2008

Asiatic Herpetological Research, Vol. 11

pp.13–16

Observations on the Ovipositional Behavior of the Crest-lessLizard Calotes liocephalus (Reptilia: Agamidae) in theKnuckles Forest Region of Sri Lanka
A. A. THASUN AMARASINGHE1, 2 AND D. M. S. SURANJAN KARUNARATHNA1, 3
1

The Young Zoologists' Association of Sri Lanka, National Zoological
Gardens, Dehiwala, Sri Lanka.
Corresponding authors E-mails: 2aathasun@gmail.com; 3dmsameera@gmail.com
Abstract.- A mature female Calotes liocephalus lying on the ground in Pitawala in the Knuckles Forest Region of SriLanka. This is the first described observation of the ovipositing of Calotes liocephalus. The ovipositional behavior
consisted of digging a hole to lay eggs, laying the eggs, scraping soil to bury the eggs, filling of the spaces between
the eggs, the tight compression of the soil and camouflaging the nest.
Keywords.- Agamidae, Calotes, egg-laying behavior, Knuckles, Sri Lanka, conservation.

Introduction
There are eighteen species of agamid lizards in SriLanka, fifteen of them are endemic to the island (Bahirand Surasinghe, 2005; Manamendra-Arachchi et al.,2006; Samarawickrama et al., 2006). Seven species
belong to the genus Calotes. Five of them (C. ceylonensis Muller, 1887; C. liocephalus Gunther, 1872; C.
liolepis Boulenger, 1885; C. nigrilabris Peters, 1860; C.
desilvai Bahir and Maduwage, 2005) are endemic. The
remaining two Calotes (C. calotes [Linnaeus, 1758); C.
versicolor (Daudin, 1802]) are probably widespread
species throughout South East Asia. According to thepublished literature, Calotes liocephalus is a largely
arboreal species found only in parts of the Knuckles
Forest Region in Sri Lanka (Manamendra-Arachchi andLiyanage, 1994). Its conservation status is Rare andEndangered (Bahir and Surasinghe, 2005). It can be distinguished from its congeners by the presence of an
oblique fold in front of the shoulder, a lower jaw that israther short, a head without spines (or rarely a rudimentary spine above the ear), enlarged supraocular scales
and poorly-developed dorsinuchal crests on the head andlower neck (Manamendra-Arachchi, 1990). Adults havea snout to vent length of 91 mm, a head length of 37 mm,
a tail length of 261 mm and an axilla to groin length of
43 mm (Deraniyagala, 1953).
Location of observation.- Observations were madeapproximately 1 km from Matale-Pallegama Road inPitawala in the Knuckles Forest Region (altitude: 783m) in Matale District, Central Province. The habitat consisted mainly of disturbed home gardens (Ekanayakeand Bambaradeniya, 2001). The ground was covered
with small amounts of wet leaf litter and the soil was© 2008 by Asiatic Herpetological Researchsoft. There was approximately 10% canopy cover andthe undergrowth consisted primarily of grasses.
Observations of the lizard was made by the unaided eye
from 2 m away between the hours of 1420 and 1600 hrs.
The animal was not disturbed during observation. Allmeasurements were taken to the nearest 0.1 mm using
dial calipers.

Observations
A mature female Calotes liocephalus (snout to ventlength: 54.0 mm, head length: 19.4 mm, head width:11.9 mm, tail length: 156.0 mm, axilla to groin length:26.5 mm) lying on the ground, approximately 50 cm
from the road, was observed on 21 June 2006 at about
1420 hr. The temperature was 23.6°C and the humidity
93%. The weather was gloomy and the cloud cover was
8/8.
Digging the nest hole.- First, the lizard lifted the anterior part of its body using its forelimbs. It then looked
around for ~10 min. During this period it repeatedly
turned its head 180° five times, without moving its body(Fig. 1). The female then began digging into the ground
while scraping the soil with its forelimbs, which wasthrown backward under its body through its raised hindlimbs. This continued for approximately 5 minutes (Fig.
2). After that it stopped digging and looked around forapproximately 5 min. while repeatedly turning its head
180° three times, without moving its body (Fig. 3).
Again, it continued digging and this time the female dug
the hole continuously for approximately 10 min. It
stopped and looked around for about 5 min. while turning its head 180° around three times, without moving its
body as in Figure 3. After that, it continued to dig the14

Asiatic Herpetological Research, Vol. 11

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Figure 6.

Figure 7.

Figure 8.

20082008

Asiatic Herpetological Research, Vol. 11

15Figure 9a.

Figure 9b.

Figure 10.

Figure 11.

Figure 12.

Figure 13.

hole for another half an hour, stopping three more times
for 5 min. each, to rest. The hole was dug into the groundat a 45° angle. The final hole was 92.6 mm deep and79.1 mm in diameter (Fig. 4). During the rest intervals
the body was coiled inside the hole with the anterior halfbent at an angle of 90° to looking around (Fig. 5). There
was a drizzle for ~15 minutes, but the female continued
digging.

behavior was that the female removed herself slowly
from the hole without lifting her limbs while it was laying its eggs (Figs. 7–8). Eight eggs were laid at a rate ofone per minute. The eggs were pure white and elliptical,
with a mean length of 14.8 mm and a mean width 8.6
mm. After the eggs were laid, the female came out of thehole completely and started looking around (Fig. 9a, b).
Then the female crept back into the hole for 15 min. topack and place the eggs below ground level using theanterior part of its lower jaw (Fig. 10).

Laying the eggs. - After half an hour of digging, thefemale turned its body 180° clockwise, placing the posterior part of its body inside the hole. It then looked
around again (Fig. 6). The significance of this egg laying

Burying the eggs and camouflaging the nest.- Aftercoming out of the hole, the female turned 180° clock-16

Asiatic Herpetological Research, Vol. 11

wise and began to drag the soil towards the hole using its
forelimbs. The dragged soil was thrown backwards
under its body while it lifted its hind limbs (Fig. 11).
After dragging the soil for about 5 min., it turned 180°
counter-clockwise and began pressing the soil with theanterior half of its lower jaw for half an hour. The hole
was filled up to 18.4 mm below ground level (Fig. 12).
After looking around, it dragged the surrounding Albizia
saman (Family: Fabaceae) leaves over the nest site forcamouflage (Fig. 13). It remained motionless for 2 min.
and then ran towards the forest, during which time it wascaught for measurement and then released.

Discussion
The oviposition behavior of this species varies from theoviposition behavior of Calotes versicolor. According toAmarasinghe and Karunarathna (2007), C. versicolor
places its cloacal aperture over the opening of the holewhile laying its eggs, but C. liocephalus places the posterior part of the body inside the hole while laying eggs.
C. versicolor also lifts the anterior part of the body with
its forelimbs while turning its head to look around, butC. liocephalus coils its entire body inside the hole whilebending the anterior part of its body to look around. C.
versicolor makes a knocking noise while packing andplacing the eggs in the hole using its lower jaw while theC. liocephalus places them softly without making any
noise. After the observation the eggs were removed fromthe hole and the hole was subsequently examined. Thebottom was conical and the soil was soft, dark and wet.
Finally the eggs were buried in a home garden to hatch.
After approximately two and half months we observed
five small holes where the hatchlings had come out.
Unfortunately we could not observe the hatchlings.
A Few diagrams, brief descriptions and notes ofCalotes liocephalus are available in popular journals,
books and magazines but almost nothing exists on thepre and post mating behavior, egg laying behavior, captive breeding and their hatchlings. In addition Calotes
liocephalus is an endemic, rare and threatened species
and therefore it may become extinct if their population
does not increase. For such a situation to be achieved,
captive breeding methods may be needed for ex-situ
conservation of this species. In addition further observations are also needed for the conservation of Calotes liocephalus.

Acknowledgments
We wish to thank Mr. Kelum Manamendra-Arachchi(WHT – Wildlife Heritage Trust) for reviewing the manuscript and Mrs. Zeenia Nissam of the Department ofSubmitted: 10 October 2006
Accepted: 23 September 2007

2008

Zoology, Faculty of Natural Sciences of the Open
University, Sri Lanka, for her generous support for thefield visit. We also thank Mr. Niranjan Karunarathna andMr. Gayan Wijeytunga (YZA – The Young Zoologists’
Association) for assisting the fieldwork. Finally, Ms.
Debbie McCormick and Mr. F. S. Abeywickrama are
acknowledged for their help in preparing this paper.

Literature Cited
Amarasinghe, A. A. T. and D. M. S. S. Karunarathna.
2007. Beobachtungen zum Eiablageverhalten derIndischen Schönechse Calotes versicolor(Daudin, 1802) (Reptilia: Agamidae) in einem
anthropogenen Biotop in Sri Lanka. Sauria,Berlin, 29(3): 27–30.
Bahir, M. M. and T. D. Surasinghe. 2005. A conservationassessment of the agamid lizards of Sri Lanka. The
Raffles Bulletin of Zoology, Suppl. 12: 381–392.
Deraniyagala, P. E. P. 1953. A colored atlas of some vertebrates from Ceylon, Tetrapod Reptilia. National
museums of Sri Lanka.
Ekanayake, S. and C. N. B. Bambaradeniya. 2001.
Trekking in the Knuckles forest – A trekking guideto Alugallena, Dekinda and Nitre cave nature trails.
IUCN Sri Lanka.
Manamendra-Arachchi, K. 1990. A guide to theAgamids in Sri Lanka. Occ. Papers of the Young
Zoologist Association of Sri Lanka. 5: 1–8.
Manamendra-Arachchi, K. and 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.
Manamendra-Arachchi, K., A. de Silva and T.
Amarasinghe. 2006. Description of a second
species of Cophotis (Reptilia:Agamidae) from thehighlands of Sri Lanka. Lyriocephalus 6(1): 1–8.
Samarawickrama, V. A. M. P. K., K. B. Ranawana, D. R.
N. S. Rajapaksha, N. B. Ananjeva, N. L. Orlov, J.
M. A. S. Ranasinghe and V. A. P. Samarawickrama.
2006. A new species of the Genus Cophotis(Squamata: Agamidae) from Sri Lanka. Russian
Journal of Herpetology 13(3): 207–214.