Calotes liocephalus Günther, 1872

Thasun Amarasinghe, A. A., Suranjan Karunarathna, D. M. S., Hallermann, Jakob, Fujinuma, Junichi, Grillitsch, Heinz & Campbell, Patrick D., 2014, A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka, Zootaxa 3785 (1), pp. 59-78 : 68-78

publication ID	https://doi.org/ 10.11646/zootaxa.3785.1.5
publication LSID	lsid:zoobank.org:pub:A0EBB085-B938-42D9-B008-65EAEAC41043
DOI	https://doi.org/10.5281/zenodo.5697598
persistent identifier	https://treatment.plazi.org/id/03C187D6-581E-7805-FF22-BF77FE767EF3
treatment provided by	Plazi
scientific name	Calotes liocephalus Günther, 1872
status

Treatment

(Figs. 2, 4–7; Tables 1, 2)

Holotype. BMNH 1946.8.11.33, adult male, SVL 91.4 mm, collected from Sri Lanka by G. H. K. Thwaites; date unknown.

Measured voucher specimens. Male, ( NMB 3353), 78.8 mm SVL, Talawakelle-Dickoya. Male, ( BMNH 95.7.24), 85.8 mm SVL, Pundaluoya. Male, WHT6504, 89.3 mm SVL, Agrapatanas. Male, ( ZSM 219/1981), 90.1 mm SVL, Upcot (6°46′56″N, 80°37′32″E). Female, WHT6503, 79.3 mm SVL, Agrapatanas. Female, (WHT1667), 86.8 mm SVL, Moray Estate, Rajamally. Female, NMB 3354, 63.9 mm SVL, Talawakelle-Dickoya.

Description. (Based on holotype). Fig. 5 View FIGURE 5 . An adult male, 91.4 mm SVL; head moderately large (HL 35.3% of SVL), elongate (HW 66.2% of HL), narrow (HW 23.4% of SVL), distinct from neck; snout elongate (ES 22.0% of HW); eye diameter greater than snout length (ED 183.0% of ES); interorbital distance narrow (IO 14.5% of HL); eye large (ED 26.6% of HL); pupil rounded; ear opening shallow, its greatest diameter dorsoventrally, around ear with keeled scales, tympanum smaller than orbit ( TYD 57.0% of ED); two tubercle like spines above the tympanum separated from the tympanum by four smaller scale rows; diameter of eyes greater than eye to ear distance (ED 119.4% TYE); forehead concave; scales on snout smooth, similar in size to those of occipital region and forehead; scales on interorbital and supercillium area smooth; nuchal crest continuous with dorsal crest and dorsal crest rudimentary, consist of 14 spines till the level of axilla; rostral scale with equal width and height, ventroposteriorly in contact with first supralabial, contacted posteriorly 3 equal sized postrostral scales, but exclusive of prenasals; around nostrils on each side one supranasal, two postnasals, one prenasal and two subnasals; among postnasals the lower one is larger; nostrils oval located slightly more posterior in an undivided nasal plate; canthus rostralis and supraciliary edges soft; 8 scales on canthus rostralis; parietal plate larger than adjacent plates, 11 scales around the parietal plate; Mental subtriangular, lengthen posteriorly, posteriolaterally in contact with two enlarged postmentals separated by a smaller scale preventing contact between them; each postmental pair bordered posteriorly by 3 smooth scales including the medial scale, but exclusive of infralabial. Gular pouch present; throat scales keeled; mid gular scales equal in size with those besides, strongly keeled, pointed and overlapping; 3 scale rows separate orbit from supralabials; supralabials 10 (8th in mid orbit position) on left side; infralabials 9, decreasing in size towards gape; ventral scales on the neck keeled, pointed and overlapping.

Body slender (AG 45.8% of SVL); mid dorsal scales equal, keeled, with pointed dorsal scales at midbody; scales on dorsum at midbody larger in size with those of venter at same level; lateral body scales slightly keeled, smaller than dorsals; directed backwards and downwards; 46 scales around midbody; pectoral scales enlarged, carinate and overlapping; abdominal scales completely and strongly carinate, and mucronate, not enlarged, pointed, overlapped with keels forming regular, parallel, continuous ventral ridges; mid ventral scale row, 70.

Forelimbs moderately short (LAL 20.8% of SVL, UAL 16.8% of SVL); hind limbs relatively long (TBL 24.6% of SVL, FEL 64.1% of SVL); tibia comparatively long (FEL 92.0% of TBL). Dorsal scales on fore and hind limbs slightly keeled, overlapped; ventral scales on upper arm smooth and lower arm, keeled, overlapped, and pointed; scales on ventral surface of thigh slightly keeled, overlapped and pointed; keels on tibia forming a series of continuous parallel ridges. Digits elongate, slender, all bearing slightly recurved claws; claws are sharp and elongate; subdigital lamellae entire and regular, subdigital lamellae on the toe IV, 32; inter-digital webbing absent; relative length of digits (fingers) 4 ≥ 3> 2> 5> 1; (toes) 4> 3> 5> 2> 1.

Tail complete (258.0 mm); tail base swollen, ventral scales on tail base bluntly pointed, keeled, overlapped; dorsal scales on tail pointed, not elongate, overlapped, directed backwards, keels forming continuous parallel ridges; tail with subcaudals shortened, median row not enlarged, keeled, mucronate, overlapped.

Measurements in millimetres. HL, 32.3; HW, 21.4; HD, 18.2; EN, 6.2; ES, 4.7; TYD, 4.9; IN, 6.0; IO, 4.7; TBW, 9.4; SVL, 91.4; AG, 41.9; TAL, 258; ED, 8.6; TYE, 7.2; UAL, 15.4; LAL, 19.0; FEL, 20.7; TBL, 22.5; FOL, 31.0; TL1, 6.2; TL2, 8.1; TL3, 15.2; TL4, 19.3; TL5, 12.9.

Colour in preserved specimen. Head and dorsum bluish grey, posteriorly becomes brown; six “V” shaped lighter sky blue markings on the body posteriorly becoming brown, the first marking on the neck reaches the eye which is black in colour, across the tympanum; lighter cross marking on inter orbital; spines black grey and pale white in colour; limbs with darker 6–7 cross bars on each limb, but unclear markings on the hind limb; throat and gular are a pale sky blue colour; ventral surface of the limbs, pectoral region, abdomen, and ventral surface of tail pale white; shoulder pit sky blue in colour; dorsal surface of the tail base bright olive green and the rest pale grey.

Colour in life. Fig. 6 View FIGURE 6. A . Based on personal observations of five males (not collected) from Peak Wilderness (1500–1800 m elevations); head and dorsum bright olive yellow, or light olive green, posteriorly becoming brown and grey; six “V” shaped dark chocolate brown markings on the body, the first marking on the neck continues to SUP under the eye across the tympanum; black colour markings on the supranasals, temporal, supracilliars and inter orbital; spines black and olive-yellow in colour; limbs with dark brown 6–7 cross bars on each limb; throat and gular pale white or very light sky bluish-white, with black, brown, grey colour with faint blotches; ventral surface of the fore limbs bright olive yellow and pectoral region bright orange-yellow or completely brownish orange; abdomen, ventral surface of thigh and ventral surface of tail are an off white colour with faint light brown markings; tibia an off white colour with light brown cross bars on the ventral surface; digits black with brownishgrey cross markings; tail brown in colour and with grey and dark brown markings.

Variation of males. Rostral scale width greater than its height; dorsal crest consists of 10–15 spines till the level of axilla; 7–9 scales on canthus rostralis; 9–10 scales around the parietal plate; each postmental pair bordered posteriorly by 4 smooth scales including the medial scale, but exclusive of infralabial (NHMW21097 has two medial scales); supralabials, 9–10; infralabials, 9–10; 48–50 scales around the midbody; ventrals, 64–79; subdigital lamellae on the toe IV, 27–32.

Description of female. (Based on WHT1667). An adult female, 86.8 mm SVL; head moderately large (HL 33.0% of SVL), elongate (HW 59.1% of HL), narrow (HW 19.5% of SVL), distinct from neck; snout elongate (ES 60.9% of HW); snout length greater than eye diameter (ED 82.5% of ES); interorbital distance narrow (IO 16.8% of HL); eye large (ED 29.7% of HL); pupil rounded; ear opening shallow, its greatest diameter dorsoventrally; keeled scales around ear; tympanum smaller than orbit ( TYD 42.3% of ED); two tubercle like spines above the tympanum separated from the tympanum by four smaller scale rows; diameter of eyes greater than eye to ear distance (ED 132.8% TYE); forehead concave; scales on snout smooth, similar in size those of occipital region and forehead; scales on interorbital and supercillium area smooth; nuchal crest continuous with dorsal crest and dorsal crest rudimentary, consist of 10 spines till the level of axilla; rostral scale much wider than high, ventroposteriorly in contact with first supralabial, in contact posteriorly with four equal sized postrostral scales; around nostrils on each side one supranasal, two postnasals, two prenasal and a subnasals; the upper postnasals beings larger in size; nostrils round are located posteriorly in undivided nasal plate; canthus rostralis and supraciliary edges sharp; 7 canthus rostralis; parietal plate slightly larger than adjacent plates, 10 scales around the parietal plate; Mental subtriangular, lengthen posteriorly, about as long as wide, posteriolaterally in contact with two enlarged postmentals separated by a smaller scale with no contact between them; each postmental pair bordered posteriorly by three smooth scales including the medial scale, but exclusive of infralabial. No gular pouch; throat scales strongly keeled, bluntly pointed and overlaped; mid gular scales strongly keeled, similar in size to those of adjacent, pointed and overlapped; three scale rows separate orbit from supralabials; supralabials 9 (7th in mid orbit position); infralabials 8, decreasing in size towards gape; ventral scales on the neck keeled, mucronate and overlapping.

Body slender (AG 52.6% of SVL); mid dorsal scales equal, keeled, with pointed dorsal scales at midbody; scales on dorsum at midbody larger in size to those of venter at the same level; lateral body scales smooth, smaller than dorsals; directed backwards and downwards; 52 scales around the midbody; pectoral scales not enlarged, keeled, pointed and overlapping; abdominal scales partially and slightly carinate, and acuminate, and keels forming regular and parallel continuous ventral ridges; ventrals, 78.

Forelimbs moderately short (LAL 19.5% of SVL, UAL 17.8% of SVL); hind limbs relatively long (TBL 26.5% of SVL, FEL 24.8% of SVL); tibia comparatively long (FEL 93.5% of TBL). Dorsal scales on fore and hind limbs smooth, overlapped; ventral scales on upper arm smooth and on lower arm keeled, overlapped, and pointed; scales on ventral surface of thigh slightly keeled, overlapped and pointed; keels on tibia forming a series of continuous parallel ridges. Digits elongate, slender, all bearing slightly recurved claws; claws are sharp and elongate; subdigital lamellae entire and regular, subdigital lamellae on the toe IV, 36; inter-digital webbing absent; relative length of digits (fingers) 3> 4> 2> 5> 1; (toes) 4> 3> 5> 2> 1.

Tail complete (231.5 mm); tail base swollen, ventral scales on tail base bluntly pointed, keeled, overlapped; dorsal scales on tail pointed, elongate, overlapped, directed backwards, keels forming continuous parallel ridges; tail with subcaudals shortened, median row not enlarged, keeled, and overlapped.

Variation of females. Dorsal crest consists of 13 and 15 spines up to the point of the axilla; 7 and 8 scales on canthus rostralis; supralabials, 11; infralabials, 9; 44 and 53 scales around the midbody; ventrals, 87; subdigital lamellae on the toe IV, 30.

Distribution and habitat. Fig. 4 View FIGURE 4 . We have observed live specimens (not collected) from Madahinna (~ 1500 m elevation), Haramitipana (1700 m elevation), and Dharmaraja gala (1600 m elevation) of the Peak Wilderness.

The habitats where C. liocephalus occur, is home to many large canopy trees (~ 30 m) such as Adinandra lasiopetala , Bhesa ceylanica , Calophyllum trapezifolium , Cullenia ceylanica , Shorea affinis , S. gardneri , Litsea gardneri , and Palaquium rubiginosum and subcanopy level (~ 15 m) consists of Apodytes dimidiata , Artocarpus nobilis , Calophyllum walkeri , Caryota urens , Cinnamomum ovalifolium , Cryptocarya wightiana , Dillenia triquetra , Elaeocarpus amoenus , Eugenia mabaeoides , Garcinia quaesita , Gordonia speciosa , Madhuca moonii , Mesua ferrea , Oncosperma fasciculatum , Schumacheria alnifolia , Stemonoporus gardneri , S. oblongifolia , Syzygium firmum , and S. turbinatum . Also ground cover (~ 3 m) consists of Acronychia pedunculata , Agrostistachys coriacea , Alpinia abundiflora , Amomum echinocarpum , Amomum masticatorium , Amorphophallus paeoniifolius , Arundina graminifolia, Calanthes sp., Cinnamomum verum , Clusia rosea , Cyathea crinita , Hedychium coronarium , Hortonia ovalifolia , Ipsea speciosa , Macaranga indica , Neolitsea cassia , Osbeckia aspera , Osbeckia lantana , Rhodomyrtus tomentosa , Strobilanthes sp., Syzygium cordifolium , Syzygium revolutum , and Utricularia striatula . Most of the habitats had 60–70% (mean 62.4 ± 4.7%) canopy cover and the undergrowth consists of shrubs and herbs. Average annual rainfall varies from 3,000–4,500 mm and the average annual temperature is 27.9°C. The range of temperature and range of humidity were 26.4–28.6°C (mean 27.3 ± 0.5°C) and 66–78% (mean 71.4 ± 3.2%), respectively.

Natural history. This species also seems to be extremely rare, but we have observed higher numbers of this species in the ecotone than in the dense forest. In Peak Wilderness (> 1400 m elevations), C. liocephalus is sympatric with Ceratophora stoddartii and Calotes calotes , but allopatric with C. nigrilabris (see Amarasinghe et al. 2012). This species is a fast moving agamid. We have observed six ovipositioning at Peak Wilderness during October –November in 2010–2012. All the ovipositioning were observed during cool and shady weather conditions (cloud cover, 50–70%; canopy cover, 15–30%) during 10:00–15:00 hr (temperature, 26.4–28.2°C; humidity, 60– 70%; light intensity, 5.3–7.3 lux). The soil was always soft-tan colouration and the leaf litter was usually 12–18 mm thick. The nest-holes were 52–58 mm (55 ± 2 mm) deep at an angle of 40–50° to the ground. The diameters of the holes were 32–38 mm (35 ± 2 mm) and the body pits were in 30–35mm radius (32 ± 2 mm). This species lays 3– 4 eggs at a time, and the eggs were 18.5–18.9 mm (18.7 ± 0.1 mm) in length and 10.1–10.4 mm (10.2 ± 0.1 mm) in width. The weight ranged between 1.4–1.8 g (1.6 ± 0.1 g) in a range. The incubation period varies from 65–72 days. The ovipositional behaviour of C. liocephalus is not described here.

Calotes liocephalus . Male, ( BMNH 1946.8.11.33), 91.4 mm SVL, Sri Lanka. Male, ( NMB 3353), 78.8 mm SVL, Talawakelle- Dickoya. Male, ( BMNH 95.7.24), 85.8 mm SVL, Pundaluoya. Male, WHT6504, 89.3 mm SVL, Agrapatanas. Male, ( ZSM 219/1981), 90.1 mm SVL, Upcot. Male, ( NHMW 21097), 97.0 mm SVL), Ceylon. Female, WHT6503, 79.3 mm SVL, Agrapatanas. Female, (WHT1667), 86.8 mm SVL, Moray Estate, Rajamally. Female, NMB3354 , 63.9 mm SVL, Talawakelle-Dickoya .