Calotes vultuosus comb. nov. (Harlan, 1825)

Gowande, Gaurang, Pal, Saunak, Jablonski, Daniel, Masroor, Rafaqat, Phansalkar, Pushkar U., Dsouza, Princia, Jayarajan, Aditi & Shanker, Kartik, 2021, Molecular phylogenetics and taxonomic reassessment of the widespread agamid lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) across South Asia, Vertebrate Zoology 71, pp. 669-696 : 669

Calotes vultuosus comb. nov. (Harlan, 1825)

Figs 6 , 7

Agama vultuosa Harlan 1825: 296

Agama indica Hardwicke and Gray 1827: 217

Calotes cf. versicolor versicolor Manthey 2008: 88 (in part)

Calotes cf. versicolor - Pal et al. (2018) (in part)

Holotype.

INDIA • 1 ♂; West Bengal, Kolkata; 20.55°N, 88.36°E, 10 m a. s. l.; Dr R. Coates leg.; ANSP 7296

Other material (morphological vouchers). All from INDIA • 1 ♂, 1 ♀; Maharashtra, Satara, Koyna WLS; Soman and Thakar leg; BNHS 356 (both); • 1 ♂; Maharashtra, Solapur, Kurduwadi; 18.09°N, 75.43°E, 516 m a. s. l.; BNHS 368; • 2 ♂; Maharashtra, Satara, Thoseghar; 17.58880°N, 73.83805°E, 1119 m a. s. l.; 06 Jun. 2015; Gaurang Gowande and Pushkar Phansalkar leg.; GenBank MW901288-89 (16S); CESL 1004-05; • 1 ♂; Maharashtra, Satara, Patan; 17.35397°N, 73.80288°E, 587 m a. s. l.; 10 Jun. 2015; Gaurang Gowande and Pushkar Phansalkar leg.; GenBank MW901290 (16S); CESL 1007; • 1 ♂; Maharashtra, Pune, Taleghar; 19.04670°N, 73.55370°E, 925 m a. s. l.; 13 Jun. 2015; Gaurang Gowande and Pushkar Phansalkar leg.; GenBank MW901255 (16S); CESL1009; • 2 ♂; Maharashtra, Pune, Waghapur; 18.39990°N, 74.12400°E, 762 m a. s. l.; 11 Oct. 2015; Gaurang Gowande and Pushkar Phansalkar leg.; CESL 1042-43; • 1 ♂; Maharashtra, Chiplun, Guhagar; 17.49495°N, 73.18475°E, 11 m a. s. l.; 24 Oct. 2016; Gaurang Gowande and Pushkar Phansalkar leg.; CESL 1036; • 1 ♂; Rajasthan, Pali; 26.82485°N, 71.85335°E, 230 m a. s. l.; 2 Jul. 2017; Vishal Varma leg.; GenBank MW901284 (16S); CESL 1041 • 1 ♂; Maharashtra, Mumbai, Bandra; 8 May 1959; H. Abdulali leg.; BNHS 350 • 1 ♂; Karnataka, Kadra; Dec. 1914; S. Prater leg.; BNHS 354 • 1 ♂; Maharashtra, Mumbai; Sir Norman Kinnear leg.; BNHS 343 • 1 ♂; Gujarat, Kutch; P. Soman leg.; BNHS 316 • 1 ♂; Maharashtra, Aurangabad; Gaurang Gowande and Pushkar Phansalkar leg.; CESL 1065 • 3 ♂; Telangana, Hyderabad; 14 Apr. 2017; Gaurang Gowande and Pushkar Phansalkar leg.; GenBank MW901282 (16S); CESL 1068-70 • 1 ♂; Maharashtra, Latur; 16 Apr. 2017; Gaurang Gowande and Pushkar Phansalkar leg.; CESL 1071 • 1 ♀; Maharashtra, Mumbai, Borivali; 6 Jun. 1971; BNHS 1027.

Genetic diagnosis.

The least within species divergence at 16S and COI was 0%, while the greatest within species distance recorded was 2.0% and 9.5% at 16S and COI respectively. The species was at least 3.3% and 13.0% divergent from C. versicolor , 2.7% and 12.8% divergent from C. irawadi , and 3.9% and 15.5% divergent from Clade4 at 16S and COI respectively (Table 2 ). The species differed from C. calotes by sequence divergence of > 5% at 16S and 16.3% at COI from C. calotes . Phylogenetically, the species was recovered as sister to a clade comprising of C. calotes , C. versicolor , and Clade4 (Fig. 3 ).

Diagnosis and comparison.

A medium to large sized species of Calotes , adult males averaging 106 mm in SVL, females averaging 78 mm in SVL; body compressed; head relatively short; dorso-lateral scales posterodorsally oriented, large, weakly to strongly keeled, homogeneous; ventral scales smaller than the dorso-lateral scales, strongly keeled; 37-45 scales around the mid-body; anti-humeral fold absent; two distinct spines in the supratympanic region, posterior spine as long as the anterior spine, at times longer, more prominent; nuchal and dorsal crest continuous, distinct, slightly recurved; scales of the nuchal crest large, those of dorsal crest reduced to mere denticulation towards the base of the tail, generally ending in the region between the mid-body and the tail, rarely continues to the base of the tail; nuchal, dorsal and supratympanic spines more pronounced in males; limbs slender, dorsal surface of the limbs strongly keeled, ventral surface weakly keeled, that of the thighs smooth.

The species can be separated from the members of Smith’s C. versicolor group (as defined above), by a combination of characters: absence of crescent-shaped patch of granular scales at the insertion of the forelimbs (vs. present in C. emma , C. grandisquamis , C. jerdoni , C. mystaceus , and C. nemoricola ), 37-45 Mid-body scale rows (vs. 49-65 in C. emma , 27-35 in C. grandisquamis , 45-57 in C. jerdoni , 58-63 in C. maria Gray, 48-60 in C. minor , and 45-58 in C. mystaceus ); nuchal and dorsal crest scales well developed, nuchal crest scales slightly larger than the dorsal crest scales, dorsal crest scales become progressively smaller towards the base of the tail (vs. nuchal spines much longer, dorsal spines reduced in C. maria and C. nemoricola ; nuchal spines much longer than dorsal spines in C. calotes , C. emma , C. grandisquamis ); two well-separated supratympanic clusters of spine-like scales, one from each cluster enlarged, prominent to form a spine (vs. row of 3-4 compressed supratympanic spines in C. grandisquamis and C. nemoricola , 8-9 compressed spines above tympanum in C. calotes ; two parallel rows of supratympanic scales in C. jerdoni and C. maria ). The species differs from C. paulus and C. zolaiking primarily by the homogeneous scalation on the dorsolateral region (vs. heterogeneous) and a comparatively well-developed dorsal crest. From the dubious species C. bhutanensis , the species differs in possessing longer head, concave orbital region, and by the absence of a row of erect scales on the sides of the neck. From C. chincollium , C. nigriplicatus , and other members of the C. mystaceus complex ( C. bachae , C. geissleri , C. goetzi , C. mystaceus , C. vindumbarbatus , sensu Wagner et al. (2021)) the species differs by the absence of an oblique fold of skin in front of forelimbs or shoulder (vs. present). From the Sri Lankan congeners ( C. ceylonensis , C. desilvai , C. liocephalus , C. liolepis , C. manamendrai , C. nigrilabris , C. pethiyagodai ) the species differs by its posterodorsal orientation of lateral body scales (vs. posteroventral) and absence of shoulder pit (vs. present). The species differs from C. irawadi by its much larger adult male body size (average SVL 106 mm, vs. 82.4 mm), lesser number of scales around the mid-body (average 42, vs. 47 in C. irawadi ); from C. htunwini by the posterodorsal or vertical orientation of scale rows on the sides of the neck and supra-axillary area (vs. horizontal in C. htunwini ). For comparison with the subspecies elevated to species rank in this communication, see the diagnosis and comparison section for that species.

The species can be further differentiated by its southern congener C. versicolor by its slightly smaller adult body size (average male SVL 106 mm vs. 108 mm in C. versicolor , female SVL 77.5 vs. 92.2 mm in C. versicolor ), dorsal crest composed of comparatively smaller scales, which become progressively smaller to the base of the tail in both sexes (vs. dorsal crest composed of large scales, which continues to the base of the tail in C. versicolor ), supratympanic spines shorter in both sexes (vs. longer in C. versicolor ). The species has shorter crus than C. versicolor (average male CrusL 22.7 vs. 26.6 in C. versicolor ). The species differs in the overall shape of the trunk, which tapers to a lesser extent in C. vultuosus comb. nov. (average PectW/PelvW 0.81) vs. trunk tapers to a greater extent in C. versicolor (average PectW/PelvW 0.70 in C. versicolor ).

Besides, the species differs from C. versicolor in adult male coloration during the breeding season. Calotes vultuosus comb. nov. males generally attain a cream to brown body coloration, the head and the anterior two-thirds of the trunk attain orange color, which at times extends to the forelimbs; the posterior parts of the trunk and the hind limbs remain dull, whereas C. versicolor males attain yellowish overall coloration, the trunk and the orbital region turns bright orange, forelimbs and hind limbs turn dark to black. Further, the black patches under the throat extend anteriorly onto the jaw musculature, at times running along the lower jaw margin on each side, before terminating posterior to the post-mental scales, whereas, in C. versicolor , the black patches under the throat do not extend anteriorly onto the jaw muscles.

Description of the holotype ANSP 7296 (based on photographic data, Fig. 6).

A medium sized male, SVL ~77 mm. Tail complete, TailL ~160 mm. The specimen is in a damaged condition, there are multiple openings on the mid-body and near the vent, all artefacts of preservation. Further, the specimen has developed randomly distributed white patches on the scales of the head and the body. The description is based on the images of the specimen and modified from the original description; hence the mensural details should be treated with caution.

Head moderately large (HL/SVL ~0.19), snout tip blunt, rounded in dorsal perspective; loreal region between the nasals and the orbit slightly concave, acute, covered by heterogeneous, juxtaposed scales, some of which are ridged; the area comprising the loreal region and nasal shield triangular from the lateral perspective; eight CanthR scales, elongate, with their ends overlapping; supraciliary and canthal edge sharp, giving the head a flat appearance laterally from the dorsal perspective; large supraocular scales do not form shields, become parallel to convergent at the supraocular region; edges of the head divergent bordered by scales of the canthus rostralis and supraciliary; rostral broader than high; nasal shields single on each side, subtriangular, pointed anteriorly, rounded posteriorly, separated from the first Suplab by one prenasal scale, from rostral by two scales, from each other by seven SnS; nostrils round, in a single large nasal shield each, centrally placed; scales between rostral and SnS small, juxtaposed; SnS heterogeneous, the median SnS smallest, the pair bordering the median scale elongate; scales of the forehead posterior to SnS sub-imbricate, very irregular in shape and size, some rugose; HeadSLn 12 bordered anteriorly by the rostral scale, posteriorly by a single, large interparietal shield; orbit encased in a sock of granular scales, separated from the nasal shield by 6-8 scale rows, from Suplab by 3-4 scale rows; HeadSTr 14, between the posteriormost supraciliary scales on each side, just anterior to the interparietal; eye opening bordered dorsally and ventrally by two rows of non-granular scales, outer row composed of larger, square-shaped scales, inner row similar in shape, slightly smaller; eyelid scales (Eyelid) 13; pupil round, large; region between the orbit and the tympanum covered by rows of 6-9 smooth, roughly hexagonal scales; tympanum large, round, naked; its greatest diameter roughly 42% of horizontal diameter of the orbit; supratympanic scales weakly keeled; two enlarged supratympanic spines, subequal in size, separated from each other by four scales; anterior spine slender, bent parallel to the horizontal plane, posterior spine erect, prominent, its apex bent; scales in a row connecting the enlarged supratympanic spines give the posterior region of the head a serrated appearance; posterior region of the jaws swollen, bulging out, covered by subtriangular, mucronate, imbricate, postero-ventrally directed scales, upper border of the jaw muscles conceal the lower portion of the tympanum; labial scales large, sub-rectangular; Suplab 10; Inflab 10; two parallel rows of scales border the upper margin of the Suplab, lower originates above the second Suplab, separating the nasal shield and the second Suplab, terminates near the last Suplab; the upper row originates slightly posteriorly, terminates abruptly above the sixth Suplab; interparietal large, irregularly pentagonal, posterior border straight, tapers toward the anteriority, anterior border nearly pointed; interparietal bordered by 10 smooth, heterogeneous scales; nuchal crest starts 3-4 scales behind the interparietal; mental shield large, single, subtriangular, narrower than rostral; two pairs of elongate postmentals, anterior pair joint, in contact with mental, narrow; posterior pair broader, separated by two small chin scales; chin scales behind the postmental scales small, progressively become larger towards the throat, median row of gular scales enlarged, mucronate, and form a longitudinal fold.

First nuchal scale smallest, size of the nuchal scale progressively increases towards the median nuchal scale, nuchal scales beyond the median nuchal scale roughly the same size as, or slightly smaller than the median nuchal scale; nuchal crest composed of ~10 long, conical spines, scales recurved, continue into dorsal crest scales; dorsal crest composed of spines that progressively become smaller towards the middle of the back; continues as an obsolete row of vertebral scales to the base of the tail, beyond which the mid-dorsal crest row terminates at the keeled tail scale rows; paravertebral scales recurved, strongly keeled, mucronate; dorso-lateral scales sub-triangular, imbricate, moderately keeled, posterodorsally oriented; those in the supra-axillary region mostly dorsally oriented, slightly smaller than those of the mid-body; body at the mid-body to the pelvic region damaged; ventrals strongly keeled, smaller than the dorso-lateral scales, mucronate, imbricate; ventral scales between the insertion of the forelimbs weakly keeled, some with rounded apices.

Limbs long and slender, covered with keeled scales, similar to dorso-laterals in shape and size, forming parallel longitudinal rows; scales on the dorsal surfaces of the thigh weakly keeled, ventral surfaces smooth, those on the crus and sole strongly keeled; hindlimbs longer, almost 80% of SVL; relative length of fingers 4>3>2>5>1, fourth slightly longer than third; relative length of toes 4>3>5>2>1; subdigital lamellae bicarinate, keels sharp; digits and subdigital lamellae slender, digits swollen at base; tail damaged ventrally near the vent, exposing the hemipenis; tail scales large, imbricate, strongly keeled on dorsal and ventral aspects, mucronate; TailL ~160 mm.

Coloration in preservation: Dorsum generally dark brown, venter paler, light yellow; head dark brown dorsally, throat paler; orbit whitish, nasal shield, canthals and supraciliary region almost white; irregularly distributed white patches on the labials, throat, jaw muscles, axillary region, on the scales between the tympanum and the ear, the supratympanic region, and on the nuchal crest scales; the white patches continue on to the upper arm, lower arm, dorso-laterals, supraxillary region, and on the dorsal crest; white patches also present on the venter, the hindlimbs and the tail; tail scales show marks of corrosion due to preservative near the vent.

Original description.

Harlan R (1825). Journal of the Academy of Natural Sciences of Philadelphia 4(1): 296-305.

Etymology.

The specific epithet ' vultuosa ' could refer to the grim or frowning look of the species when viewed frontally, which it gets due to the flattened scales of the CanthR and the supercilium. The specific epithet Calotes vultuosa is feminine singular and is here changed to the masculine gender Calotes vultuosus in agreement with the generic epithet, Calotes , which is masculine in gender.

Variation in the material examined.

Eighteen male specimens were examined. The means for the mensural characters in mm are: HL 21.2; HW 25.3; HH 19.9; JawW 19.9; SnEye 10.9; NarEye 5.7; EyeEar 8.6; SnW 6.5; Interorb 11.1; SVL 106.0; TrunkL 47.1; TailH 16.4; TailW 13.1; PectW 18.4; PelvW 15.1; SnForeL 41.2; UpArmL 19.8; LoArmL 19.5; ForefL 19.2; 4FingLng 12.7; UpLegL 25.1; CrusL 23.1; HindfL 30.8; 4ToeLng 19.0; ForeLimbL 58.5; HindLimbL 79.1. The means for meristic characters are: SnS 7; HeadSTr 13; HeadSLn 14; CanthR 7; Eyelid 12; Suplab 11; Inflab 11; TempSp 2; Dorsal 48; Mid-body 42; 4FingLm 21; 4ToeLm 26. Additionally, two female specimens were examined. The means for the mensural characters in mm are: HL 17.4; HW 13; HH 12.4; JawW 13; SnEye 8.1; NarEye 4.1; EyeEar 4.8; SnW 4.5; Interorb 8.7; SVL 77; TrunkL 36.8; TailH 6.3; TailW 5.9; PectW 11.5; PelvW 9.6; SnForeL 28.9; UpArmL 16.6; LoArmL 13.5; ForefL 14.7; 4FingLng 10.7; UpLegL 17.6; CrusL 17.9; HindfL 23.7; 4ToeLng 16.3; ForeLimbL 44.3; HindLimbL 59.3. The means for meristic characters are: SnS 6; HeadSTr 13; HeadSLn 13; CanthR 7; Eyelid 10; Suplab 10; Inflab 10; TempSp 2; Dorsal 45; Mid-body 40; 4FingLm 18; 4ToeLm 25. The ranges for each of these characters are given in Table 4 .

Distribution.

The species appears to be widely distributed, occupying parts of the Dharwar Craton till Brahmagiri, Deccan Volcanic Province, the Central Highlands, the Gangetic Plains, and the Indian Deserts (Fig. 1 ).

Figures

Figure 1. A map of south and Southeast Asia representing the localities from which morphological and / or primary or secondary genetic data were collected. Red circles indicate the localities from which C. versicolor sensu stricto (Clade 1) originate, yellow samples indicate those of C. irawadi sensu lato (Clade 2), blue circles represent the localities of C. vultuosus comb. nov. (Clade 3), green circles indicate Clade 4, with C. farooqi stat. nov. (77, 83, 85, 86) and C. aff. farooqi (78, 79). A single sequence of Calotes irawadi from Huanan Province, China (locality 73) is not marked on the map, since it was very distant to all other localities, as also those whose precise localities were unknown. Triangles indicate type localities of the respective species. See supplementary Table S 1 for details on molecular and / or morphological vouchers used, and their respective localities.

Figure 3. Maximum Likelihood 16 S gene tree based on the ~ 416 base pair sequence data revealing the position of the species of the Calotes versicolor complex within the genus Calotes. Numbers at the nodes indicate bootstrap values. The tree was rooted using the Mantheyus phuwuanensis (Manthey and Nabhitabhata, 1991) as the out group. The tree was analysed under the TIM 2 + F + I + G 4 model of sequence evolution, implementing 1000 ultrafast bootstraps on the web platform of IQ-TREE. Species delimited in this study are colour coded: Green - C. farooqi stat. nov.; Yellow - C. irawadi; Red - C. versicolor; Blue - C. vultuosus comb. nov.

Figure 6. Holotype of Calotes vultuosus comb. nov. ANSP 7296. A. full body ventro-lateral, B. full body lateral, C. head dorsal, D. head ventral, E. head lateral. Photographs by Ned Gilmore.

Figure 7. An uncollected male of Calotes vultuosus comb. nov., from Burdwan, West Bengal, India, in life. Photograph by Ayan Mondal.