Calotes versicolor (Daudin, 1802)

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 versicolor (Daudin, 1802)

Figs 4 , 5

Agama versicolor Daudin, 1802

Agama tiedemanni Kuhl, 1820

Calotes versicolor - Duméril and Bibron, 1837: 405

Calotes cristatus Jaquemont, 1844

Calotes viridis Gray, 1846: 429

Calotes gigas Blyth, 1853: 648 (nomen dubium fide Gowande et al. 2016)

Calotes versicolor major Annandale, 1921: 321 (nomen nudum fide Gowande et al. 2016)

Calotes versicolor - Smith 1935; Taylor 1963; Manthey and Grossmann 1997; Cox et al. 1998; Manthey and Schuster 1999; Amarasinghe et al. 2009

Calotes cf. versicolor versicolor Manthey 2008: 86 (in part)

Calotes cf. versicolor - Mahony et al. 2009; Pal et al. 2018 (in part)

Neotype (designated by Gowande et al. 2016).

INDIA • 1 ♂; Puducherry, Kalapet, Pondicherry University Campus; 12.02909°N, 79.85034°E; 34 m a. s. l.; 20 Mar. 2016; Anurag Mishra and Zeeshan Mirza leg.; GenBank: MW901307 (16S), MZ489209 (COI); NCBS AT102.

Note.

The neotype designation by Gowande et al. (2016) stands valid sensu Article 75, and we reject the invalidation suggested by Chaitanya et al. (2017). Further comparisons and justifications are included herein.

Other material (morphological vouchers).

All from INDIA • 1 ♂; Kerala, Thiruvananthapuram, Peppara Wildlife Sanctuary; 8.62448°N, 77.13646°E, 132 m a. s. l.; 15 Apr. 2010; Saunak Pal and Mrugank Prabhu leg.; GenBank MH844713 (16S), MZ489207 (COI); CESL 036; • 1 ♂; Kerala, Periyar Wildlife Sanctuary, Ranni Forest Division, Sabarimala; 9.40692°N, 77.06780°E, 181 m a. s. l.; 21 Feb. 2011; Saunak Pal and Mrugank Prabhu leg.; GenBank MH844729 (16S); CESL 182 • 1 ♂; Kerala, Pooppara, Mathikettan Shola National Park; 9.97136°N, 77.23283°E, 1088 m a. s. l.; 27 Mar. 2011; Saunak Pal leg.; GenBank MH844730 (16S); CESL 190 • 1 ♂; Kerala, Kottappadi part, Chembra; 11.55390°N, 76.08328°E, 1085 m a. s. l.; 10 Jun. 2010; Saunak Pal and Mrugank Prabhu leg; CESL 046; • 1 ♂; Tamil Nadu, Thiruchitrambalam, Aranya Forest and Sanctuary; 11.96644°N, 79.76334°E, 28 m a. s. l.; 14 May 2017; Gaurang Gowande, Zeeshan Mirza and Vishal Verma leg.; CESL 1072 • 11 ♂; Puducherry; 11.93°N, 79.88°E; 1 m a. s. l.; Shekhar Dattatri leg.; CMNH152047-CMNH152051, CMNH152053, CMNH152054, CMNH152066, CMNH152068, CMNH152069, CMNH152071 • 1 ♂; Kerala, Palakkad District ; Jan. 1961; DN Mathew leg; BNHS 362 • 1 ♂; Tamil Nadu, Masinagudi; Jun. 1975; Rezakhan leg; BNHS 1260 • 1 ♂, 3 ♀; Tamil Nadu, Jamestown; May-Jun. 1965; RB Grubh leg; BNHS 728, BNHS 810, BNHS 811, BNHS 813 • 1 ♂; Tamil Nadu, Courtallam; 19 Jun. 1966; JC Daniel leg; BNHS 858 • 2 ♂; Odisha, Chilika Lake ; Jan. 1967; BNHS 779, BNHS 781 • 1 ♂; Tamil Nadu, Papanasam Wildlife Sanctuary; 3 Apr. 1978; H Abdulali leg; BNHS 1257 • 1 ♀; Kerala, Mananthavady; 1 May 1975; PBSL Mahadik leg; BNHS 1129 .

Genetic diagnosis.

The within species genetic divergence across all the examined sequences is up to 1.6% at 16S and 0.2-7.6% at COI. The species is at least 3.3% and 13.0% divergent from Clade3, at least 2.7% and 12.6% divergent from C. irawadi and at least 3.5% and 17.1% divergent from Clade4 at 16S and COI respectively. From C. calotes , the species differs by sequence divergence of at least 5.1% at 16S and 16.7% at COI (Table 2 ). The species was recovered as sister to a clade comprising of C. calotes and Clade4 in the phylogenetic analyses (Fig. 3 ).

Diagnosis and comparison.

A medium to large sized species of Calotes , adult males averaging 108 mm in SVL, females averaging 90 mm in SVL; body compressed; head relatively long; dorso-lateral scales posterodorsally oriented, large, weakly to strongly keeled, homogeneous; ventral scales smaller than the dorso-lateral scales, strongly keeled, mucronate; 36-46 scales around the mid-body; anti-humeral fold absent; two distinct, elongated spines in the supratympanic region on each side of the head, anterior spine longer and more prominent; nuchal and dorsal crest continuous, distinct, slightly recurved; scales of the nuchal crest long, those of dorsal crest slightly shorter, ending at the top of the base of the tail; nuchal, dorsal and supratympanic spines more pronounced in males; limbs slender, dorsal surface strongly keeled, ventral surface moderately keeled.

The species can be separated from all the members of Smith’s C. versicolor group, which includes the species C. calotes , C. emma Gray, 1845, C. grandisquamis , C. jerdoni Günther, 1870, C. maria Gray, 1845, C. minor (Hardwicke and Gray, 1827), C. mystaceus Duméril and Bibron, 1837, C. nemoricola 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, and C. nemoricola ), 36-46 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 , 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 (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 conical scales, one scale 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 , single well-developed postorbital spine in C. emma ). The species differs from C. paulus (Smith, 1935) and C. zolaiking Giri, Chaitanya, Mahony, Lalrounga, Lalrinchhana, Das, Sarkar, Karanth and Deepak, 2019 primarily by the homogeneous scalation on the dorsolateral region (vs. heterogeneous) and a comparatively well-developed dorsal crest. From the dubious species C. bhutanensis Biswas, 1975, 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 Vindum, 2003, C. nigriplicatus Hallermann, 2000 and other members of the C. mystaceus complex ( C. bachae Hartmann, Geissler, Poyarkov, Ihlow, Galoyan, Rödder and Böhme, 2013, C. geissleri Wagner, Ihlow, Hartmann, Flecks, Schmitz and Böhme, 2021, C. goetzi Wagner, Ihlow, Hartmann, Flecks, Schmitz and Böhme, 2021, C. mystaceus , C. vindumbarbatus Wagner, Ihlow, Hartmann, Flecks, Schmitz and Böhme, 2021, sensu Wagner et al. (2021)) C. versicolor 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 Müller, 1887, C. desilvai Bahir and Maduwage, 2005, C. liocephalus Günther, 1872, C. liolepis Boulenger, 1885, C. manamendrai Amarasinghe and Karunarathna, 2014, C. nigrilabris Peters, 1860, C. pethiyagodai Amarasinghe, Karunarathna and Hallermann, 2014) the species differs by its posterodorsal orientation of lateral body scales (vs. posteroventral) and absence of shoulder pit (vs. present). The species most closely resembles, and differs from C. irawadi by its much larger adult male body size (average SVL 108 mm in C. versicolor , vs. 82.4 mm, in C. irawadi ), lesser number of dorsal crest scales (average 39.8 in C. versicolor vs. 48.9 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 ). Detailed comparison with the species resurrected and elevated in this communication is presented in the respective diagnosis and comparison sections of those species.

Original description.

Daudin (1802). Histoire naturelle, génerale et particulière des reptiles, ouvrage faisant suite, a l’histoiure naturelle, génerale et particulière composée par Leclerc de Buffon, et redigée par C.S. Sonnini, vol. III. Paris, F. Dufart, 452 pp.

Etymology.

The specific epithet is an adjective in Latin referring to variable or to turn (versi, derived from versare) and color (color) of the species in life.

Variation in the material examined.

Twenty-three male specimens were examined. The means for the mensural characters in mm are: HL 26.5; HW 26.2; HH 19.6; JawW 21.3; SnEye 10.3; NarEye 5.8; EyeEar 8.6; SnW 6.6; Interorb 12.5; SVL 110; TrunkL 49.0; TailL 280.4; TailH 16.8; TailW 13.6; PectW 19.3; PelvW 13.7; SnForeL 45.0; UpArmL 19.2; LoArmL 20.2; ForefL 18.9; 4FingLng 12.8; UpLegL 24.6; CrusL 26.4; HindfL 34.5; 4ToeLng 19.9; ForeLimbL 58.6; HindLimbL 85.2. The means for meristic characters are: SnS 7; HeadSTr 13; HeadSLn 15; CanthR 8; Eyelid 12; Suplab 11; Inflab 11; TempSp 2; Dorsal 40; Mid-body 42; 4FingLm 21; 4ToeLm 27. Seven female specimens were examined. The means for the mensural characters in mm are: HL 21.4; HW 17.1; HH 14.7; JawW 15.3; SnEye 9.0; NarEye 5.2; EyeEar 6; SnW 5.3; Interorb 10.5; SVL 90; TrunkL 42.3; TailL 228; TailH 10.3; TailW 9.6; PectW 15.6; PelvW 10.8; SnForeL 33.2; UpArmL 17.7; LoArmL 16.4; ForefL 15.2; 4FingLng 11.5; UpLegL 20.4; CrusL 21.4; HindfL 27.2; 4ToeLng 16.6; ForeLimbL 44.5; HindLimbL 64. The means for meristic characters are: SnS 7; HeadSTr 15; HeadSLn 13; CanthR 8; Eyelid 13; Suplab 11; Inflab 9; TempSp 2; Dorsal 45; Mid-body 42; 4FingLm 18; 4ToeLm 25. The ranges for each of these characters are given in Table 3 .

Distribution.

The species appears to be endemic to India, occurring largely on the Southern Granulite Terrain and the eastern coast of India. The species was found in the Southern Western Ghats, the south-west coast of India, the southern Eastern Ghats, the eastern coast of India till Mahanadi basin in the north, and in the low-elevation areas of Peninsular India between the Eastern and the Western Ghats during this study. The species also has been introduced to the Maldives (Figs 1 - 3 ).

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 4. Neotype of Calotes versicolor NCBS AT 102. A. full body dorsal, B. full body ventral, C. full body lateral, D. head dorsal, E. head ventral, F. head lateral. Photographs by Zeeshan Mirza.

Figure 5. Calotes versicolor (CESL 036), Peppara WLS, Thiruvananthapuram, Kerala, India, in life. Photograph by Saunak Pal.