Calotes pethiyagodai, Amarasinghe, Karunarathna & Hallermann, 2014

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 : 62-78

Calotes pethiyagodai sp. nov. Amarasinghe, Karunarathna & Hallermann

( Figs. 1–4 , 7 ; Tables 1, 2)

Holotype. Male, WHT6211 , 91.8 mm SVL, near Midland Estate, Knuckles , Sri Lanka, 7°31’N, 80°44’ E, alt. 915 m, coll. M. M. Bahir, A. Silva & K. Maduwage, 24 IX 2004.

Paratypes. Male, WHT6154A, 91.3 mm SVL, Midlands Estate-Knuckles, alt. 915 m, coll. M. M. Bahir & M. Meegaskumbura, 5 VI 2004; Male, WHT6241, 88.9 mm SVL, Cobet’s Gap-Knuckles, alt. 1,000, coll. K. Manamendra-Arachchi & M.M. Bahir; Males, ZSM 215/1981/3–4, 59.4 mm SVL (sub adult), 76.8 mm SVL, Gammaduwa-Knuckles, W. Erdelen, 21 VI 1980; Male, ZSM 216/1981/1, 86.1 mm SVL, Cobet’s Gap-Knuckles, W. Erdelen, 27 VI 1980; Male, ZSM 258/1979, 76.3 mm SVL, Cobet’s Gap-Knuckles, W. Erdelen, 0 1 IV 1979; Males, ZSM 218/1981/1–3, 81.5 mm SVL, 85.5 mm SVL, 76.7 mm SVL, Midcar-Knuckles, W. Erdelen, 0 5 XII 1979; Female, WHT 6154B, 80.8 mm SVL, Midlands Estate-Knuckles, alt. 915 m, coll. M. M. Bahir & M. Meegaskumbura, 5 VI 2004; Female, WHT106A, 78.9 mm SVL, Gammaduwa Estate-Knuckles, alt. 915 m, coll. K. Manamendra-Arachchi & D. Gabadage, 23 X 1993; Female, WHT1435, 77.2 mm SVL, Midlands Estate- Knuckles, coll. D. Gabadage & M.M. Bahir, 3 IX 1996. Females, ZSM 215/1981/1–2, 75.8 mm SVL, 78.3 mm SVL, Gammaduwa-Knuckles, W. Erdelen, 21 VI 1980; Female, ZMH R06165, 76.5 mm SVL, Gammaduwa- Knuckles, W. Erdelen, 7°34'00"N 80°42'00"E, coll. W. Erdelen, 1972; Female, ZSM 217/1981, 66.5 mm SVL, Cobet’s Gap-Knuckles, W. Erdelen, 19 VI 1979; Females, ZSM 218/1981/4–5, 71.8 mm SVL, 55.8 mm SVL (sub adult), Midcar-Knuckles, 0 5 XII 1979.

Diagnosis. Males of C. pethiyagodai sp. nov. differ from the males of C. liocephalus by the absence of a gular pouch (vs. present); mid gular scales smaller (vs. equal or larger) than the scales besides; scales on snout larger (vs. smaller) than the scales on occipital and forhead; pectoral scales not enlarged (vs. enlarged); subcaudals elongated (vs. shortened); abdominal scales partially and slightly carinate, and acuminate (vs. completely and strongly carinate, and mucronate); scales on venter somewhat larger in size than those on dorsum at same level (vs. smaller); tubercle like spine above the tympanum, one (vs. two); axila–groin, tibia, forth toe, upper arm, and snout, longer (151.4–153.3%, 69.4–73.7%, 63.0–75.7%, 51.4–53.9%, and 36.6–37.9% of HL respectively) vs. shorter (109.4–129.7%, 65.5–67.9%, 51.9–59.7%, 47.7–51.3%, and 27.8–33.7% of HL respectively); larger eye, diameter 28.7–29.5% of HL (vs. smaller, diameter 26.6–28.5% of HL); mid body scales, 50–54 (vs. 46–50); and subdigital lamellae, 36–38 (vs. 27–32). C. pethiyagodai sp. nov. further differs from congeners by the following opposing characters of Sri Lankan and Indian species of the genus Calotes ; C. andamanensis : enlarged keeled scales on ventral surface of thigh; C. aurantolabium : shoulder pit absent; C. rouxii : two small groups of spines on each side of the neck; C. calotes , C. grandisquamis , C. jerdoni , C. nemoricola , C. versicolor : lateral scales directed backwards and upwards; and C. ceylonensis , C. desilvai , C. elliotti , C.emma , C. liolepis , C. maria , C. nigrilabris : well-developed spines above the tympanum.

Description. (Based on holotype). Fig. 1 . An adult male, 91.8 mm SVL; head moderately large (HL 35.6% of SVL), elongate (HW 57.8% of HL), narrow (HW 20.6% of SVL), distinct from neck; snout elongate (ES 65.1% of HW); snout length greater than eye diameter (ED 77.2% of ES); interorbital distance narrow (IO 15.0% of HL); eye large (ED 29.0% of HL); pupil rounded; ear opening shallow, its greatest diameter dorsoventrally; smooth scales around ear; tympanum smaller than orbit ( TYD 40.0% of ED); one tubercle like spine above the tympanum separated from the tympanum by three smaller scale rows; diameter of eyes greater than eye to ear distance (ED 128.4% TYE); forehead concave; scales on snout smooth, larger in size those of occipital region and forehead; scales on interorbital and supercillium area smooth; nuchal crest continue with dorsal crest and dorsal crest rudimentary, consist of 15 spines till the level of axilla; rostral scale width greater than its height, ventroposteriorly in contact with first supralabial, also in contact posteriorly with three equal sized postrostral scales; around nostrils on each side one supranasal, two postnasals, a prenasal and two subnasals; among postnasals the lower one being larger; nostrils oval and located slightly more posterior in an undivided nasal plate; canthus rostralis and supraciliary edges soft; 7 scales on canthus rostralis; parietal plate larger than adjacent plates, 11 scales around the parietal plate; Mental subtriangular, lengthen posteriorly, its length approximately equal to its width, posteriolaterally in contact with two enlarged postmentals separated by a smaller scale with no contact between them; each postmental pair bordered posteriorly by 3 smooth scales including the medial scale, but exclusive of infralabial. No gular pouch; throat scales slightly keeled, bluntly pointed and overlapped; mid gular scales strongly keeled, smaller in size than those of besides, pointed and overlapped; five scale rows separate orbit from supralabials; supralabials 10 (7th in mid orbit position); infralabials 9, decreasing in size towards gape; ventral scales on the neck keeled, mucronate and overlapping.

Body slender (AG 53.9% of SVL); mid dorsal scales equal, smooth, with pointed dorsal scales at midbody; scales on dorsum at midbody smaller in size than those of venter at same level; lateral body scales keeled, smaller than dorsals; directed backwards and downwards; 54 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, 88.

Forelimbs moderately short (LAL 19.7% of SVL, UAL 18.4% of SVL); hind limbs relatively long (TBL 24.7% of SVL, FEL 24.4% of SVL); tibia comparatively long (FEL 98.7% of TBL). Dorsal scales on fore and hind limbs keeled, overlapped; ventral scales on upper and lower arm keeled, overlapped, and pointed; scales on ventral surface of thigh strongly 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 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 (216.1 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 elongated, median row not enlarged, keeled, and overlapped.

Measurements in millimetres. HL, 32.7; HW, 18.9; HD, 16.3; EN, 6.6; ES, 12.3; TYD, 3.8; IN, 7.4; IO, 4.9; TBW, 10.7; SVL, 91.8; AG, 49.5; TAL, 216.1; ED, 9.5; TYE, 7.4; UAL, 16.9; LAL, 18.1; FEL, 22.4; TBL, 22.7; FOL, 31.0; T1, 5.2; T2, 10.4; T3, 14.1; T4, 23.6; T5, 10.8.

Colour in preserved specimen. Dorsum greenish blue; 8 “V” shaped black markings along the vertebral, incomplete black cross bars on limbs; tail brownish, light coloured 8 cross bars; tail base olive green; venter light bluish white, throat white; palm cream colour; tympanum sky blue; shoulder pit brownish black.

Colour in life. Fig. 3 . Based on personal observations of five males (not collected) from Riverstone-Knuckles; have dorsum bright green and colour can be bluish, or light brown in hind parts of the body, tail base olive green; around eight “V” shaped light blue or black coloured markings along the dorsal surface, incomplete greenish brown or black cross bars on limbs; two to six black cross bands between eyes; labials sky blue or yellowish green; tympanum light yellow; tail brownish, cream coloured or black 8–10 cross bars; venter bright yellow or creamy white, throat bright yellow, yellowish green or sky blue; palm dirty white. When aggressive: supralabials, around eye and tympanum darken and the rest of the body becomes dark brown. Juveniles are generally bright green in colour with brown tails, when disturbed: body colour they become dark brown.

Variation in male paratypes. dorsal crest consists of 6–13 spines up to the point of the axilla; 7–8 scales on canthus rostralis; 12 scales around the parietal plate; each postmental pair bordered posteriorly by 3 and 4 smooth scales including the medial scale, but exclusive of infralabial; supralabials, 8–11; infralabials, 8–10; mid-ventral scale row, 65–94; subdigital lamellae on the toe IV, 36–38.

Description of female. (Based on paratype WHT 6154B). An adult female, 80.8 mm SVL; head moderately large (HL 32.5% of SVL), elongate (HW 55.5% of HL), narrow (HW 18.1% of SVL), distinct from neck; snout elongate (ES 71.2% of HW); snout length greater than eye diameter (ED 70.2% of ES); interorbital distance narrow (IO 17.5% of HL); eye large (ED 27.7% of HL); pupil rounded; ear opening shallow, its greatest diameter dorsoventrally; keeled scales around ear; tympanum smaller than orbit ( TYD 54.8% of ED); one tubercle like spine above the tympanum separated from the tympanum by three smaller scale rows; diameter of eyes greater than eye to ear distance (ED 114.0% TYE); forehead concave; scales on snout keeled, larger in size those of occipital region and forehead; scales on interorbital and supercillium area keeled; nuchal crest continuous with dorsal crest and dorsal crest rudimentary, consist of 14 spines till the level of axilla; rostral scale width much greater than its height, ventroposteriorly in contact with first supralabial, posteriorly in contact with three equal sized postrostral scales; around nostrils on each side one supranasal, two postnasals, a prenasal and a subnasals; among postnasals the upper one is larger; nostrils oval located slightly more posterior in an undivided nasal plate; canthus rostralis and supraciliary edges soft; 8 canthus rostralis; parietal plate larger than adjacent plates, 11 scales around the parietal plate; Mental subtriangular, lengthen posteriorly, the width and length of which equal, posteriolaterally in contact with two enlarged postmentals separated by a smaller scale with no contact between them; each postmental pair bordered posteriorly by 3 smooth scales including the medial scale, but exclusive of infralabial. No gular pouch; throat scales slightly keeled, bluntly pointed and overlaped; mid gular scales slightly keeled, pointed and overlapped; three scale rows separate orbit from supralabials; supralabials 9 (7th in mid orbit position); infralabials 9, decreasing in size towards gape; ventral scales on the neck keeled, mucronate and overlapping.

Body slender (AG 48.8% of SVL); mid dorsal scales equal, smooth, with pointed dorsal scales at midbody; scales on dorsum at midbody smaller in size to those of venter at same level; lateral body scales keeled, 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. Keels forming regular and parallel continuous ventral ridges; mid-ventral scale row, 78.

Forelimbs moderately short (LAL 20.0% of SVL, UAL 16.5% of SVL); hind limbs relatively long (TBL 26.8% of SVL, FEL 23.4% of SVL); tibia comparatively long (FEL 88.3% of TBL). Dorsal scales on fore and hind limbs slightly keeled, overlapped; ventral scales on upper and lower arm keeled, overlapped, and pointed; scales on ventral surface of thigh strongly 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 elongated, median row not enlarged, keeled, and overlapped.

Variation in female paratypes. dorsal crest consists of 5–13 spines up to the point of the axilla; 7–9 scales on canthus rostralis; supralabials, 8–11; infralabials, 8–10; mid-ventral scale row, 65–94; subdigital lamellae on the toe IV, 36–38.

Etymology. The species epithet is an eponym latinized in the genitive singular, honouring Tilak Rohan David Pethiyagoda (Pethiyagoda, R.), a Rolex awarded conservationist and the founder of the Wildlife Heritage Trust of Sri Lanka (WHT), for his dedication and contribution to biodiversity conservation in Sri Lanka; his leading contribution to herpetological and ichthyological explorations in the Indian Subcontinent; and the great work he has done in order to restore and preserve the forests in the central highlands is highly commendable. His contributions are extremely important and they undoubtedly inform the many new taxonomists emerging out of India and Sri Lanka, especially at a time such as now when new taxonomists are very much in demand. Suggested English name: Pethiyagoda’s Crestless Lizard; Sinhala (local) name: Pethiyagodagë Nosilu Katussa; Tamil (local) name: Pethiyagodavin Oonan.

Distribution and habitat. Fig. 4 . We have observed a number of live specimens (not collected) from Riverstone (1200 m elevation), Dotalugala (~ 1500 m elevation), Gammaduwa (~ 900 m elevation), Kobonilagala (1400 m elevation), Rangala (1400 m elevation), Cobet’s Gap (1000 m elevation) and Thangappuwa near Cobet’s Gap (1000 m elevation) of the Knuckles massif.

The habitats where C. pethiyagodai sp. nov. occurs, is home to many medium canopy trees (~ 8 m) such as Creteava religiosa, Phyllanthus indica , Sterculia foetida , Bombax ceiba , Dimocarpus longan , Palaquium hinmolpedda and Vitex altissima species. The subcanopy level (~ 5 m) consists of Breynia vitisidea , Miliusa indica , Pavetta indica and Streblus asper species. Ground cover (~ 2 m) consists of Begonia hirtella , Carex filicina , Carex jakiana , Curculio orchioides and Procris crenata species. The Knuckles forest range is extremely wet throughout the year, with an average annual rainfall> 4000 mm, though the lower eastern slopes are much drier. Most of the habitats had 50–65% (mean 57.7 ± 7.5%) canopy cover and the undergrowth consists of shrubs and herbs. The range of temperature and range of relative humidity are 25.5–26.8°C (mean 26.2 ± 0.7°C) and 65–74% (mean 69.0 ± 4.6%), respectively. The evergreen submontane forests represent the major natural vegetation type in the Riverstone area.

Natural history. This species seems to be extremely rare, but we have observed higher numbers of this species in the ecotone than in the dense forest. C. pethiyagodai sp. nov. is sympatric with Calotes cf. liolepis (see Amarasinghe et al. 2014), Ceratophora tennentii (see Pethiyagoda & Manamendra-Arachchi 1998) and Cophotis dumbara (see Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006). During the mid-day time, most individuals prefer to inhabit trees (> 3 m from the ground level) in shady areas, in the early morning and at night they are found on shrubs (~ 2 m). Juveniles are usually found on shrubs (~ 1 m), especially basking in the sun between 9:00–11:00 hr as bright sunlight is not very common in the Knuckles. We observed predation on dragonflies, butterflies, and moths. This species is a fast moving agamid, well adapted to climbing trees and it also jumps from one shrub to another (~ 1 m distance). The ovipositional behaviour of C. liocephalus described by Amarasinghe & Karunarathna (2008) in the Knuckles also refers to C. pethiyagodai sp. nov.

Calotes pethiyagodai . Male (WHT6211), 91.8 mm SVL, near Midland Estate, Knuckles. Male, (WHT6154A), 91.3 mm SVL, Midlands Estate-Knuckles. Male, (WHT6241), 88.9 mm SVL, Cobet’s Gap-Knuckles. Female, (WHT 6154B), 80.8 mm SVL, Midlands Estate-Knuckles. Female, (WHT106A), 78.9 mm SVL, Gammaduwa Estate-Knuckles. Female, (WHT1435), 77.2 mm SVL, Midlands Estate-Knuckles. Male, ( ZSM 215/1981/3–4), 59.4 mm SVL, 76.8 mm SVL, Gammaduwa-Knuckles. Male, ( ZSM 216/1981/1), 86.1 mm SVL, Cobet’s Gap-Knuckles. Male, ( ZSM 258/1979), 76.3 mm SVL, Cobet’s Gap-Knuckles. Males, ( ZSM 218/1981/1–3), 81.5 mm SVL, 85.5 mm SVL, 76.7 mm SVL, Midcar- Knuckles. Female, (WHT 6154B), 80.8 mm SVL, Midlands Estate-Knuckles. Female, (WHT106A), 78.9 mm SVL, Gammaduwa Estate-Knuckles. Female, (WHT1435), 77.2 mm SVL, Midlands Estate-Knuckles. Females, ( ZSM 215/ 1981/1–2), 75.8 mm SVL, 78.3 mm SVL, Gammaduwa-Knuckles. Female, ( ZMH R06165), 76.5 mm SVL, Gammaduwa- Knuckles. Female, ( ZSM 217/1981), 66.5 mm SVL, Cobet’s Gap-Knuckles. Females ( ZSM 218/1981/4–5), 71.8 mm SVL, 55.8 mm SVL, Midcar-Knuckles. Juvenile ( ZSM 214/1981), Gammaduwa-Knuckles. Juvenile, ( ZSM 216/1981/2), Cobert’s Gap.

Figures

FIGURE 1. Holotype male (86.6 mm SVL) of Calotes pethiyagodai sp. nov., WHT 6211; a, dorsal head; b, lateral head; c, ventral head; d, pectoral scales: note greater snout scales and smaller occipital scales (a); longer snout (b); smaller mid gular scales (c), and not-enlarged pectoral scales (d), (Photos: M. De Silva).

FIGURE 2. Scales on abdomen of; a, Calotes pethiyagodai sp. nov., WHT 6211, holotype (86.6 mm SVL) and; b, Calotes liocephalus, BMNH 1946.8. 11.33, holotype (91.4 mm SVL): note partially and slightly carinate, and acuminate abdominal scales of C. pethiyagodai sp. nov. and completely and strongly carinate, and mucronate scales of C. liocephalus; subcaudals of; c, Calotes pethiyagodai sp. nov., WHT 6211, holotype (86.6 mm SVL) and; d, Calotes liocephalus, BMNH 1946.8. 11.33, holotype (91.4 mm SVL): note elongate subcaudal scales of C. pethiyagodai sp. nov. and shortened scales of C. liocephalus (Illustration: A. A. T. Amarasinghe).

FIGURE 3. A live adult male (86.6 mm SVL) of C. pethiyagodai sp. nov. (not collected) at 1300 m elevation of the Knuckles (Photo: V. Weeratunge).

FIGURE 4. Distribution map of C. liocephalus (red), and C. pethiyagodai sp. nov. (blue) based on examined specimens and personal observations made over the past 10 years.

FIGURE 7. Habitat of; a, C. pethiyagodai sp. nov. at high elevations of the Knuckles massif; b, C. liocephalus at mid elevations of Peak Wilderness (Photos: W. M. S. Botejue).