Connectivity in Heterogeneous Landscapes: Analyzing the Effect of Topography

Animal response to landscape heterogeneity directs dispersal and affects connectivity between populations. Topographical heterogeneity is a major source of landscape heterogeneity, which is rarely studied in the contexts of movement, dispersal, or connectivity. The current study aims at characterizing and quantifying the impacts of topography on landscape connectivity. We focus on ‘hilltopping’ behavior in butterflies, a dispersal-like behavior where males and virgin females ascend to mountain summits and mate there. Our approach integrates three elements: an individual-based model for simulating animal movements across topographically heterogeneous landscapes; a formula for the accessibility of patches in homogenous landscapes; and a graphical analysis of the plots of the simulation-based vs. the formula-based accessibility values. We characterize the functional relationship between accessibility values and landscape structure (referred to as ‘accessibility patterns’) and analyze the influence of two factors: the intensity of the individuals’ response to topography, and the level of topographical noise. We show that, despite the diversity of topographical landscapes, animal response to topography results in the formation of two, quantifiable accessibility patterns. We term them ‘effectively homogeneous’ and ‘effectively channeled’. The latter, in which individuals move toward a single summit, prevails over a wide range of behavioral and spatial parameters. Therefore, ‘channeled’ accessibilities may occur in a variety of landscapes and contexts. Our work provides novel tools for understanding and predicting accessibility patterns in heterogeneous landscapes. These tools are essential for linking movement behavior, movement patterns and connectivity. We also present new insights into the practical value of ecologically scaled landscape indices.