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Variables that were tested reflect mainly boldness, exploration and activity of the tested animals.
Different behaviours were correlated within and between tests and a cluster analysis showed three possible behavioural syndrome-axes, which we name boldness, exploration and activity.
Individual level variation in behaviour can be understood within the context of the animal personality literature, which focuses on broad trait categories including boldness, exploration behaviour, risk taking, aggression and sociability (Bell, Hankison, & Laskowski, 2009; Reale, Reader, Sol, McDougall, & Dingemanse, 2007).
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These behavioural tendencies, personality traits or behavioural syndromes (such as activity, shyness-boldness, exploration, and aggressiveness [30] [31]) can greatly determine how these species respond to changes such as those produced by human development [16], [31] [32].
Scientists have described five major axes of personality variation: shyness-boldness, exploration-avoidance, activity, sociability and aggressiveness.
This result supports the idea that female helpfulness is part of an aggressiveness-boldness-exploration continuum ('behavioural syndrome'), where females of these various behavioural types might follow alternative life-history trajectories correlating with their propensity to provide help [42], [43].
The genes that control melanogenesis in vertebrates pleiotropically influence physiology and behaviour, including boldness and exploration.
Moreover, boldness and exploration relate directly to vulnerability to fishing gear (Alós et al. 2012, 2015; Härkönen et al. 2014; Diaz Pauli et al. 2015).
Consistent among-individual behavioral variation has classically been studied in traits such as boldness-shyness, exploration, aggressiveness, or activity.
Moreover, any changes in energy allocation might be related to metabolic changes (e.g. routine metabolism) or changes in energy acquisition patterns related to behavior (exploration and boldness; Enberg et al. 2012).
It could be that exploration and boldness can be adapted more easily to current environmental conditions like predator presence or food availability, which may have differed between conditions in the wild, which animals experienced prior to the first testing and in the enclosures, prior to the second testing.
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