Human-macaque encounters in Ifrane National Park, Morocco: behavioural, spatial, and temporal coping strategies of the Barbary macaque (Macaca sylvanus)

Wild animals in human-dominated landscapes are exposed to a broad range of human activities and infrastructure that have the potential to alter their fitness and behaviour. Understanding wildlife responses to human disturbance is an important conservation goal, particularly when the fate of many wildlife populations depends on their capacity to coexist with humans. Most field studies have focussed on the effects of one type of disturbance only (i.e. a single disturbance type per study), however in heavily human-modified habitats animals must often deal with numerous different types of human activity and infrastructure, and less is known about whether different types differentially affect animal behavioural responses. This study examined the responses of 5 wild Barbary macaque (Macaca sylvanus) groups in Ifrane National Park, Morocco, to a range of human activities and infrastructure on a short-term behavioural scale (individual focal follows) and a longer-term spatio-temporal scale (individual scan sampling). Using a series of GLMMs I first compared pre-, during-, and post- human-macaque encounter levels of escape, affiliative, and self-directed behaviours (behaviours that may serve as components of a coping strategy) to determine whether different encounter types differentially affect macaque behavioural responses. Using logistic regression to estimate resource selection functions I then examined the behaviour-specific habitat use of macaques to determine whether (and how) different types of human activity and infrastructure influence habitat selection within home ranges. Two broadly consistent responses to human encounters were observed; the first in association with potentially threatening encounters (those that involved/potentially involved dogs), and the second with potentially rewarding encounters (those that involved human provisioning). In response to potentially threatening encounters macaques made extensive use of escape behaviours, varied use of affiliative behaviours, and limited use of self-directed behaviours. In response to potentially rewarding encounters, macaques made use of escape, affiliative, and self-directed behaviours. Three broadly consistent patterns of habitat selection/avoidance were observed in response to human activities and infrastructure within macaque home ranges. Macaques exhibited a general spatial preference for areas close to roads and a general spatial avoidance of both open areas (i.e. with no tree cover) and areas close to herding routes. Macaques also selected/avoided potentially threatening and rewarding areas (i.e. shepherd herding routes and provisioning loci) on a flexible temporal scale, only using potentially threatening areas when the probability of a human encounter was low, and preferentially using rewarding areas only when the risk/benefit balance associated with human provisioning was most heavily in their favour. The results of this study highlight the value of considering the effects of multiple different types of human activity and infrastructure (at different spatio-temporal scales) on wildlife behaviour and welfare, and of assessing human impact on habitat selection separately for different types of behaviour. Such detailed studies on the effects of human activities on wild animal populations can enhance our understanding of, and ability to manage, the impacts of increasing human expansion into wildlife habitat.