Hemiphyllodactylus tonywhitteni, Grismer & Wood Jr & Kyaw Thura & Zin & Quah & Murdoch & Grismer & Li & Kyaw & Lwin, 2017

Grismer, L. Lee, Wood Jr, Perry L., Kyaw Thura, Myint, Zin, Thaw, Quah, Evan S. H., Murdoch, Matthew L., Grismer, Marta S., Li, Aung, Kyaw, Htet & Lwin, Ngwe, 2017, Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar, Journal of Natural History 52 (13 - 16), pp. 881-915 : 891-898

Hemiphyllodactylus tonywhitteni sp. nov.

Phapant dwarf gecko

( Figures 5 and 6 )

Holotype

Adult male ( LSUHC 13026 ) collected on 18 October 2016 at 1600 hours by Evan S. H. Quah, Perry L. Wood, Jr., Matthew L. Murdoch, Thaw Zin, Myint Kyaw Thura, Htet Kyaw, Marta S. Grismer, and L. Lee Grismer from Phapant Cave , 25.2 km north-east of Taunggyi, Taunggyi District, Shan State, Myanmar (21°11.472N, 96°33.214E; 1270 m).

Paratypes

Adult females ( LSUHC 13027 and 13030) and juvenile female ( LSUHC 13028 ) and juvenile male ( LSUHC 13029 ) bear the same data as the holotype .

Diagnosis

Hemiphyllodactylus tonywhitteni sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 38.8 mm; 5–8 chin scales; enlarged postmentals; 3–5 circumnasal scales; 2–4 scales between supranasals (=postrostrals); eight or nine supralabials; eight infralabials; 13–16 longitudinally arranged dorsal scales at midbody contained within one eye diameter and 7–9 ventral scales; varied digital formulae ( Table 3); three subdigital lamellae on the first finger; three or four subdigital lamellae on the first toe; 20–26 continuous pore-bearing femoroprecloacal scales; no plate-like subcaudal scales; dark postorbital stripe not extending onto trunk; pairs of paravertebral light spots on trunk; dorsal body pattern not unicolour; postsacral marking bearing light-coloured anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all species of Hemiphyllodactylus from clades 3 and 4 ( Table 3).

Description of holotype

Adult male; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.33); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernible; snout moderate, rounded in dorsal profile; eye large; ear opening round, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by small supranasals; three internasals (=postnasal); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 8 (R,L) rectangular supralabials tapering to below posterior margin of orbit; 8 (R,L) rectangular infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single large, sublabial; seven chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals.

Body somewhat elongate (Trunk/SVL 0.48), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 14 dorsal scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, seven ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totalling 26; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II–V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits II–V 4-4-4-4 (R,L); three transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I well developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II–V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits II–V 4-4-4-4 (R,L); three transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; dorsal caudal scales small, square, subimbricate; tail regenerated, covered with flat imbricate scales. Morphometric data are presented in Table 5.

Coloration before preservation ( Figure 5 )

Top of head, body, limbs, and tail grey, overlain with darker, broken bands on trunk appearing as paravertebral markings highlighted posteriorly by light-coloured, diffuse blotches; poorly defined dark, lineate markings extend from occipital region to shoulder; spotting or striping on trunk absent; diffuse, dark, preorbital stripe; dark, postorbital stripe irregularly shaped, extending to shoulder region; limbs bearing irregularly shaped, dark markings; tail generally unicolour; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; and pigmentation density increases posteriorly with the abdomen being generally grey.

Variation ( Figures 5 and 6 )

The colour patterns of the paratypes generally match that of the holotype. LSUHC 13027 is darker overall and the colour pattern is less distinct . The light-coloured, paravertebral blotches in LSUHC 13029 are salmon coloured . The dark dorsal pattern of LSUHC 13028 is more speckled and that of the adult female LSUHC 13030 is more reticulate . LSUHC 13030 has an original tail lacking enlarged subcaudal plates and bearing a weak, ventrolateral fringe and a distinct banding pattern . The tail is oval in cross-section and the underside is dull orange. The intensity of coloration and contrast in pattern changes with mood and activity. Differences in scales counts are presented in Table 5.

Distribution

Hemiphyllodactylus tonywhitteni sp. nov. is known only from the type locality of Phapant Cave , Taunggyi District, Shan State, Myanmar ( Figure 1 ) .

Natural history

Phapant Cave is a complex of three caves situated around a small depression along a narrow river. The karstic ridge and outcroppings surround a small monastery which incorporates the caves for worship. The hilly area connecting the caves is composed of highly eroded limestone walls bearing many cracks and pores. Large limestone boulders that have broken away from the cliff face line the base of the shallow escarpment ( Figure 7 ). We believe Hemiphyllodactylus tonywhitteni sp. nov. is a karst-adapted species. A specimen of H. tonywhitteni sp. nov. was found just inside a small opening of one of the caves nearly 4 m above the cave entrance. More specimens were found on the boulders at the base of the cliff and one on one of the cement buildings of the monastery. Syntopic with H. tonywhitteni sp. nov. on both the karst outcroppings and the cement building was an undescribed species of Hemidactylus . Hemidactylus sp. nov. was also found on wooden structures and vegetation where H. tonywhitteni sp. nov. was absent.

Etymology

This specific epithet ‘ tonywhitteni ’ honours Dr Tony Whitten of Fauna & Flora International who has championed a broad range of conservation efforts in Indonesia and the Asia Pacific for well over a quarter of a century. His tireless efforts to conserve and help manage karst ecosystems have been a great inspiration to the senior author (LLG) herein.

Comparisons

The molecular analyses indicate that Hemiphyllodactylus tonywhitteni sp. nov. is embedded within clade 4 of the typus group and is the sister species of H. montawaensis sp. nov. It can be distinguished from H. jinpingensis , H. chiangmaiensis and the species of clade 3 by lacking dark, dorsolateral stripes on the trunk and transverse, dorsal blotches. The PCA analysis shows that it occupies a unique morphospace with respect to H. montawaensis sp. nov. and H. linnwayensis sp. nov. with PC1 and PC2 accounting for 49% of the variation in the concatenated dataset ( Figure 3 ). PC1 accounted for 29% of the variation and loaded most heavily for trunk length and the number of subdigital lamellae on the first toe ( Table 6). PC2 accounted for an additional 20% of the variation and loaded most heavily for the number of dorsal scales. The first four components of the PCA were retained for the DAPC which shows that not only are all three species distinct but all individuals of each species fall very close to or within the 95% confidence ellipses ( Figure 4 ). Uncorrected pair-wise sequence divergence between H. tonywhitteni sp. nov. and all other species of clades 3 and 4 ranges from 6.4–18.7% ( Table 7). Hemiphyllodactylus tonywhitteni sp. nov. is most similar to its sister species H. montawaensis sp. nov. but differs in having more femoroprecloacal pores (20–26 versus 19–21) and a relatively wider head (0.17–0.19 versus 0.16–0.17) throughout its growth trajectory ( Figure 8 ) and a statistically significantly wider head (p <0.24, n = 5) as an adult.

Figures

Figure 1. Distribution of Hemiphyllodactylus montawaensis sp. nov., H. tonywhitteni sp. nov., H. linnwayensis sp. nov., and H. cf. linnwayensis in the Shan Hills of eastern Myanmar.

Figure 3. Principal component analysis (PCA) of the species Hemiphyllodactylus montawaensis sp. nov., H. tonywhitteni sp. nov., and H. linnwayensis sp. nov., showing their distribution in morphospace along the first two principal components.

Figure 4. Discriminant analysis of principal components (DAPC) of the species Hemiphyllodactylus montawaensis sp. nov., H. tonywhitteni sp. nov., H. linnwayensis sp. nov. emphasizing between-group variance and minimizing within-group variance based on retention of the first four components.

Figure 5. (a) Juvenile male paratype (LSUHC 13029) of Hemiphyllodactylus tonywhitteni sp. nov. from the type locality of Phapant Cave 25.2 km north-east of Taunggyi, Taunggyi District, Shan State, Myanmar. (b) Adult female paratype (LSUHC 13030) from the type locality.

Figure 6. Type series of Hemiphyllodactylus tonywhitteni sp. nov. from the type locality of Phapant Cave 25.2 km north-east of Taunggyi, Taunggyi District, Shan State, Myanmar.

Figure 7. Karst wall microhabitat of Hemiphyllodactylus tonywhitteni sp. nov. from the type locality.

Figure 8. Comparative growth trajectories of head width between Hemiphyllodactylus montawaensis sp. nov., H. tonywhitteni sp. nov. and H. linnwayensis sp. nov. Shaded areas represent the 95% confidence intervals.