Leiolepis glaurung, Wanchai & Rujirawan & Murdoch & Aksornneam & Promnun & Kaatz & Gregory & Nguyen & Iderstein & Quah & Grismer & Grismer & Aowphol, 2024

Wanchai, Pratyaporn, Rujirawan, Attapol, Murdoch, Matthew L., Aksornneam, Akrachai, Promnun, Pattarapon, Kaatz, Amanda, Gregory, Jeren J., Nguyen, Eddie, Iderstein, William Van, Quah, Evan S. H., Grismer, L. Lee, Grismer, Jesse L. & Aowphol, Anchalee, 2024, The description of the first rock-dwelling species of butterfly lizard Leiolepis Cuvier, 1829 (Squamata, Agamidae) from the Khorat Plateau in northeastern Thailand, ZooKeys 1210, pp. 299-324 : 299-324

Leiolepis glaurung sp. nov.

Figs 5 , 6 , 9 , 11 ; Table 1 ; Suppl. materials 1, 2 Suggested English name: Khorat Plateau Butterfly Lizard

Type material.

Holotype. Adult male ( THNHM 30909 ; Fig. 5 A ) collected from just northeast of Wat Phu Noi in Kaeng Kheng Subdistrict, Kut Khaopun District, Ubon Ratchathani Province, Thailand (15 ° 48 ' 10.7 " N, 105 ° 09 ' 24.9 " E) on 20 March 2023, at 10: 00 am by Pratyaporn Wanchai, Anchalee Aowphol, Attapol Rujirawan, Akrachai Aksornneam, Jesse L. Grismer, L. Lee Grismer, Evan S. H. Quah, and Matthew L. Murdoch . Paratypes. Adult female ( THNHM 30908 ; Fig. 5 B ) and two adult males ( THNHM 30910 –30911; Fig. 5 C, D ) bear the same locality and collectors as the holotype . Three adult males ( THNHM 30902 –30903, THNHM 30907 ) and three adult females ( THNHM 30904 –30906; Fig. 6 ) were collected by Pratyaporn Wanchai between 4–10 July 2020 from the same locality as the holotype .

Diagnosis.

Leiolepis glaurung sp. nov. can be diagnosed from all sexual species of Leiolepis by having a black gular region with a wide medial yellow stripe, a yellow ventrum with black mottling, bright red to orange ventral tail coloration, having reduced to no expandable flanks, and having a maximum of one black transverse bar on the flank (Suppl. material 1). Leiolepis glaurung sp. nov. can be further diagnosed from all other sexual species by having the combination of a AG of 61.0–88.0 mm; BL of 11.0– 16.8 mm; CW of 22.3–33.1 mm; DN of 4.3–6.7 mm; FA of 14.4–19.0 mm; HE of 5.6–7.1 mm; HH of 14.0– 21.3 mm; HL of 28.8–41.9 mm; HW of 19.2–29.7 mm; PW of 10.5–17.0 mm; TIB of 24.34–29.0 mm; TE of 14.0–18.0 mm, RH of 9.7–14.3 mm; three dorsal stripes; 28–29 ventral scales; 21–24 dorsal caudal scales; 19–26 non-pore bearing scales between the pore-bearing femoral scales across the pelvis; maximum number of seven keeled scale rows across the forearm; 17–20 femoral pores per side; 6–8 scales across the frontal bone; maximum of nine subtibial scales; seven scales long the dorsal surface of the first toe; three enlarged subdigital lamellae on the third toe; and 32–34 scales along the dorsal surface of the fourth toe (Figs 7 , 8 ).

Description of holotype.

Head large, (HL 41.9 mm; HL / SVL 0.25) obtusely rounded in lateral profile, triangular in dorsal profile (HW 29.7 mm; HW / HL 0.71); interorbital, frontal region, and rostrum, convex (HH 21.3 mm; RH 14.3 mm; RH / HH 0.67), sloped anteriorly, covered with small, undifferentiated keeled scales; occipital and supraorbital regions covered with 19 keeled, granular scale rows half the size as dorsal head scales; canthus rostralis short, rounded; dorsal head scales strongly keeled, eight keeled scales across the frontal bone between supraorbital regions; rostral large, triangular (wider than long), bordered posteriorly by six smaller scales; external nares large, set wide apart (DN 6.7 mm; DN / HW 0.23) rounded, directed laterally, set in single, oval, nasal scale surrounded by several small scales; elongate, keeled, large fused, suborbitals (on five right side; five on the left) extend from anterior margin of eye to posterior margin of eye; superciliary scales elongate, keeled, imbricate, continuous with canthal scales; eyelid scales granular; tympanum naked, deeply set, surrounded by granular scales; temporal scales keeled, small, slightly raised; nine rectangular supralabials whose contact with one another produces an distinct labial margin, bordered ventrally by small granular scales; mental longer than wide, pointed posteriorly, larger than adjacent infralabials; two large postmentals in contact medially, being first of a series of 14 enlarged scales along the angle of jaw (left side: right side was damaged); 10 rectangular infralabials; gular scales small, rounded, granular; two distinct anterior and posterior gular folds present; dewlap absent; antehumeral fold continuous with posterior gular fold (Figs 5 A , 6 A ).

Body elongate (AG 88.0 mm; AG / SVL 0.52) somewhat dorsoventrally compressed; expandable flanks reduced to absent; body scales small, granular, slightly keeled; 43 scales between dorsolateral stripes; 20 scales between vertebral stripe and dorsolateral stripes at widest point of trunk; scales of flanks abruptly transition into much larger, flat scales of belly and pectoral region; 28 scales across middle of belly contacting apex of the umbilical scar; precloacals smooth and much smaller than ventral scales; forelimbs short, robust (FA 19.0 mm; BL 16.8 mm); dorsal surface of forelimbs and posterior surface of brachia covered with large, keeled, imbricate scales; six rows of enlarged, keeled scales across forearm; ventral surface of forelimbs covered with granular scales; plantar scales small, granular; subdigital lamellae of fingers are composed of a single wide transversely elongated scales; claws long; hind limbs relatively long (FM 33.6 mm; TIB 27.2 mm); dorsal scales on hind limbs small, weakly keeled; scales on anterior surface of thighs large, flat, weakly keeled, imbricate; those on forelegs slightly enlarged, keeled; postfemoral scales small, granular; eight longitudinal rows of large, smooth, flat, imbricate subtibial scales; 36 total femoral pores; each pore set in larger scale; 25 non-pore-bearing scales between pore-bearing-femoral scales across the pelvis; plantar scales small, raised; subdigital lamellae of toes bicarinate, 32 enlarged, plate-like scales along dorsal surface of the fourth toe; three enlarged, triangular scales on posterior surface at base of third toe (ESL 2.6 mm; ESL / TE 0.16); seven enlarged, plate-like scales along length of first toe; tail dorsoventrally compressed, noticeably wider at base, constricted at its contact point with body, covered dorsally with small, keeled scales grading ventrally into larger, flat, weakly keeled, subcaudals; caudal scales in transverse rows encircle the tail; and the last 110 mm of the tail is regenerated (Fig. 5 A ).

Coloration in life.

Dorsal ground color of head, body, limbs, and tail is grey to almost black (when animal is cold base color is black); anterior portion of the head pale grey with no pattern; three white lines radiate from the posterior region of the orbit with one extending posteriorly to the parietal region, one to the tympanic region, one to the corner of the mouth; four yellow stripes extending posteriorly from the parietal region of the head, the two central stripes connect on the nape of the neck forming a yellow Y-shape and extend posteriorly as a vertebral stripe that terminating at the anterior margin of the pelvis, the two lateral stripes extend dorsolaterally from the parietal region running the entire length of the body terminating at the anterior margin of pelvis; dorsal pattern is composed of three yellow dorsal stripes (mentioned above) separated by distinct darker regions with yellow spots; the base color of the flanks are yellowish orange with one black transverse bar in the axillary region, followed by pale yellow transverse bands composed of small yellow spots, three (L) and four (R); the forearms have prominent yellow and white spots; the hind limbs have small white ocelli with diffuse edges; the dorsal caudal coloration is composed of the same small ocelli as on the hind limbs and the coloration turns into dark transverse caudal bars approximately 30 % down the length of the tail; the gular region is black with a wide medial yellow strip; pectoral region is yellow with black mottling; the limb ventral color is yellow; subcaudal coloration is bright red to orange and extends laterally to the dorsolateral coloration of the tail (Figs 5 A , 6 , 9 , 11 )

Etymology.

The specific epithet glaurung is in reference to the large, terrestrial, golden-colored, non-winged dragon, Glaurung in Middle-earth – a character created by J. R. R. Tolkien in The Silmarillion (1977). Glaurung the Golden is the father of all dragons and tunneled into the sides of mountains forming burrows. The reduced expandable lateral flanks, yellow ventral and dorsal colors, with the construction of burrows beneath rocky outcrops is similar to the descriptions of Glaurung mentioned above, from ‘ The Silmarillion’ and ‘ The Children of Húrin’ ( Tolkien 1977, Tolkien and Tolkien 2007). Additionally, in Thailand the word “ Yae ” is used to refer to Leiolepis belliana , L. ocellata , and L. rubritaeniata . However, on the Khorat Plateau the populations of Leiolepis glaurung sp. nov. are called “ Yarb ”.

Distribution.

Leiolepis glaurung sp. nov. is currently known from three locations on the Khorat Plateau (Fig. 1 ). We have only collected specimens from the type locality in Ubon Ratchathani Province (Fig. 1 ) from the Phu Phan Formation ( Jin-geng and Meesook 2013), however a population from Chaiyaphum Province (15 ° 54 ' 7.92 " N, 102 ° 9 ' 50.4 " E) in the western region of the Khorat Plateau was observed and photographed, but no specimens were collected (AK, AR, and PP pers. obs.), and from southern extent of the plateau on the border of Cambodia (PW pers. obs.) in Ubon Ratchathani Province (Fig. 1 ). Lizards from both populations are very similar in appearance to those from the type locality.

Variation.

Differences in scale counts and measurements are presented in Suppl. material 1. The coloration of the males THNHM 30911 and THNHM 30910 are similar to the holotype but their subcaudal coloration is a much more vibrant coral-red and they have complete original tails (Fig. 5 C, D ). The coloration of the yellow stripes and spots that composed the dorsal pattern, and the flank coloration are not as pronounced in the female specimens THNHM 30904–30906 and 30908 as they are in the males (Figs 6 , 9 ; Suppl. material 1). Additionally, the dorsal caudal pattern is only faintly visible in the female specimens (Figs 6 , 9 ).

Natural history.

Similar to other species of Leiolepis , Leiolepis glaurung sp. nov. constructs subterranean tunnels. However, given that they live exclusively in rocky habitats, individuals make compressed and shallow burrows in patches of loose soil underneath rocks or rockpiles (Figs 10 – 12 ). Males tend to forage during the hottest part of the day and the feces we found appeared to be mostly composed of vegetation and arthropods. Leiolepis glaurung sp. nov. is a food source for local people in the area (as many Leiolepis species are across Indochina; Grismer and Grismer 2010) and talking with local collectors, they report that individuals are highly philopatric. The collectors say that if the burrow is open that means they are out foraging and if the burrow is plugged with soil they are inside. Additionally, local people say that Leiolepis glaurung sp. nov. is mostly active during the dry season (November – March) and estivates during the rainy season (April – November) but will come out of their burrows after rains to eat arthropods. Lastly, all individuals observed and collected (specimens THNHM 30908–30911) were found during the day at 36 ° C and 77 % humidity. Other species of reptiles observed sharing the same habitat at the type locality included Calotes versicolor (Daudin, 1802) , Dixonius siamensis (Boulenger, 1899) , Gekko petricolus Taylor, 1962 , G. gecko (Linnaeus, 1758) , Scincella melanosticta (Boulenger, 1887) , S. rupicola (Smith, 1916) , Eutropis macularia (Blyth, 1853) , Chrysopelea ornata (Shaw, 1802) , Lycodon laoensis Günther, 1864 and Calloselasma rhodostoma (Kuhl, 1824) .

Figures

Figure 1. The distribution of all Leiolepis species and the localities of specimens used in this study. Question marks indicate an area where new populations could possibly occur. The different shades of the green circles for Leiolepis peguensis correspond to their phylogenetic positions in Fig. 2.

Figure 5. Dorsal and ventral views of the holotype and paratypes of Leiolepis glaurung sp. nov. A male holotype THNHM 30909 B female paratype THNHM 30908 C male paratype THNHM 30910, and D male paratype THNHM 30911.

Figure 6. Dorsal and ventral views of paratypes THNHM 30906, THNHM 30907, THNHM 30904, THNHM 30902, THNHM 30905, and THNHM 30903 Leiolepis glaurung sp. nov. Specimen tags bear field numbers.

Figure 7. Comparisons of the normalized morphometric characters showing the ranges, frequencies, mean (white dots), and 50 % quartiles represented by black bars.

Figure 8. Comparisons of statistically different meristic characters among well-supported clades of Leiolepis. White circles are means and the black horizontal bars are medians.

Figure 9. A Female paratype THNHM 30908 B male paratype THNHM 30911 of Leiolepis glaurung sp. nov.

Figure 10. The rocky habitat of the Khorat Plateau from the type locality.

Figure 11. A Male paratype THNHM 30911 displaying the dorsal and lateral display colors and its rocky habitat in the background B adult male Leiolepis rubritaeniata from southeastern Cambodia C male paratype THNHM 30911 of Leiolepis glaurung sp. nov.

Figure 12. A Two burrow entrances going beneath a large rock B multiple small burrow entrances in the soil underneath and around a rock pile C a single entrance hole to a burrow beneath a large rock D the rock from panel C lifted to show the shallow subterranean burrow of an individual Leiolepis glaurung sp. nov.