Pseudocalotes viserion, 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 : 480-484

Pseudocalotes viserion sp. nov.

Viserion’s False Garden Lizard Figs. 3 , 6 , 10

Holotype. Adult female LSUHC 12227 collected on 26 March 2015 by Evan S. H. Quah at 1000 hrs crossing the radar tower road at Ulu Kali at Genting Highlands, Pahang, Peninsular Malaysia (03° 26.166 N 101° 47.021 E; 1754 m in elevation).

Paratype. Adult male LSUHC 12141 found dead on the radar tower road at Ulu Kali at Genting Highlands, Pahang, Peninsular Malaysia (03° 26.12 N 101° 47.345 E; 1750 m in elevation) by L. Lee Grismer, Perry L. Wood, Jr., Hayden R. Davis, Matthew L. Murdoch, Brandon R. Burch, and Anthony J. Cobos.

Diagnosis. Pseudocalotes viserion sp. nov. can be separated from all other species of Psuedocalotes by having a combination of three postrostrals; 10 circumorbitals; four or five canthals; 5–7 superciliaries; rostral and nasal in contact; supralabials contacting nasal; six or seven supralabials; seven or eight infralabials; two or three postmentals; three enlarged chinshields; 47 or 48 smooth, flat, gular scales; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; 7–9 nuchal crest scales lacking gaps and not extending beyond midbody; weakly keeled and plate-like scales on flanks; 35–38 scales around midbody; ventrals smaller than dorsals; 22 or 23 subdigital lamellae on fourth finger; 26 or 27 subdigital lamellae on fourth toe; preaxial scales on third not modified; subdigital lamellae not unicarinate; HW/HL 0.62; no white marking below the eye; dewlap in males yellow; and no elbow or knee patches. These characters or a subset of them are scored across all species in Table 3.

Description of holotype. Adult female SVL 71 mm; head relatively large, triangular in lateral and dorsal profile; HW/HL 0.62; HL/SVL 0.30; interorbital and frontal regions flat; snout not elongate but convex, sloped anteriorly, canthus rostralis sharp, composed of four scales; supraorbital scales enlarged, keeled; single row of 10 enlarged, circumorbital scales bordering medial margins of supraorbital region, posteriormost of series enlarged and tuberculate; five, flat, imbricate superciliary scales; scales of orbit (surrounding eye) granular; three enlarged postorbital scales, posteriormost scale acuminate, keeled; dorsal, head scales weakly acuminate; rostral low, rectangular, bordered laterally by first supralabials, dorsolaterally by nasal scale, and posteriorly by three smaller scales; external nares in slightly elevated single, rectangular, large, nasal scale; three enlarged, acuminate, median scales in prefrontal region; parietal, occipital, and upper postorbital region covered with slightly enlarged, irregularly shaped, weakly acuminate scales; an enlarged, keeled parietal scale on right side of midline followed by a large, raised occipital scale; enlarged parietal on left side of midline broken up and adjacent occipital scale not as large or raised as the corresponding scale on right; interparietal small, bearing eyespot; temporal scales irregular in size; single enlarged, raised postorbital scale; two, raised, enlarged, keeled, posttemporal scale; tympanum naked, scales on dorsal margin not enlarged; six (R,L) supralabials; nine (R,L) postnasal-supraorbital scales; mental triangular; two large postmentals in medial point contact forming first in a series of three enlarged chinshields separated from infralabials by three anteriorly and 10 posteriorly rows of smaller scales; 7 (R), 8(L) infralabials; gular scales smooth, flat, directed posteromedially, 47 at midline; dewlap small; transverse gular and antehumeral folds weak.

Seven, large, lanceolate nuchal scales beginning on occiput, reaching maximum height on nape, tapering to a low, vertebral crest on body composed of a single row of keeled, slightly enlarged scales that extend onto base of tail where they are largest; body round in cross-section; dorsal body and flank scales weakly keeled, subimbricate, rectangular, arranged in indistinct, transverse rows; flank scales larger than dorsal body scales, plate-like; ventrals smaller than dorsals; scales of pectoral region, belly, and precloacal regions strongly keeled; limbs moderate in stature, covered dorsally with keeled scales; five digits on pes and manus; hind limbs larger, more robust than forelimbs; subdigital lamellae of toes I, II, IV, and V bicarinate; 22 lamellae beneath fourth finger; 27 lamellae beneath fourth toe; preaxial scales of third toe unmodified; tail laterally compressed, not swollen at base, covered with keeled scales, 1.68 times SVL.

Coloration in life ( Fig. 10 ). Dorsal ground color of body and tail dull yellow; labials, lateral regions of throat, proximal region of brachia and anterior portion of flanks white; ground color of top of head light-green, randomly mottled with small, irregularly shaped, dark markings; four very faint, wide, diffuse, dark vertebral patches between forelimb and hind limb insertions extending onto dorsal margins of flanks; ground color of forelimbs yellow-green, mottled with darkly edged scales; ground color of hind limbs dingy green, limbs faintly banded; wide, diffuse faint bands on anterior region of tail fading posteriorly; gular region beige, bearing a series of faint, diffuse, obliquely oriented, brown stripes in lateral margins; gular pouch yellow; subcaudal region gray; rest of venter beige, generally immaculate; bottoms of hands and feet slightly darker.

Variation ( Fig. 10 ). The male paratype (LSUHC 12141) resembles the female holotype (LSUHC 12227) in aspects of color pattern but the overall ground color is brown, not dull-yellow. It also has a greatly swollen tail base. Differences in scalation are presented in Table 5 .

Distribution. Pseudocalotes viserion sp. nov. is known only from the radar tower road at Ulu Kali at Genting Highlands, Pahang Peninsular Malaysia ( Fig. 2 ).

LSUHC LSUHC LSUHC LSUHC LSUHC ZRC FMNH Etymology. The specific epithet viserion refers to this species’ resemblance in form and color to the yellowish dragon, Viserion—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. The holotype (LSUHC 12227) was collected during mid-morning at 1000 hrs under broken cloud cover as it was crossing the radar tower road at Ulu Kali at Genting Highlands. The paratype (LSUHC 12141) was found freshly killed on the same road at 1500 hrs. This short stretch of road runs along the crest of Ulu Kali and winds through a mossy cloud forest between 1,700 and 1,800 m in elevation ( Fig. 11 ). The holotype was a gravid female that had just ovulated indicating that the reproductive season of this species extends through March.

Comparisons. Pseudocalotes viserion sp. nov. is readily differentiated from all other species of Pseudocalotes except it sister species P. flavigula by having enlarged, plate-like scales on the flanks. It can be separated from P. flavigula by having larger, plate-like scales on the flanks (compare Figs. 10 and 12 ) as evidenced by having fewer midbody scale rows (35–38 versus 41–44); having more gular scales (47 or 48 versus 40–46); fewer subdigital lamellae on the fourth finger (22 or 23 versus 22–28); and fewer subdigital lamellae on the fourth toe (26 or 27 versus 26–30) ( Table 5 ). There appears to be less yellow coloration in the gular region of P. f l a v i g ul a in that the yellow does not extend onto the region of the throat anterior to the forelimb insertions as it does in P. viserion sp. nov. and the skin between the scales is white in P. flavigula as opposed to being yellow in P. viserion sp. nov. (Figs. 10,12). Additionally, these two species share a 22% uncorrected pairwise sequence divergence between them. 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 3. Lateral view of the head of Pseudocalotes viserion sp. nov. (LSUDPC 9909) from Genting Highlands, Pahang, Peninsular Malaysia illustrating descriptive scale characteristics. Photograph by Kurt Orion.

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 10. Pseudocalotes viserion sp. nov. from the radar tower road, Genting Highlands, Pahang, Peninsular Malaysia. Left column: adult females LSUHC 12227 (upper, photograph by L. Lee Grismer), LSUDPC 9907 in light phase (middle, photo by Kurt Orion), and LSUDPC 9908 in dark phase (lower, photo by Kurt Orion). Right column (photographs by L. Lee Grismer): adult female LSUHC 9951 (upper), adult male LSUHC 9484 (middle and lower).

FIGURE 11. Microhabitat of Pseudocalotes viserion sp. nov. along the radar tower road at Ulu Kali at Genting Higlands, Panhang, Peninsular Malaysia. Photograph by L. Lee Grismer.

FIGURE 12. Pseudocalotes flavigula from Cameron Highlands, Pahang, Peninsular Malaysia. Upper left: Juvenile from Tanah Rata (LSUDPC 9902, photo by Zhaharil Dzulkafly). Middle left; Adult female from Brinchang Trail (LSUDPC 4350, photo by Leong Tzi Ming). Lower left and right: Adult male from Tanah Rata (LSUHC 12420, photo by L. Lee Grismer). Upper right and middle, respectively: Adult females from Tanah Rata (LSUHC 12581) and Brinchang Trail (LSUHC 12580); photos by Evan S. H. Quah.

Tables

TABLE 5. Diagnostic morphological and color pattern characters separating Psedocalotes flavigula and P. v i s e r i on sp. nov.

	12114 12227	12420	12580	12581	2.5162	143903
	paratype holotype viserion sp. nov.	flavigula
Postrostrals Interparietal	3 3 No Yes	5 No	6 No	4 No	7 No	/ No
Cicrcumorbitals Canthals	10 10 5 4	10 4	10 4	10 5	10 4	10 5
Superciliaries Scale between	7 5	5	6	7	7	7
rostral and nasal Supralabials contacting nasal	No No Yes Yes	No Yes	Yes Yes	No Yes	No Yes	/ Yes
Supralabials	7 6	6	7	6	6	7
Infralabials postnasal-suborbitals	7 7,8 8 9	7 9	7 8	6 8	7 /	8 9
Postmentals	2 3	3	3	4	3	3
Chinshields Gulars	3 3 48 47	4 46	5 40	6 40	5 /	6 42
Gulars smooth or keeled Gulars granular or flat Transverse gular fold	Smooth Smooth Flat Flat Weak Weak	Smooth Flat Weak	Weak Flat Weak	Smooth Flat Weak	Smooth Flat Weak	Smooth Flat Weak
Antehumeral fold	Weak Weak	Weak	Weak	Weak	Weak	Weak
Enlarged scales between eye and ear	2 3	2	3	2	3	3
Enlarged posttemporals Enlarged supratympanics	2 2 1 1	2 1	2 1	2 1	2 1	2 1
Enlarged postrictals Nuchal crest	No No 9 7	No 7	No 9	No 8	No 6	No 10
Gap in crest Enlarged crest scales	No No No No	No No	No No	No No	No No	No No
Beyond midbody
Flank scales keeled Flank scales plate-like	Weak Weak Yes Yes	Weak Yes	Weak Yes	Weak Yes	Weak Yes	Weak Yes
Scales around midbody Midventrals smaller than dorsals 4th finger lamellae	35 38 Yes Yes 23 22	41 Yes 27	44 Yes 25	41 Ye s 26	/ Yes 24,28	44 Yes 22
4th toe lamellae Preaxial lamellae	26 27 No No	30 No	27 No	30 No	/ No	28 No
modified on toe III HL/SVL	/ 0.3	0.31	0.3	0.29	/	0.1
TL/SVL Male dewlap color	1.91 1.68 Yellow /	1.91 Yellow	1.77 /	1.58 /	/ /	1.79 Yellow
Sex	Male Female	Male	Female	Female	Female	Male
SVL	7 9 7 1	7 9	7 6	7 7	8 0	7 3