Herpetology Notes, volume 12: 783-786 (2019) (published online on 15 July 2019)

Territorial behaviour in Oplurus grandidieri (Mocquard, 1900), a rock dwelling lizard from Madagascar

Javier Lobón-Rovira1,*, Francesco Belluardo1, Angelica Crottini1, and Gonçalo M. Rosa2,3

Territorial behaviour has been reported in numerous fight, whereby the intensity of the fight is thought to be lizards, with a range of different displays and tactics inversely correlated to the size of the territory to defend documented (Bustard, 1967; Sheldahl and Martins, or conquer: the smaller the available space the more 2000; Huyghe et al., 2005; Stapley and Whiting, aggressive the interactions (Brillet, 1982). Intraspecific 2006; Fernández et al., 2018). Anoles and iguanids are aggressive behaviour among males in natural conditions particularly well studied and exhibit a wide variety of is reported for Oplurus quadrimaculatus Duméril and aggressive behaviours that correspond to territoriality Bibron, 1951 and Oplurus cyclurus (Merrem, 1820), (Johnson and Wade, 2010), predominantly in relation where the area controlled by a single male can exceeded to defending suitable habitats or accessing reproductive 100 m2 (Brillet, 1982). Aggressive interactions have females (DeCourcy and Jenssen, 1994; Lailvaux et been observed also in arboreal species, such as Oplurus al., 2004; Johnson et al., 2009; Johnson and Wade, cuvieri (Gray, 1831) (Randriamahazo and Mori, 1999), 2010). Some agonistic displays seem to be shared and which could suggest territorial behaviour in this species commonly observed across many iguanid lizards, such too. Here we describe an aggressive territorial interaction as head-bobbing, lateral flattening of the back, and between two individuals of Oplurus grandidieri distension of the gular region (Evans, 1953; Murphy, (Mocquard, 1900), a rock-dwelling iguanid from the 1969; Debusk and Glidewell, 1972). highlands of south-central Madagascar. In Madagascar, there are 8 recognized species of During a herpetological survey at the Andringitra iguanids, all of which belong to the endemic family massif (Haute Matsiatra region, Madagascar), two Opluridae, which contains two genera, Chalarodon adult male O. grandidieri were observed displaying Peters, 1854 and Oplurus Cuvier, 1829 (Glaw and aggressive behaviour. Species identity was confirmed Vences, 2007; Münchenberg et al., 2008; Miralles et using samples from two other individuals collected a few al., 2015; Uetz et al., 2019). Territorial behaviour in hundred meters apart from the site of the observation, Opluridae has been first described in captive animals and then sequenced for a fragment of around 650 bp (Blanc and Capenter, 1969; Brillet, 1982; Balcar, 2006), of the standard barcoding region of the cytochrome with contests between individuals, on occasion, resulting oxidase I gene (Nagy et al., 2012), following the in fatalities (Hofstra, 2004). Territorial availability protocol described in Cocca et al. (2018). Sequences seems to be an important determining trigger of a were deposited in GenBank (MN031243, MN031242). These sequences had 3% genetic distance to a sample of O. grandidieri from Isalo (MH063380). The episode took place on 28 November 2018 in Iantaranombe (22°07’46.9”S 46°50’50.7”E, 1588 m 1 CIBIO Research Centre in Biodiversity and Genetic a.s.l.) close to the limit of the Andringitra National Park. Resources, InBIO, Universidade do Porto, Campus Agrário When our team encountered the two males (around de Vairão, Rua Padre Armando Quintas 7, 4485-661 Vairão, 08:00 am), the individuals were standing on a sunlit part Vila do Conde, Portugal. of a large boulder, approximately half a metre apart, 2 Institute of Zoology, Zoological Society of London, Regent’s displaying a typical head-bobbing behaviour (Brillet, Park, London NW1 4RY, UK. 3 Centre for Ecology, Evolution and Environmental Changes 1982; DeCourcy and Jenssen, 1994). The two males (CE3C), Faculdade de Ciências da Universidade de Lisboa, were identified as either the intruder or the resident Campo Grande, 1749-016 Lisboa, Portugal. based on their position when we first saw them. We * Corresponding author. E-mail: [email protected] considered as resident the individual staying in a more 784 Javier Lobón-Rovira et al.

Figure 1. Territorial behaviour of Oplurus grandidieri in Andringitra massif, south-eastern Madagascar: A. lateral flattening (displaying the longitudinal vertebral band) of the resident male, while the intruder head-bobs; B. males approaching face-to-face with their open mouths in intimidation behaviour; C. laterally positioned with vertebral band display; D. resident male lowering head perpendicular to intruder; E. face-to-face with open mouths before physical attack; F. first simultaneous attack with physical aggression; G. vertebral band display; H. intruder retreating while resident male head-bobs. Photos by Gonçalo M. Rosa and Javier Lobon-Rovira.

elevated spot – likely ideal for signalling and basking backs to one another and presenting their light grey/ – and as intruder the one that was approaching the greenish longitudinal vertebral band, while slightly opponent. Both animals placed themselves laterally moving their tail up and down (Fig. 1.A). Both lizards – the resident from an upper position and the intruder kept moving towards each other until they were 5 cm from a lower, flat part of the boulder – side lifting their apart, standing face-to-face with their open mouths (Fig. Territorial behaviour in Oplurus grandidieri from Madagascar 785

1.B). After a few seconds with no physical interaction, for Science and Technology) support the Investigador FCT grant they both retreated to a distance of 30-35 cm, positioned to AC (IF/00209/2014), and the PhD fellowships through the themselves laterally and displayed their backs once BIODIV Doctoral Programme of JLR (PD/BD/140808/2018) and FB (PD/BD/128493/2017). A special thanks to C. Sergeant for his more (Fig. 1.C). This posture included lateral flattening comments on the manuscript. with extension of the leg on the raised body side, while flexing the opposite leg. The resident male then stood References perpendicular to the opponent, raising his shoulders while lowering his head open-mouthed (Fig. 1.D). Balcar, M. (2006): Breeding of madagascan iguanian lizard. Zoo The same display was adopted by the other individual Report Profi 4: 1–4. Blanc, C.P., Carpenter, C.C. (1969): Studies on the Iguanidae of (Fig. 1.E), as Blanc and Carpenter (1969) observed in Madagascar III. Social and reproductive behavior of Chalarodon Chalarodon madagascarensis (Peters, 1854). After a fast, madagascariensis. Journal of Herpetology 3: 125–134. simultaneous attack with physical aggression (Fig. 1.F), Brillet, C. (1982): Contribution a l’étude des relations entre the two lizards retreated once again to approximately individus chez cinq espéces d’iguanes malgaches du genre 30-35 cm apart and positioned themselves laterally (Fig. Oplurus. Revue d’Ecologie (Terre Vie) 36: 79–148. 1.G). The attempted bites by both lizards were directed Bustard, H.R. (1967): The comparative behavior of chameleons: to the head and neck of the opponent. A new display of fight behavior in Chameleo gracilis Hallowell. Herpetologica, the vertebral band led to a face-to-face confrontation, 23: 44–50. Cocca, W., Rosa, G.M., Andreone, F., Aprea, G., Eusebio Bergò, followed by another physical attack. This sequence of P., Mattioli, F., Mercurio, V., Randrianirina, J.E., Rosado, D., displays was observed four times, interspersed with a Vences, M., Crottini, A. (2018): The herpetofauna (Amphibia, gap of a few minutes while individuals moved laterally Crocodylia, Squamata, Testudines) of the Isalo Massif, and leaned to one side to show their dorsal markings. Southwest Madagascar: combining morphological, molecular Head-bobbing and gular expansion, where individuals and museum data. Salamandra 54: 178–200. revealed the orange region of the throat, were often Debusk, J., Glidewell, J.R. (1972): Social dominance in the South displayed during this period. After holding the same American iguanid lizard Plica plica. Journal of Herpetology 6: position for some minutes, the intruder retreated and 139–141. Decourcy, K.R., Jenssen, T.A. (1994): Structure and use of male abandoned its position on the boulder (Fig. 1.H). The territorial headbob signals by the lizard Anolis carolinensis. animals did not bear any visible injuries after the Animal Behaviour 47: 251–262. attacks. Evans, L.T. (1953): Tail display in an iguanid lizard, Liocephalus We described an aggressive encounter between two carinatus coryi. Copeia 1953: 50–54. adult male O. grandidieri. Both males actively displayed Fernández, J.B., Bastiaans, E., Medina, M., de la Cruz, F.R.M., their longitudinal vertebral band to the rival, probably as Sinervo, B.R., Ibargüengoytía, N.R. (2018): Behavioral and a gesture of intimidation. Whilst this behaviour has been physiological polymorphism in males of the austral lizard Liolaemus sarmientoi. Journal of Comparative Physiology A observed in Phelsuma day geckos (Gehring et al., 2010), 204: 219–230. it has not previously been reported for the iguanids of Gehring, P.-S., Crottini, A., Glaw, F., Hauswaldt, S., Ratsoavina, Madagascar. No conspicuous tail displays, typical of F.M. (2010): Notes on the natural history, distribution and other iguanids (e.g. coiling or tail raising; Evans, 1953), malformations of day geckos (Phelsuma) from Madagascar. were recorded during this male-male interaction. This Herpetology Notes 3: 321–327. observation can possibly provide some clues on the Glaw, F., Vences, M. (2007): A Field Guide to the Amphibians and desirable architecture that makes a good quality habitat Reptiles of Madagascar, Third Edition. Köln, Vences & Glaw for this species. The attractive elevated spot with direct Verlag. Hofstra, G. (2004): Oplurus cuvieri cuvieri. European StudyBook sunlight and no apparent visual obstruction may have Foundation: 1–10. served as a trigger for the interaction here described. Huyghe, K., Vanhooydonck, B., Scheers, H., Molina-Borja, M., The report contributed to provide new information on Van Damme, R. (2005): Morphology, performance and fighting O. grandidieri behaviour in the wild. capacity in male lizards, Gallotia galloti. Functional Ecology 19: 800–807. Acknowledgments. We thank local guides Boba Joseph and Johnson, M.A., Revell, L.J., Losos, J.B. (2009): Behavioral Francky Rasolondraibe for assisting with fieldwork. We are convergence and adaptive radiation: effects of habitat use on grateful to MICET and the Malagasy authorities, in particular territorial behavior in Anolis lizards. Evolution 64: 1151–1159. the Ministère de l’Environnement et des Eaux et Forêts, for Johnson, M.A., Wade, J. (2010): Behavioural display systems providing the necessary research permits. This work was funded across nine Anolis lizard species: sexual dimorphisms in by the National Geographic Society (grant number EC-50656R- structure and function. Proceedings of the Royal Society B: 18). The Portuguese National Funds through FCT (Foundation Biological Sciences 277: 1711–1719. 786 Javier Lobón-Rovira et al.

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Accepted by Anamarija Zagar