Abstract
Abstract A crucial element in the “the landscape of fear” concept is that prey animals are aware of varying levels of predation risk at a spatial scale. This often leads to a negative spatial relationship between prey and predator in which prey avoid the most risky sites in the landscape. In this paper, we argue that our understanding of large carnivore-ungulate interactions is biased by studies from highly heterogeneous landscapes (e.g. the Yellowstone National Park). Due to a high availability of refuges and foraging sites in such landscapes, prey are able to reduce predation risk by showing habitat shifts. Besides the spatial heterogeneity at the landscape scale, the ungulate response to predation risk can be affected by the hunting mode (stalking vs. cursorial) of the predator. We propose that prey cannot easily avoid predation risk by moving to less risky habitats in more homogenous landscapes with concentrated food resources, especially where the large carnivores’ assemblage includes both stalking and cursorial species. No distinct refuges for prey may occur in such landscapes due to equally high accessibility to predators in all habitats, while concentrated resources make prey distribution more predictable. We discuss a model of a densely forested landscape based on a case study of the Białowieża Primeval Forest, Poland. Within this landscape, ungulates focus their foraging activity on small food-rich forest gaps, which turn out to be “death traps” as the gaps are primarily targeted by predators (stalking lynx and cursorial wolf) while hunting. No alternative of moving to low predation risk areas exist for prey due to risk from wolves in surrounding closed-canopy forest. As a result, the prey is exposed to constant high predation pressure in contrast to heterogeneous landscapes with less concentrated resources and more refuge areas. Future research should focus on explaining how ungulates are coping with predation risk in these landscapes that offer little choice of escaping predation by considering behavioural and physiological (e.g. metabolic, hormonal) responses.
Highlights
Avoidance of predation risk by animals is a widely accepted concept of the dynamic interactions between prey and their predators (e.g. Brown 1999; Verdolin 2006; Hammond et al 2007; Hochman and Kotler 2007; Valeix et al 2009b; Thaker et al 2011; Burkepile et al 2013; Laundré et al 2013; Venter et al 2014)
As prey species do not always have the chance to directly confront their predators, perception of predation risk should often rely on indirect cues
If the strategies of predators’ detection and avoidance by prey as well as searching for and acquiring prey by predators are to be evolutionarily stable (Maynard Smith and Price 1973; Kriva and Cressman 2009), ideally the prey should seek patches richest in food and lowest in predation risk, and the predator should focus on sites where prey is most abundant and/or most vulnerable to predation (Sih 2005)
Summary
Avoidance of predation risk by animals is a widely accepted concept of the dynamic interactions between prey and their predators (e.g. Brown 1999; Verdolin 2006; Hammond et al 2007; Hochman and Kotler 2007; Valeix et al 2009b; Thaker et al 2011; Burkepile et al 2013; Laundré et al 2013; Venter et al 2014). If the strategies of predators’ detection and avoidance by prey as well as searching for and acquiring prey by predators are to be evolutionarily stable (Maynard Smith and Price 1973; Kriva and Cressman 2009), ideally the prey should seek patches richest in food and lowest in predation risk, and the predator should focus on sites where prey is most abundant and/or most vulnerable to predation (Sih 2005) What emerges from these ostensibly conflicting strategies is that it often leads to a negative relationship between the prey and predators’ spatial distribution (Kunkel and Pletscher 2000; Orrock et al 2004; Creel et al 2005; Thaker et al 2011). We argue that prey cannot escape predators in space as shown by many published studies from highly heterogeneous landscapes
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