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

pyinyaungensis View in CoL sp. nov. peguensis annandalei *

Mean 8.0 7.5 /

SD 0 0.50 /

Range 8 7 or 8 8–12

N 5 23

Precloacal pores (PP)

Mean 8 7.5 /

SD 0 0.50 /

Range 8 7 or 8 11 or 12

N 2 23 Post-precloacal scale rows ( PPS)

Mean 2 3 /

SD 0 0 /

Range 2 3 /

N 3 3 /

Body bands (BB)

Mean 5.8 6 /

SD 0.45 0 /

Range 5 or 6 6 5

N 5 23

Light caudal bands (LCB)

Mean 10.8 12 /

SD 1.71 0 /

Range 9–13 12 11

N 4 13

Dark caudal bands (DCB)

Mean 11.0 12 /

SD 1.63 0 /

Range 9–13 12 11 or 12

N 4 13

Morphology

Body tubercles low, weakly keeled No Yes No

Body tubercles raised, moderately to strongly keeked Yes No Yes

Tubercles extend beyond base of tail No No No

Enlarged femoral scales present Yes No No

Enlarged femoral and precloacal scales continuous No / /

Pore-bearing femoral and precloacal scales continuous / / /

Enlarged proximal femoral scales ~1/2 size of distal No / /

femorals

Medial subcaudal scales two or three times wider than Yes Yes Yes long

Medial subcaudal extend onto lateral surface of tail No No No

Colour pattern

Nuchal loop divided medially No No No

Nuchal loop with anterior azygous notch No No No

Posterior border of nuchal loop Straight Straight Straight Band on nape Yes Yes Yes

Dorsal banding with paravertebral elements Yes Yes No Paravertebral elements of dorsal bands confluent Yes No /

Dorsal bands wider than interspaces Yes Yes No

Dorsal bands bearing lightened centres No Yes No

Dorsal bands edged with light-coloured tubercles Yes Yes No

Shape of dorsal bands Hourglass Paired blotches Diagonal Dark markings in dorsal interspaces No No No

/, data were unobtainable or not applicable.

*Data from Bauer (2003).

Nonetheless, five specimens were observed in two nights. BYU 52234 View Materials was collected during October of the previous year (2016) approximately 0.3 m above the ground on the side of a karst boulder. The collection site is situated at the lower end of a steeply sloping valley composed of disturbed secondary forest, scattered outcroppings of karst boulders and shallow lateral drainages feeding into the valley floor at 642 m elevation. Only one specimen was seen in this habitat in four nights of searching. These observations suggest that C. pyinyaungensis sp. nov. is a habitat generalist in hilly areas bearing rocky substrates. One hatchling was observed on 29 March 2017 but no gravid females were found .

Comparisons: Cyrtodactylus pyinyaungensis sp. nov. is part of a large clade of Indo-Burmese species within which it is closely related to C. annandalei from which it differs in having six vs. ten infralabial scales; 16–19 vs. ten total subdigital lamellae on the fourth toe; 30–36 vs. 43 ventral scales; 17 or 18 vs. ten or 11 femoral pores; eight vs. 11 or 12 precloacal pores; the presence vs. the absence of enlarged femoral scales; the dorsal bands being wider as opposed to narrower than the interspaces and edged with light-coloured tubercles; the dark caudal bands being wider than the light caudal bands; and the top of the head being blotched as opposed to patternless ( Table 7). The sister population of C. pyinyanugensis sp. nov. from Popa Mountain approximately 120 km to the west was misidentified as C. feae (USNM 559805; Wood et al., 2012 and Agarwal et al., 2014) and was reported to L.L.G. to be C. peguensis (G. R. Zug, unpubl. data). Although we do not have access to this specimen, a comparison of its sister species, C. pyinyaungensis sp. nov., to one of the syntypes of C. peguensis (BMNH 1946.8.23.10) shows that although they are similar in overall colour pattern ( Fig. 13), they differ in that the former has fewer rows of longitudinal tubercles (11 vs. 14); fewer ventral scales (30–36 vs. 43–45); enlarged femoral scales; and femoral pores ( Table 7); and dorsal pattern composted of confluent paravertebral markings as opposed to large, separate, paravertebral spots ( Fig. 13). Genetic distances among the species of this group range from 11.3 to 15.1%.

The intermedius group

The monophyletic intermedius group is the sister lineage to the linnwayensis group ( Fig. 9) and ranges from Hon Tre and Phu Quoc islands and isolated mountainous regions in the Mekong Delta of southern Vietnam west through the Cardamom Mountains of southern Cambodia and south-eastern Thailand ( Grismer et al., 2015b; Fig. 2) and is composed of Cyrotdactylus intermedius (Smith) , C. phuquocensis Ngo, Grismer & Grismer , and C. hontreensis Ngo, Grismer & Grismer. The intermedius group is defined by the following characters ( Ngo, Grismer & Grismer, 2008, 2010): 9–13 supralabials; 9–12 infralabials; 14–22 longitudinal rows of dorsal tubercles; 38–50 ventral scales; 15–19 subdigital lamellae on the fourth toe; no enlarged femoral not contiguous with enlarged precloacal scales; 7–10 precloacal pores in males; post-preclocal scales large; wide, transverse caudal scales; top of head unicolour to sparsely blotched; no anterior azygous notch in nuchal loop or first body band; no band on nape; 3–5 regularly shaped body bands with lightened centres and edged with light tubercles or markings; anterodorsal margins of thighs, brachia and ventrolateral fold not pigmented; and maximum SVL 72.3–78.0 mm.

The linnwayensis group

The linnwayensis group is a monophyletic group composed of the sister species Cyrtodactylus linnwayensis sp. nov. and C. shwetaungorum sp. nov. ( Fig. 9) and range through the western regions of the Shan Hills of the Mandalay Region and Shan State ( Fig. 2). This group is defined by the following range of characters 7–9 supralabials; 6–8 infralabials; dorsal body tubercles low, weakly keeled, not extending beyond base of tail; 25–35 paravertebral tubercles; 13–21 longitudinal rows of tubercles; 33–42 ventral scales; 20–23 subdigital lamellae on the fourth toe; enlarged femoral and precloacal scales continuous; 24–32 enlarged femoral scales nearly the same size throughout; 15–22 femoral pores in males; 8–12 enlarged precloacal scales; 6–11 precloacal pores in males; three or four post-precloacal scale rows; transverse caudal scales twice as wide as long, not extending onto lateral subcaudal regions; top of head mottled and bearing light-coloured, reticulated pattern; no band on nape; three or four regular (vs. zigzagged) body bands with lightened centres and edged with light tubercles; anterodorsal margins of thighs, brachia and ventrolateral fold pigmented; eight or nine light caudal bands; 7–9 dark caudal bands; and maximum SVL 101.7–102.2 mm ( Table 7). The description and diagnosis of each species follows.