Placing linkages among fragmented habitats: do least-cost models reflect how animals use landscapes?

P>1. The need to conserve and create linkages among fragmented habitats has given rise to a range of techniques for maximizing connectivity. Methods to identify optimal habitat linkages face trade-offs between constraints on model inputs and biological relevance of model outputs. Given the popularity of these methods and their central role in landscape planning, it is critical that they be reliable and robust. 2. The most popular method used to inform habitat linkage design, least-cost path (LCP) analysis, designates a landscape resistance surface based on hypothetical 'costs' that landscape components impose on species movement, and identifies paths that minimize cumulative costs between locations. 3. While LCP analysis represents a valuable method for conservation planning, its current application has several weaknesses. Here, we review LCP analysis and identify shortcomings of its current application that decrease biological relevance and conservation utility. We examine trends in published LCP analyses, demonstrate the implications of methodological choices with our own LCP analysis for bighorn sheep Ovis canadensis nelsoni, and point to future directions in cost modelling. 4. Our review highlights three weaknesses common in recent LCP analyses. First, LCP models typically rely on remotely sensed habitat maps, but few studies assess whether such maps are suitable proxies for factors affecting animal movement or consider the effects of adjacent habitats. Secondly, many studies use expert opinion to assign costs associated with landscape features, yet few validate these costs with empirical data or assess model sensitivity to errors in cost assignment. Thirdly, studies that consider multiple, alternative movement paths often propose width or length requirements for linkages without justification. 5. Synthesis and applications. LCP modelling and similar approaches to linkage design guide connectivity planning, yet often lack a biological or empirical foundation. Ecologists must clarify the biological processes on which resistance values are based, explicitly justify cost schemes and scale (grain) of analysis, evaluate the effects of landscape context and sensitivity to cost schemes, and strive to optimize cost schemes with empirical data. Research relating species' fine-grain habitat use to movement across broad extents is desperately needed, as are methods to determine biologically relevant length and width restrictions for linkages.