Current status of Calotes liocephalus Günther, 1872 (Reptilia: Agamidae) of Sri Lanka
A.A. Thasun Amarasinghe1, D.M.S. Suranjan Karunarathna2 & Dinesh E. Gabadage3
1,3 Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka
2 IUCN - Sri Lanka Country office, No. 53, Horton place, Colombo
07, Sri Lanka
Email:
1 aathasun@gmail.com ; 2 dmsameera@gmail.com
Date of publication (online): 26
November 2009
Date of publication (print): 26
November 2009
ISSN 0974-7907 (online) |
0974-7893 (print)
Editor: Aaron Bauer
Manuscript details:
Ms # o2216
Received 25 May 2009
Final received 28 October 2009
Finally accepted 29 October 2009
Citation: Amarasinghe, A.A.T., D.M.S.S. Karunarathna& D.E. Gabadage (2009). Current status of Calotes liocephalus Günther, 1872 (Reptilia: Agamidae) of Sri Lanka. Journal of Threatened Taxa 1(11): 553-557.
Copyright: © A.A. Thasun Amarasinghe, D.M.S. Suranjan Karunarathna & Dinesh E. Gabadage 2009. Creative
Commons Attribution 3.0 Unported License. JoTT allows unrestricted use of this article in any medium
for non-profit purposes, reproduction and distribution by providing adequate
credit to the authors and the source of publication.
Author Details: A.A. Thasun Amarasinghe, the chairman
of the Taprobanica Nature Conservation Society is a
Sri Lankan herpetologist. He is the
editor-in-chief of the Taprobanica journal, is an
editorial board member of Herpetotropicos, and a
member of CSG/SSC/IUCN; currently involved in biodiversity conservation projects in Sri Lanka.
D.M.S.
Suranjan Karunarathna,
an instructor of the reptile study group in Young Zoologists’ Association of
Sri Lanka and currently works as a project assistant for the IUCN, Sri Lanka.
Also, he is a member of CSG/SSC/IUCN.
Dinesh
E. Gabadage,
the secretary of the Taprobanica Nature Conservation
Society is a fieldbiologist working on Sri Lankan herpetofauna and
promoting conservation awareness of the value of biodiversity among Sri
Lankans.
Author Contributions: All authors
have equally contributed to this study and in writing the paper.
Acknowledgments:We
sincerely thank Mohomed M. Bahirfor comments; Colin McCarthy for measurements and photographs of Calotes liocephalus holotype; Rohan Pethiyagoda, Sudath Nanayakkara (WHT) for assistance and Kasun Benaragama & Kasun Ekanayake for photographs.
Abstract: Calotes liocephalus Günther, 1872 is an endemic and threatened, arboreal
agamid lizard of Sri Lanka, which is found only in the forests of the Knuckles
Mountain Range and Peak Wilderness up to 1850m. This work is mainly based on examination of the type specimen and WHT
specimens as well as published literature and our observations of ecology, behaviour and threats on Calotes liocephalus.
Keywords:Behaviour, biology, Calotes liocephalus, conservation, ecology, natural
history, taxonomy, Sri Lanka.
For Figures & Images –
click here
Eighteen species
of agamid lizards, family Agamidae, have been
reported from Sri Lanka (Manamendra-Arachchi et al. 2006;Somaweera & Somaweera2009), representing one subfamily: Draconinae (Macey et al. 2000; Manthey2008). The genus Calotes extends through southern Asia to most of the East Indian Archipelago
(Taylor 1953).
Seven species ofCalotes occur in Sri Lanka. Five of them (C. nigrilabrisPeters, 1860; C. liocephalus Günther, 1872; C. liolepis Boulenger, 1885; C. ceylonensis Müller, 1887; C. desilvai Bahir & Maduwage,
2005) are endemic. The remaining
two, C. calotes (Linnaeus, 1758) and C. versicolor (Daudin, 1802) are
widespread throughout South and South East Asia. Calotes liocephalus is an arboreal species found only in the
forests of the Knuckles Range and Peak Wilderness up to 1850m (Erdelen 1984; Manamendra-Arachchi& Liyanage 1994; Das & de Silva 2005; de
Silva et al. 2005). It is a threatened
species (Manamendra-Arachchi & Liyanage 1994; Bahir & Surasinghe 2005; IUCNSL & MENR 2007).
The examined holotype is at the Natural History Museum, London, UK
(BMNH) and others are with the Wildlife Heritage Trust of Sri Lanka,
Colombo, Sri Lanka (WHT). Diagnoses and
descriptions are based on morphology. The locality record for each species includes WHT specimen data,
published information and our observations during the past decade.
All measurements
were taken with dial calipers to the nearest 0.1mm.
Scale counts: SUP - Supralabials were counted from the first scale anterior to
that at the angle of gape, not including the median scale (when present); INF -infralabials were counted from the first scale
posterior to mental, to the angle of gape; DS - dorsal spines were counted from
the first spine to the last of mid-dorsal row; CR - canthus rostralis (counted from rostralscale along scale row passing over nostril to posterior end of supraciliary ridge); MDS - mid dorsal scales (counted from
scale behind the rostral to the posterior margin of
the thigh); MBS - mid body scales were counted from center of the mid-dorsal
row forwards and downwards across ventrals (this
count is unreliable, by the unequal size and uneven arrangement of the lateral
scales); MVS - mid ventral scales were counted from the first scale posterior
to mental, to the last scale anterior to vent; SAT - Spines around tympanum
were counted from the first spine to the last above tympanum.
External
measurements(in mm): SVL - snout–vent length (distance between tip of snout to anterior
margin of vent); HL - head length (distance between posterior edge of mandible
and tip of snout); HW - head width (maximum width of head); DHL - dorsal head
length (distance between posterior edge of cephalic bone and tip of snout); NFE
- nostril – front eye length (distance between anterior most point of orbit and
middle of nostril); UAL - upper-arm length (distance between axilla and angle of elbow); LAL - lower-arm length
(distance from elbow to wrist with both upper arm and palm flexed); FL - finger
length (distance between tip of claw and the nearest fork); FEL - femur length
(distance between groin and knee); TBL - tibia length (distance between knee
and heel, with both tibia and tarsus flexed); TL - toe length (distance between
tip of claw and nearest fork); AG - axilla–groin
length (distance between axilla and groin); SA -
snout–axilla length (distance between tip of snout
and axilla); TAL - tail length (measured from
anterior margin of vent to tail tip); PAL - palm length (taken from posterior
most margin of palm and tip of longest finger); FOL - foot length (distance
between heel and tip of longest toe, with both foot and tibia flexed); TBW -
width of tail base (greatest width of the tail base); IOW - inter orbital width
(minimum distance between the upper margins of orbits); ED - eye diameter
(horizontal diameter of orbit); SFE - snout–front eye length (distance between
anterior most point of orbit and tip of snout); SBE - snout–back eye length
(distance between posterior most point of orbit and tip of snout); SFT -
snout–front tympanum length (distance between anterior most point of tympanum
and tip of snout); TD - Tympanum diameter (minimum distance between the inner
margins of tympanum).
Calotes liocephalus Günther, 1872
Günther(1872), Annals and Magazine of Natural History.
Boulenger(1885). Catalogue of the Lizards in the British Museum (Nat. Hist.).
Boulenger(1890), The Fauna of British India, including Ceylon and Burma.
Smith (1935), The fauna of British India including Ceylon and Burma.
Deraniyagala(1953), A Colored Atlas of some vertebrates from Ceylon.
Taylor (1953), A review of the lizards of Ceylon.
Wermuth(1967). Liste der rezenten Amphibienund Reptilien. Agamidae.
Erdelen(1984).The genus Calotes (Sauria: Agamidae) in Sri Lanka: distribution patterns.
Manamendra-Arachchi& Liyanage (1994), Conservation and distributions
of the agamid lizards of Sri Lanka with illustrations of the extant species.
Manthey& Schuster (1999). Agamen - 2. Aufl. Natur und Tier Verlag (Münster).
Das & de
Silva (2005), Snakes and other Reptiles of Sri Lanka.
Manthey(2008), Terralog - Agamid Lizards of Southern Asia, Draconinae - 1.
Amarasinghe et al. (2009), The original descriptions and figures of Sri Lankan Agamid
Lizards (Squamata: Agamidae)
of the 18th and 19th centuries.
Somaweera& Somaweera (2009). Lizards of Sri Lanka.
English Name:
Crestless Lizard; Sinhala Name: Nosilu Katussa
Holotype: Male (90.0mm
SVL); Cat.no. BMNH 1946.8.11.33; Loc. Peradeniya District, Ceylon (= Sri Lanka); Coll. G. H.K. Thwaites; Date. Unknown (Images 1 &
2).
Other material
examined: WHT
1667, male, 87.4mm, Sri Pada; WHT 6241, male, 85.4mm,Corbetts Gap; WHT 6211, male, 90.8mm, Knuckles; WHT
6154, male, 89.4mm, midlands Est.-Knuckles; WHT 0106A, female, 83.6mm, Maussakanda-Gammaduwa; WHT 6504, male, 89.1mm, Agra-Bopath; WHT 6503, female, 82.5mm, Torrington Est.-Agra-Bopath.
Diagnosis: Head without
spines or rarely a rudimentary spine above the tympanum; dorsinuchalcrest poorly developed; lateral scales directed backwards and downwards.
Description: [Based on Holotype and WHT collection: (Images 1-2; Figs. 1-3) the
range of morphometric data for holotypeand WHT specimen shown at the end of the description]. Head without spines or
rarely a rudimentary spine above ear, an oblique fold in front of shoulder,
lower jaw rather short. Canthus rostralis not very prominent, its outline straight. Six scales from eye to nostril. Tympanum about half size of the orbit. Supraocular scales enlarged. Dorsinuchal crest
poorly developed and the spines strongest on head, low on neck and rudimentary
on body, scales on sides point backwards and downwards; dorsal scales smaller,
feebly keeled, about as large as the ventrals; gular sac undeveloped; gularscales as large as or larger than the ventrals;
faintly carinate. Body compressed; Body scales faintly carinate;Ventrals strongly carinate. Limbs moderate; third and fourth fingers
equal or fourth finger a little longer than the third. Relative
length of fingers; 1<5<2<3<4. Fourth toe distinctly
longer than the third. Relative length of toes; 1<2<5<3<4. Hind limb
reaches to the eye or nearly as far. In
a fully-grown male the head is larger and tail is markedly swollen at the base,
with large, thick, keeled scales. SVL:
82.5-90.0; HL: 27.2-32.4; HW: 17.1-22.0; DHL: 21.4-26.8; NFE: 7.0-8.2; UAL:
16.0-23.5; LAL: 15.5-17.3; FL I: 5.2-6.4; FL II: 7.4-9.6; FL III: 11.4-13.1; FL
IV: 11.5-13.4; FL V: 6.8-7.7; FEL: 20.6-27.3; TBL: 19.6-24.2; TL I: 5.4-6.8; TL
II: 8.1-10.2; TL III: 13.8-16.6; TL IV: 15.5-19.7; TL V: 11.3-14.0; AG:
40.7-51.0; SA: 34.1-45.0; TAL: 241-267; PAL: 15.6-18.1; FOL: 27.9-30.9; TBW:
6.5-11.2; IOW: 1.3-2.3; ED: 5.2-8.1; SFE: 11.4-14.2; SBE: 16.7-19.1; SFT:
22.4-25.8; TD: 4.0-4.4; SUP: 9-10; INF: 9-9; MDS: 70-78; CR: 8-13; MBS: 45-56;
MVS: 68-78; DS: 13-25; SAT: 0-1.
Remarks: [Based on WHT
collection and other observations on live specimens: (Images 3 & 4)]. The body colour is
green or bluish-green dorsally with five or six transverse dark cross
bars. The throat is
yellowish-green. Three black cross bands
from eye to eye. A black band runs along
posterior half or upper lip through the tympanum. Sides of the lower jaws are with black spots,
sometimes dark cross bars on the top of the head; base of the tail light olive
brown, the rest of it alternately banded with light and dark; below
greenish-white.
Distribution and
habitat:Calotes liocephalus appears to be rare and has been recorded only from the forests of the
Knuckles Range (from 800-1850 m) and Peak Wilderness (from 800-2000 m) and this
species is endemic to Sri Lanka. This
species is largely arboreal and its diet comprises of young leaves, buds,
insects and worms (de Silva et al. 2005).
Hemipenis morphology: The hemipenis of Calotes liocephalus (Image 5) is well developed. The pedicel is slightly shorter than the
head; below the head, it is broadened out in to two shallowly concave
shoulders; there are no spines. The head
is quadrangular, shallowly divided longitudinally into four lobes, two being
slightly larger. Surface of the head is reticulatedly pitted, the pits being larger on the outside,
diminishing in size towards the divisions between the lobes.
Nest hole and
eggs:A female dug a hole in the ground 92.6mm deep and 79.1mm in diameter. 3-8 eggs
were deposited in June. The bottom of
the hole is conical and is dug at 450 angle. The eggs were white and elliptic, with a mean
length of 14.8mm and 8.6mm width (n=8). While laying eggs C. liocephalus places
the posterior part of its body inside the hole (Fig. 4) and it does not lift
the posterior part of the body to lay eggs. Its entire body curls inside the
hole, bending the anterior part of its body to look around (see Amarasinghe & Karunarathna2008).
Other behaviour: This species is active during daytime,
largely arboreal and sometimes on trees 5m above ground level. C. liocephalusis widespread within moderately open riverine-forested
areas and uncommon in home gardens. Hatchlings typically ambush their prey on Strobilanthes bushes, camouflaging in light
green. On threat, these hatchlings creep
in to the bush. Adults usually perch
vertically on branches. In the Knuckles Range we located this species on Mallotus tetracoccus(Euphorbiaceae) trees. When threatened the lizard rapidly climbs on
a tree or shrub and flattens against a branch. They also change their body colour to black.
According to Manamendra-Arachchi et al. (2006) the lowlands (elevation
~500m) of the Mahaweli River, which separates the Dumbara Hills (= Knuckles Hills) from the Central
Mountains, appears to have served as a barrier to the dispersion of highland
species. According to Günther (1872) the holotype of Calotes liocephalus (BMNH
1946.8.11.33), was collected from “PeradeniyaDistrict” presumably, Peradeniya (alt. ~500m) in the
Kandy District. However, Peradeniya is on the margin of the Knuckles and Central
Hills and the Mahaweli River flows across Peradeniya. If the
report is correct, C. liocephalus was once
distributed throughout the low elevations of Central and Knuckles
Mountains. We could not locate any Calotes liocephalusspecimen in Peradeniya. Two isolated populations in the Central Hills
and Knuckles Region occur and these have not been compared critically (Fig.
5). Therefore, a morphological and morphometric analysis of the two isolated C. liocephalus populations is needed. If the Knuckles population is a distinct morphospecies, its range would be restricted to the
Knuckles massif and it would be at risk of extinction before it is even named.
References
Amarasinghe, A.A.T. & D.M.S.S. Karunarathna(2008). Observations on the ovipositional behavior of the Crest – Less Lizard Calotes liocephalus(Reptilia: Agamidae) in the
Knuckles Forest Region of Sri Lanka. Asiatic Herpetological Research 11:
13–16.
Amarasinghe, A.A.T., U. Manthey, E. Stöckli, I. Ineich, S. O Kullander, F. Tiedemann, C. McCarthy & D.E. Gabadage (2009).The original descriptions and figures of Sri Lankan Agamid Lizards (Squamata: Agamidae) of the 18th
and 19th centuries. Taprobanica 1 (1): 2-15+4
pls.
Bahir,
M.M. & T.D. Surasinghe (2005). A conservation assessment of the
agamid lizards of Sri Lanka, pp. 407-412. In: Yeo,
D.C.J., P.K.L. Ng & R. Pethiyagoda (eds.). Contributions to Biodiversity
Exploration and Research in Sri Lanka. The Raffles
bulletin of Zoology Supplement 12.
Boulenger,
G.A. (1885). Catalogue
of the Lizards in the British Museum (Nat. Hist.) - I. Geckonidae,Eublepharidae, Uroplatidae,Pygopodidae, Agamidae. London, 450pp.
Boulenger,
G.A. (1890). The Fauna of British
India, including Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis,
London, xviii+541pp.
Das,
I. & A. de Silva (2005). Snakes and other Reptiles of Sri Lanka. New Holland Publishers, UK. 144pp.
Deraniyagala(1953). A Colored Atlas of Some Vertebrates from Ceylon, Tetrapod Reptilia Vol - 2. National Museums of Sri
Lanka, 101pp.
de
Silva, A., A.M. Bauer, S. Goonewardene, J. Drake, S.
Nathanael, W.P.R. Chandraratne, S.A.U.S. Somathilaka (2005). Status of the agamids in the
Knuckles massif with special reference to Calotes liocephalus Günther,
1872 and Cophotis ceylanicaPeters, 1861. Lyriocephalus 6(1&2):
43–71.
Erdelen, W. (1984).The genus Calotes (Sauria: Agamidae) in Sri Lanka: distribution patterns. Journal
of Biogeography 11: 515-525
Günther,
A. (1872). Descriptions
of some Ceylonese Reptiles and Batrachians. Annals
and Magazin of Natural History (4)9: 85-88.
IUCNSL & MENR (2007). The 2007 Red List of threatened Fauna and Flora of
Sri Lanka. Colombo, IUCN Sri Lanka, 148pp.
Macey, J.R., J.A. Schulte & A. Larson (2000). Evolution and phylogeneticinformation content of mitochondrial genomic structural features illustrated
with acrodont lizards. Systematic Biology49(2): 257-277.
Manamendra-Arachchi, K., A. de Silva & T. Amarasinghe (2006). Description of a second species of Cophotis (Reptilia:Agamidae) from the highlands of Sri Lanka. Lyriocephalus 06: Supplement 1: 1-8.
Manamendra-Arachchi, K. & S. Liyanage (1994). Conservation and distributions of
the agamid lizards of Sri Lanka with illustrations of the extant species. Journal of South Asian Natural History 1(1): 77-96.
Manthey, U. & N.
Schuster (1999). Agamen - 2. Aufl. Natur und Tier Verlag (Münster),
120pp.
Manthey, U. (2008). Terralog- Agamid Lizards of Southern Asia, Draconinae - 1. Edition Chimaira, 160pp.
Smith, M.A.
(1935). The Fauna of British India including Ceylon and Burma, Reptiliaand Amphibia, Volume II - Sauria,
144-206pp.
SomaweeraR. & N. Somaweera (2009). Lizards of
Sri Lanka: A Colour Guide with Field Keys.Edition Chimaira, Germany, 303pp.
Taylor, E.H.
(1953). A review of the lizards of Ceylon. The University
of Kansas Science Bulletin 35 (12): 1525-1585.
Wermuth, H. (1967). Liste der rezenten Amphibien und Reptilien. Agamidae. Das Tierreich - 86. Walter de Gruyter, Berlin, xiv+127pp.