Russian Journal of Herpetology

Vol. 18, No. 2, 2011, pp. 111 118

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 including 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 and Surasingha, 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 genus 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 nationally threatened (IUCNSL and MENR, 2007). The non-endemics are the C. calotes and C. versicolor, which are relatively common (Erdelen, 1978, 1984, 1988; Haly, 1887; Nevil, 1887).
1

2 3

Nature Exploration & Education Team, No. B-1/G-6, De Soysapura, Morauwa 10400, Sri Lanka. E-mail: dmsameera@gmail.com The Young Zoologists Association, Department of National Zoological Gardens, Dehiwala, Sri Lanka.

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 separated localities restricted to the mountain forests of Sri Lanka, mainly in open shrub and grassland areas of forests in the wet zone and home gardens above 1500 m elevation 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 agamid 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

112 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 February 6, 2010, without making any disturbance. The specimen was captured after oviposition and examined carefully to record scale patterns and external measurements, and photographed before releasing back to the same habitat. Scales were observed using a 10 hand lens and external 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 during the observation. Geographic coordinates are derived from 1:50,000 topographical map series of the Survey Department of Sri Lanka. The diagnostic keys and characters given by Smith (1935), Manamendra-Arachci (1990), Boulenger (1890), Gnther (1864), Somaweera and Somaweera (2009), Deraniyagala (1953), Manthey

D. M. S. S. Karunarathna et al. (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 distance 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 disturbed 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

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 Year Month Climate condition Time duration Soil and it color Leaf-litter thickness, mm Canopy cover, % Cloud cover, % Temperature, C Humidity, % Body pit diameter, mm Angle of the hole, deg. Depth of the hole, mm Diameter of the hole, mm The number of eggs in clutch Hakgala-1 2009 March Cool and sunny 12:25 13:10 Wet-dark-soft 10 0 50 21.5 58 70 45 50 40 Hakgala-2 2010 February Mist rainy 11:38 14:24 Wet-dark-soft 15 0 60 22.1 62 80 45 55 33 4 Seetha-eliya 2010 February Mist rainy 10:45 11:20 Wet-dark-soft 15 0 50 20.3 65 80 48 55 30 3 Nuwara-eliya 2009 February Cool and shady 12:10 13:26 Wet-dark-soft 10 0 60 22.5 71 75 50 50 38 Horton-1 2010 March Mist rainy 10:50 12:15 Wet-dark-soft 10 0 80 20.2 68 85 45 58 35 4 Horton-2 2010 March Mist rainy 11:20 13:35 Wet-dark-soft 10 0 80 20.4 60 80 50 55 34 3

TABLE 2. Egg Measurements of the C. nigrilabris (n = 14) February Character (1) Hakgala-2 (2) (3) (4) (5) Seetha-eliya (6) (7) (8) 17.1 10.5 27.6 Horton-1 (9) 17.5 10.4 27.9 (10) 17.3 10.4 27.7 (11) 17.7 10.6 28.3 (12) 19.5 10.2 29.7 March Horton-2 (13) 17.5 10.1 27.6 (14) 17.8 10.8 28.6 17.66 10.31 27.98 Mean

EL, mm 17.4 17.6 17.3 17.5 18.1 17.6 17.4 EW, mm 10.1 10.3 10.2 10.1 10.2 10.1 10.4 ToL 27.5 27.9 27.5 27.6 28.3 27.7 27.8 Note. EL, egg length; EW, egg width; ToL, total of egg width and long.

Ovipositional Behavior of the Calotes nigrilabris in Sri Lanka 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 Gunatilleke (1990) the major vegetation type in the study area is tropical 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 Calotes 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 its body. 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 continuously 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 approximately 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 intervals the tail was coiled around the nest hole with the head

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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 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. It then looked around a span of 300 angle, thrice, spending ~6 min. The lizard then laid eggs in the hole without lifting 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 locations (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 process. Burying eggs and covering the nest. After the resting 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 anterior 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 noises were herd during each pressing. After filling approximately 3/4 of the hole in about 35 min, it turned 120 clockwise and moved to a distance of 150 mm. Then it collected few small pieces of grass leaves for camouflaging 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

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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 grass leaves 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).

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 surrounding to cover the nest site. It took 15 min for camouflaging 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.

Ovipositional Behavior of the Calotes nigrilabris in Sri Lanka DISCUSSION This is the first detailed record of the ovipositional behavior of Calotes nigrilabris which is one of the endemic and uncommon agamid lizards in Sri Lanka. We believe this notes will not only provide useful information 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 released 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 of C. nigrilabris were observed in 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 February 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 observations, it can be concluded that there is a significant behavioral uniformity in nesting among different females of C. 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 behavior of C. nigrilabris described here varies considerably from the ovipositional behavior of C. ceylonensis, C. liocephalus, and C. versicolor, according to Amarasinghe and Karunarathna (2007 and 2008), Karunarathna et al. (2009), Erdelen (1986), and Pradeep and Amarasinghe (2009). According to published ligatures, C. ceylonensis places posterior part of its body over the hole while laying 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.

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liolepis (Fig. 2f). Few other behavioral comparisons in oviposition of C. nigrilabris and other Sri Lankan Calotes lizards are discussed below. C. calotes, C. liocephalus, 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 Gupta, 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 recorded to throw the soil backward under its body through its raised hind limbs, in a way similar to other species except 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 (Amarasinghe 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 Karunarathna, 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 of C. 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. ceylonensis (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 compressing the soil on the nest-hole using its lower jaw (Karunarathna et al., 2009). C. nigrilabris also makes a knocking noise while packing and placing the eggs in the hole using its lower jaw, it is similar to C. calotes and C. versicolor (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 established 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 behaviors, such as coiling of the tail around the outer margin of

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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 by Thasun Amarasinghe).

the hole and making a knocking noise while compressing the soil on the nest-hole. By thus far the ovipositional behavior of six out of seven species in the Genus is been described, 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 individuals (males + females + juveniles) per hectare. Our personal observations suggest its populations are at a decline. 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 for C. 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 information.
Acknowledgments. First we would like to thank Dr. Channa Bambaradeniya and Dr. Sandun Perera for critical reviewing and corrections. We are grateful to Mr. Dishad Jemzeed (YZA) for excellent photographs and Mr. Thasun Amarasinghe (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 discussions; Mr. Panduka Silva, Mr. Asanka Udayakumara, Mr. Toshan Peries, Mr. Chamila Soysa, Mr. Anushka Kumarasinghe, Mr. Tiran Abeywardene and members of the YZA for helped

Ovipositional Behavior of the Calotes nigrilabris in Sri Lanka

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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