Colour Change As an Anti-Predatory Mechanism in Calotes

Home , Calotes, Calotes calotes, Dendrelaphis tristis, Otocryptis wiegmanni

Herpetology Notes, volume 11: 675-678 (2018) (published online on 24 August 2018)

Colour� �� � � ��r���or� ��� �� Calotes calotes (Linnaeus, 1758) (Squamata: Agamidae) sighted at the �� r��, Sr�� Lanka

Dulan Jayasekara1 and Chathuranga Dharmarathne1,*

Survival and reproduction are the main aspects the predator and prey species to which it must appear of any life form. When it comes to survival, animals camouflaged need to be considered to better understand are continuously exposed to different threats and these strategies (Endler, 1978; Kelman et al., 2008). dangers from nature. Most animals face the challenge Calotes calotes (Linnaeus, 1758) (Common Green of dealing with multiple predators and their level of Forest Lizard) is an agamid lizard found in the forests of survival depends on their ability to detect predators, the Western Ghats and the Shevaroy Hills in India, and various predator avoidance mechanisms and anti- Sri Lanka. It is a considerably large species of agamid, predatory �ehaviours. Most prey species ad�� measuring up to 41 cm in length, including the tail. The behaviour or appearance in response to their predators body colouration of C. calotes is generally green. There (Hopper 2001; Templeton and Shriner, 2004; Stuart- is no record on contrastingly different colour changes in Fox et al., 2006; Langridge et al., 2007; Rundus et al., 2007). Body colouration is the main characteristic that determines the appearance of an animal, which comes in the forms of camouflage, conspicuous/ disruptive colouration and mimicry against predators (Stevens, 2007). Physiological colour change is used by prey species for rapid colour changes (Stuart-Fox and Moussalli, 2008). Colour change not only enables animals to match different backgrounds but also enables prey to change their appearance in the face of different predators (Langridge et al., 2007) and potentially adjust their camouflage depending on the visual capabilities of the predators. Many animals use colour change for camouflage as well as communication. Colour patterns in animals exploit different predator avoidance or antipredator mechanisms including disruptive camouflage or motion dazzle (stripes), intimidation of predators or deflecting attention towards the tail and background matching (Stevens, 2007; Stevens et al., 2008). The visual perception mechanisms of the animal, as well as

1 Department of Zoology, Faculty of Applied Sciences, University of Sri Jayewardenepura, Sri Lanka. *Corresponding author. E-mail: Figur� 1. Calotes calotes in general green body colouration at [email protected] Wasgamuwa National Park, Sri Lanka. 676 Dulan Jayasekara & Chathuranga Dharmarathne this species apart from adult males changing their head Here we present some field notes on the observation of and gular sac colouration from yellowish green into a prey-predator interaction among D. tristis and C. calotes red or dark red hue during breeding (Somaweera and at the Wasgamuwa National park. Wasgamuwa National Somaweera, 2009). The underside is a pale green, the Park is home to several agamid species including tail is light brown. the common C. calotes as well as C. ceylonensis and Dendrelaphis tristis (Daudin, 1803), Common Otocryptis wiegmanni (Senarathna, 2005). Bronzeback Tree Snake, is a long, slender, graceful tree Attack incident.—On 30thJuly 2015 around 16:00 snake that is non-venomous. This species is distributed when we were inside the National park, we observed in the wet, intermediate and dry zone lowlands of Sri a C. calotes being attacked by a D. tristis (Figure 2a). Lanka and can be found in India, Pakistan and Nepal as This lizard was on the ground among the “Mana” grass well (Das and de Silva, 2005). This diurnal and arboreal (a tall grass species) bushes. The initial colour of this snake is known to forage on land and along the edge of C. calotes was green with 5 or 6 white/cream colour water. According to Das and de Silva (2005) they can transverse stripes. make long jumps between trees, up to distance of 25m. The initial attack took place at 16:08. We observed Diet comprises frogs, lizards, bird eggs and insects. the D. tristis jumping towards the lizard through the (Das and de Silva, 2005). “Mana” grasses. The Snake was rapid and the lizard was

Figur� 2. (a) C. calotes being attacked by the D. tristis. (b) (c) Swallowing stages of C. calotes by D. tristis; Lizard has turned black with white stripes while the head has turned bright red (d) Colour change of attacked C. calotes from black to its general green body colouration Colou��r Calotes calotes 677 gripped from its head. The lizard, a well grown male, change has the potential to startle the predator and attempted several times to escape. In its defense to the deviate its focus. When it comes to visual perception snake, we observed the colouration of lizard changing of snakes, Gracheva et al. (2010) describe the ability of from a greenish dorsal colouration to a dark black snakes to detect infrared (IR) radiation which enables dorsal colouration and the appearance of broad, white the generation of a ‘thermal image’ of prey. However, transverse stripes (Figure 2b). The colour of the head IR detection is more accurately and frequently used by of the lizard changed from green to bright red/orange. nocturnal snakes and pit-bearing snakes (Roelke and There were 17 white stripes clearly visible from head Childress, 2007; Gracheva et al., 2010). Therefore, C. to the tail end. In addition, bands of white spots were calotes may be using this anti-predatory colour change present on the forelimbs and the hind limbs. The colour against other predators as well. Against a different change was complete by 16:10 and it took less than 120 predator it might have more impact and a higher chance seconds to change the general body colouration into the of survival. contrasting defense colouration against the predator. Despite the change of colour and continuous Acknowledgements. We would like to thank Mr. Sarath, Mr. escaping efforts, the lizard appeared to be defeated Supun and field staff of Sri Lanka Wildlife Conservation Society and immobilized (probably as a result of the strong (SLWCS) Wasgamuwa. We extend our gratitude to Mr. C. head restraint of lizard by the snake) at 16:21, eighteen Fernando (Ecologist), Ms. K. Butler (University of Newcastle, Australia) and Prof. W.A.D. Mahaulpatha (USJP) for their support minutes after the initial attack. The snake began to work on and off the field. We would like to thank Mr. D. Dharmarathne, the lizard around and was swallowing the lizard head Mrs. B. Rodrigo, and Mr. D. Jayakody. We appreciate the efforts first. At 16:29, the dark black colour of the C. calotes of Department of Wildlife Conservation for the protection of our started to change back to the previous green colour. It natural habitats. took nine minutes to change the colour back into the general green body colouration (Figure 2c – Figure 2d). R efer��� By this time the lizard was probably dead. Therefore, Cott, H.B. (1940): Adaptive coloration in animals. Methuen; it can be concluded that the lizard actively changed its London. colou��r Das, I., De Silva, A. (2005): A photographic guide to snakes and can be very aggressive towards snakes since they are other reptiles of Sri Lanka. New Holland. relatively large agamids themselves. However, in this Dodd Jr, C.K.. (1981): Infrared reflectance in chameleons case the non-venomous D. tristis easily overpowered (Chamaeleonidae) from Kenya. Biotropica 13(3): 161–164. the C. calotes. Endler, J.A. (1978): A predator’s view of animal color patterns. In: Hopefully, this field record from Wasgamuwa Evolutionary biology, p. 319–364. US, Springer. Gracheva, E.O., Ingolia, N.T., Kelly, Y.M., Cordero-Morales, National Park may help to understand a less known and J.F., Hollopeter, G., Chesler, A.T., Sánchez, E.E., Perez, J.C., a relatively complex anti-predatory �� Calotes Weissman, J.S., Julius, D. (2010): Molecular basis of infrared lizards. Our observations suggest a deviation from the detection by snakes. Nature 464: 1006. general camouflage strategies of lizards in which they Hopper, K. R. (2001): Flexible antipredator behavior in a dragonfly blend with their background to avoid the predator. In species that coexists with different predator types. Oikos 93: fact, the lizard changed its color as a response to the 470–476 attack by the snake. Therefore, it seems that C. calotes Kelman, E. J., Osorio, D., Baddeley, R. J. (2008): A review of cuttlefish camouflage and object recognition and evidence for lizards tend to distract their predator when the attack is depth perception. Journal of Experimental Biology 211: 1757– unavoidable. Despite such behaviours being uncommon 1763. among�� Langridge, K. V., Broom, M., Osorio, D. (2007): Selective signaling strategy more often (Cott, 1940; Dodd Jr, 1981; Stuart- by cuttlefish to predators. Current Biology 17: 1044–1045. Fox et al., 2006). Several explanations can be given for Roelke, C.E., Childress, M.J. (2007): Defensive and infrared this unusual rapid �� reception responses of true vipers, pitvipers, Azemiops and distract the non-venomous D. tristis by mimicking the colubrids. Journal of Zoology 273(4): 421–425. colouratio��n Bungarus caeruleus Rundus, A. S., Owings, D. H., Joshi, S. S., Chinn, E., Giannini, N. (2007): Ground squirrels use an infrared signal to deter (Schneider, 1801) (�� rattlesnake predation. Proceedings of the National Academy of similar pattern of white bands on dark body; 2. Deflecting Sciences of the United States of America 104: 14372–14376. the attention away from the head by interfering with Senarathna, P.M. (2005): “Wasgamuwa”. Sri Lankawe Wananthara, the visual perception of the predator. The rapid colour 1st Edition. Sarasavi Publishers. 678 Dulan Jayasekara & Chathuranga Dharmarathne

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Accepted by Hendrik Müller