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A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka

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A new species of agamid lizard, of the genus Calotes, is described based on morphological evidence. This species is restricted to the Knuckles massif (>900 m elevation) of Sri Lanka. The genus Calotes consists of seven species in Sri Lanka, five of which appear to form an endemic radiation. The new species most closely resembles C. liocephalus Günther, 1872 which has an isolated population in the central highlands and is only known from Pundaluoya (~1000m), Dickoya (~1200m), Upcot (~1400m), Agrapatanas (1665m) and Peak Wilderness (Sri Pada) (>1400m). The populations from Pundaluoya and Dickoya appear to be locally extinct from the wild and are known only from museum specimens collected over 120 years ago. Males of the new species are different from males of C. liocephalus because of the absence of a gular pouch; by having mid gular scales smaller in size than those of its counterpart; scales on the snout which are larger in size than those on the occipital and forehead; pectoral scales which are not enlarged; elongated subcaudal scales; slightly carinate and acuminate abdominal scales; and scales on venter which are somewhat larger in size than those on dorsum at the same level. Finally, we also redescribe Calotes liocephalus, and provide a key to the Sri Lankan species of genus Calotes.
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Accepted by S. Carranza: 25 Feb. 2014; published: 2 Apr. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3785 (1): 059078
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http://dx.doi.org/10.11646/zootaxa.3785.1.5
http://zoobank.org/urn:lsid:zoobank.org:pub:A0EBB085-B938-42D9-B008-65EAEAC41043
A new species of the genus Calotes (Squamata: Agamidae) from high elevations
of the Knuckles Massif of Sri Lanka
A. A. THASUN AMARASINGHE1,6, D. M. S. SURANJAN KARUNARATHNA2, JAKOB HALLERMANN3,
JUNICHI FUJINUMA4, HEINZ GRILLITSCH5 & PATRICK D. CAMPBELL6
1Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424, Indonesia
2Nature Exploration and Education Team, No. B–1 / G–6, De Soysapura Housing Scheme, Moratuwa 10400, Sri Lanka
3Zoologisches Museum Hamburg, Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
4Graduate School of Environmental Science, Hokkaido University, Sapporo, Hokkaido, Japan
5 Natural History Museum, Herpetological Collection, Burgring 7, A–1010 Wien, Austria
6Department of Life Sciences, Darwin Centre, Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD,
England
6Corresponding author. E-mail: thasun.taprobanica@gmail.com
Abstract
A new species of agamid lizard, of the genus Calotes, is described based on morphological evidence. This species is re-
stricted to the Knuckles massif (>900 m elevation) of Sri Lanka. The genus Calotes consists of seven species in Sri Lanka,
five of which appear to form an endemic radiation. The new species most closely resembles C. liocephalus Günther, 1872
which has an isolated population in the central highlands and is only known from Pundaluoya (~1000m), Dickoya
(~1200m), Upcot (~1400m), Agrapatanas (1665m) and Peak Wilderness (Sri Pada) (>1400m). The populations from Pun-
daluoya and Dickoya appear to be locally extinct from the wild and are known only from museum specimens collected
over 120 years ago. Males of the new species are different from males of C. liocephalus because of the absence of a gular
pouch; by having mid gular scales smaller in size than those of its counterpart; scales on the snout which are larger in size
than those on the occipital and forehead; pectoral scales which are not enlarged; elongated subcaudal scales; slightly car-
inate and acuminate abdominal scales; and scales on venter which are somewhat larger in size than those on dorsum at the
same level. Finally, we also redescribe Calotes liocephalus, and provide a key to the Sri Lankan species of genus Calotes.
Key words: biogeography, Calotes liocephalus, conservation, Reptilia, systematics, taxonomy
Introduction
The agamid genus Calotes Cuvier, 1817, currently consists of 24 species (Hartmann et al. 2013; Uetz &
Hallermann 2013). Seven are represented in Sri Lanka; five of which appear to form an endemic radiation (Macey
et al. 2000; de Silva 2006; Somaweera & Somaweera 2009; Wickramasinghe 2012): C. nigrilabris Peters, 1860; C.
liocephalus Günther, 1872; C. liolepis Boulenger, 1885; C. ceylonensis Müller, 1887; and C. desilvai Bahir and
Maduwage, 2005. Calotes calotes (Linnaeus, 1758) is distributed in India, and C. versicolor (Daudin, 1802) is
widely spread throughout tropical Asia (Smith 1935; Deraniyagala 1953; Taylor 1953; Moody 1980; Manthey
2008; Das 2010; Venugopal 2010). Both complexes in Sri Lanka are in need of further taxonomic clarification (see
Zug et al. 2006; Hartmann et al. 2013).
Calotes liocephalus is considered to be restricted to the Knuckles massif (Erdelen 1984; Manamendra
Arachchi & Liyanage 1994) even though there were some older records from the central highlands (Deraniyagala
1953; Smith 1935; Taylor 1953). In 1981 one specimen (ZSM 219/1981) was collected from Upcot (~1400 m
elevation) and in 2005 two specimens (WHT6503, WHT6504) were collected from Agrapatanas (1665 m
elevation) in the central highlands. In addition, Amarasinghe et al. (2009) recorded C. liocephalus from the Peak
Wilderness. After examining several museum specimens at the BMNH, NMB, and ZSM we found three specimens
from Pundaluoya (~1000 m), Dickoya (~1200 m) and Upcot (~1400 m) respectively, in the central highlands of Sri
AMARASINGHE ET AL.
60 · Zootaxa 3785 (1) © 2014 Magnolia Press
Lanka. The species, C. liocephalus was described using a single male specimen (BMNH 1946.8.11.33) by Günther
(1872) without a precise location; therefore we had to compare the holotype with museum specimens and live
specimens (not collected) from the Knuckles massif and central highlands, where both areas are known to be world
heritage sites (UNESCO, 2013). After identifying the specimens,we saw that the holotype resembled the central
highlands populations. Hence, here we describe the population distributed in the Knuckles massif as a distinct
species with a redescription of C. liocephalus based on the holotype.
Material and methods
Museum acronyms follow Sabaj Pérez (2013). Specimens were examined at the Natural History Museum, London,
UK (BMNH); Museum national d’Histoire naturelle, Paris, France (MNHN); Naturhistorisches Museum, Basel,
Switzerland (NMB); National Museum of Sri Lanka, Colombo, Sri Lanka (NMSL); Naturhistorisches Museum
Wien, Vienna, Austria (NHMW); Wildlife Heritage Trust, Colombo, Sri Lanka (WHT); Zoologische
Staatssammlung München, Munich, Germany (ZSM); and Zoological Museum Hamburg, Hamburg, Germany
(ZMH). The WHT collection is now deposited at NMSL (not yet catalogued). Morphometric data for species
comparisons were obtained from examined specimens (Appendix I). We used a Leicawild M3Z and a ZEISS DCR
dissecting microscope to examine the external morphology of specimens and a Cannon EOS 7D SLR digital
camera to take Photographs. The map was made by using Arc Gis 10.1 (ESRI© 1995-2012) software. The
conservation status of the new species was evaluated based on IUCN Red List Categories and Criteria (2001) and
IUCN Standards and Petitions Subcommittee (2013): Versions 3.1, 4.0, and 10.1 to assess their risk of extinction.
Sex was determined by the presence or absence of hemipenes or hemipenal bulges.
The following characters were measured with a Mitutoyo digital caliper to the nearest 0.1 mm and on the left
side of the body for symmetrical characters: eye diameter (ED), horizontal diameter of orbit; eye–nostril length
(EN), distance between anteriormost point of orbit and middle of nostril; snout length (ES), distance between
anteriormost point of orbit and tip of snout; tympanum–eye length (TYE), distance between anterior most margin
of tympanum and posterior most margin of eye; tympanum diameter (TYD), longest diameter of the tympanum;
internarial distance (IN), shortest distance between the inner margins of nares; interorbital width (IO), shortest
distance between upper margins of orbits; head length (HL), distance between posterior edge of mandible and tip of
snout; head width (HW), maximum width of head; head depth (HD), distance between occiput and throat; upper
arm length (UAL), distance between axilla and angle of elbow; lower arm length (LAL), distance from elbow to
wrist with both upper arm and palm flexed; palm length (PAL), distance between wrist (carpus) and tip of longest
finger, with both palm and lower arm flexed; femur length (FEL), distance between groin and knee; tibia length
(TBL), distance between knee and heel, with both tibia and tarsus flexed; foot length (FOL), distance between heel
and tip of longest toe, with both foot and tibia flexed; toe length (TL), distance between tip of claw and nearest
fork; axilla–groin length (AG), distance between axilla and groin; snout–vent length (SVL), measured from tip of
snout to anterior margin of vent; tail base width (TBW), largest diameter of the tail base; tail length (TAL),
measured from anterior margin of vent to tail tip; Meristic characters were taken as follows: supralabials (SUP) and
infralabials (INF), first labial scale to last labial scale towards gape, which is distinctly larger than the granular
scales at gape; canthus rostralis (CR), counted from first scale posterior to supranasal, to end of supraciliary ridge;
mid body scales (MBS), counted from center of mid dorsal row forwards and downwards across ventrals;
dorsonuchal crest spines (DS), counted from first spine of nuchal crest to the level of axilla; mid ventral scale row
(MVS), counted from first scale posterior to mental, to last scale anterior to vent; subdigital lamellae on toe IV
(SDL), from first proximal enlarged scansor wider than twice the width of the largest palm scale, to distalmost
lamella at tip of digit. All the measurements were normalised to the percentage of HL (HL itself is given as a
percentage of SVL), because we are confident that HL is more accurate than SVL, especially when measuring
older museum specimens.
Natural history observations were made by looking at the animal at a distance of at least 3–4 m away being
careful not to make a disturbance. The eggs were measured with a Mitutoyo digitmatic caliper to the nearest 0.1
mm, and the eggs were carefully deposited back in the original nest hole. A standard thermometer, hygrometer and
lux meters were used to record the environmental parameters during the observations.
Standard morphometric and meristic statistics are presented in Table 1. Statistically informative tests could not
be performed because of the lack of C. liocephalus specimens.
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A NEW SPECIES OF THE GENUS CALOTES FROM SRI LANKA
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Taxonomy
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, ZSM258/1979, 76.3 mm SVL, Cobet’s Gap-Knuckles, W. Erdelen, 01 IV 1979;
Males, ZSM 218/1981/1–3, 81.5 mm SVL, 85.5 mm SVL, 76.7 mm SVL, Midcar-Knuckles, W. Erdelen, 05 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, 05 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
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A NEW SPECIES OF THE GENUS CALOTES FROM SRI LANKA
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.
FIGURE 1. Holotype male (86.6 mm SVL) of Calotes pethiyagodai sp. nov., WHT6211; 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).
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FIGURE 2. Scales on abdomen of; a, Calotes pethiyagodai sp. nov., WHT6211, 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., WHT6211, 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).
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
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A NEW SPECIES OF THE GENUS CALOTES FROM SRI LANKA
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.
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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).
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A NEW SPECIES OF THE GENUS CALOTES FROM SRI LANKA
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.
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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.
TABLE 2. External morphological character variation in Sri Lankan Calotes based on examined specimens (modified
after Bahir & Maduwage 2005)
TABLE 2. (Continued).
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
character C. calotes C. versicolor C. ceylonensis C. liolepis
gular pouch present present absent present
lateral body-scale orientation
(backwards)
upwards upwards straight downwards
Shoulder pit present absent present present
supratympanic spines single continuous row two clusters two clusters two clusters
size of ventral scales relative to dorsals similar smaller smaller smaller
scales on ventral thigh keeled keeled smooth keeled
character C. desilvai C. pethiyagodai sp. nov. C. liocephalus C. nigrilabris
gular pouch present absent present present
lateral body-scale orientation
(backwards)
downwards downwards downwards downwards
Shoulder pit present present present present
supratympanic spines two clusters absent absent single continuous row
size of ventral scales relative
to dorsals
smaller larger smaller larger
scales on ventral thigh smooth keeled keeled keeled
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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, NMB3354, 63.9 mm SVL, Talawakelle-Dickoya.
FIGURE 5. Holotype male (91.4 mm SVL) of Calotes liocephalus, BMNH 1946.8.11.33; a, dorsal head; b, lateral head; c,
ventral head; d, pectoral scales: note greater snout scales and occipital scales (a); shorter snout (b); larger mid gular scales (c),
and enlarged pectoral scales (d) respectively (Photos: A.A.T. Amarasinghe).
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,
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70 · Zootaxa 3785 (1) © 2014 Magnolia Press
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.
FIGURE 6. A live adult male (104.9 mm SVL) of C. liocephalus (not collected) at 1500m elevations of Peak Wilderness
(Photo: D. Samarasinghe).
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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).
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72 · Zootaxa 3785 (1) © 2014 Magnolia Press
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. 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 brownish-
grey 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.
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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.
Discussion
Günther (1872), in his original type description was obviously not absolutely clear of its precise locality,
documenting that the specimens appear to have been collected chiefly in the neighbourhood of the locality named
“Peradeniya district”. All subsequent museum documents (the various catalogues including that of Boulenger and
the specimen label jar itself) relating to this specimen lists the locality as “Ceylon” only. There are no further
details for locality recorded in the museum. The collector, George Henry Kendrick Thwaites was appointed
superintendent of the botanical gardens at Peradeniya, Sri Lanka from March 1849 until he retired in 1879 (Lee
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1898). There remains the possibility of collecting a specimen around Peradeniya. Peradeniya “District” was
restricted to a smaller administration division during 1849–1872. On the other hand, it is possible that the C.
liocephalus holotype was actually collected somewhere else in the central highlands. There is no other record
confirming C. liocephalus’distribution around Peradeniya except Deraniyagala (1953). We could not find any
specimens collected from Peradeniya deposited either at NMSL or at BMNH. It is possible that Deraniyagala
(1953)’s location in Peradeniya is anecdotal to Günther’s (1872) statement. It is also a possibility is that Günther
made the presumption that the specimen was collected from Peradeniya District, as it had been sent by G. H. K.
Thwaits who worked in Peradeniya District, but in actual fact the specimen may have been collected from
somewhere else. In 1860, G. H. K. Thwaites established chinchona nurseries at Hakgala (~1500 m elevation in
central highlands) and he sent many specimens to England around 1869 (Lee 1898). The older catalogue number of
the holotype of C. liocephalus (71.12.16.4) indicates the year of first registration 1871, Therefore there is a
possibility that the specimen was collected around Hakgala.
Also it has been claimed that the species described from Kandy during the 19th century [e.g. Adenomus
kandianus (Günther, 1872)], is currently not found in Kandy, but at higher elevations of Peak Wilderness in the
central highlands (see Wickramasinghe et al. 2012). The morphological characters and morphometrics show that
the holotype of C. liocephalus belongs to the central highland population, and not to the Knuckles range. Therefore
we can confirm that the holotype of C. liocephalus was collected from central highland, but that the exact location
or elevation cannot be confirmed for certain. Smith (1935) and Deraniyagala (1953) observed this species from
Pandalu Oya (~1000 m), and Agrapatanas (~1500 m) in central highlands. However Erdelen (1984), and
Manamendra-Arachchi & Liyanage (1994) recorded C. liocephalus (now C. pethiyagodai sp. nov.) only from the
Knuckles range. In 1981, one specimen was collected from Upcot (ZSM219/1981) and in 2005, two specimens
from Agrapatanas (WHT6503 and WHT6503). In addition, Amarasinghe et al. (2009) reported C. liocephalus
from 800–2000 m at Peak Wilderness (the elevation should be corrected as above 1400 m). Even though, Erdelen
(1984) recorded C. liocephalus (now C. pethiyagodai sp. nov.) at 800 m elevations, we failed to record any below
900m. This may be due to habitat destruction in the Knuckles over the past three decades. In addition, the
population in Pundaluoya, which is known from BMNH95.7.24, Smith (1935), and Deraniyagala (1953)] and
Dickoya (known from NMB3353–4) are known to be locally extinct the past 60 years.
It is also argued that “There may sometimes be stronger faunal differentiations between wet, dry, and cloud
forest zones within Sri Lanka than between that island’s dry zone and the dry region of South India” (Helgen &
Groves 2005). Also the lowlands (~500m) of the Mahaweli River, appears to have served as a barrier (see Fig. 4)
separating the central highlands from the Knuckles massif (Manamendra-Arachchi et al. 2006), and the Knuckles
population is well separated from the river Mahaweli from the central mountain chain of the country (see the Fig.1
in Fernando et al. 2007). However, two isolated populations of C. liocephalus in the central highlands and the
Knuckles massif occur and these have not been compared critically (Amarasinghe et al. 2009). Therefore, here we
compare the specimens based on few available museum collections and based on the observations of live
specimens from the Knuckles massif and Peak Wilderness. C. liocephalus and C. pethiyagodai sp. nov. are
completely allopatric and isolated to small populations in two mountain ranges. This isolation has led to a
significant morphological and morphometric variation between these two species.
We only had nine male specimens from the Knuckles and five males from the central highland to conduct this
study. However, we feel it is useful to describe this species, C. pethiyagodai sp. nov. as it is significantly different,
and we hope that future exploration will produce more individuals, and so we also recommend that this rare species
should receive immediate conservation attention. Due to the lack of specimens to conduct enhanced statistically
informative tests, the first two authors of this manuscript had requested permission twice from the Department of
Wildlife Conservation (DWC). Permission was requested to collect only 3 individuals from the Knuckles massif
and Peak Wilderness, but the applications were rejected (due to reasons other than the rarity of the species). The
limited scientific capacity of DWC (Bahir & Gabadage 2009; Pethiyagoda et al. 2007), suggest that wildlife
officials be immediately educated and that the department should be furnished with biology graduates
(Pethiyagoda et al. 2007). Because of the reasons above, we found it absolutely necessary to describe this new
species based on nine available male museum specimens (Appendix I) at WHT (now at NMSL), ZSM and ZMH,
we only had five male specimens of C. liocephalus for the comparison. The massive effort we undertook tracking
down European museum specimens which were collected from central highland proved unfruitful, despite the fact
that the central highland population was first discovered nearly 150 years ago. Therefore, here we highlight the
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importance of extensive field work. We also suggest that proper phylogenetic studies be carried out to further
understand the biogeography of the genus Calotes. This is particularly necessary for the C. liolepis and C.
liocephalus complexes (Erdelen 1984).
The results of the application of the IUCN Red List criteria (2001: Versions 3.1, 4.0, and 2013: version 10.1)
shows that C. pethiyagodai sp. nov. is Endangered (EN) and C. liocephalus is Critically Endangered (CR). C.
pethiyagodai sp. nov. is restricted to an area of occupancy (AOO) <25 km2 (six spotting sites) and extent of
occurrence (EOO) <180 km2 in Knuckles massif [Applicable criteria is B2-b (iii)] with the same forest area.
Calotes liocephalus is restricted to an area of occupancy (AOO) <15 km2 (four spotting sites during the last 50
years) and extent of occurrence (EOO) <100 km2 in Peak Wilderness, Upcot, and Agrapatanas [Applicable criteria
is B1-b (i)] with the same area. See the map (Fig. 4) for distribution data of C. pethiyagodai sp. nov. and C.
liocephalus in separate forest reserves.
Chena cultivations, illegal timber felling, encroachments, manmade fire, soil erosion, garbage dumping,
habitat destruction, unplanned constructions, rock exploitations, illegal gem mining, and land fillings are all
identified as main reasons for habitat loss and fragmentation in the Knuckles region (Amarasinghe & Karunarathna
2010; Lindström et al. 2012). These threats to Calotes pethiyagodai sp. nov. could be exacerbated by the
surrounding cardamom cultivations that indiscriminately use pesticides (Bahir & Surasinghe 2005). Road kills are
an additional threat to C. pethiyagodai sp. nov. (personal observations during the past decade in the Riverstone
area). Specimens of C. pethiyagodai sp. nov. have been recorded by Amarasinghe et al. (2009) that were dead for
reasons unknown. There are several previously unknown species awaiting description from the Knuckles massif
(Amarasinghe et al. 2014) and they would be at risk of extinction before they are even described fully
(Amarasinghe & Karunarathna 2010). Threats to the unique recognised biodiversity and the challenges to its
conservation (Pethiyagoda 2012), demand urgent international and national level scientific attention, policy and
planning (Pethiyagoda et al. 2007; Bahir & Gabadage 2009; Amarasinghe & Karunarathna 2010). Therefore, it is
our own responsibility to conserve our natural heritage without any further delay.
Key to Sri Lankan species of genus Calotes
1. Shoulder pit present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Shoulder pit absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. versicolor
2. Row of well-developed, compressed spines above tympanum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Well separated, rudimentary or prominent spines above the tympanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Ventral scales larger than dorsal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nigrilabris
Ventral scales smaller than dorsal scales. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. calotes
4. Well separated spines prominent and developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Well separated spines rudimentary and tubercle like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
5. Lateral scales pointing backwards, and downwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
- Lateral scales pointing backwards, and straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ceylonensis
6. Scales on ventral surface of thigh smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. desilvai
Scales on ventral surface of thigh keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. liolepis
7. Abdominal scales mucronate, pectoral scales enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. liocephalus
- Abdominal scales acuminate, pectoral scales not enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pethiyagodai sp. nov.
Acknowledgments
We thank Wolfgang Böhme, Colin Groves,Ulrich Manthey, Rohan Pethiyagoda, Kelum Manamendra-Arachchi,
Madhava Meegaskumbura, Mohamed Bahir, Sudath Nanayakkara, Anslem de Silva, Mendis Wickramasinghe,
Ruchira Somaweera, Pradeep Samarawickrama, Dinesh Gabadage and Deepani Jayantha helped in diverse ways to
enrich this work. Edi Stöckli and Frank Glaw for kindly sending us specimens from NMB and ZSM respectively.
Nanda Wickramasinghe, Sanuja Kasthuri-Arachchi, Manori Nandasena, Chandrika Munasinghe, M.B.
Vaidyasekara, C. Kotalawala, A. Samanthika, P. Gunasiri and R. Wickramanayake at NMSL for assisting while
examining collections under their care. I. Nuwan Bandara, Dushantha Kandambi, Prasanna Samarawickrama,
Majintha Madawala, Indika Peabotuwage, Madhava Botejue, Dulan Vidanapathirana, Amila Chanaka and the field
AMARASINGHE ET AL.
76 · Zootaxa 3785 (1) © 2014 Magnolia Press
staffs of the DF and DWC for their support during the field works. Kelum Manamendra-Arachchi and Dinesh
Gabadage for personal communications. Colin McCarthy, Franz Tiedemann, Richard Gemel, Ivan Ineich, Sven O.
Kullander and Bodil Kajrup for kindly examining collections under their care. We would like to thank Vimukthi
Weeratunge, Mahesh de Silva, Dinal Samarasinghe, and Madhava Botejue for providing excellent photographs and
members of the Young Zoologists’ Association of Sri Lanka (YZA) for various assistance during the study.
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APPENDIX I. Specimens examined.
Calotes calotes. Male, (NRM 106), 112.0 mm SVL, Sri Lanka. Male, (WHT 108A), 85.8 mm SVL, Laggala–Knuckles. Male,
(WHT 108B), 83.0 mm SVL, Laggala–Knuckles. Male, (WHT 182), 107.4 mm SVL, Puttalam. Male, (WHT 187), 128.4
mm SVL, Yatapatha (Lihinigala). Male, (WHT 188), 100.8 mm SVL, Yatapatha (Lihinigala). Male, (WHT 381A), 114.2
mm SVL, Warnagalla (near Erathne). Male, (WHT 616), 123.0 mm, Koskulana near Panapola.
Calotes ceylonensis. Male, (NMB 3340), 81.9 mm SVL, Kumbukkan Oya. Male, (NMB 3341), 67.6 mm SVL, Northeastern
Sri Lanka. Male, (WHT 7397), 59.1 mm SVL, Wasgamuwa. Male, (WHT 7514), 74.8 mm SVL, Giritale. Male, (WHT
1427A), 79.6 mm SVL, Wasgamuwa. Male, (WHT 1427B), 75.0 mm SVL, Wasgamuwa. Male, (WHT 1428), 80.3 mm
SVL, Tatugala–Bulupitiya. Male, (WHT 0515), 72.9 mm SVL, Pallegama. Male, (WHT 1625A), 77.2 mm SVL,
Wasgamuwa. Male, (WHT 1625B), 71.9 mm SVL, Wasgamuwa. Male, (WHT 0511), 82.5 mm SVL, Konketiya–Buttala.
Male, (WHT 0522), 73.2 mm SVL, Konketiya–Buttala. Male, (WHT 0519), 73.0 mm SVL, Yala. Male (WHT 0520), 73.4
mm SVL, Yala. Male, (WHT 1624A), 65.1 mm SVL, Wasgamuwa.
Calotes desilvai. Female, (WHT 1412), 59.2 mm SVL, Morningside. Female, (WHT 5998), 73.7 mm SVL, Morningside.
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, (NHMW21097), 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.
Calotes liolepis. Female, (BMNH 69.7.24.2), 76.7 mm SVL, Sri Lanka. Female, (NMB 3345), 75.8 mm SVL, Peradeniya.
Female, (WHT 491B), 64.8 mm SVL, Kottawa. Female, WHT 495, 66.8 mm SVL, Kottawa. Female, (WHT 491A), 65.9
mm SVL, Kottawa. Female, (WHT499B), 69.8 mm SVL, Mederipitiya. Female, (WHT192), 76.6 mm SVL,
Batadomabalena near Kuruwita. Female, (WHT1415), 63.1 mm SVL, Richmond Hill near Galle. Female, (WHT489),
68.8 mm SVL, Hapugala near Galle. Female, (WHT 176), 59.5 mm SVL, Dimbula near Kotagala. Female, (WHT 6162),
73.4 mm SVL, Kumaradola Group, Moneragala. Female, (WHT 6186), 77.4 mm SVL, Puwakpitiya. Female, (WHT
6184), 75.6 mm SVL, Puwakpitiya.
Calotes nigrilabris. Male, (NHMW 23355), 99.8 mm SVL, Nuwara Eliya. Male, (WHT 380C), 87.9 mm SVL, Horton Plains.
Male, (WHT 1555), 84.3 mm SVL, Hakgala. Male, (WHT 2262), 91.8 mm SVL, Hakgala.
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, (ZSM258/1979), 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.
Calotes versicolor. Male, (WHT 165), 95.0 mm SVL, Mousakanda–Knuckles. Male, (WHT 105), 81.5 mm SVL, Pallegama–
Knuckles. Male, (WHT 384), 80.5 mm SVL, Peradeniya. Male, (WHT 382), 86.0 mm SVL, Warnagalla near Kuruwita.
Male, (WHT 104), 94.6 mm SVL, Laggala–Knuckles. Male, (WHT 164A), 98.0 mm SVL, Mahapelassa near Kirinda.
Male, (WHT 181), 119.0 mm SVL, Nagagamuwa–Puttalam. Male, (WHT 204), 125.3 mm SVL, Bundala–Hambantota.
Male, (WHT 205), 101.0 mm SVL, Siribopura–Hambantota. Male, (WHT 199), 66.5 mm SVL, Pannipitiya–Maharagama.
... Diagnosis: Ceyloncalotes subgen. nov. is readily separated from the other subgenera of Calotes by the following suite of characters: One or other of: 1/ An oblique fold or pit covered with small granular scales in front of the shoulder, not extending across the throat; dorsal scales as large as or smaller than ventrals; lateral scales pointing downwards and backwards; no spines whatsoever on the head (Calotes (Ceyloncalotes) liocephalus Günther, 1872 and C. (Ceyloncalotes) pethiyagodai Amarasinghe, Karunarathna and Hallermann, 2014); or: 2/ An oblique fold or pit covered with small granular scales in front of shoulder, not extending across throat; dorsal scales larger than ventrals, smooth or nearly so; lateral scales pointing upwards and backwards, or straight backwards; 60 scales round middle of body (Calotes (Ceyloncalotes) ceylonensis Müller, 1887). Ceyloncalotes subgen. ...
... Tail round, the scales of its basal part very large and hard, those of the median upper row forming a serrated edge. Pale olive-green, above with transverse bands of a darker green; four angular reddish-brown cross bands on the back; a dark streak from the eye to above the tympanum; limbs and tail with alternate lighter and darker cross bands (Calotes (Ceyloncalotes) liocephalus Günther, 1872 and C. (Ceyloncalotes) pethiyagodai Amarasinghe, Karunarathna and Hallermann, 2014); or: 2/ Upper head-scales smooth, imbricate, considerably enlarged on the supraorbital region; two small spines on each side above the tympanum; diameter of the tympanum half that of the orbit. No gular sac; gular scales very strongly keeled, much smaller than dorsals. ...
... Etymology: Named in recognition of the location that the subgenus occurs (Ceylon), and the genus from which the species have been traditionally placed. Content: Calotes (Ceyloncalotes) liocephalus Günther, 1872 (type species); C. (Ceyloncalotes) ceylonensis Müller, 1887; C. (Ceyloncalotes) pethiyagodai Amarasinghe, Karunarathna and Hallermann, 2014. SUBGENUS TAMILNADUCALOTES SUBGEN. ...
Article
A number of recent molecular studies have highlighted divergences between south-east Asian agamid lizards within the Draconinae previously thought to be sufficiently close as to be placed in the same genera. Consolidating recently published studies and further investigations into relevant taxa, incorporating available molecular, morphological and geological evidence some genera as presently recognized are re-arranged to better reflect the relationships of species. For some of these taxa, names are available and as a result of this review they are formally resurrected from synonymy. Where names for taxa don’t exist, the species groups are formally named and defined herein according to the Zoological Code (Ride et al. 1999). Well established genera that are in effect split up include the following: Gonocephalus, Kaup, 1825 is divided into three genera. The nominate genus is also split into five subgenera, all named for the first time except for (Dilophyrus Gray, 1845) and the nominate subgenus. Japalura Gray, 1853 is divided into three genera, one of which is named for the first time. Two genera are further divided into two subgenera. Calotes Daudin, 1802 is divided into three genera, two also divided into subgenera. The taxon originally described as “Calotes andamanensis Boulenger, 1891” is also placed in a new monotypic genus as it also clearly sits ouside the two genera in which it has been recently placed (Calotes and Pseudocalotes Kaup, 1827). Ceratophora Gray, 1835, is divided in three, with the two most divergent taxa, C. aspera Günther, 1864 and C. karu Pethiyagoda and Manamendra-Arachchi, 1998 each placed in a new monotypic genus. The remaining Ceratophora are split into two obvious subgenera. Bronchocela Kaup, 1827 is divided into two subgenera. The taxon currently known as Bronchocela cristatella Kuhl, 1820 from the island of Halmahera, Indonesia is formally described as a new species. Other potential new species are identified. Phoxophrys Hubrecht, 1881 is divided into three subgenera, one formally named for the first time. Aphaniotis Peters, 1864 is subdivided into two subgenera as is ptyctolaemus Peters, 1864. Salea Gray, 1845 is divided into two subgenera for which a names are already available. Draco Linnaeus, 1758 is herein conservatively divided nine ways into subgenera of which five are formally named for the first time. It is likely that other taxonomists may treat these divisions as full genera. Furthermore, Draconinae taxonomy and nomenclature at the level between subfamily and genus is tidied up. The result is the formal erection of ten tribes and the addition six subtribes formally defined and named. As a result of this scientific reorganisation of the subfamily, this paper presents a list of all recognized Draconinae species in correct tribes, subtribes, genera and subgenera. Keywords: Taxonomy; nomenclature; genera; Gonocephalus; Japalura; Cophotis; Calotes; Ceratophora; Diploderma; Aphaniotis; Phoxophrys; Bronchocela; Lophocalotes; Dendragama; Otocryptis; Ptyctolaemus; Mictopholis; Salea; Sitana; Otocryptis; Pseudocalotes; Paracalotes; Draco; Dilophyrus; Oriotiaris; Pelturagonia; Lyriocephalus; Coryphophylax; Ptyctolaemus; Mantheyus; Acanthosaura; new genera; Daraninagama; Doongagama; Maxhoseragama; Crottyagama; Skrijelus; Notacalotes; Pethiyagodaus; Manamendraarachchius; subgenera; Dracontoides; Rhacodracon; Pterosaurus; new subgenera; Honlamagama; Mantheysaurus; Denzeragama; Eksteinagama; Jamesschulteus; Rubercalotes; Ghatscalotes; Laccadivecalotes; Ceyloncalotes; Tamilnaducalotes; Freudcalotes; Khasicalotes; Amboncalotes; Ferebronchocela; Olorenshawagama; Proboscisagama; Mindatagama; Macguiredraco; Philippinedraco; Engannodraco; Somniadraco; Spottydraco; new species; Harradineus. new tribes; Dracoiini; Maxhoseragamiini; Crottyagamiini; Daraninagamaiini; Pethiyagodaiini; Japaluraiini; Lophocalotesiini; Phoxophryiini; Mantheyiini; Dendragamaiini; new subtribes; Maxhoseragamiina; Sitanaiina; Acanthosauriina; Saleaiina; Pethiyagodaiina; Doongagamaiina.
... Lizards of the genus Calotes Cuvier, 1817, belonging to the family Agamidae in the order Squamata, currently include 26 species distributed from eastern Iran through south China to Sumatra, Indonesia (Vindum et al. 2003;Zug et al. 2006;Krishnan 2008;Hartmann et al. 2013; and Fujinuma 2014; Amarasinghe, Karunarathna, Hallermann et al. 2014). The forest crested lizard, Calotes emma Gray,1845, one of the typical members of the genus Calotes, is widely distributed across from southern China, India, Vietnam, Burma, Lao PDR, Thailand, Cambodia to Peninsular Malaysia (Zhao et al. 1999;Chan-Ard et al. 2015). ...
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The whole mitogenome can prove useful tools for phylogenetic reconstruction and efficiently recover with reasonable taxon sampling. Calotes emma is widely distributed and arboreal in habits. However, studies of C. emma are still very limited, including population genetics and evolutionary biology. In this study, we reported the complete mitochondrial genome of the C. emma by next-generation sequencing for future more researches on systematics and evolution of C. emma from the perspective of mitochondrial DNA. The length of mitogenome was 17,688 bp, including 13 protein-coding genes (PCGs), 2 ribosomal RNA (rRNA) genes, 22 tRNA genes and a control region. The phylogenetic tree recovered the monophyly of the Calotes and revealed that newly sequenced C. emma well supported as the sister taxon to C. mystaceus by very high posterior probabilities (1.0). The complete mitochondrial genome of C.emma in this study will be helpful for understanding the phylogenetic systematics and relationships, and molecular evolution of Calotes in Agamidae.
... Except Cruziohyla sylviae, with its four confirmed range states, all these species are endemic, i.e., restricted to one single range state. Some species are limited to very small Areas of Occupancy (AOO), such as Gekko lauhachindai with an AOO of less than 10 km 2 [57], Eurydactylodes occidentalis with an AOO of only 2.5 km 2 [58], and Calotes pethiyagodai with an AOO of less than 25 km 2 [51]. For T. nebulosum no AOO has been defined but its Extent of Occurrence (EOO) is considered to be only 940 km 2 [54]. ...
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Direct exploitation is one of the five main reasons for the loss of biodiversity, and collections for the international pet trade are an ongoing threat for many reptiles and amphibians. The European Union and in particular Germany have a central role as a hub and destination for exotic pets from all over the world. Rare species of reptiles and amphibians especially are in the focus of collectors. Rarity on the market may be either caused by rarity of a species in the wild or by a limited availability for sale, e.g., due to national protection measures in the range state or remote localities. The present study identified 43 species that are not listed by the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) and were only recently described, but have already entered the European pet trade. Ten of these species were selected as case studies, representing species from different geographic regions and illustrating the marketing mechanisms. Many such species that are new to science are neither assessed by the International Union for Conservation of Nature (IUCN) Red List of Threatened Species nor are they covered by international legislation, even though in several countries, where such internationally sought-after species are caught, national protection measures are in place. This paper analyses the challenges and opportunities for the protection of potentially threatened and newly described reptile and amphibian species against over-exploitation for the pet trade.
... Deraniyagala (1931) did not provide the exact locality from where he collected or received the specimen. Many other species reported in the mid-20th century from Peradeniya have also not been subsequently recorded from this region (e.g., Calotes ceylonensis (Müller), C. liocephalus Günther, C. nigrilabris Peters, Chalcidoceps thwaitesi (Günther) [Bahir and Surasinghe, 2005;Amarasinghe et al., 2014; personal observation]). Deraniyagala (1931) gave 38 scales around mid-body for his specimen. ...
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As a part of an ongoing revision of Sri Lankan scincid lizards, I review the genus Lankascincus Greer on the basis of their morphology and morphometric data. I demonstrate that the long-disputed Sphenomorphus megalops (Annandale) is in fact a Lankascincus species, and a neotype is designated. Lankascincus deraniyagalae Greer and L. munindradasai Wickramasinghe, Rodrigo, Dayawansa, and Jayantha are shown to be junior subjective synonyms of L. fallax (Peters) and L. taprobanensis (Kelaart), respectively. Lankascincus deignani (Taylor) is confined to the forests around Kandy and does not extend to Nuwaraeliya and its environs. Previous records of L. deignani from Nuwaraeliya and surrounding localities are reidentified as L. sripadensis Wickramasinghe, Rodrigo, Dayawansa, and Jayantha. Three groups of Lankascincus species are identified on the basis of their morphology and breeding biology: the fallax, taprobanensis, and dorsicatenatus groups. Each group has a unique combination of characters. Lankascincus taprobanensis is restricted to high-elevational regions (1,500 m above mean sea level). Previous records from lower elevations (Sinharaja, Knuckles Range, and Peradeniya) are based on misidentified specimens of other Lankascincus spp. Lankascincus fallax has a wide distribution in Sri Lanka. Lankascincus deignani is restricted to Gannoruwa Forest Reserve and the Ambagamuwa area and is the most endangered skink in Sri Lanka. Ecologically, Lankascincus comprises a largely forest-dwelling species group (L. deignani, L. dorsicatenatus (Deraniyagala), L. gansi Greer, L. greeri Batuwita and Pethiyagoda, L. megalops new combination, and L. taprobanensis) and a group of species that live in altered habitats (L. sripadensis, L. fallax, and L. taylori). The present study confirms that the genus Sphenomorphus Fitzinger is not represented in Sri Lanka. Because of a lack of comprehensive genetic data to support a previously described new family, Ristellidae Hedges for Lankascincus and Ristella Gray, here I assign both genera into a new tribe, Ristellini. Phylogenetic relationships of Lankascincus and Ristella remain unresolved.
... Saltwater Crocodiles are distributed in a wide variety of saline and freshwater habitats, including rivers and creeks, coastlines, coastal flood plains, lagoons, swamps, river and canal outfalls (e.g., see Webb & Manolis 1989; Letnic & Connors 2006). Although popularly referred to as 'salties' in Sri Lanka, a high proportion of the Saltwater Crocodile population exists in freshwater habitats in the western and southern parts of the country (e.g., Nilwala, Bentota, Kelani, Maha Oya; de Silva 2013), including Colombo, the capital (Devapriya 2004; Jayawardene 2004; Porej 2004; Samarasinghe 2014). Under Sri Lankan legislation the Saltwater Crocodile is listed as 'endangered' (MOE 2012). ...
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Human-wildlife conflict occurs when human requirements encroach on those of wildlife populations, with potential costs to both humans and wild animals. As top predators in most inland waters, crocodilians are involved in human-wildlife conflicts in many countries. Here we present findings of a 5-year survey on human-crocodile conflict on the island of Sri Lanka and relate the results to improving management practices. We aimed to quantify and understand the causes of human-crocodile conflict in Sri Lanka, and propose solutions to mitigate it. Visual encounter surveys were carried out to estimate the population size of Saltwater Crocodiles. We recorded 778 sightings of Saltwater Crocodiles at 262 of 400 locations surveyed, and estimate the total population to comprise more than 2000 non-hatchlings and to have increased at an average rate of 5% p.a. since 1978. We propose four crocodile vigilance zones within the wet zone and one crocodile vigilance zone within the dry zone of the country. Specific threats to Saltwater Crocodiles identified in crocodile vigilance zones were: habitat destruction and loss; illegal killing and harvesting (17 killings out of fear, ~200 incidents of killing for meat and skins, ~800 eggs annually for consumption); unplanned translocations; and, interaction with urbanization (10 incidents of crocodiles being run over by trains/vehicles and electrocution). Additionally, 33 cases of crocodile attacks on humans were recorded [8 fatal, 25 non-fatal (minor to grievous injuries)] and more than 50 incidents of attacks on farm and pet animals.
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Jaffna peninsula is quite an unexplored area of Sri Lanka's lowland dry zone. We constructed a species checklist for all herpetofauna of this area based on a short-term field survey, a comprehensive literature review, museum specimens, and observations made by field herpetologists. Based on 200 × 10 m belt transects, we surveyed herpetofauna both during day and night time, in 10 different types of habitats. The species checklist we compiled comprised 44 species of reptiles (including three nationally threatened, one globally threatened, and eight endemic species) and 15 species of amphibians (including one nationally threatened and three endemic species). Based on published literature, museum specimens, expert opinions, and current field survey, we documented 85 species of herpetofauna in this area. Of this entire list, we were unable to record the presence of 25 species through our field survey. Our field survey documented 18 species that were not previously reported from Jaffna Peninsula. Our study revealed that inland water bodies, cultivated lands, home gardens, and coastal beaches are of high importance for native herpetofauna of Jaffna peninsula. Many human disturbances, such as habitat alterations, vengeful killing, consumption overexploitation, and road mortality are the key threats encountered by herpetofauna in Jaffna. Our intention of this study is to compile baseline information on diversity of amphibians and reptiles to support more detailed studies in future and assist conservation and management decisions within the region. We believe that our study will provide a basic foundation for conservation planning and future research.
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The montane agamid lizard genus Oriocalotes is currently considered monotypic, represented by the species, O. paulus. The systematic status of this taxon has remained questionable since its initial descriptions in the mid-1800s. A detailed molecular and morphological study was carried out to assess the validity of this genus, and its systematic position within the Asian agamid subfamily, Draconinae. Freshly collected and historical museum specimens from the type locality of O. paulus were examined morphologically, along with additional samples collected from localities in Mizoram state, Northeast India. Utilising newly generated molecular sequences (two mitochondrial and three nuclear genes), combined with those previously published for representative genera from the subfamilies Draconinae and Agaminae, Maximum Likelihood and Bayesian phylogenetic trees were constructed. Phylogenetic results suggest that Oriocalotes is part of the widespread South and Southeast Asian radiation of Calotes. Comparative morphological studies (including external morphology, hemipenis and osteology) between Oriocalotes and related genera further support this systematic placement. Oriocalotes is herein regarded as a junior subjective synonym of Calotes. Calotes paulus comb. nov. is also assigned a lectotype and given a detailed redescription based on the lectotype, paralectotypes and additional topotypic material. Furthermore, the specimens collected from Mizoram populations are found to be morphologically and genetically distinct from Calotes paulus comb. nov., and are described herein as a new species, Calotes zolaiking sp. nov.
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We describe a new species, Aspidura desilvai sp. nov., closely resembling A. trachyprocta and the last addition to the genus A. ravanai, from Knuckles massif, Matale District, of Sri Lanka. The species represents the ninth species of the genus known from Sri Lanka, and is readily distinguished from all other congeners by its colour pattern, the scale nature in the ischiadic region, and morphometric characteristics specially from its ratio between the snout to eye distance to its eye width. The species is currently known only from the type locality.
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Lizards of the genus Calotes are geographically restricted to South Asia, Indo-China and parts of Southeast Asia. The greatest diversity of the genus is from the biodiversity hotspots in South Asia: Western Ghats (Peninsular India), Sri Lanka and Indo-Burma. Here, we present a systematic revision of members of the genus Calotes from Peninsular India using a combination of molecular phylogeny, geographical distribution and morphological characters. We show that Calotes from the Western Ghats is paraphyletic and consists of three major clades, one of which is widely distributed in South and Southeast (SE) Asia, while the others are restricted to Peninsular India. The Peninsular Indian clade is composed of two sister clades: Psammophilus, with a wider distribution and a second clade, composed of two extant species, Calotes rouxii and Calotes ellioti and two new species, all restricted to the Western Ghats region. Based on morphological differences, we retain the generic status of Psammophilus and assign its sister clade to a new genus Monilesaurus gen. nov. and transfer the following species, C. rouxii and C. ellioti, to this new genus. We also provide diagnoses and descriptions for two new species recognized within Monilesaurus gen. nov. In addition, Calotes aurantolabium from the Western Ghats was observed to be deeply divergent and to share a sister-relationship with the clade composed of Calotes, Monilesaurus gen. nov., and Psammophilus. Based on its phylogenetic position and morphological attributes, we assign this species to a new genus Microauris gen. nov. These new discoveries highlight the evolutionary significance of the Western Ghats in housing novel lizard diversity.
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Brachysaura is a monotypic genus of agamid lizard found in the Indian subcontinent; the identity and systematic position of B. minor has been long debated, and it has at times been subsumed into Agama, Charasia and Laudakia, with some authors suggesting affinities to Calotes. We constructed nuclear and mitochondrial phylogenetic trees including Brachysaura and allied agamid genera to resolve its phylogenetic position. We also compared osteology and external morphology with the genera Agama, Calotes and Laudakia. Hemipenial morphology was compared with Calotes and some other agamids from South Asia. Both nuclear and mitochondrial phylogenies demonstrate that Brachysaura is nested within the widespread South and Southeast Asian genus Calotes, with which it also shares certain external morphological, osteological and hemipenial characters. Adaptations to ground dwelling in Brachysaura minor has resulted in unique modifications to its body plan, which is likely why generic allocation has been long confused. This study also highlights the need for an integrated systematic approach to resolve taxonomic ambiguity in Asian agamids.
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The paper describes a new frog of the genus Nannophrys from Sri Lanka. The new species is distinguished from all other Nannophrys species by the following combination of morphological characters; distal subarticular tubercle and penultimate subarticular tubercles of 4th toe are well separated (vs. these two tubercles close in all other species of the genus). Supernumerary tubercle absent or smaller than penultimate subarticular tubercle on 4th toe. A hard sharp narrow symphysial knob and close pair of sharp apophyses on anterior edge of mandible and symphysial knob lower than two apophyses (vs. all other species with a blunt wider symphysial knob and wider pair of apophyes on anterior edge of mandible, and symphysial knob of same height or higher than two apophyses. Palmar tubercles comparatively smaller, inner palmar tubercle separated from outer palmar tubercle (vs. other relatives possess palmar tubercles comparatively large, inner palmar tubercle connected with outer palmar tubercle). Ecological and distributional notes for the new species are given.
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Over the past two decades many checklists of reptiles of India and adjacent countries have been published. These publications have furthered the growth of knowledge on systematics, distribution and biogeography of Indian reptiles, and the field of herpetology in India in general. However, the reporting format of most such checklists of Indian reptiles does not provide a basis for direct verification of the information presented. As a result, mistakes in the inclusion and omission of species have been perpetuated and the exact number of reptile species reported from India still remains unclear. A verification of the current listings based on distributional records and review of published checklists revealed that 199 species of lizards (Reptilia: Sauria) are currently validly reported on the basis of distributional records within the boundaries of India. Seventeen other lizard species have erroneously been included in earlier checklists of Indian reptiles. Omissions of species by these checklists have been even more numerous than erroneous inclusions. In this paper, I present a plea to report species lists as annotated checklists which corroborate the inclusion and omission of species by providing valid source references or notes.
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Adenomus kandianus Günther (1872) was previously known only from two specimens both deposited in the British Museum, the holotype BMNH1947.2.20.63, and the syntype of A. kelaarti BMNH1947.2.20.62. The only record of A. kandianus since the initial description in 1872 was by Ferguson in 1876, who mentions two specimens resembling Bufo kandianus in his collection, making A. kandianus the world's rarest toad. The species had not been reported since, and was considered extinct. Here we report on its rediscovery.
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The island of Sri Lanka has a heritage of astonishing biodiversity of which comparatively little remains, restricted to small forest islands. Conservation efforts have been aided by many high-quality publications, while on the other hand inaccurate reports can present obstacles to effective efforts. Here we discuss inaccuracies in a report prepared for the Department of Wildlife Conservation of Sri Lanka and the Ministry of Environment and Natural Resources, from which it is apparent that the consultants dominating conservation science do not appreciate the necessity of identifying species accurately, citing appropriate references, or updating their knowledge of current nomenclature and distribution of taxa. Conservation is an important national and international issue, and it is incumbent upon educators, conservation managers, legal advisors, funding agencies, officials and policy makers to work along with research scientists to ensure that inaccurate information does not endanger efforts to safeguard Sri Lanka’s remaining endangered biodiversity treasures. Towards this end, procedures for the conduct of conservation studies should be revised to incorporate input from researchers familiar with current knowledge and methods.