Pseudocalotes rhaegal, Grismer, L. Lee, Quah, Evan S. H., Wood, Perry L., Anuar, Shahrul, Muin, Abdul, Davis, Hayden R., Murdoch, Matthew L., Grismer, Jesse L., Cota, Michael & Cobos, Anthony J., 2016

Grismer, L. Lee, Quah, Evan S. H., Wood, Perry L., Anuar, Shahrul, Muin, Abdul, Davis, Hayden R., Murdoch, Matthew L., Grismer, Jesse L., Cota, Michael & Cobos, Anthony J., 2016, Dragons in the mist: three new species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the sky island archipelago of Peninsular Malaysia, Zootaxa 4136 (3), pp. 461-490 : 477-479

Pseudocalotes rhaegal sp. nov.

Rhaegal’s False Garden Lizard Figs. 5 , 6 , 8 , 9

Holotype. Adult female LSUHC 12178 collected on 18 March 2015 by L. Lee Grismer, Evan S. H. Quah, Shahrul Anuar, Mohd A. Muin, Perry L. Wood, Jr., Hayden R. Davis, Matthew L. Murdoch, Brandon R. Burch, and Anthony J. Cobos at 2030 hrs at Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia (04° 43.283 N 101° 23.129 E; 1411 m in elevation).

Paratypes. Adult female LSUHC 12179 bears the same locality collecting data as the holotype. Adult female LSUHC 12000 bears the same collecting locality and collectors but was collected on 4 September 2014.

Diagnosis. Pseudocalotes rhaegal sp. nov. is differentiated from all other Psuedocalotes by having the combination of a convex rostrum; 6–8 postrostrals; an interparietal; nine or 10 circumorbitals; five canthals; 7–10 superciliaries; one or two scales between the rostral and nasal scales; eight or nine supralabials; seven or eight infralabials; 11 or 12 postnasal-suborbital scales; four postmentals; 4–6 chinshields; 40–45 smooth, wide, gular scales; no transverse gular fold; weak antehumeral fold; three or four enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; no large scales bordering the upper margin of the ear opening or in the pretympanic region; 6–8 enlarged nuchal crest scales not separated by a gap; enlarged vertebral scales extending to base of tail; weakly keeled, non-plate-like scales on flanks; 52–58 scales around midbody; midventrals smaller than dorsals; 19–21 subdigital lamellae on fourth finger; 22–26 subdigital lamellae on fourth toe; preaxial scales on third toe enlarged and rounded; subdigital lamellae not unicarinate; HW/HL 0.50–0.54; HL/SVL 0.28–0.30; no elbow or knee patches; and female dewlap bearing a purple base. These characters or a subset of them are scored across all species in Table 3.

Description of holotype. Adult female SVL 83.7 mm; head relatively large subtriangular in lateral and dorsal profile; HW/HL 0.54; HL/SVL 0.30; interorbital region flat; rostrum convex, sloped anteriorly, canthus rostralis sharp, composed of five large canthal scales; supraorbital scales slightly enlarged, keeled; single row of nine or 10 enlarged, circumorbital scales bordering medial margins of supraorbital region; eight flat, imbricate to subimbrictae, superciliary scales; scales of orbit (surrounding eye) granular; short row of three (R) four (L) enlarged, acuminate, postorbital scales extend to above anterior margin of tympanum; dorsal, head scales acuminate; rostral low, rectangular, bordered laterally by first supralabials, posteriorly by eight smaller scales; external nares in slightly elevated, rectangular, large, nasal scale; seven enlarged, acuminate, median scales in prefrontal region forming a Y-shaped series; parietal and occipital regions covered with enlarged, elevated, acuminate to keeled scales; an enlarged parietal scale on each side of midline; interparietals small, irregularly shaped, centralmost bearing a small eyespot; temporal scales unequal with small scales intermixed with enlarged, weakly keeled scales; one, enlarged, keeled, scale in temporal region followed by a single, enlarged, spinose, posttemporal scale; tympanum naked, not bordered dorsally by large supratympanic scales; nine (R,L) supralabials; 12 (R), 11 (L) large, postnasal-suborbital scales; mental triangular, larger than adjacent infralabials; four larger postmentals, separated medially by two small scales and forming first of a series of five (R) four (L) enlarged chinshields separated from infralabials by one anteriorly and two posteriorly rows of smaller scales; seven (R,L) infralabials; gular scales wide, smooth, directed posteromedially, 40 at midline; dewlap small; transverse gular and antehumeral folds weak.

Nuchal crest composed of six, large, contiguous, lanceolate scales beginning on occiput, reaching maximum height on nape, tapering to a low, vertebral crest on body composing a single row of slightly enlarged, keeled vertebral scales extending to base of tail; body laterally compressed; dorsal body and flank scales weakly keeled, subimbricate, rectangular, arranged in indistinct, transverse, and posteroventrally oriented rows; flank scales larger than dorsal body scales; ventrals same size as dorsals; 52 scales around midbody; scales of pectoral region, belly, and precloacal regions strongly keeled; limbs slender, covered dorsally with keeled scales; five digits on pes and manus; hind limbs larger, slightly more robust than forelimbs; subdigital lamellae of toes I, II, IV, and V bicarinate; preaxial lamellae on toe III enlarged and rounded ( Fig. 5 ); 20 lamellae beneath fourth finger; 25 lamellae beneath fourth toe; and tail laterally compressed, slightly swollen at base, covered with keeled scales but lacking an enlarged vertebral row, 1.67 times SVL.

Coloration in life ( Fig. 9 ). Dorsal ground color of head, body, and limbs, brown, mottled with shades of slightly darker brown; linear patch of light-colored suborbital scales beneath eye; four very faint, diffuse, dark bands between forelimb and hind limb insertions not extending to base of flanks; lighter interspace between bands mottled; scattered greenish scales in postorbital, temporal, lower flank, and pelvic regions; scales of forearms greenish with dark bands; seven dark caudal bands anteriorly separated by dull-green interspaces, dark bands fade on posterior one-third of tail; gular region beige with irregularly shaped, brown, thin, submandibular bands; gular pouch small, base of pouch cyan with purple center.

Variation ( Fig. 9 ). LSUHC 12179 very closely approximates the holotype in overall body coloration and pattern, except for being lighter overall. LSUHC 12000 bears the same general color pattern as the holotype and LSUHC 12178 but differs greatly from them in being overall more green in coloration and having four dark, diffuse, dorsal bands between the forelimb and hind limb insertions that are more prominent. Additionally, the tail is distinctly banded for nearly its entire length. Variation in squamation is presented in Table 4 .

Distribution. Pseudocalotes rhaegal sp. nov. is known only from Robinson Falls, Cameron Highlands, Pahang Peninsular Malaysia ( Fig. 2 ) but is expected to range more widely along the Banjaran Titiwangsa.

Etymology. The specific epithet rhaegal refers to this species’ resemblance in form and color to the greenish dragon, Rhaegal—one of three dragons born in the Dothraki Sea and commanded by Daenerys Targaryen—the Mother of Dragons—in George R. R. Martin’s fictional work Game of Thrones.

Natural history. All Pseudocalotes rhaegal sp. nov. were collected at night between at 2000 and 2300 hrs while sleeping on thin, horizontal branches of small trees in the vicinity of a river flowing through the hill dipterocarp forest at Robinson Falls ( Fig. 9 ). All were gravid females collected during mid-March or mid- September, indicating that this species may breed year-round.

Comparisons. Pseudocalotes rhaegal sp. nov. can be differentiated from all other species of Pseudocalotes by having a cyan dewlap with a purple center (females) and the unique combination of numerous other characteristics ( Table 3). With the exception of Pseudocalotes flavigula and P. viserion sp. nov., P. rhaegal sp. nov. most closely resembles the other geographically proximate Thai-Malay Peninsula species P. khaononensis , P. larutensis , P. drogon sp. nov., and P. dringi . Pseudocalotes rhaegal sp. nov. can be differentiated from P. flavigula and P. viserion sp. nov., by several characteristics the most notable of which is the lack of enlarged, plate-like scales on the flanks (Figs. 9,10). From Pseudocalotes khaononensis , P. rhaegal sp. nov. can be differentiated by having a maximum SVL of 85.2 mm versus 104.5 mm; five as opposed to six canthals; seven or eight as opposed to nine infralabials; smooth as opposed to acuminate gular scales; three or four versus one enlarged scale between the eye and the ear; enlarged vertebral scales extending beyond midbody as opposed to not extending beyond midbody; 52–58 as opposed to 72–75 scales around midbody; ventral scales being smaller as opposed to being larger than dorsal scales; 22–26 as opposed to 27 subdigital lamellae on the fourth toe; having as opposed to lacking enlarged, rounded, preaxial lamellae on the third toe; and having a cyan dewlap with a purple center (female) versus a purple dewlap in males ( Fig. 8 ). From P. larutensis , P. rhaegal sp. nov. can be differentiated by having nine or 10 versus 11–14 circumorbitals; 40–45 versus 55–69 gulars; smooth as opposed to weakly keeled gular scales; enlarged vertebral scales extending beyond midbody as opposed to not extending beyond midbody; preaxial scales on third toe bearing enlarged, rounded, scales versus being unmodified; having a cyan dewlap with a purple tip (females) versus a yellow dewlap with a purple, horizontal, centrally positioned marking ( Fig. 8 ). From P. dringi , P. rhaegal sp. nov. is separated by having a maximum SVL of 85.2 mm versus 70.3 mm; 6–8 versus five postrostrals; four as opposed to two postmentals; a weak, antehumeral fold as opposed to lacking a fold; having three or four versus two enlarged scales between eye and ear; enlarged vertebral scales extend beyond midbody as opposed to not extending beyond midbody; having as opposed to lacking enlarged, spinose, preaxial lamellae on the third toe; 52–58 scales around midbody versus 48–52; 23 versus 26 fourth toe subdigital lamellae; and having a cyan dewlap with a purple tip (females) versus a purple dewlap in males ( Fig. 8 ). From P. drogon sp. nov., P. rhaegal sp. nov. can be differentiated by having nine or 10 as opposed to 11 circumorbitals; 11 or 12 as opposed to 10 postnasalsuborbitals; 40–45 as opposed to 47 gular scales; a row of three or four versus two enlarged scales between the ear and eye; lacking as opposed to having three large supratympanic scales; pretympanic scales small as opposed to large; a convex versus a flat rostrum; lacking versus having enlarged vertebral scales on the tail; 52–58 versus 51 midbody scales; having enlarged, rounded, preaxial scales on the third toe as opposed to enlarged preaxial scales on the third toe being spinose; shorter snout (HL/SVL = 0.28–0.30 versus 0.31); and having a white patch of suborbital scales as opposed to lacking a white patch of suborbital scales ( Table 4 ). Differences from other species are listed in Table 3.

Figures

FIGURE 2. Distribution of the species of Pseudocalotes on the Thai-Malay Peninsula. Starred circles are type localities. Pseudocalotes khaonanensis from Kaho Nan National Park, Nakkon Si Thammarat Province, Thailand. Pseudocalotes sp. nov. from the Hala-Bala Wildlife Sanctuary, Yala Province, Thailand. Pseudocalotes dringi from the type locality at Gunung Tahan, Pahang and Gunung Lawit, Terengganu, Peninsular Malaysia. Pseudocalotes larutensis from Bukit Larut, Perak, Peninsular Malaysia. Pseudocalotes rhaegal sp. nov. from Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia. Pseudocalotes drogon sp. nov. from Fraser’s Hill, Pahang, Peninsular Malaysia. Pseudocalotes flavigula from Gunung Brinchang, Cameron Highlands, Pahang, Peninsular Malaysia. Pseudocalotes viserion sp. nov. from Genting Highlands, Pahang, Peninsular Malaysia.

FIGURE 5. Lateral view of preaxial scales on the third toe of the left hind limb in P. drogon sp. nov. (LSUHC 12223), P. rhaegal sp. nov. (LSUHC 12178), and P. flavigula (LSUHC 12420) showing the degrees of modification. Photos by L. Lee Grismer.

FIGURE 6. Maximum Likelihood tree topology showing the relationships of the new species Pseudocalotes drogon sp. nov., P. rhaegal sp. nov., and P. viserion sp. nov. to selected other species of Pseudocalotes. Numbers at the nodes are Bayesian posterior probabilities (BI) / Maximum Likelihood (ML) bootstrap values.

FIGURE 8. Dewlap coloration in male Pseudocalotes drogon sp. nov. (LSUDPC 9497; photograph by L. Lee Grismer), P. larutensis (LSUDPC 9400; photograph by L. Lee Grismer), P. khaonanensis (LSUDPC 9910; photograph by Micheal Cota), and female P. rhaegal sp. nov. (LSUDPC 9488; photograph by Mohd. A. Muin).

FIGURE 9. Type series of Psedocalotes rhaegal sp. nov. from Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia and habitat at the type locality. Upper left: LSUHC 12179, photograph by L. Lee Grismer. Upper right: LSUHC 12000, photograph by L. Lee Grismer. Lower left: LSUHC 12178, photograph by Mohd A. Muin. Lower right: Montane forest microhabitat at Robinson Falls, Cameron Highlands, Pahang.