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The importance of wild populations in studies of animal temperament

Archard, Gabrielle A. and Braithwaite, V. A. 2010. The importance of wild populations in studies of animal temperament. Journal of Zoology 281 (3) , pp. 149-160. 10.1111/j.1469-7998.2010.00714.x

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Animal temperament describes behavioural differences between individuals that are consistent across time and contexts. Variation in animal temperament is rapidly gaining interest and attention within behavioural and evolutionary ecology. If we are to understand the causes and consequences of temperament variation within and between populations we need to determine the selection pressures that affect temperament in natural environments. To date, however, the vast majority of temperament studies have been carried out on captive-bred individuals. This review highlights potential problems that arise from using captive animals to elucidate the ecological and evolutionary functions of temperament in wild populations. For example, development, learning and environmental variability can all affect behaviour. Thus, both environment and gene-by environment interactions can affect the fitness functions of different temperaments, and hence selection. We stress the need for measurements of repeatability and heritability, and the importance of biological and ecological validation of temperament tests in wild animals. We describe the limited evidence from wild populations of the fitness consequences of temperament variation, and the use of intra- and inter-specific comparisons to prove adaptation. To identify multiple axes of behavioural variation, and how these interact with environments that vary spatially and temporally, we need long-term studies on wild populations – yet few studies of this nature currently exist. Finally, and perhaps counter-intuitively, we suggest that there is much to be gained from incorporating some of the approaches and statistics employed in the much longer established field of human personality. Consistent differences in behaviour between individuals, known as temperament (or personality), have long been studied by psychologists, using both humans and other animals as subjects (Gosling, 2001; Brosnan, Newton-Fisher & van Vugt, 2009). Research on non-humans has typically occurred in the laboratory, using rodent models to extend knowledge of human temperament and its effects on well-being and health. For example, there are many studies that have addressed addiction (Olmstead, 2006) and anxiety (Pawlak, Ho & Schwarting, 2008). Efforts to improve the production and welfare of domesticated animals have also had an interest in individual differences (Price, 1999). Recently there has been a surge in interest in the effects that temperament has in studies of behavioural ecology. While the existence of individual differences has long been recognized – for example, Krebs & Davies (1987) discussed different behavioural strategies such as ‘producers’ or ‘scroungers’ or individuals that respond differently to risk – the impact that such differences have on behaviour has only recently become of interest for behavioural and evolutionary ecologists. Two main reasons for this have been the lack of consistent terminology (e.g. ‘personality’, ‘temperament’, ‘coping styles’ and ‘behavioural syndrome’ are all found in the literature), and the lack of an ecological and evolutionary framework for temperament studies (Wilson et al., 1994; Gosling, 2001; Sih, Bell & Johnson, 2004a,b; Réale et al., 2007). Now, as has been highlighted in recent reviews, there is increasing awareness that individuals within a population are not always ecologically equivalent (Magurran, 1993; Bolnick et al., 2003), and the fact that temperament affects fitness and selection has been widely recognized (Wilson, 1998; Dall, Houston & McNamara, 2004; Dingemanse & Réale, 2005; Réale et al., 2007; Smith & Blumstein, 2008). These reviews have emphasized the potential evolutionary causes and functions of temperament (e.g. Dall et al., 2004; Sih et al., 2004a,b; Smith & Blumstein, 2008), and have linked temperament studies to other areas of interest to behavioural ecologists, such as behavioural genetics (e.g. van Oers et al., 2005), conservation biology (e.g. McDougall et al., 2006; Réale et al., 2007; Smith & Blumstein, 2008) and behavioural plasticity (e.g. Réale et al., 2007; Dingemanse et al., 2010). Studies of temperament in wild populations are noted in some of these reviews (e.g. Sih et al., 2004a; Réale et al., 2007). However, none have fully discussed the important role that field studies with natural populations play (but, see Dingemanse & Réale (2005) for discussion of the fitness consequences of temperament variation in wild populations). It is well established that individuals within wild populations may show distinct and non-changeable alternative behavioural strategies, which are adaptive and confer different fitness benefits (Magurran, 1993; Gross, 1996; Smith & Skúlason, 1996). In contrast, non-categorical inter-individual variation in behaviour was traditionally seen as noise around averages (Wilson, 1998; Dall et al., 2004), and not itself functional in terms of fitness. Individual variation is, however, a consequence of natural selection (Wilson, 1998; Dingemanse & Réale, 2005; van Oers et al., 2005). Lack of recognition of this fact has led to neglect in the behavioural literature in terms of how and why individuals differ in their behaviour (Wilson, 1998; Dall et al., 2004; Sih et al., 2004a), that is, the fitness functions and evolutionary consequences of the individual variation. Compared with the large number of studies on captive, usually captive-bred, populations, temperament has only rarely been studied in the wild. If the ecological and evolutionary function of temperament is to be elucidated, we need information on the selection pressures that affect temperament in natural environments (Dingemanse & Réale, 2005; Smith & Blumstein, 2008). There may also be important gene by environment interactions that help to shape temperament in natural populations (Merilä & Sheldon, 1999; Nussey, Wilson & Brommer, 2007; Dingemanse et al., 2009) that studies of captive individuals will never reveal. Furthermore, knowledge of how ecological pressures act on temperament in wild populations may aid the maintenance and health of captive populations, for example in aquaculture (Huntingford et al., 2006) and conservation (McDougall et al., 2006). Emphasizing the importance of temperament studies on wild, preferably non-captive, animals is the aim of this review.

Item Type: Article
Date Type: Publication
Status: Published
Schools: Biosciences
Subjects: Q Science > QH Natural history > QH301 Biology
Q Science > QL Zoology
Uncontrolled Keywords: environmental variation; fitness; personality; phenotypic plasticity; selection; temperament; wild populations
Publisher: Wiley-Blackwell
ISSN: 0952-8369
Last Modified: 23 Oct 2017 08:04

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