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The Role of Tusks, Musth and Body Size in Male-Male Competition Among

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The Role of Tusks, Musth and Body Size in Male-Male Competition Among Animal Behaviour 86 (2013) 1207e1214 Contents lists available at ScienceDirect Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav The role of tusks, musth and body size in maleemale competition among Asian elephants, Elephas maximus Karpagam Chelliah, Raman Sukumar* Centre for Ecological Sciences, Indian Institute of Science, Bangalore, India article info The evolution of sexually dimorphic, elaborate male traits that are seemingly maladaptive may be driven e Article history: by sexual selection (male male competition and or female mate choice). Tusk possession in the Asian Received 30 April 2013 elephant is sexually dimorphic and exaggerated but its role in maleemale competition has not yet been Initial acceptance 18 June 2013 determined. We examined the role of the tusks in establishing dominance along with two other known Final acceptance 3 September 2013 maleemale signals, namely, body size and musth (a temporary physiologically heightened sexual state) Available online 10 October 2013 in an Asian elephant population in northeastern India with equal proportions of tusked and tuskless MS. number: 13-00362R males. We observed 116 agonistic interactions with clear dominance outcomes between adult (>15 years) males during 458 field days in the dry season months of 2008e2011. A generalized linear mixed- Keywords: effects model was used to predict the probability of winning as a function of body size, tusk possession Elephas maximus and musth status relative to the opponent. A hierarchy of the three maleemale signals emerged from this maleemale competition analysis, with musth overriding body size and body size overriding tusk possession. In this elephant male secondary character e sexual selection population tusk possession thus plays a relatively minor role in male male competition. An important tuskless male elephant implication of musth and body size being stronger determinants of dominance than tusk possession is that it could facilitate rapid evolution of tuskless males in the population under artificial selection against tusked individuals, which are poached for ivory. Ó 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. Extravagant, ornamental and seemingly useless male traits Antlers are elaborately branched, the tips are often curved back and abound in nature across many taxa, extant and extinct, examples of hardly effective in striking an injurious blow to the opponent the latter being the highly curved tusks of adult mammoths and the (Barrette 1977). Alternatively, weapons and badges of dominance immense antlers of the Irish elk (Gould 1974). In addition to the could have evolved originally in the context of maleemale combat theory of natural selection, Darwin (1871) proposed the mecha- and, subsequently, into ornamental forms through female choice nism of sexual selection to account for the evolution of such (Berglund et al. 1996). exaggerated male traits. The two mechanisms of sexual selection as The evolution and function of tusks in elephants (the African originally proposed by Darwin are maleemale competition and savannah elephant, Loxodonta africana, the African forest elephant, female mate choice. In the former, males compete with each other Loxodonta cyclotis, and the Asian elephant) pose similar challenges to mate with females and the male trait may function as a weapon to biologists as do mammalian horns and antlers. Tusks are elon- or as a signal of fighting ability between males. In the latter, females gated second upper incisors and have been the norm in the pro- actively choose the most splendidly ornamented male with which boscidean fossil record (Osborn 1936, 1942). Male and female to mate. These two mechanisms could act on the same trait and African elephants possess long tusks whereas female Asian ele- affect the strength and direction of selection (Hunt et al. 2009). phants are tuskless or with vestigial tusks that are barely visible. The best-studied examples in mammals of male secondary traits Male elephants may also be tuskless; although extremely rare in as possible weapons in maleemale combat are the horns of bovids male African elephants, the percentage of tuskless males (called (Geist 1966; Bro-Jørgenson 2007) and antlers of cervids (Clutton- makhnas) in Asian elephants varies from about 5% to over 90% Brock 1982, 1987). A fundamental problem in accepting that ant- across different populations (Sukumar 1989; Kurt et al. 1995). lers are weapons in maleemale combat is their inefficient design. The functional role of tusks in the elephant has not been empirically determined but there are anecdotal observations of elephants using their tusks for digging mud for minerals and debarking trees. Such functions, however, could be merely by- * Correspondence: R. Sukumar, Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560012, India. products of, and not the driver of, tusk evolution. No trait acts in E-mail address: [email protected] (R. Sukumar). isolation in interanimal interactions (Hoem et al. 2007) and this is 0003-3472/$38.00 Ó 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.anbehav.2013.09.022 1208 K. Chelliah, R. Sukumar / Animal Behaviour 86 (2013) 1207e1214 clearly evident in maleemale competition in the African elephant using morphological attributes mentioned above to facilitate (L. africana) population of Amboseli, Kenya (Poole 1989a; Hollister- searching and identifying individuals in the database. A comput- Smith et al. 2007). Sexually active males ranging in age from 15 to erized version of the database was implemented in R (R 55 years engage in agonistic interactions to establish dominance. Development Core Team 2013), with Graphical User Interface Two traits, body size and the state of musth, are important de- provided by the package ‘tctltk’. terminants of dominance. Generally, the larger male in a contest Shoulder height was used as a surrogate for age up to 20 years for wins except when the opponent is in a state of musth (Poole 1987, males according to an ageeheight relationship derived from captive 1989a, b; Briffa et al. 2013). Musth is a temporary but intense sexual elephants of known ages and suitably corrected for wild elephants state, expressed by both the African and the Asian elephant (Sukumar et al. 1988; Sukumar 1989). Shoulder heights were (Sukumar 2003), physiologically characterized by sharply elevated measured using three different methods described elsewhere (see plasma testosterone levels and signalled through pheromone- Appendix). Elephants older than 15 years of age were considered as laden secretions from an active temporal gland and pungent adults and classified with some degree of subjectivity into five broad urine dribble (Jainudeen et al. 1972; Poole 1982, 1987; Rasmussen age classes (15e20, 20e30, 30e40, 40e50 and 50þ years) based on a et al. 1984; Ganswindt et al. 2005; Hollister-Smith et al. 2008). combination of features such as ear fold, wrinkled skin, temporal Being in a state of musth is positively associated with high domi- depression and size of the cranium relative to the body. nance status and high reproductive success in the male African elephant (Poole 1989a, b; Hollister-Smith et al. 2007; Rasmussen et al. 2007). Behavioural Observations The role of the tusks in establishing dominance and conse- fi e quently improving male reproductive success, however, has not We de ned competitive male male interactions between a pair ’ been investigated in elephants. It would be difficult to delineate the of males as follows: both males are apparently aware of each other s ’ effects of body size/age and tusk possession as these traits are presence or at least one of the two males is aware of the other s positively correlated with each other. An ideal study population presence and one or more of the behavioural repertoire listed in would have both tusked and tuskless adult males and body size and Table 1 is observed. If two adult males were encountered within tusk length would be uncorrelated. Such populations are found in about 500 m of each other, engaged in activities such as feeding, northeastern India where roughly equal proportions of the two drinking, resting, wallowing in water or mud bathing, they were phenotypes can be found (Sukumar 1989). We therefore examined the correlation between dominance outcome of adult maleemale Table 1 e agonistic interactions and the three presumed male male signals, Behavioural repertoire of adult maleemale agonistic interactions in Asian elephants namely tusks, musth and body size in an Asian elephant population Behaviour Definition at Kaziranga in northeastern India. Nontactile dominant behaviour METHODS Approach An individual pauses its current activity and moves towards another individual Charge An individual suddenly breaks into a run towards the other Study Site and Elephant Population with ears spread out Follow An individual follows another that is walking or running Kaziranga National Park (KNP; 26610e26720N, 93190e away Chase An individual runs behind another individual that is walking 93480E; 993 km2), Assam, India, is located along the floodplains of or running away the Brahmaputra, and comprises mainly riverine habitat, tall and Circular head Vigorous shake of the head from side to side in a plane short grassland, water bodies and patches of tropical semi- shake perpendicular to the ground evergreen forest (Kushwaha 2008). KNP has
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