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Calotes versicolor (DAUDIN, 1802)

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Higher TaxaAgamidae (Draconinae), Sauria, Iguania, Squamata (lizards)
Common NamesE: Eastern Garden Lizard, Oriental Garden Lizard, Indian Garden Lizard, Common Garden Lizard, Bloodsucker, Changeable lizard
G: Verschiedenfarbige Schönechse, Blutsaugeragame
Chinese: 变色树蜥 
SynonymAgama versicolor DAUDIN 1802: 395
Agama tiedemanni KUHL 1820
Agama vultuosa HARLAN 1825: 296
Calotes versicolor — FITZINGER 1826
Agama indica GRAY 1827: 217
Calotes versicolor — DUMÉRIL & BIBRON 1837: 405
Calotes cristatus JAQUEMONT 1844
Calotes viridis GRAY 1846: 429
Calotes gigas BLYTH 1853: 648 (nomen dubium fide GAWANDE et al. 2016)
Calotes versicolor — ANDERSON 1872
Calotes versicolor — BOULENGER 1885: 321
Calotes versicolor — DE ROOIJ 1915: 124
Calotes versicolor major ANNANDALE 1921 (nom. subst. pro C. gigas)
Calotes versicolor — SMITH 1935: 189
Calotes versicolor — TAYLOR 1963: 891
Calotes versicolor — MANTHEY & GROSSMANN 1997: 163
Calotes versicolor — COX et al. 1998: 96
Calotes versicolor — MANTHEY & SCHUSTER 1999: 35
Calotes versicolor versicolor — MANTHEY 2008: 86
Calotes cf. versicolor — MAHONY et al. 2009
Calotes versicolor — SPAWLS et al. 2018: 241
Calotes cf. versicolor — KARTHIK et al. 2018
Calotes versicolor — GOWANDE et al. 2021: 677
Calotes vultuosus — GOWANDE et al. 2021 
DistributionSE Iran, Afghanistan, Pakistan, Nepal, Bhutan,
India (incl. Tamil Nadu, Assam, Gujarat, Maharashtra, Madhya Pradesh, Kerala, Himachal Pradesh, Mizoram, Nagaland, Tripura, Andaman Islands, Jammu and Kashmir, Punjab, Odisha), Sri Lanka (Ceylon), Bangladesh, Burma (= Myanmar), Thailand, W Malaysia (incl. Pulau Pangkor), Vietnam (Binh Thuan, Son La etc.), e.g. Pulo Condore Island, Cambodia, Laos,
S China (Fujian, Guangdong, Guangxi, Honkong, Hainan Island), Maldives
Indonesia (Sumatra, Natuna islands), Singapore

Introduced to Celebes, Maldives, Seychelles
Mauritius (Reunion, Rodrigues; fide Glaw, pers. comm. and ROGNER 2006)
Introduced to the Arabian peninsula in Oman (fide VAN DER KOOIJ 2001, Grossmann & Kowalski 2019)
Introduced to Asia in Singapore, Borneo (DAS et al. 2009).
Introduced to Africa in SW (coastal) Kenya (Spawls et al. 2018)
Introduced to USA (Florida, fide SSAR checklist)

Type locality: “India” fide Auffenberg & Rehman 1993; restricted to “Pondicherry” by KUHL (fide Auffenberg & Rehman 1993) but also designated by SMITH 1935. Neotype locality: Pondicherry University Campus, Kalapet, Pondicherry, India.  
Reproductionoviparous. Temperature‐dependent sex determination (TSD) fide Cornejo-Páramo et al. 2020 (and references therein). 
TypesNeotype: NCBS AT102, male from Pondicherry University Campus, Kalapet, Pondicherry, India (12.029097° N, 79.850343° E, elevation 34m). Collected by Anurag Mishra and Zeeshan A. Mirza on 20 March 2016. Neotype designated by Gowande et al. 2016 but invalidated by Chaitanyia et al. 2017.
Holotype: lost (Auffenberg & Rehman 1993). The specimen MNHN 2548, collected from Pondicherry by Leschenault, cannot be a syntype of C. versicolor Daudin 1802, because Leschenault visited Pondicherry only after 1816. According to Daudin’s original description (1802) one of his two syntypes measured 81 mm SVL, 202.5 mm tail length and 283.5 mm total length.
Holotype: ANSP 7296 [Agama vultuosa]
Holotype: unknown fide Gowande et al. 2016 [Calotes gigas]
Syntype: MNHN 2548 from Pondicherry, collected by Leschenault [Agama tiedemanni] 
DiagnosisDiagnosis and comparison. A medium to large sized Calotes (SVL up to 99.4 mm) characterized by the posterodorsal orientation of lateral scales; antehumeral fold absent; 38-44 midbody scale rows; nuchal and dorsal crest well developed, continuous, extending till the start of tail in males; two well separated supratympanic spines; dorsal and lateral scales large, more or less distinctly keeled, ventral scales strongly keeled, mucronate; paired postmentals, first pair separated by 1–2 median scales; 15–22 subdigital lamellae under fourth finger, 21–25 subdigital lamellae under fourth toe; 10–12 supralabials and 9–11 infralabials; uniform, light brown or greyish above, juveniles and females with darker cross bars or blotches on lateral side, often with paired lighter dorsolateral stripes, gular pouch brick red and overall body reddish in breeding males.
From other members of Smith’s C. versicolor group (C. nemoricola, C. grandisquamis, C. calotes, C. minor, C. maria, C. jerdoni, C. emma and C. mystaceus) C. cf. versicolor can be distinguished by a combination of the following characters: smaller body size: adult SVL 73.8–99.4 mm, n=9 (vs. C. nemoricola, adult SVL 112–134.8 mm, n=7, n=5; and C. grandisquamis, adult SVL 110.0–136.5 mm, n=5); 38–44 midbody scale rows (vs. 27–35 in C. grandisquamis; 30–35 in C. calotes, n=3; 36–43 in C. nemoricola; 58–63 in C. maria; 45–57 in C. jerdoni; 49– 65 in C. emma; 48–58 in C. mystaceus and 48–60 in C. minor ); crescent-shaped patch of granular scales in front of the shoulder absent (vs. present in C. nemoricola, C. grandisquamis, C. emma, C. mystaceus and C. jerdoni); nuchal and dorsal crest well developed, composed of almost equal spines (vs. nuchal spines much longer, dorsal crest reduced in C. nemoricola and C. maria; nuchal spines much longer than dorsal spines in C. grandisquamis, C. calotes and C. emma); two well separated supratympanic spines (vs. row of 3–4 compressed supratympanic spines in C. nemoricola and C. grandisquamis; 8–9 compressed spines in C. calotes; two parallel rows of compressed supratympanic scales in C. maria and C. jerdoni; single well developed postorbital spine in C. emma). Calotes cf. versicolor can be distinguished from its other Sri Lankan congeners based on posterodorsal orientation of lateral scales and absence of oblique fold or pit in front of the shoulder. (Pal et al. 2018: 422) 
CommentSynonymy partly after BOULENGER 1885, SMITH 1935, and WERMUTH 1967. Note that Calotes versicolor nigrigularis AUFFENBERG & REHMANN 1993 is a junior primary homonym of Calotes nigrigularis OTA & HIKIDA 1991 (i.e. it has the same name but is a different species).

Distribution: Very common in Sri Lanka (BAHIR & SURASINGHE 2005). See map in SMID et al. 2014 for distribution in Iran. Gowande et al. 2021 (Fig. 1) restricted C. versicolor to S and NW peninsular India, at least based on their sampling. Much of central India represents a different clade, C. vultuosus. However, Gowande et al. did not sample huge swaths of N India and the himalayas, so their conclusions seem a bit preliminary. Possibly in Yunnan (China) but records from there are questionable (Wang et al. 2022). See map in Gowande et al. 2021 (Fig. 1) for both versicolor and vultuosus.

Types: The species was first described as L’Agama arlequin, a deux raies, which translates to Harlequin Agama with two stripes, by Daudin (1802), based on two specimens from Muséum national d'Histoire naturelle, Paris (MNHN), however he was not aware of the origin of these specimens, and thus described the species without type locality. Kuhl (1820) was the first author to restrict Pondicherry as the type locality of Calotes versicolor, and Leschenault, a French Botanist and Ornithologist, as the collector of the specimens examined by Daudin, while Kuhl himself described Agama tiedemanni from the same locality. As of 2016, MNHN hosts only three specimens from Pondicherry, of which only one is collected by Leschenault, which most probably is the syntype of Kuhl’s Calotes tiedemanni (Amarasinghe et al. 2009), the second deposited by Maindros in 1881 and third being deposited almost a century after description of the species, and thus these cannot be the type specimens. An illustration by Seba cannot be the type (Gowande et al. 2016).

NCBI taxonID: 2824826 [farooqi]
NCBI taxonID: 2824827 [vultuosa] 
EtymologyFrom versi = variable and colour (color), referring to the variability of the species. 
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