Calotes paulus ( Smith, 1935 ) Giri & Chaitanya & Mahony & Lalrounga & Lalrinchhana & Das & Sarkar & Karanth & Deepak, 2019

Giri, Ad. B., Chaitanya, R., Mahony, Stephen, Lalrounga, Samuel, Lalrinchhana, C., Das, Abhijit, Sarkar, Vivek, Karanth, Praveen & Deepak, V., 2019, On the systematic status of the genus Oriocalotes Günther, 1864 (Squamata: Agamidae: Draconinae) with the description of a new species from Mizoram state, Northeast India, Zootaxa 4638 (4), pp. 451-484 : 460-464

Calotes paulus ( Smith, 1935) comb. nov.

Figs. 5 and 6 ; Table 2 and 5.

Calotes minor Gray, 1845:244 (partim: “Affghanistan” & “Khassia hill”).

–– Oriocalotes minor Günther 1864:147 (partim: “ Afghanistan ”, “Khasya” & “Sikkim”).

–– Oreocalotes [sic] minor Jerdon 1870:77 .

–– Charasia minor Theobald 1876:113 (partim: “ Sikkim ”).

–– Acanthosaura minor Boulenger 1885:304–305 (partim: “ Afghanistan ”, “Khasia” & “Sikkim”).

Oriocalotes paulus Smith, 1935:166–167 , 226.

Lectotype by present designation. Adult male, BMNH 1946.8.11.28 (previously BMNH xxiii.30.B), from “Affghanistan” (in error, likely to be Khasi Hills [in Meghalaya state, Northeast India], fide Günther 1864; type locality referred to as “Khasi Hills” by Smith 1935 without discussion), presented by the East India Company.

Paralectotypes by present designation. Adult male, BMNH 1946.8 .11.27 (previously BMNH xxiii.30.A), and juvenile, BMNH 1946.8 .11.29 (previously BMNH xxiii.30.C), from “Affghanistan” (see comment for lectotype), presented by the East India Company; adult male, BMNH 1946.8 .11.35 (previously BMNH xxiii.30.D), from “Khassia hill”, Meghalaya state, Northeast India, presented by the East India Company .

Additional material. Subadult female, NCBS-AQ 197, and adult female, NCBS-AQ 198, from Sohra (previously known as Cherrapunjee, 25°16′39.38′′N; 91°41′48.95′′E, 1370 m a.s.l.), East Khasi Hills District, Meghalaya state, Northeast India, collected by Vivek Sarkar on 27 October 2014; an adult female and juvenile, BMNH 70.11.29.43, from ‘Khassya’, Meghalaya state, Northeast India, presented by T.C. Jerdon; adult male, BMNH 60.3.19.1378, from“Himalaya” (given as “ Sikkim ” [Northeast India] in Boulenger 1885]), collected by Messrs. von Schlagintweit; adult male, BMNH 72.4.17.328, and adult female, BMNH 72.4.17.329, from unknown localities (given as “ Sikkim, [Northeast India] in Boulenger 1885]), presented by T.C. Jerdon; adult male, BMNH 60.3.19.1029, from “Afghan.?” (in error, likely to be “Khasya” [Meghalaya state, Northeast India], fide Günther 1864), presented by Dr. Griffith.

Diagnosis. A medium sized Calotes , snout to vent length averaging 57.5 ± 12.5 mm, and maximum to at least 71.0 mm. Body feebly compressed with a weak dorsal crest; scales on top of head highly heterogeneous, keeled, those above orbits are largest, scales surrounding parietal are more or less similar in size; three spines on each side of head, one above orbit, one above tympanum and one on temporal region on each side; dorsal pholidosis heterogeneous, composed of medium sized, weakly pointed, strongly keeled scales intermixed with similar but slightly larger scales which are distinct on flanks; upper rows directed backwards and upwards and a few lower rows directed backwards; 42–46 midbody scale rows; a weakly developed fold anterior to forelimb insertion having granular scales; tympanum small, typically covered by a single large scale; tail rounded; seven to nine supralabials and seven to nine infralabials; lamellae bicarinate, 18–23 on fourth finger and 20–26 on fourth toe. Presence of heterogeneous scales on dorsum and weakly developed dorsal crest of Calotes paulus comb. nov. distinguish it from all known congeners.

Description of lectotype (BMNH 1946.8.11.28). Adult male. Mensural and meristic data is summarised in Table 2. General habitus slightly laterally compressed. Head relatively long (HL/SVL 0.32), broad (HW/HL ratio 0.66), not depressed (HD/HL ratio 0.55), slightly broader than neck ( Fig. 5A ). Snout short (SE/HL ratio 0.39), longer than orbital diameter (OD/SE ratio 0.85). Orbit large (OD/HL ratio 0.33); pupil round. Snout bluntly pointed in profile ( Fig. 5C ), rostral rectangular, approximately two times longer (1.8 mm) than deep (0.8 mm), contacted by first supralabials and four scales dorsally. Nostril oval, laterally positioned, nasal large, pentagonal in shape, bordered by seven scales including supralabials one and two ( Fig. 5C ). Nine smooth rectangular supralabials, and seven infralabials. Loreal region slightly concave, scales of loreal region small, flat, dominated by a single row of enlarged scales extending from posterior edge of nasals, below orbits, to posteriormost supralabial on each side, enlarged scale row runs parallel with supralabials (though mostly separated from them by a single row of narrow scales; Fig. 5C ). A row of four enlarged longitudinally keeled temporal scales extending from posterior edge of orbit to above anterior edge of tympanum. Orbital scales small, rounded, granular. Tympanum covered by a single scale ( Fig. 5C ). Canthals enlarged, overlapping slightly. Scales on dorsal surface of snout are heterogeneous in size and shape, smaller scales weakly tubercular, larger scales with distinct keels. Supraorbital scales weakly keeled, increase in size from supraciliaries to inner edges of orbits, of which enlarged scales follow curvature of orbits posterolaterally. Two small sized scale rows divide enlarged scales of inner orbits at narrowest point of frontal ( Fig. 5B ). Posterior mid-dorsal surface of the head (surrounding parietal scale) is dominated by a patch of slightly enlarged scales. Parietal scale moderately enlarged, longitudinally subrectangular without visible pineal eye (but with distinct depression) ( Fig. 5B ). Temporal spine longer than nuchal spines, orbital spine shorter, and supra-tympanic spine small ( Fig. 5B ). Mental subtriangular, approximately as wide as long, bordered posteriorly by a ventrolateral row of five elongated chin shields gradually changing shape and size posteriorly to blend with surrounding rows of gular scales. First pair of post-mental chin shields in contact laterally with infralabials, remaining shields are separated from infralabials by one to two rows of small gular scales ( Fig. 5D ). Remaining gular scales keeled (increasingly posteriorly), subimbricate, mid-gular scales smallest, increasing in size slightly posterolaterally and again decreasing in size dorsally over jowls. No distinct gular pouch present, and transverse gular fold absent ( Fig. 5D ). Ventral scales imbricate, strongly keeled, heterogeneous in size and arranged in approximate rows, 45 scales around mid-body. Nuchal crest is composed of nine short laterally compressed spines, largest on mid-nuchal region, decreasing in size anteriorly and posteriorly. Remaining vertebral scales are enlarged relative to adjacent rows, and possess a more pronounced median keel providing a serrated appearance in profile, 28 mid-dorsal scales in total. Dorsal and lateral scales are heterogeneous in size and shape ( Fig. 5A ), all scales with a moderate median keel, arranged into approximate rows, keels on those of upper flanks are mostly oriented obliquely upward, horizontal on mid-flanks, and obliquely downward on lower flanks. A distinct shallow oblique fold in front of and curving around anterior forelimb insertion ( Fig. 5C ). Scales of forelimbs and ventral hindlimbs form approximate rows, those of dorsal hindlimbs do not form regular rows and are heterogeneous in size. Dorsal scales larger than ventral scales on forelimbs and hindlimbs, all moderately keeled dorsally, and very weakly keeled ventrally. Forelimbs moderately long and thin ( Fig. 5A ); hindlimbs long and thin. Digits slender, each with a strong, slightly curved claw. Lamellae entire, bicarinate, 23 on fourth finger, and 26 on fourth toe. Tail rounded in cross section, approximately twice as long as SVL (TL/SVL ratio 1.98), slightly enlarged posterior to cloaca; scales on dorsal and ventral surfaces of tail are regularly arranged, strongly keeled, imbricate. Precloacal and femoral pores absent.

Coloration (in preservative). Dorsal surface of head mostly pale grayish/yellowish-brown with dark brown speckling on snout ( Fig. 5B ); lateral surfaces of snout pale grayish-white with brown blotches on lips; broad brown stripe extends from below eye to slightly posterior of tympanum ( Fig. 5C ). Dorsal and lateral surfaces of anterior two thirds of body pale grayish-brown, posterior half mostly dark brown mottled with pale grayish-brown ( Fig. 5A ). Dorsal surfaces of upper forelimbs pale grayish-brown, lower forelimbs mostly dark brown mottled with pale grayish-brown; dorsal surfaces of hindlimbs mottled light and dark brown ( Fig. 5A ). Tail primarily pale grayish-brown, with faint darker brown bands. Ventral surfaces of head, body and limbs pale grayish-white, with faint irregular stripes radiating from the central gular region ( Fig. 5D ).

Coloration in life (based on specimens not collected). Head, body, limbs and tail mostly mid to dark brown, with darker brown mottling, and occasionally green mottling. Area on lateral snout white, or white and green. Refer to Figure 6 for finer details of markings and coloration.

Variation. Most specimens examined correspond well with description of lectotype. Females generally larger in size (female SVL 57.9–71.3 mm, N = 4 versus male SVL 55.5–66.6 mm, N = 5). When hind limb is extended anteriorly along body, it typically reaches between posterior edge of tympanum and posterior edge of orbit on all specimens except BMNH 72.4 .17.328 (reaches rear of jaw) and BMNH 72.4 .17.329 (reaches anterior forelimb insertion). Transition from nuchal crest spines to strongly keeled dorsal scales is not clearly defined on some specimens (e.g., BMNH 72.4 .17.328 and BMNH 60.3.19.1029). Tympanum covered by a single scale on all specimens, bar one ( BMNH 60.3.19.137), which possess one to two rows of smaller scales bordering a single large scale anteriorly. Spinose scales in general poorly developed on juveniles. Finer details of coloration and markings vary between individuals, with some specimens having a distinct broad dark brown transverse cross bar on posterior half of dorsum. Further details of morphometric and meristic variation are presented in Table 2 .

Suggested common name. Khasi Hills Forest Lizard.

Distribution and natural history. Calotes paulus comb. nov. is known with certainty only from the the Khasi Hills, Meghalaya state, Northeast India. The syntype series included three specimens with the associated collection locality “Affghanistan”, which has been considered in error by several authors (e.g., Günther 1864; Boulenger 1885), and as such Afghanistan has been omitted from the distribution of the species by others (e.g., Anderson 1879; Smith 1935). Additional specimens in the NHMUK collected by T.C. Jerdon and the Schlagintweit brothers, have the associated collection locality “Sikkim”. These collectors travelled to both the Darjeeling/Sikkim region and the Khasi Hills. Jerdon (1870) notes that he was aware that several of the localities given by the brothers Schlagintweit were erroneous “probably from the displacement of labels”. Furthermore, collection localities associated with some of Jerdon’s specimens from “Sikkim” have also been demonstrated to have been mislabeled for species otherwise found only in the Khasi Hills ( Biju et al. 2016; Mahony et al. 2018). Jerdon (1870) also admits to not having seen this species, despite presenting specimens to the BMNH (NHMUK) soon after. By 1870, Jerdon had completed his tour to Sikkim, thus it is most likely that his specimens were collected from the Khasi Hills postscript. Smith (1935) clearly questioned Sikkim in the distribution of this species, as do we, since it has not been reported since these early specimens. References to the presence of this species in Xizang (Tibet), China by Zhao and Adler (1993) and Li et al. (2010, and references therein) require further confirmation.

Natural history observations of Calotes paulus comb. nov. were made by one of the authors (VS) over a period of one year at Sohra (previously known as Cherrapunjee), on the Shillong Plateau, East Khasi Hills District, Meghalaya state, Northeast India ( Fig. 1 ). The remaining natural habitat in the vicinity of Sohra predominantly consists of grasslands with isolate trees and small patches of tropical moist evergreen forest ( Champion & Seth 1968) on hill slopes and around perennial hill streams ( Fig. 7 ) Calotes paulus comb. nov. is one of the common agamids found in this area, where it has been observed along the fringes of forests, or on isolated tree trunks or large bushes. An adult female (BMNH 70.11.29.43) contained six large eggs (two measured, 14.1 X 8.2 mm and 14 X 8.5 mm). One individual of Calotes paulus comb. nov. was observed feeding on a cicada ( Gudaba sp.).

Figures

FIGURE 1. Map showing type locality of Calotes paulus comb. nov. and Calotes zolaiking sp. nov. in North East India.

FIGURE 5. Lectotype of Calotes paulus comb. nov. (BMNH 1946.8.11.28). A. Dorsal view of body. B. dorsal view of head, C. lateral view of head, D. ventral view of head. Scale bar: 10 mm.

FIGURE 6. Calotes paulus comb. nov. showing live coloration, photographed in-situ from Sohra, East Khasi Hills District, Meghalaya state, Northeast India.

FIGURE 7. Habitat where Calotes paulus comb. nov. were found in Sohra, East Khasi Hills District, Meghalaya state, Northeast India.