A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka Author Thasun Amarasinghe, A. A. Author Suranjan Karunarathna, D. M. S. Author Hallermann, Jakob Author Fujinuma, Junichi Author Grillitsch, Heinz Author Campbell, Patrick D. text Zootaxa 2014 2014-04-02 3785 1 59 78 journal article http://dx.doi.org/10.11646/zootaxa.3785.1.5 66e8e8e7-9324-451f-9464-2d5b739516b1 1175-5326 285817 A0EBB085-B938-42D9-B008-65EAEAC41043 Calotes liocephalus Günther, 1872 (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, WHT 6504 , 89.3 mm SVL, Agrapatanas. Male, ( ZSM 219 / 1981 ), 90.1 mm SVL, Upcot ( 6 ° 46 ′ 56 ″N, 80 ° 37 ′ 32 ″E). Female, WHT 6503 , 79.3 mm SVL, Agrapatanas. Female, (WHT 1667 ), 86.8 mm SVL, Moray Estate, Rajamally. Female, NMB 3354 , 63.9 mm SVL, Talawakelle-Dickoya. Description. (Based on holotype ). Fig. 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 ( 8 th 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; TL 1 , 6.2 ; TL 2 , 8.1 ; TL 3 , 15.2 ; TL 4 , 19.3 ; TL 5 , 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 . 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 ( NHMW 21097 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 WHT 1667 ). 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 ( 7 th 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 . 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, WHT 6504 , 89.3 mm SVL, Agrapatanas. Male, ( ZSM 219 / 1981 ), 90.1 mm SVL, Upcot. Male, ( NHMW 21097 ), 97.0 mm SVL), Ceylon . Female, WHT 6503 , 79.3 mm SVL, Agrapatanas. Female, (WHT 1667 ), 86.8 mm SVL, Moray Estate, Rajamally. Female , NMB 3354 , 63.9 mm SVL, Talawakelle-Dickoya .