Behavioural responses of a large, heat-sensitive mammal to climatic variation at multiple spatial scales.

Climate warming creates energetic challenges for endothermic species by increasing metabolic and hydric costs of thermoregulation. Although endotherms can invoke an array of behavioural and physiological strategies for maintaining homeostasis, the relative effectiveness of those strategies in a climate that is becoming both warmer and drier is not well understood. In accordance with the heat dissipation limit theory which suggests that allocation of energy to growth and reproduction by endotherms is constrained by the ability to dissipate heat, we expected that patterns of habitat use by large, heat-sensitive mammals across multiple scales are critical for behavioural thermoregulation during periods of potential heat stress and that they must invest a large portion of time to maintain heat balance. To test our predictions, we evaluated mechanisms underpinning the effectiveness of bed sites for ameliorating daytime heat loads and potential heat stress across the landscape while accounting for other factors known to affect behaviour. We integrated detailed data on microclimate and animal attributes of moose Alces alces, into a biophysical model to quantify costs of thermoregulation at fine and coarse spatial scales. During summer, moose spent an average of 67.8% of daylight hours bedded, and selected bed sites and home ranges that reduced risk of experiencing heat stress. For most of the day, shade could effectively mitigate the risk of experiencing heat stress up to 10°C, but at warmer temperatures (up to 20°C) wet soil was necessary to maintain homeostasis via conductive heat loss. Consistent selection across spatial scales for locations that reduced heat load underscores the importance of the thermal environment as a driver of behaviour in this heat-sensitive mammal. Moose in North America have long been characterized as riparian-obligate species because of their dependence on woody plant species for food. Nevertheless, the importance of dissipating endogenous heat loads conductively through wet soil suggests riparian habitats also are critical thermal refuges for moose. Such refuges may be especially important in the face of a warming climate in which both high environmental temperatures and drier conditions will likely exacerbate limits to heat dissipation, especially for large, heat-sensitive animals.

Spatial-social familiarity complements the spatial-social interface: evidence from Yellowstone bison.

Social animals make behavioural decisions based on local habitat and conspecifics, as well as memorized past experience (i.e. 'familiarity') with habitat and conspecifics. Here, we develop a conceptual and empirical understanding of how spatial and social familiarity fit within the spatial-social interface-a novel framework integrating the spatial and social components of animal behaviour. We conducted a multi-scale analysis of the movements of GPS-collared plains bison (Bison bison, n = 66) residing in and around Yellowstone National Park, USA. We found that both spatial and social familiarity mediate how individuals respond to their spatial and social environments. For instance, individuals with high spatial familiarity rely on their own knowledge as opposed to their conspecifics, and individuals with high social familiarity rely more strongly on the movement of conspecifics to guide their own movement. We also found that fine-scale spatial and social phenotypes often scale up to broad-scale phenotypes. For instance, bison that select more strongly to align with their nearest neighbour have larger home ranges. By integrating spatial and social familiarity into the spatial-social interface, we demonstrate the utility of the interface for testing hypotheses, while also highlighting the pervasive importance of cognitive mechanisms in animal behaviour. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.