Testing for the presence of coping styles in a wild mammal Caterina Ferrari a, b , Cristian Pasquaretta c , Claudio Carere d , Elena Cavallone e , Achaz von Hardenberg b , Denis Réale a, * a Canada Research Chair in Behavioural Ecology, Département des Sciences Biologiques, Université du Québec à Montréal, Montréal, QC, Canada b Alpine Wildlife Research Centre, Gran Paradiso National Park, Valsavarenche, Italy c Dipartimento di Scienze della Terra e dell’Ambiente (DISTA), Università di Pavia, Pavia, Italy d Dipartimento di Scienze Ecologiche e Biologiche, Università degli Studi della Tuscia, Viterbo, Italy e Dipartimento di Scienze Cliniche Veterinarie, Università degli Studi di Milano, Italy article info Article history: Received 22 June 2012 Initial acceptance 21 August 2012 Final acceptance 11 February 2013 Available online xxx MS. number: A12-00482R Keywords: breathing rate cortisol docility heart rate MCMCglmm open-field test personality variance components Coping styles represent alternative response patterns in reaction to a stressor. The coping style model provides a set of predictions about correlations between behavioural and neurophysiological reactions to a stressful situation. According to this model, high levels of activity should be correlated with high levels of aggressiveness at the behavioural level, and to high sympathetic reactivity, low parasympathetic reactivity (higher heart rate levels) and low hypothalamusepituitaryeadrenocortical (HPA) axis reac- tivity (low production of glucocorticoids in response to a stressor). More recent versions of the model, however, are challenging this view and consider the possibility of independent axes of coping style and stress reactivity. The coping style model has mainly been tested on artificially selected or inbred lines in laboratory settings. Such a situation restricts its generalization to a larger number of species and there is a need for studies testing it in the wild under more natural situations. Here, we test the predictions of the coping style model in a wild alpine marmot, Marmota marmota, population. We show that several behavioural (i.e. exploration in an open field, impulsivity and docility) and neurophysiological traits (i.e. heart rate, breathing rate and cortisol production) assumed to represent individual differences in coping style were significantly repeatable over 2e3 years. Not all the correlations between traits predicted by the coping style model were found in marmots, which supports the more recent two-axes model. Furthermore, most correlations were observed at the between-individual level, and the within- individual correlations (i.e. phenotypic plasticity) were weaker. Overall, our results support the predic- tion of the coping style model, but highlight the fact that the association between traits found in artificial conditions may be weaker in a more natural setting. Ó 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. Coping styles are defined as alternative response patterns to a stressor (Koolhaas et al. 1999, 2010). Conceptual studies on coping style generally assume that the coping style model is represented by a set of strongly intertwined traits representing two different strategies: the proactive and the reactive type (Koolhaas et al. 1999; Groothuis & Carere 2005; Ellis et al. 2006; Cockrem 2007). Ac- cording to these studies, we should find strong correlations be- tween physiological and behavioural reactions to a stressor. Compared to reactive individuals, proactive individuals are fast explorers, highly aggressive, actively avoid stressors form routines easily and show little flexibility and sensitivity to cues in their environment (Koolhaas et al. 1999, 2010). Links between the expression of coping styles at the behavioural and neurophysio- logical levels have been described for several model species (e.g. Mus musculus and Rattus norvegicus: Koolhaas et al. 1999; Veenema et al. 2003), and more recently in artificially selected strains of wild species (e.g. Parus major: Drent et al. 2003; Carere et al. 2003; Stöwe et al. 2010; Taeniopygia guttata: Roberts et al. 2007; Onco- rhynchus mykiss: Øverli et al. 2005). Proactive individuals (i.e. bold, active and aggressive animals) are assumed to show high sympa- thetic reactivity and low parasympathetic reactivity, leading to an increase in noradrenaline concentration in the blood, and a high but consistent heart rate under stressors (Koolhaas et al. 1999; Ellis et al. 2006). Proactive individuals are thus assumed to actively cope with the source of stress through a ‘flight-or-fight’ response. In contrast, reactive individuals (i.e. passive, shy and minimally aggressive) show the opposite physiological patterns and tend to react by freezing (Koolhaas et al. 1999). Compared to reactive in- dividuals, proactive ones are assumed to show lower hypothalamuse pituitaryeadrenal (HPA) axis activity and reactivity (Koolhaas * Correspondence: D. Réale, Canada Research Chair in Behavioural Ecology, Département des Sciences Biologiques, Université du Québec à Montréal, 8888 Montréal, QC H3C 3P8, Canada. E-mail address: reale.denis@uqam.ca (D. Réale). Contents lists available at SciVerse ScienceDirect Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav 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.2013.03.030 Animal Behaviour xxx (2013) 1e12 Please cite this article in press as: Ferrari, C., et al., Testing for the presence of coping styles in a wild mammal, Animal Behaviour (2013), http:// dx.doi.org/10.1016/j.anbehav.2013.03.030