Functional responses, seasonal variation and thresholds in behavioural responses of moose to road density

Understanding the consequences of environmental change on populations is an essential prerequisite for informed management of ecosystems and landscapes. In lieu of quantifying fitness effects directly, which is often difficult, behavioural functional responses provide insight into how animals balance trade-offs, and into thresholds in responses to environmental change. Here, we explore this principle using the response of moose Alces alces L. to roads and restricted-access tracks as a case study. Because roads are associated with the conversion of conifer to mixed deciduousconifer forest that provides better foraging opportunities, moose in Ontario favour areas of moderate road density at a landscape scale. At a finer scale, however, moose avoid roads. These opposing effects indicate a costbenefit trade-off. We quantified behavioural responses of moose to roads using road-crossing rate. An expected distribution of crossing rates was derived from correlated random walk null model simulations. Moose exhibited a seasonally variable, nonlinear functional response in road-crossing rate at the within seasonal range scale. A pronounced response to roads was observed when road density reached approximate thresholds of 0 center dot 2 and 0 center dot 4kmkm2 in summer and winter respectively. Road-crossing rate was proportional to road density, though crossing rates were higher in summer than winter. Crossing rates were best explained by the interaction between mean movement rate and road density. Seasonal differences in road-crossing rate arise from seasonal differences in movement rate and seasonal range area, but not road density within seasonal ranges. Within the protected park, moose did not appear to respond to tracks. Our analysis implies that for the majority of the landscape outside of protected areas the response of moose to roads is pronounced. Synthesis and applications. Identifying thresholds in nonlinear responses to landscape modification is a key management objective as they represent transition zones where small changes can have disproportionately large effects on wildlife populations. We establish these thresholds for moose and roads, but find no response to tracks, implying that the effects of tracks can be mitigated by restricting access to them. We discuss the implications of this work on the problem of moosevehicle collisions.