Ovipositional behaviour of Calotes nigrilabris (Sri Lanka)

7/31/2019 Ovipositional behaviour of Calotes nigrilabris (Sri Lanka)

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FIRST REPORT ON THE OVIPOSITIONAL BEHAVIOR OF Calotes nigrilabris

PETERS, 1860 (REPTILIA: SAURIA: AGAMIDAE)

FROM THE CENTRAL MASSIF OF SRI LANKA

D. M. S. S. Karunarathna,1,2 W. A. A. D. G. Pradeep,3

P. I. K. Peabotuwage,3

and M. C. De Silva3

Submitted May 26, 2010.

Calotes nigrilabris (Black-cheek lizard) is an endemic, range-restricted and threatened lizard species in Sri Lanka.

This paper provides the first detailed description of the ovipositional behavior of Calotes nigrilabris in Sri Lanka,

recorded from the periphery of Hakgala Strict Nature Reserve in the Central massif. The observations described in

this article would be useful in conservation planing for this species which is restricted to the central highlands of

the island. But ovipositional behavior of the critically endangered Calotes desilvai still not document.

Keywords: Agamidae; Calotes; conservation; decline; ovipositional behavior; Sri Lanka.

INTRODUCTION

Sri Lanka harbors a rich biodiversity, which includes

high herpetofaunal diversity (Bossuyt et al., 2004;

Gunawardene et al., 2007; Meegaskumbura et al., 2002;

Myers, et al. 2000). There are 96 species of saurians in-

cluding 72 (75%) species that are endemic to the island.

Among them are 18 species of agamid lizards including 3

relict and endemic genera (Ceratophora, Cophotis,

Lyriocephalus) and 15 (83%) endemic species (Bahir andSurasingha, 2005; De Silva, 2006; Somaweera and

Somaweera, 2009). These 18 native species belong to the

subfamily Draconinae and consists of 6 genera; Calotes,

Ceratophora, Cophotis, Lyriocephalus, Otocryptis, and

Sitana (Deraniyagala, 1953; Macey et al., 2000;

Manamendra-Arachchi, 1990; Manthey, 2008). The ge-

nus Calotes consist of seven species, of which five are

endemic to the island; these are C. ceylonensis, C.

desilvai, C. liocephalus, C. liolepis, and C. nigrilabris

(Bahir and Maduwage, 2005; Das and De Silva, 2005;

Manamendra-Arachchi, 1998), all of which are nation-

ally threatened (IUCNSL and MENR, 2007). Thenon-endemics are the C. calotes and C. versicolor, which

are relatively common (Erdelen, 1978, 1984, 1988; Haly,

1887; Nevil, 1887).

Calotes nigrilabris, the Black-cheek lizard known as

Kl kopul katss in Sinhalese (Das and De Silva,

2005), has been recorded from only a few widely sepa-

rated localities restricted to the mountain forests of Sri

Lanka, mainly in open shrub and grassland areas of for-

ests in the wet zone and home gardens above 1500 m ele-

vation above sea level (a.s.l.) (Manamendra-Arachchi

and Liyanage, 1994). However, Deraniyagala (1953) had

reported a specimen from Peradeniya (~650 m a.s.l.),way below its other localities. This species is considered

as nationally threatened, and placed under the vulnerable

category (IUCNSL and MENR, 2007), even though,

Bahir and Surasingha (2005) have listed it as endangered.

This sub-arboreal species is much dark green in color

than other Calotes in Sri Lanka and unusual among aga-

mid lizards due to its hissing sound when alarmed

(Deraniyagala, 1931; De Silva, 1991; Fernando, 1998).

Average adult snout to vent length (SVL) is 86 mm

(range 84.3 101.5 mm) in males and 74 mm (range

64.4 78.9 mm) in females, head length (HL) is 32 mm

(range 29.1 35.7 mm) in males and 24 mm (range

21.3 24.8 mm) in females, and the tail length (TL) is

245 mm (range 101 300 mm) in males and 228 mm

(range 205 270 mm) in females. Somaweera and

Somaweera (2009) record different color variations of

this species.

1026-2296/2011/1802-0111 2011 Folium Publishing Company

Russian Journal of Herpetology Vol. 18, No. 2, 2011, pp. 111 118

1 Nature Exploration & Education Team, No. B-1/G-6, De Soysapura,Morauwa 10400, Sri Lanka.

2 E-mail: [email protected] The Young Zoologists Association, Department of National Zoolog-

ical Gardens, Dehiwala, Sri Lanka.


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MATERIAL AND METHODS

Observations of the lizard were made by the naked

eye at a distance of 2.5 m from 11:38 to 14:24 on Febru-

ary 6, 2010, without making any disturbance. The speci-

men was captured after oviposition and examined care-

fully to record scale patterns and external measurements,

and photographed before releasing back to the same habi-

tat. Scales were observed using a 10 hand lens and ex-

ternal measurements were taken to the nearest 0.1 mm

using a manual vernier caliper (accuracy 0.005 mm)

and a 1-m measuring tape. Measurements of the eggs

were also obtained (Tables 1 and 2) through the vernier

caliper and the eggs were carefully deposited back in the

original nest hole. A thermometer and a hygrometer were

used to record the temperature and relative humidity dur-

ing the observation. Geographic coordinates are derivedfrom 1:50,000 topographical map series of the Survey

Department of Sri Lanka. The diagnostic keys and char-

acters given by Smith (1935), Manamendra-Arachci

(1990), Boulenger (1890), Gnther (1864), Somaweera

and Somaweera (2009), Deraniyagala (1953), Manthey

(2008), Amarasinghe et al. (2009), Das and De Silva

(2005), and Taylor (1953) were used for the identification

of species. The plant nomenclature is based on Senaratna

(2001) and identifications are based on Ashton et al.

(1997).

STUDY AREA AND HABITAT

Observation was made approximately 500 m dis-

tance from Hakgala Strict Nature Reserve (SNR) in

Nuwara-Eliya District, Central Province, Sri Lanka

(65532 N and 804926 E, 1850 m a.s.l.). There was

no canopy cover at the ovipositional location, but the

canopy cover 3 m towards the north of the location was

about 50%. The undergrowth was also very poor. The

surrounding area was covered with small grasslands and

Rhododendron arboretum bushes about 1 m tall. The dis-

turbed habitat is abandoned potato cultivation, and the

degraded land was covered with a layer of leaf litters as

thin as 4 mm. The soil was wet, soft and it contains

blackish brown earth loosely-bound particles. The air

112 D. M. S. S. Karunarathna et al.

TABLE 1. Ecological Parameters During Oviposition and Nest-Hole Description of the C. nigrilabris at Six Sites in the Central Massif of Sri

Lanka (table followed by Karunarathna et al., 2009)

Ecological data Hakgala-1 Hakgala-2 Seetha-eliya Nuwara-eliya Horton-1 Horton-2

Year 2009 2010 2010 2009 2010 2010

Month March February February February March MarchClimate condition Cool and sunny Mist rainy Mist rainy Cool and shady Mist rainy Mist rainy

Time duration 12:25 13:10 11:38 14:24 10:45 11:20 12:10 13:26 10:50 12:15 11:20 13:35

Soil and it color Wet- dark-sof t Wet-dar k- soft Wet-dar k- soft Wet-dark-soft Wet-dark-soft Wet-dark-soft

Leaf-litter thickness, mm 10 15 15 10 10 10

Canopy cover, % 0 0 0 0 0 0

Cloud cover, % 50 60 50 60 80 80

Temperature, C 21.5 22.1 20.3 22.5 20.2 20.4

Humidity, % 58 62 65 71 68 60

Body pit diameter, mm 70 80 80 75 85 80

Angle of the hole, deg. 45 45 48 50 45 50

Depth of the hole, mm 50 55 55 50 58 55

Diameter of the hole, mm 40 33 30 38 35 34

The number of eggs in clutch 4 3 4 3

TABLE 2. Egg Measurements of the C. nigrilabris (n = 14)

Char-

acter

February March

MeanHakgala-2 Seetha-eliya Horton-1 Horton-2

(1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14)

EL, mm 17.4 17.6 17.3 17.5 18.1 17.6 17.4 17.1 17.5 17.3 17.7 19.5 17.5 17.8 17.66

EW, mm 10.1 10.3 10.2 10.1 10.2 10.1 10.4 10.5 10.4 10.4 10.6 10.2 10.1 10.8 10.31

ToL 27.5 27.9 27.5 27.6 28.3 27.7 27.8 27.6 27.9 27.7 28.3 29.7 27.6 28.6 27.98

Note. EL, egg length; EW, egg width; ToL, total of egg width and long.


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temperature was measuring ~22.1C and the humidity

~62%. The weather on the day of the observations was

misty with a little bit of sunshine, and the cloud cover

was 60% and notably, there was a heavy rain during the

previous day. According to the Gunatilleke and Gunatil-

leke (1990) the major vegetation type in the study area istropical mountain forest dominant with Calophyllum,

Syzygium, and Walkeria community mixed with wet

pathana grasslands. The mean annual rainfall varies from

3500 to 5000 mm (mainly during South-west monsoon),

while the mean annual temperature of the area varies

from 15.4 to 20.7C.

RESULTS AND OBSERVATIONS

Excavating of the nest hollow. A mature female Ca-

lotes nigrilabris (SVL = 73 mm, HL = 22.6 mm,HW = 13.4 mm, TL = 157 mm, AGL = 43.6 mm) was

lying on the ground in the Hakgala, about 1 m away from

a secondary gravel road, on February 6, 2010, at about

11:38. The lizard showed a body color of green and the

tail was fully dark brownish, lacking camouflage. First,

the lizard was digging with vigilant for about 14 min.

Sometimes it used only one forelimb for digging while

standing on the other. Then it tilted its body (~45) to

make a small body pit (~80 mm in diameter and ~12 mm

high). After about 8 min it stopped the activity to have a

look-around. During the preparation of body pit it turned

its head around for about a 60 twice, without moving itsbody. The bottom of the body pit was flat and large

enough for the female to lie in while laying eggs. Another

super male C. nigrilabris was observed on a small bush

~2 m away from the female, about 1 m above the ground

level. The male was very colorful, and it was continu-

ously at watch on the nest building female on the ground

(Fig. 1a g).

Then the lizard started excavating the ground and at

the same time scraping the soil with its forelimbs one

hand after the other, in this process turning its body

clockwise and counterclockwise 13 times during this

process. The scraped soil was thrown backward under its

body through its raised hind limbs. Then it compacted the

soil inside the nest-hole using the forehead. This activity

was repeated for about 10 times, spending ~2 min a time,

and looked around for about five times, spending approx-

imately 5 min a time without moving its body. During

this activity it changed the body color along its dorsal line

to become darker matching well with the ground color.

While digging, it was observed stopping after 18 23

scraps for ~1 min each, to rest. During the resting inter-

vals the tail was coiled around the nest hole with the head

been turned around for an angle of 110 to look around. It

finished the digging in 22 min and looked around for

~15 min without any movement. The female took almost

one hour to finish the nest-hole and it was dug into the

ground at an angle of 45. The nest-hole was 50 mm deep

and 45 mm in diameter.

Laying of eggs. After about an hour of digging, it

turned its body 150 counterclockwise; placing the poste-

rior part of its body at the mouth of the nest-hole and the

entire tail was coiled at the outer margin of the hole. It

then looked around a span of 300 angle, thrice, spending

~6 min. The lizard then laid eggs in the hole without lift-

ing its forelimbs but with a slight lift in hind limbs. The

hind limbs were placed at the top-opposite side of the

body pit. The gular area and breast were well gripping the

ground during the whole egg laying period, but the snout

was slightly lifted at an angle of ~40. It always moved

its eyes and kept watching when crows were flying over.

Four eggs were laid at a rate of approximately one per

2 min. The eggs were pure white and much elliptical with

a thin pliable shell: mean length = = 17.45 mm and mean

width = 10.17 mm (n = 4). By pooling these data with

later observations of C. nigrilabris from few other loca-

tions (n = 14) the mean egg size was 17.66 10.31 mm;

with a range of (17.1 19.5) (10.1 10.8) mm. The

lizard placed down its snout on the ground and remained

motionless for 12 min, resting after the egg-laying pro-

cess.

Burying eggs and covering the nest. After the rest-ing time, the lizard turned 210 counterclockwise and

moved back into the nest-hole and spent 4 min packing

and placing the eggs below ground level using the ante-

rior part of its snout tip. It then turned 180 clockwise and

began dragging wet soil towards the hole using its

forelimbs. The excavated soil was pulled back by using

its hind limbs and it continued to fill the hole for 52 min,

burying its eggs. During this time, it stopped the activity

13 times for ~1 min each, to rest. Thereafter, the lizard

started compressing the soil and a knocking noise was

produced as the lizard hit the substrate 18 times with its

snout tip to compress the soil. Eight to 12 knock noiseswere herd during each pressing. After filling approxi-

mately 3/4 of the hole in about 35 min, it turned 120clockwise and moved to a distance of 150 mm. Then it

collected few small pieces of grass leaves for camouflag-

ing the nest and these grass leaves were dragged

and pulled using its hind limbs one after the other

(Fig. 1b, c).

Then the nest-hole was completely filled up to the

ground level, and thereafter the lizard turned its body

180 counterclockwise to start compressing the soil again

Ovipositional Behavior of the Calotes nigrilabris in Sri Lanka 113


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with the snout tip for nearly 11 min. Then it looked

around by turning around its head for an angle of 160

and started dragging fallen grass leaves from the sur-

rounding to cover the nest site. It took 15 min for camou-

flaging the nest and then remained motionless for

~10 min, while changing its body color into light green.

Then it moved slowly towards to the grassland and lied

there for ~5 min. After that, the specimen was caught for

measurement, and then released to the same place with

minimal disturbance. The eggs were also removed from

the nest-hole smoothly for examination and deposited

back the same way, as soon as possible.


a b

d

f

c

e

Fig. 1. Ovipositional behavior of the C. nigrilabris: a, female digging the nest; b, female filling the nest-hole with soil; c, female scraping the grassleaves for camouflage the nest; d, super male being watch full; e, the nest hole with the 4 eggs; f, 7 eggs were observed from different locations

(photographs taken by Dilshad Jemzeed in Horton Plains National Park).


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DISCUSSION

This is the first detailed record of the ovipositional

behavior of Calotes nigrilabris which is one of the en-

demic and uncommon agamid lizards in Sri Lanka. We

believe this notes will not only provide useful informa-tion on the nesting behavior of the lizard, but also have

conservation implications (Karunarathna et al., 2009).

After egg-laying was over and the female lizard was re-

leased after measurements, it moved to the one meter tall

grassland mixed with bushes. Surprisingly then the male

resting on the nearby bush rushed down and griped the

female at its mid-body area using its forelimbs and

mouth. Even though the female tried to escape initially,

they copulated for nearly 15 min. This incident was much

similar to the observation on C. calotes (Gabadage et al.,

2009). Subsequent to this detailed observation several

other egg laying episodes ofC. nigrilabris were observedin six different occasions between 10:45 hrs to 14:24 hrs

in the month of February and March (see Table 1 for

comparisons and Table 2 for egg measurements).

These observations suggest that C. nigrilabris prefers

to lay eggs in open areas lacking canopy, with egg laying

done during the daytime, and during the Febru-

ary March period of the year. We can conclude that

C. nigrilabris lays 3 to 4 eggs, builds a holes are 30

40 mm diameter (average 35 mm) and hole depths are

50 58 mm (average 53.8 mm). From all our observa-

tions, it can be concluded that there is a significant be-

havioral uniformity in nesting among different females ofC. nigrilabris. The oviposition behaviors of Sri Lankan

Agamids has received a recent attention, documenting

these behaviors for most in the genus Calotes in Sri

Lanka (Gabadage et al., 2009; Karunarathna et al., 2009),

except only the Calotes desilvai. The ovipositional be-

havior of C. nigrilabris described here varies consider-

ably from the ovipositional behavior of C. ceylonensis,

C. liocephalus, and C. versicolor, according to

Amarasinghe and Karunarathna (2007 and 2008), Karu-

narathna et al. (2009), Erdelen (1986), and Pradeep and

Amarasinghe (2009).

According to published ligatures, C. ceylonensisplaces posterior part of its body over the hole while lay-

ing eggs and also stretch the posterior part of its body

(Fig. 2a), C. liocephalus places the posterior part of the

body inside the hole while laying eggs (Fig. 2b ) and C.

versicolor places its cloacal aperture over the opening

while laying eggs (Fig. 2c). It was significantly different

in C. nigrilabris, where it placed the posterior part of its

body at the mouth of the nest-hole and the entire tail was

coiled at the outer margin of the hole, while hind limbs

were lifted a little, but not stretched horizontally

(Fig. 2d), somewhat similar to C. calotes (Fig. 2e) and C.

liolepis (Fig. 2f). Few other behavioral comparisons in

oviposition of C. nigrilabris and other Sri Lankan

Calotes lizards are discussed below. C. calotes, C. lioce-

phalus, and C. versicolor do not create a body pit to dig

the nest-hole (Amarasinghe and Karunarathna, 2007,

2008; Gabadage et al., 2009; Rathinasabapathy and Gup-ta, 1997). However, C. ceylonensis, C. liolepis, and C.

nigrilabris finely create their body pits (Karunarathna et

al., 2009; Pradeep and Amarasinghe, 2009).

The body pits of C. nigrilabris are relatively small

(70 85 mm), in comparison to that of C. ceylonensis

and C. liolepis (120 150 mm). C. nigrilabris was re-

corded to throw the soil backward under its body through

its raised hind limbs, in a way similar to other species ex-

cept C. liolepis. C. nigrilabris as same as the C.

liocephalus, C. versicolor, and C. ceylonensis dug their

nest-holes with an angle of 45. However, C. calotes dug

the nest-hole into the ground at an angle of 35 and C.liolepis was observed digging its nest-hole at 90

(Gabadage et al., 2009). During oviposition C. versicolor

usually lifts the anterior part of the body with its

forelimbs while turning its head to look around (Amara-

singhe and Karunarathna, 2007) and C. liocephalus coils

its entire body inside the hole while bending the anterior

part of its body to look around (Amarasinghe and Karu-

narathna, 2008). But C. liolepis coiles the tail inside the

nest-hole with the head turned at an angle of 90 to look

around (Karunarathna et al., 2009).

Although in the case ofC. calotes the tail is coiled at

the outer margin of the nest-hole (Gabadage et al., 2009;Kannan and Bhupathy, 1996), much similar to C. ceylo-

nensis (Pradeep and Amarasinghe, 2009). When it comes

to C. nigrilabris the entire tail was coiled around the

outer margin of nest-hole and looked around at an angle

of 300. C. ceylonensis and C. liocephalus places the

eggs softly without making any noise (Amarasinghe and

Karunarathna, 2008; Pradeep and Amarasinghe, 2009)

but C. liolepis makes a knocking noise while compress-

ing the soil on the nest-hole using its lower jaw (Karuna-

rathna et al., 2009). C. nigrilabris also makes a knocking

noise while packing and placing the eggs in the hole us-

ing its lower jaw, it is similar to C. calotes and C. versico-lor (Amarasinghe and Karunarathna, 2007; Gabadage et

al., 2009; Karthikeyan, 1993; Prasad and Jayanth, 1991).

According to the phylogenetic analysis of Macey et al.

(2000) and hemipenes analysis of agamid lizards by

Maduwage et al. (2008), relationships have been estab-

lished for two clades; C. ceylonensis, C. liocephalus, and

C. liolepis in one, and C. calotes and C. nigrilabris the

other, sister to the first clade.

We found complementary observations that further

support the above relationships, in the oviposition behav-

iors, such as coiling of the tail around the outer margin of



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the hole and making a knocking noise while compressing

the soil on the nest-hole. By thus far the ovipositional be-

havior of six out of seven species in the Genus is been de-

scribed, but nothing is known for that of the Calotes

desilvai. But according to Bahir and Maduwage (2005)

one of the paratypes of C. desilvai has two white eggs

measuring 16 8.5 mm size. Hence, ecological studies

on the critically endangered Agamid species C. desilvai.is envisaged. Erdelen (1984, 1988) records C. nigrilabris

as a mountain specific species with an average of 220 in-

dividuals (males + females + juveniles) per hectare. Our

personal observations suggest its populations are at a de-

cline. In addition, we could record Sri Lanka whistling

thrush (Myophonus blighi), jungle crows (Corvus

macrorhynchos), and feral cats (Felis catus) in our study

areas, both species recorded to be opportunistic feeders

in human habitations and predate on lizards

(Karunarathna and Amarasinghe, 2008; De Silva, 2006,

2007; Warakagoda, 1997). Also several road kills were

found at the Horton plains National park and its outside.

Hence, conservation planning forC. nigrilabris is of

an immediate need and we believe captive breeding

methods may be needed for ex situ conservation of the

species, for which this report will provide vital informa-

tion.

Acknowledgments. First we would like to thankDr. Channa Bambaradeniya and Dr. Sandun Perera for critical

reviewing and corrections. We are grateful to Mr. Dishad Jem-

zeed (YZA) for excellent photographs and Mr. Thasun Amara-

singhe (TNCS) for excellent drawings. This study was start in

2008 by Mr. Niranjan Karunarathna (YZA), who provided

guidance and financial support it helped improve the data. We

are grateful to Mr. Mendis Wickramasinghe (HFS), Mr. Kelum

Manamendra-Arachchi (WHT), Mr. Mohomad Bahir (TNCS),

and Dr. Anslem De Silva (ARROs) for the valuable discus-

sions; Mr. Panduka Silva, Mr. Asanka Udayakumara, Mr. To-

shan Peries, Mr. Chamila Soysa, Mr. Anushka Kumarasinghe,

Mr. Tiran Abeywardene and members of the YZA for helped


Fig. 2. Oviposition behavior patterns of the 6 species of the genus Calotes: a, C. ceylonensis Mller, 1887; b, C. liocephalus Gnther, 1872; c, C.

versicolor (Daudin, 1802); d, C. nigrilabris Peters, 1860; e, C. calotes (Linnaeus, 1758); f, Calotes liolepis Boulenger, 1885 (drawings made byThasun Amarasinghe).


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various ways to enrich this works. Finally, we would like to

thank Dr. Wolfgang Bhme and Dr. Natalia Ananjeva for their

suggestions and reviewing.

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Ovipositional behaviour of Calotes nigrilabris (Sri Lanka)