Corresponding author: Gaurang Gowande ( gaurang.gowande@gmail.com ) Academic editor: Uwe Fritz
© 2021 Gaurang Gowande, Saunak Pal, Daniel Jablonski, Rafaqat Masroor, Pushkar U. Phansalkar, Princia Dsouza, Aditi Jayarajan, Kartik Shanker.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gowande G, Pal S, Jablonski D, Masroor R, Phansalkar PU, Dsouza P, Jayarajan A, Shanker K (2021) Molecular phylogenetics and taxonomic reassessment of the widespread agamid lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) across South Asia. Vertebrate Zoology 71: 669-696. https://doi.org/10.3897/vz.71.e62787
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Abstract
The genus Calotes Cuvier, 1817 (Agamidae: Draconinae) is highly diverse, with species occurring in South and Southeast Asia, and Oceania. Most species of the subfamily except C. versicolor have narrow geographic distributions. Calotes versicolor is distributed from western Iran in the west to south China and Indonesia in the east and has been introduced to parts of Africa and North America. The species has had a complicated taxonomic history; multiple species and subspecies related to C. versicolor were described from India and adjoining regions, which were synonymized in subsequent revisions. However, a study of Burmese C. versicolor yielded two new species, C. htunwini and C. irawadi, indicating that C. versicolor is a species complex. Such integrative taxonomic studies have not been carried out in India, the supposed type locality of C. versicolor. Hence, we studied C. versicolor sensu lato from the Indian subcontinent and generated sequences of mitochondrial 16S and COI fragments from tissues sampled from multiple localities in the region, including the type localities of its synonyms. Phylogenetic analyses revealed four well-supported, deeply-divergent lineages, supported by morphological data. These lineages represent (i) C. versicolor sensu stricto, from South India and parts of the east coast, (ii) C. irawadi sensu lato from northeast India and Southeast Asia, (iii) a synonym from the eastern Indo-Gangetic Plains which we resurrect here, and (iv) a subspecies from Pakistan which we elevate to species level. We provide re-descriptions for the resurrected or elevated species, and a diagnostic key to the species of the C. versicolor complex. The study shows that C. versicolor sensu stricto is endemic to parts of southern and eastern India, and not widely distributed, though it may have been introduced to other parts of the world.
16S, COI, distribution, mtDNA, phylogenetics, species complex, species delimitation, systematics
Widely distributed species often display intraspecific molecular and morphological variation (
The genus Calotes includes species which are characterized by the presence of largely homogeneous, regularly arranged dorso-lateral scales and an exposed tympanum. Until recently, the genus included more than 25 species, with the greatest species diversity occurring in the Western Ghats of India, Sri Lanka, and Indo-Burmese biodiversity hotspots (
The Oriental Garden Lizard, Agama versicolor, was described based on two specimens housed at the
Therefore, as per the provisions of ICZN (1999),
Following
Calotes versicolor has been shown to be a species complex. Two species of the complex, C. htunwini Zug and Vindum, 2007 and C. irawadi Zug, Brown, Schulte and Vindum, 2007 were described from the Burmese regions adjoining northeast India (
Presently, only one species of the C. versicolor complex is known to occur in the Indian subcontinent with certainty, namely C. versicolor. Literature (
Systematic sampling was carried out throughout Peninsular India, the adjoining Indo-Gangetic Plains of India, the Indian deserts and northeast India (Fig.
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 (Clade1) originate, yellow samples indicate those of C. irawadi sensu lato (Clade2), blue circles represent the localities of C. vultuosus comb. nov. (Clade3), green circles indicate Clade4, 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 S1 for details on molecular and/or morphological vouchers used, and their respective localities.
Total genomic DNA was extracted from the collected tissue samples using the Phenol-Chloroform-Isoamyl Alcohol method (
A fragment of the mitochondrial 16S rRNA gene (16S) was amplified using PCR, using 16Sar (5’-CGCCTGTTTATCAAAAACAT-3’) and 16Sbr (5’-CTCCGGTTTGAACTCAGATCA-3’) (
The amplified PCR products were checked using a 2% Agarose Gel and viewed under the AlphaDigiDoc RT2 system. The PCR samples were purified using QIAquick® PCR Purification Kit. Sequencing was performed using a 3130xl Genetic Analyzer. This resulted in the generation of 67 16S sequences, and 19 COI sequences. Additionally, 16S and COI sequences were generated for one C. calotes Linnaeus, 1758 sample from south India, which was used as an outgroup. Laboratory work was largely carried out at CES and sequencing was outsourced to Barcode Biosciences (Bengaluru, India), except for Pakistani samples, which were processed by DJ in Slovakia and their sequencing of PCR products was outsourced to the Macrogen Europe (Amsterdam, The Netherlands). The generated sequences were checked manually in MEGA v.6 (
Maximum Likelihood (ML) and Bayesian Inference (BI) methods of phylogenetic analyses were implemented to determine the clustering pattern of the sequences of the C. versicolor complex generated in this study, as well as those of the related species, including C. calotes and C. irawadi. The non-coding 16S region (522 bp) was not partitioned by codon positions, whereas the coding COI region (608 bp) was partitioned by codon position. COI sequences were translated to their protein products using the vertebrate mitochondrial code table to check for premature stop-codons in the alignment and to rule out the possibility of having sequenced pseudogenes. Maximum Likelihood trees were analysed using W-IQ-TREE (
The uncorrected genetic “p-distances” were calculated separately for 16S (522 bp) and COI (608 bp) in MEGA v.6 (Table
We further retained one or two representative sequences of each of the species which we here delimit as distinct from the C. versicolor complex revealed by the analyses, and inserted these into the larger 16S dataset used in
A summary of the models of sequence evolution used for each partition in the ML and BI molecular phylogenetic analyses. *COI 2nd was combined with 16S in BI, as per the output of PartitionFinder analysis.
Partition | Codon position | Model | |
ML | BI | ||
1 | 16S | TPM+F+I+G4 | HKY+I |
2 | COI 1st | TN+F+G4 | GTR+G |
3 | *COI 2nd | TIM3e+G4 | – |
4 | COI 3rd | F81+F | F81 |
Table representing the least and the greatest un-corrected percent pairwise genetic p-distances within and between species recovered in this study. Values above the diagonal represent divergence on COI, whereas those below the diagonal represent divergence on 16S. Values in bold along the diagonal represent distance within species, written as COI/16S. *Does not include Calotes aff. farooqi.
Species | Calotes farooqi stat. nov.* | Calotes irawadi | Calotes versicolor | Calotes vultuosus comb. nov. | Calotes calotes |
Calotes farooqi stat. nov.* | –/0.2–0.4 | 16.9–17.5 | 17.1–17.9 | 15.5–16.9 | 19.2–20.3 |
Calotes irawadi | 3.3–6.0 | 0.0–4.5/0.2–2.4 | 12.6–15.1 | 12.8–15.9 | 17.2–18.1 |
Calotes versicolor | 3.5–5.2 | 2.7–4.7 | 0.2–7.6/0–1.5 | 13.0–15.5 | 17.0–18.7 |
Calotes vultuosus comb. nov. | 3.9–5.1 | 2.7–4.7 | 3.3–4.6 | 0.0–9.5/0–2.0 | 16.1–17.4 |
Calotes calotes | 4.9–6.3 | 5.9–6.9 | 5.1–6.1 | 5.3–6.9 | –/0.8 |
Mensural and meristic data from the sampled specimens as well as those deposited at museums in India and abroad, including the available type material of the species under the synonymy of C. versicolor, were recorded for use in systematics. Published literature (
Data for the following characters were recorded following
We followed a modified version of species delimitation strategy employed by
CES = Centre for Ecological Sciences, Indian Institute of Science, Bengaluru, Karnataka, India.
CUDZ = Comenius University, Department of Zoology (collection of Daniel Jablonski).
Maximum Likelihood and Bayesian Inference (Fig.
The ML phylogenetic output representing the ML+BI phylogeny based on the 1130 bp long 16S+COI combined dataset representing the clustering pattern of the sequences of the Calotes versicolor species complex. Values at the nodes indicate ultrafast ML bootstrap values/BI posterior probabilities. Colour code follows that of Figure
Clade1 included samples from south India, largely from the Southern Granulite Terrain, roughly to the south of 12°N latitude, including the southern Central and the Southern Western Ghats, along with the samples from the east coast of India (Puducherry and Bhubaneshwar). A single sequence from Huraa Islands, Maldives also belonged to this clade. This clade includes genetic sequences (MW901307: 16S, MZ489209: COI) from the type locality (discussed below) of C. versicolor, and hence we designate this clade as Calotes versicolor sensu stricto. Within the clade, the p-distance was up to 1.5% at 16S and between 0.2%–7.5% at COI, whereas the clade was at least 2.7% and 12.6% divergent from the other three clades at 16S and COI respectively.
Clade2 was composed of samples from northeast India and the adjoining Southeast Asian regions, including samples from Cambodia, China, Myanmar, and Vietnam. This clade contained the topotypic sequence of C. irawadi. Thus, we confirm the presence of this taxon in India and refer to this clade as Calotes irawadi sensu lato. This clade was the most heterogeneous at 16S, as the within clade p-distance ranged between 0.2%–2.4%; however, at COI, the within clade p-distance was not as high due to the absence of COI sequences from localities where 16S sequences were most divergent. The clade was at least 2.7% and 12.6% divergent from other clades at 16S and COI respectively. This clade likely contains additional undescribed species distributed across Southeast Asia. Given the heterogeneous nature of this clade, the clade was comparatively poorly supported in the phylogenetic analyses (ML bootstrap values 88, BI posterior probabilities 0.82).
Clade3 included samples from the Indian deserts (Jaisalmer), the Gangetic Plains (West Bengal), the Central Highlands (Bhedaghat, Chhattisgarh), and parts of Peninsular India (Deccan Volcanic Province) to the north of Southern Granulitic Terrain. The within clade divergence for Clade3 was up to 2.0% on 16S and up to 9.5% at COI, whereas the clade was at least 2.7% and 12.8% divergent from all other clades at 16S and COI respectively. This clade included a sequence from Burdwan near Kolkata, West Bengal, the type locality of Agama vultuosa (MW901298: 16S). Since this clade represents a genetically and morphologically distinct lineage, we resurrect Agama vultuosa from the synonymy of C. versicolor, provide diagnosis, and re-describe the taxon. Clade2 and Clade3 appear to be separated by the rivers Ganga and Brahmaputra, with Clade2 distributed on the eastern side of the rivers, whereas Clade3 occurs on the western side (Fig.
Lastly, the samples from the northern hilly regions and southern plains of Pakistan formed a distinct clade, Clade4, which was at least 3% divergent on 16S from all the other clades recovered in this study. Within this clade, the genetic distance was between 0.8–2.7% at 16S and 11.5% at COI, whereas the clade was at least 3.7% and 15.5% divergent from all other clades at 16S and COI respectively. This clade included sequences from near the type locality of the subspecies C. v. farooqi (MW901312–14: 16S, MZ489214: COI; Khyber Pakhtunkhwa and Punjab Provinces, Pakistan). Our results revealed that the population in the hilly regions of northern Pakistan is genetically and morphologically highly divergent from C. versicolor sensu stricto, C. irawadi and Clade3. Additionally, our analyses revealed the existence of yet another lineage (referred here as Calotes aff. farooqi, p-distance up to 2.7% at 16S and 11.5% at COI) distributed in the southern hilly regions as well as the southern plains (localities 78, 79 in Fig.
Three of the four phylogenetically distinct clades, Clade1, Clade2, and Clade3, fulfilled our species delimitation criteria with the integration of morphology and geography, and the identity of these clades as distinct species is established here. Clade4, on the other hand, included two divergent lineages, of which, the one from northern hilly regions of Pakistan, including the sequences from near the type locality of C. v. farooqi, is dealt with herein.
The 16S gene tree analysed using the sequences of representative species of the subfamily Draconinae suggested the relationships of the recovered lineages within the subfamily (Fig.
Maximum Likelihood 16S 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 TIM2+F+I+G4 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.
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
Calotes versicolor major
Annandale, 1921: 321 (nomen nudum fide
Calotes versicolor
–
Calotes cf. versicolor versicolor
Calotes cf. versicolor– Mahony et al. 2009;
The neotype designation by
Mensural and meristic characters of the type and non-type specimens of Calotes versicolor and its synonym Agama tiedemanni, mensural characters recorded as a ratio of SVL. Note that the morphometric data for the types of other synonyms of C. versicolor were not available.
Species | Calotes versicolor | ||
Specimens measured | Neotype Calotes versicolor NCBS AT102 |
Syntype Agama tiedemanni (fide |
Other morphological vouchers (n=29) |
Mensural characters | Range | ||
HeadL/SVL | 0.23 | 0.22 | 0.22–0.28 |
HeadW/SVL | 0.23 | 0.20 | 0.15–0.28 |
JawW/SVL | 0.17 | – | 0.15–0.21 |
HeadH/SVL | 0.17 | – | 0.14–0.21 |
SnEye/SVL | 0.10 | – | 0.08–0.11 |
NarEye/SVL | 0.05 | – | 0.04–0.07 |
EyeEar/SVL | 0.07 | – | 0.06–0.09 |
Snw/SVL | 0.07 | 0.06 | 0.05–0.07 |
InterOrb/SVL | 0.13 | 0.13 | 0.10–0.14 |
TrunkL/SVL | 0.46 | – | 0.38–0.50 |
PectW/SVL | 0.19 | – | 0.11–0.23 |
PelvW/SVL | 0.11 | – | 0.09–0.16 |
SnForeL/SVL | 0.35 | – | 0.34–0.41 |
UpArmL/SVL | 0.21 | – | 0.14–0.23 |
LoArmL/SVL | 0.20 | – | 0.16–0.20 |
ForefL/SVL | – | – | 0.14–0.21 |
4fingLng/SVL | 0.11 | – | 0.10–0.18 |
UpLegL/SVL | 0.24 | – | 0.17–0.25 |
CrusL/SVL | 0.26 | – | 0.19–0.28 |
HindfL/SVL | 0.31 | – | 0.28–0.39 |
4toeLng/SVL | 0.18 | – | 0.15–0.26 |
ForeLimbL/SVL | – | – | 0.48–0.59 |
HindLimbL/SVL | – | – | 0.69–0.91 |
ForeLimbL/HindLimbL | – | – | 0.61–0.73 |
PelvW/PectW | 0.60 | – | 0.5–1.0 |
Meristic characters | |||
SnS | 7 | – | 6–8 |
HeadSTr | 13 | – | 11–17 |
HeadSLn | 17 | – | 11–18 |
CanthR | 8 | – | 6–9 |
Eyelid | 13 | – | 10–15 |
Suplab | 11 | – | 9–14 |
Inflab | 12 | 12 | 8–12 |
TempSp | 2 | 2 | 2 |
Dorsal | 44 | – | 31–51 |
Midbody | 42 | – | 36–46 |
4FingLm | 21 | – | 15–24 |
4ToeLm | 28 | – | 21–30 |
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;
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
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
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.
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
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
We did not include this species in our analyses; however, the species is distinct from members of the C. versicolor group due to the horizontal orientation of the scale rows of the neck and the supra-axillary region. The species occurs throughout the lower elevations of Myanmar’s Central Dry Zone (
MYANMAR • 1 ♀; Sagaing Division, Chatthin Wildlife Sanctuary; 23.5743°N, 95.7376°E; ca. 110 m a. s. l.; 22 May 1998; Htun Win leg.;
The species can be diagnosed by its posteriorad or vertical orientation of the scale rows of the neck and the supra-axillary region, this separates the species from C. htunwini; detailed comparisons with C. versicolor, the species resurrected and elevated in this communication are provided in the diagnoses and comparisons sections of those species. The species was thought to have a narrower distribution in the Central Dry Zone of Myanmar, in comparison to C. htunwini (
MYANMAR • 1 ♂; Sagaing Division, Chatthin Wildlife Sanctuary; 23.5743°N, 95.7376°E; ca. 110 m. a. s. l.; 17 Jul. 1997;
Agama vultuosa
Agama indica
Calotes cf. versicolor versicolor
Calotes cf. versicolor
–
INDIA • 1 ♂; West Bengal, Kolkata; 20.55°N, 88.36°E, 10 m a. s. l.; Dr R. Coates leg.;
Other material (morphological vouchers). All from INDIA • 1 ♂, 1 ♀; Maharashtra, Satara, Koyna WLS; Soman and Thakar leg;
Mensural and meristic characters of the type and non-type specimens of Calotes vultuosus comb. nov. and Calotes farooqi stat. nov., mensural characters recorded as a ratio of SVL. Note that the morphometric data for the types of A. indica were not available, whereas those for C. farooqi have been sourced from
Species | Calotes vultuosus comb. nov. | Calotes farooqi stat. nov. | ||
Specimens measured |
Holotype |
Other morphological vouchers (n=20) |
Holotype |
Other morphological vouchers (n=3) |
Mensural characters | Range | Range | ||
HeadL/SVL | 0.19 | 0.16–0.30 | 0.27 | 0.23–0.25 |
HeadW/SVL | – | 0.16–0.26 | 0.24 | 0.19–0.21 |
JawW/SVL | – | 0.17–0.21 | – | 0.17–0.20 |
HeadH/SVL | – | 0.16–0.21 | 0.20 | 0.14–0.16 |
SnEye/SVL | – | 0.08–0.11 | – | 0.09–0.11 |
NarEye/SVL | – | 0.04–0.06 | – | 0.04–0.05 |
EyeEar/SVL | – | 0.06–0.09 | – | 0.07 |
Snw/SVL | – | 0.05–0.08 | – | 0.06 |
InterOrb/SVL | – | 0.09–0.13 | – | 0.12–0.15 |
TrunkL/SVL | 0.49 | 0.37–0.50 | 0.50 | 0.48–0.52 |
PectW/SVL | – | 0.12–0.20 | – | 0.14–0.20 |
PelvW/SVL | – | 0.09–0.17 | – | 0.11–0.13 |
SnForeL/SVL | – | 0.34–0.45 | – | 0.31–0.35 |
UpArmL/SVL | – | 0.14–0.23 | 0.20 | 0.13–0.18 |
LoArmL/SVL | – | 0.16–0.23 | – | 0.16–0.17 |
ForefL/SVL | – | 0.14–0.20 | – | 0.17–0.18 |
4fingLng/SVL | – | 0.10–0.15 | – | 0.12–0.13 |
UpLegL/SVL | – | 0.18–0.29 | – | 0.21–0.22 |
CrusL/SVL | – | 0.19–0.25 | – | 0.22 |
HindfL/SVL | – | 0.28–0.34 | – | 0.30–0.35 |
4toeLng/SVL | – | 0.15–0.23 | – | 0.13–0.20 |
ForeLimbL/SVL | – | 0.47–0.59 | – | – |
HindLimbL/SVL | – | 0.68–0.85 | – | – |
ForeLimbL/HindLimbL | – | 0.65–0.81 | – | – |
PelvW/PectW | – | 0.74–1.0 | – | 0.60–0.86 |
Meristic characters | ||||
SnS | 7 | 5–7 | 6 | 6 |
HeadSTr | 14 | 10–16 | 14 | 11 |
HeadSLn | 12 | 12–17 | 12 | 13–14 |
CanthR | 7 | 7–8 | 8 | 8 |
Eyelid | 13 | 11–14 | 9 | 9–11 |
Suplab | 10 | 9–13 | 13 | 11–13 |
Inflab | 10 | 9–13 | 13 | 12 |
TempSp | 2 | 2 | 2 | 2 |
Dorsal | – | 35–62 | – | 40–44 |
Midbody | – | 37–45 | 51 | 41–44 |
4FingLm | – | 19–22 | 21 | 17–18 |
4ToeLm | – | 23–28 | 23 | 24–25 |
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
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
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.
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.
Harlan R (1825). Journal of the Academy of Natural Sciences of Philadelphia 4(1): 296–305.
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 vultuosa is feminine singular and is here changed to the masculine gender vultuosus in agreement with the generic epithet, Calotes, which is masculine in gender.
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
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.
Calotes versicolor nigrigularis Auffenberg and Rehman, 1993
Calotes versicolor farooqi Auffenberg and Rehman, 1995 (nom. nov. for C. v. nigrigularis)
Pakistan • 1 ♂; Khyber Pakhtunkhwa, Mansehra, Shargal (corrected to Sarhan); 34.3°N, 73.4°E, 1077 m a.s.l; 15 Jun. 1990;
The analysis included a total of three samples (MW901312–14: 16S, MZ489214: COI) from the localities Tangora, Gulbandi Buner (localities 85 and 86 in Figure
A medium to large species of Calotes, adult males ranging from 94–99 mm in SVL, body moderately 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; anti-humeral fold absent; two distinct spines in the supratympanic region; nuchal and dorsal crest continuous, distinct; scales of the nuchal crest large, those of dorsal crest slightly smaller, slightly recurved, ending at the top of the base of the tail, males with distinct black patches on both sides of the lower jaw, extending into the forebody, in the breeding season.
The species can be separated from all the members of the C. versicolor group 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), 41–51 Mid-body scale rows (vs. 49–65 in C. emma, 27–35 in C. grandisquamis, 58–63 in C. maria Gray, 48–60 in C. minor Hardwicke and Gray, and 45–58 in C. mystaceus); nuchal crest scales well-developed, dorsal crest scales much smaller, more or less equal in size (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 spines (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 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 members of the C. mystaceus complex (C. bachae, C. geissleri, C. goetzi, C. mystaceus, C. vindumbarbatus, sensu
The species is most similar in appearance to C. versicolor and C. vultuosus comb. nov., however, can be differentiated from C. versicolor by its smaller adult male body size (average male SVL 97 mm, vs. 108 mm in C. versicolor, female SVL 82 vs. 92 in C. versicolor), dorsal crest scales composed of comparatively shorter scales, which become shorter progressively to the base of the tail (vs. dorsal crest composed of longer scales, dorsal crest continues to the base of the tail in C. versicolor), lesser number of eyelid scales, Eyelid 9–11 (vs. 10–15 in C. versicolor, 11–14 in C. vultuosus comb. nov.), the shape and the size of the scales between the nasal shield and the orbit (large, <6 in a row between the nasal shield and the orbit in C. farooqi stat. nov., vs. small, >6 in C. versicolor and C. vultuosus comb. nov.), by the acuteness of the region between the nostril and the orbit (more acute in C. farooqi stat. nov., less acute in C. vultuosus comb. nov. and C. versicolor). C. farooqi stat. nov. also has slightly lower number of SnS (6), in comparison to C. versicolor and C. vultuosus comb. nov. (generally 7).
The species also differs from C. versicolor and C. vultuosus comb. nov. in terms of breeding coloration of the adult males. The head and the forebody of the males of C. farooqi stat. nov. attain grey to black colour, except the vertebral region and the parts of the head above the jaw muscles; the lower portion of the orbit turns black, the black colour extends to the outer surfaces of the forelimbs, the inner surfaces of the forelimbs and the region of the venter intervening the limbs turn greyish black. In contrast, C. versicolor males attain a yellowish colour, trunk and orbital region turn orange, forelimbs and hind limbs turn dark to black, the black patches under the throat do not extend anteriorly onto the jaw muscles; C. vultuosus comb. nov. males attain a cream to brown body colour, the head and the anterior two-thirds of the trunk attain orange colour, which may extend on to the forelimbs, the posterior parts of the body remain duller.
A large-sized male, SVL 94 mm, tail complete, 245 mm. The specimen is in a mediocre condition, the head is bent to the left, tail to the right; forelimbs adpressed to the body. A vertical incision on the ventral surface between the insertion of the forelimbs.
Head large (HL/SVL 0.27), snout tip pointed in dorsal perspective; region between the nasals and the orbit slightly concave, acute, covered by heterogeneous, juxtaposed scales; loreal region from the nasal shield triangular, the anterior border of the nasal shield at the vertex of the triangle; 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; dorsal edges of the forehead divergent, bordered by scales of the canthus rostralis and supraciliary region, rostral broader than high; nasals 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 six 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; scales of the forehead posterior to SnS sub-imbricate, very irregular in shape and size, some rugose; HeadSLn 12, bordered anteriorly by the rostral, and posteriorly by the single, large interparietal shield; orbit large, encased in a sock of granular scales, separated from the nasal shield by rows of 4–5 scales, from Suplab by three scale rows; HeadSTr 14, between the posteriormost supraciliary scales on each, just anterior to the interparietal; eye opening bordered 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; region between the orbit and the tympanum covered by rows of 6–7 smooth, irregular in shape; tympanum large, round, naked; 3.1 mm at its greatest height; supratympanic scales smooth to weakly keeled; two enlarged supratympanic spines, separated from each other by 3–4 scales; anterior spine slender, shorter, posterior spine prominent; posterior region of the jaws swollen, bulges out, covered by subtriangular, imbricate, postero-ventrally directed scales, some of these mucronate; upper border of the jaw muscles conceal the lower portion of the tympanum; labial scales large, sub-rectangular; Suplab 12; Inflab 11; 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 ninth Suplab; interparietal large, irregularly pentagonal, bordered by 9–10 smooth, heterogeneous scales; nuchal crest starts 3–4 scales behind the interparietal; mental shield large, single, subtriangular, , mental narrower than rostral; two pairs of elongate postmentals, anterior pair narrow, separated by a single small chin scale posterior to the mental; posterior pair broader, separated by three small scales; chin scales posterior to the postmental scales small, progressively become larger towards the throat; scale rows of the posterior region of the throat large, mucronate.
First nuchal scale smallest, size of the nuchal scale increases modestly towards the median nuchal scale; beyond which, the size reduces slightly, nuchal crest composed of lanceolate, recurved spines, which continue into dorsal crest scales; dorsal crest scales slightly smaller than nuchal crest scales, composed of spines that progressively become smaller towards the middle of the back; continues to the base of the tail, beyond which the mid-dorsal crest row terminates at the keeled tail scales; paravertebral scales recurved, strongly keeled, mucronate; dorso-lateral scales sub-triangular, imbricate, distinctly keeled, posterodorsally oriented; those in the supra-axillary region slightly smaller than those of the mid-body; ventrals strongly keeled, smaller than the dorso-lateral scales, mucronate, imbricate; ventral scales between the insertion of the forelimbs weakly keeled.
Limbs long and slender, covered with keeled scales, similar to dorso-laterals in shape and size, forming parallel longitudinal rows; scales on the dorso-ventral surfaces of the thigh weakly to moderately keeled, those on the crus and sole strongly keeled; first finger shortest, third and fourth almost equal, fifth longer than first; relative length of toes 4>3>5>2>1; subdigital lamellae bicarinate, keels sharp, 20 under the fourth finger, 24 under the fourth toe; tail slender, swollen at base, TailL 245 mm.
Coloration in preservation: Anterior two-thirds of the body generally greyish-tan, dorsum distinctly black till mod-body, black coloration fades away in the posterior half, vertebral and paravertebral row of scales paler; venter greyish black at the pectoral region, slightly duller towards the vent, colour abruptly changes to greyish-cream; a dark ventral midline runs from the pectoral region to the vent; dorsal surface of the forelimbs dark, lower surfaces slightly paler; hindlimbs and tail greyish-cream; supra-axillary region greyish-black, the black patches continue anteriorly along the lower surface of the jaws and gular region, except the mid-gular region which is strikingly lighter, pinkish in colour; head generally greyish above, paler laterally and posteriorly; eyelids light grey anteriorly, a small black patch near the anterior corner of the eye, a large black patch near the posterior corner of the eye.
Auffenberg W, Rehman H (1993). Asiatic Herpetological Research 5: 14–30.
The specific epithet is a patronym in genitive singular case, dedicated to Farooq Ahmed, former Director, Zoological Survey Department, Pakistan.
One male specimen (CUDZ DJ7902) was examined. The mensural characters in mm are: HL 28.1; HW 21.0; HH 16.9; SVL 99.0; TrunkL 49.2; UpArmL 16.6; The meristic characters are: SnS 6; HeadSTr 11; HeadSLn 13; CanthR 8; Eyelid 9; Suplab 11; Inflab 12; TempSp 2; Mid-body 41; 4FingLm 17; 4ToeLm 24. Additionally, two female specimens were examined. The means for the mensural characters in mm are: HL 20.2; HW 16.8; HH 12.4; JawW 15.7; SnEye 8.6; NarEye 3.6; EyeEar 5.6; SnW 5.1; Interorb 11.8; SVL 82; TrunkL 41.3; TailH 7.3; TailW 7; PectW 14; PelvW 10.5; SnForeL 17.6; UpLegL 17.5; CrusL 17.9; HindfL 25.3; 4ToeLng 13.4; HindLimbL 60.5. The means for meristic characters are: SnS 6; HeadSTr 11; HeadSLn 14; CanthR 8; Eyelid 11; Suplab 12; Inflab 12; TempSp 2; Dorsal 42; Mid-body 43; 4ToeLm 25. The ranges for each of these characters are given in Table
The species has been reported from the northern hilly regions of Pakistan, while the samples from the southern plains represent a hitherto undescribed species (localities 78, 79 in Fig.
1a | Orientation of the scale rows on the neck and the supra-axillary region lateral, Southeast Asia | C . htunwini |
1b | Orientation of the scale rows on neck and supra-axillary region oblique or vertical | 2 |
2a | Average adult male SVL < 90 mm, supratympanic or temporal spines and crest scales short, average number of mid-dorsal scales ~49, northeast India and Southeast Asia | C . irawadi |
2b | Average adult male SVL > 90 mm, average number of mid-dorsal scales < 49 | 3 |
3a | Nuchal and dorsal crest scales long, dorsal crest continues to the base of the tail; supratympanic spines long, slender, southern Peninsular India and east coast, males in breeding season yellowish, trunk and orbital region orange, limbs dark to black, a black patch across the neck | C. versicolor |
3b | Nuchal and dorsal crest scales short, dorsal crest shortens progressively at mid-body; supratympanic spines short | 4 |
4a | Scales between the nasal shield and the orbit large, < 6; Eyelid 9–11; head long, northern hilly regions of Pakistan, males during breeding season grey to black, except the vertebral region and upper head, orbits, inner and outer surface of forelimbs and anterior and venter between the forelimbs dark to black | C . farooqi stat. nov. |
4b | Scales between the nasal shield and the orbit small, > 6; Eyelid 11–14; head short, Peninsular India to the north of 13° latitude, Indian deserts, males during breeding season cream to brown, head and anterior two-thirds of trunk orange | C . vultuosus comb. nov. |
Integrated investigations into the south Asian “Calotes versicolor” demonstrate that C. versicolor is not a widely distributed single species, as was previously considered, but harbours multiple lineages. Molecular phylogenetic analyses revealed the existence of four distinct species distributed across South Asia: (i) Calotes versicolor sensu stricto, (ii) C. irawadi, (iii) C. vultuosus comb. nov. and, (iv) C. farooqi stat. nov., which allowed us to revise its taxonomy. This study also addressed the taxonomic predicaments in revising the nomenclature of the species complex.
Daudin’s illustration of C. versicolor as well as the measurements of one of the syntypes presented in the original description (
It is impossible to comment with objectivity on where the specimens of Agama versicolor were collected from; nevertheless, a review of the herpetofaunal species described by Daudin (from 1801 to 1803) suggests that almost all the species described by him were based on the material collected from along the east coast of India (Coromandel Coast), especially from the localities Tranquebar, Puducherry, Vishakhapatnam, or rarely from Bengal (
The study led to the revalidation of C. vultuosus comb. nov., which represents a lineage genetically and morphologically distinct from C. versicolor sensu stricto. Agama vultuosa was described based on a specimen brought by Dr R. Coates from Calcutta (now Kolkata), West Bengal, which according to the original description (
Thus, we here resurrect the nomen Agama vultuosa Harlan, 1825 from its synonymy as Calotes vultuosus comb. nov. (Harlan, 1825). A related species A. indica Hardwicke and Gray, 1827 (type locality Kolkata) was described, from the same locality as C. vultuosus comb. nov. The species was diagnosed on the basis of its green coloration, dimensions of the scales of the head, body, tail and limbs, two bundles of spines above the ears, and a crest composed of simple, compressed spines that extends only to the middle of the back, and tail that is nearly thrice as long as the body. A majority of these characters, such as the dimensions of scales of the head, body, tail and limbs, and the extension of the dorsal crest, overlap with those of C. vultuosus comb. nov. Similarly, Harlan’s description of A. vultuosa precedes that of A. indica, and thus, the name C. vultuosus comb. nov. is prioritized, and A. indica is transferred to the synonymy of C. vultuosus comb. nov. Calotes vultuosus comb. nov., the northern Peninsular Indian representative of the complex, was observed to be phenotypically variable during the study, at least in terms of coloration and a select few mensural and meristic characters. For example, the specimen CESL 1041 from the deserts differs from the other examined specimens of the species in certain morphological aspects, such as the length of the head and the size of the dorso-lateral body scales; however, it clusters phylogenetically with members of C. vultuosus comb. nov., with very shallow genetic divergence from other examined sequences of the species. Similarly, there is a certain variation in the sexual characters such as the size and the number of nuchal and dorsal crest scales and the extent and the brightness of the orange patch in breeding males. But, as
Results from molecular phylogenetic analyses revealed that the sequences generated from Southeast Asia, including those from northeast India, Myanmar, Vietnam, Cambodia and far eastern China fell into the C. irawadi clade (Clade2). We tentatively assign the Southeast Asian populations morphologically resembling C. versicolor in terms of the obliquely posteriorad or vertical orientation of the neck and the supra-axillary area to C. irawadi, pending further investigation, and suggest that records from Southeast Asia labelled as C. versicolor should be treated with caution. The within clade p-distance for this clade was high, the greatest recorded being 2.4% at 16S and 4.5% at COI (Table
The p-distances between the species recognized in this study ranged between 2.7% (between C. versicolor and C. irawadi; and C. vultuosus comb. nov. and C. irawadi) and 6.0% (between C. farooqi stat. nov. and C. irawadi) at 16S, and between 12.6% (between C. versicolor and C. irawadi) and 20.4% (between C. calotes and C. farooqi stat. nov.) at COI. Comparable genetic divergences were observed in the other, previously defined species, such as those in the genus Monilesaurus, between the species pair C. grandisquamis-C. nemoricola (p-distance 3.7%–5.1%), and species of the C. mystaceus complex (
The ML tree analysed using additional representatives of the subfamily Draconinae based on the 16S dataset revealed the tentative positions of the species of the C. versicolor species complex (Fig.
With the recognition of C. farooqi stat. nov. and C. vultuosus comb. nov., the genus Calotes now includes 30 species (
This study provides further evidence to support the hypothesis that many widely distributed species represent species complexes (
An integrated molecular-morphological investigation demonstrates that C. versicolor on the Indian subcontinent is composed of multiple lineages. It is not as widely distributed as was earlier believed, but occurs in southern India and on the east coast, with the possibility of its occurrence on the island of Sri Lanka. The study led to the revalidation of the species C. vultuosus comb. nov., a species potentially endemic to the Deccan Volcanic Province, Central Highlands, arid and semi-arid regions of India and Gangetic Plains, which was under the synonymy of C. versicolor; and the promotion of the subspecies C. versicolor farooqi from Pakistan to species rank, as C. farooqi stat. nov. We further confirm the presence of C. cf. irawadi in northeast India. The taxonomic status of the populations introduced to Maldives was also resolved. Our results lend further support to the argument that many widely distributed species represent species complexes, demanding urgent taxonomic attention.
Fieldwork was supported by the Rufford Small Grants for Nature Conservation to GG. Molecular analysis was supported by the DBT-IISc Partnership Programme. DJ was supported by Slovak Research and Development Agency under contract nos. APVV-15-0147 and APVV-19-0076.
Conflict of Interest: None declared.
We are grateful to the respective forest departments for granting the necessary permissions for sample collection. We thank numerous colleagues and associates who helped by providing samples, specimens from various regions of the study area, for accompanying us in the field, and help in the laboratory: Anurag Mishra, Atish Gawai, Ayan Mondal, Ashok Kumar Mallik, Chandramouli S.R., Harshal Bhosale, Harshil Patel, Jana Poláková, K.P. Dinesh, Mandar Sawant, Mrugank Prabhu, Pranav Mhaisalkar, Shubhankar Deshpande, Varun Kher, Varun Torsekar, Vedant Dixit, Vedant Thite, Vishal Varma and Zeeshan Mirza. GG is indebted to Salvador Carranza and Varad Giri for encouraging him to pursue this study, and to V. Deepak for discussions during the initial phase of the study. GG is also grateful to Maria Thaker for supporting the administration of the grant. The phylogenetic analyses would have been incomplete without the topotypic C. irawadi sequences provided by Jens Vindum, Lauren Scheinberg and Anna Sellas, and without the morphological details of the C. versicolor specimens at CMNH from Puducherry provided by George Zug. Ned Gilmore of