Behavior at Habitat Boundaries Can Produce Leptokurtic Movement Distributions

There is increasing recognition that behavioral ecology should be integrated with landscape ecology for a better understanding of the effects of landscape properties on populations (Lima and Zollner 1996; Roitberg and Mangel 1997; Zollner and Lima 1999; Morales and Ellner 2002). In particular, movement behavior affects the way in which individuals redistribute themselves over space and thus has the potential to affect many ecological processes (Kareiva 1990; Tilman and Kareiva 1997; Turchin 1998). Many of these notions come from theoretical studies where the landscape is considered to be homogeneous and connectivity between habitat patches or points in space is either fixed or a simple function of distance. However, landscape ecology is rich in descriptions of land-cover types and spatial patterns that may affect movement (Turner 1989; Wiens et al. 1993; Gustafson and Gardner 1996). It is not well understood how the spatial arrangement of different habitat patches in the landscape matrix affects the movement of individuals. Behavior at habitat boundaries further complicates modeling movement in heterogeneous landscapes. Habitat boundaries may bias turning behavior, reducing permeability of edges to dispersing animals (Stamps et al. 1987). This in turn can affect species interactions (Remer and Heard 1998; Fagan et al. 1999), corridor use, and dispersal (Tischendorf and Wissel 1997; Haddad 1999; Kindvall 1999; Schultz and Crone 2001). Understanding how habitat edges alter ecological processes is believed to be of