection on the fourth toe, 12 or 13 unmodified distal subdigital lamellae, 20–22 total subdigital lamellae; low, weakly keeled, dorsal body tubercles; body tubercles not extending beyond base of tail; enlarged femoral and precloacal scales not continuous; 14–20 enlarged femoral scales; 8–16 femoral pores in males; 9–12 enlarged precloacal scales; seven or eight precloacal pores in males; two rows of enlarged post-precloacal scales; median subcaudal scales three times as wide as long, extending onto lateral surface of tail; top of head diffusely mottled, no yellow reticulum; nuchal loop not divided medially, lacking an anterior, azygous notch, posterior border straight or sinuous; five (rarely four), dark, regularly shaped, dorsal bands not wider than interspaces, nearly always with lightened centres, not edged with white tubercles; band on nape present; dark markings in dorsal interspaces; ventrolateral fold not whitish; anterodorsal margin of thighs and brachia unpigmented; 7–10 light caudal bands bearing dark markings, not encircling tail; 7–10 dark caudal bands wider than light caudal bands; and regenerated tail not spotted.

Description of holotype: Adult male SVL 70.7 mm; head moderate in length (HL/SVL 0.32), wide (HW/HL 0.65), flat (HD/HL 0.41), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region moderately concave, canthus rostralis flattened; snout elongate (ES/HL 0.44), rounded in dorsal profile, flat in lateral profile; eye large (ED/HL 0.25); ear opening round, moderate in size (EL/HL 0.11); 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, one azygous internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial; 10(R,L) square supralabials extending to below midpoint of eye; 8(R)7(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 not intermixed with tubercles; dorsal superciliaries not elongate or keeled; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting medially for 50% 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 0.43) with well-defined ventrolateral folds; dorsal scales small, raised and interspersed with moderately sized, subconical, low, semi-regularly arranged, weakly keeled tubercles; tubercles extend from nape to base of tail but no farther; tubercles absent on nape, tubercles on posterior portion of body largest and more heavily keeled; approximately 14 longitudinal rows of body tubercles; 28 paravertebral tubercles; 37 flat, subimbricate, ventral scales larger than dorsal scales; 11 enlarged, precloacal scales; eight precloacal pores; two rows of large post-precloacal scales; and 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, not interspersed with tubercles; palmar scales slightly raised flat; digits well-developed, relatively long, inflected at basal, interphalangeal joints; digits much more narrow distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale at base; hindlimbs more robust than forelimbs, moderate in length (TBL/SVL 0.21), covered dorsally by raised scales intermixed with larger tubercles, bearing flat, slightly larger scales anteriorly; ventral scales of thigh flat, imbricate, larger than dorsal scales, one row of 12(R)8(L) enlarged femoral scales not in contact with enlarged precloacal scales; 8(R,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 low, slightly raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints; 8(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection, 13(R,L) unmodified subdigital lamellae distal to inflection, 21 total subdigital lamellae; and claws well-developed, base of claw sheathed by a dorsal and ventral scale.

Tail original, moderate in proportions, 110.0 mm in length, 7.2 mm in width at base, tapering to a point; dorsal caudal scales flat; transversely expanded, median subcaudal scales three times as wide as long, extending onto lateral surface of tail; two enlarged postcloacal tubercles at base of tail on hemipenal swellings; and postcloacal scales flat.

Coloration in life ( Fig. 24): Dorsal ground colour of head body, limbs and tail magenta; top of head and rostrum diffusely mottled, no yellow reticulum; superciliary scales dull-yellow; dark-brown nuchal band unnotched anteromedially, bearing a sinuous posterior margin; short, hourglass-shaped, band on nape bordered laterally and anterolaterally by diffuse, dark markings; four generally evenly bordered bands on body bearing lightened centres, not wider than interspaces; one sacral band; interspaces bearing distinct dark markings; limbs diffusely and irregularly banded; anterodorsal margins of thighs and brachia not darkly pigmented; ventrolateral body folds whitish; dark caudal bands with lightened centres, wider than light caudal bands; light caudal bands bear dark markings, do not encircle tail; ventral surfaces of head body and limbs beige with stippled lateral edges; and subcaudal region darker, variegated.

Variation ( Fig. S7): The colour pattern of the paratypes varies little from that of the holotype. LSUHC 12885 is a juvenile and has bolder body bands, lacks a nuchal band and has immaculate dorsal interspaces indicating some ontogenetic change in colour pattern is present. LSUHC 12875 is much darker and the interspaces more heavily mottled. The posterior border of the nuchal loop in BYU 52223–24 , LSUHC 12875– 80 , 12887 and 12890 is straight as opposed to sinuous. The lateral sections of the dorsal bands in LSUHC 12886 are quite faded on the flanks. BYU 52221 , 5223 , LSUHC 12881 and 12883 have partially regenerated tails. Meristic and mensural differences are presented in Table 14 .

Distribution: Cyrtodactylus sanpelensis sp. nov. is known only from Sanpel Cave 21.3 km south-east of Mawlamyine, Mawlamyine District, Mon State, Myanmar ( Fig. 20).

Etymology: The specific epithet, sanpelensis , is a noun in apposition in reference to the type locality of Sanpel Cave.

Natural history: Sanpel Cave is situated at the north end of a small karst ridge approximately 53 m wide, 189 m long and 194 m high. The ridge is surrounded by paddy fields and is adjacent to a small, ephemeral river to the east. The opening of the cave is small (~ 8 m high and ~ 6 m) and approximately 20 m above the base of the hill. The interior of the cave is complex with many side passages that extend to great depths below the entrance. There are several stalactites, cracks in the walls and side passages that lizards utilize as retreats. Just inside the entrance of the cave are several religious statues on which a number of lizards were collected. Lizards were abundant inside the cave both day and night. After dark, lizards were extremely common on the outside karst hillside that also had an abundance of cracks and holes ( Fig. 25). Lizards were even found in well-illuminated areas near the monastery. In essence, there was no rocky habitat that lizards did not utilize. No lizards were seen on vegetation. The juvenile LSUHC 12885 was found on a brick wall. Gekko gecko was observed on the outside of the cave.

Comparisons: Cyrtodactylus sanpelensis sp. nov. is part of the sadanensis group. See Comparisons section for C. sadansinensis sp. nov. and C. pharbaungensis sp. nov.

The sinyineensis group

The monophyletic sinyineensis group is composed of the sister species Cyrtodactylus sinyineensis sp. nov. and C. welpyanensis sp. nov., and the sister species C. dammathetensis sp. nov. and C. aequalis Bauer ( Fig. 9). This group ranges from the uplands of the southern section of the Shan Hills in Kayin State to the lowland flood plain of the Salween River Basin in Kayin and Mon and states ( Fig. 20). It could be argued that C. tigroides belongs in this group given it is the sister lineage. However, the deep genetic divergence between C. tigroides and the sinyinensis group is commensurate with that between the Thai sister lineages of the other Burmese species groups and it is likely that when the phylogenetic relationships of additional Thai species are known, C. tigroides will likely align with them in a separate species group as is the case for the intermedius group and the zebraicus group ( Fig. 9). The sinyineensis

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

group is defined by the flowing range of characters: 8–10 supralabials; 6–8 infralabials; dorsal body tubercles raised, moderately to strongly keeled, usually extending beyond base of tail; 30–36 paravertebral tubercles; 13–19 longitudinal rows of body tubercles; 19–30 ventral scales; 19–22 total subdigital lamellae; enlarged femoral and precloacal scales continuous; 25–36 enlarged femoral scales nearly the same size throughout; 13–36 femoral pores in males; 5–13 enlarged precloacal scales; 4–9 precloacal pores in males; three post-precloacal scale rows; transverse, median subcaudal scales usually twice as wide as long, not extending onto lateral surface of tail; top of head bearing a dark mottled pattern; no anterior, azygous notch in nuchal loop; dark band on nape; five or six variably shaped body bands generally lacking lightened centres; anterodorsal margins of thighs and brachia darkly pigmented; 9–11 light caudal bands encircling tail; 9–11 dark caudal bands; and maximum SVL 69.3– 91.6 mm ( Table 8). The description and diagnosis of each new species follows.