Topographic determinants of mobile vertebrate predator hotspots: current knowledge and future directions.

Despite being identified as a driver of mobile predator aggregations (hotspots) in both marine and terrestrial environments, topographic complexity has long remained a challenging concept for scientists to visualise and a difficult parameter to estimate. It is only with the advent of high-speed computers and the recent popularisation of geographical information systems (GIS) that terrain attributes have begun to be quantitatively measured in three-dimensional space and related to wildlife dynamics, making the well-established field of geomorphometry (or 'digital terrain modelling') a discipline of growing appeal to biologists. Although a diverse array of numerical metrics is now available to describe the shape, geometry and physical properties of natural habitats, few of these are known to, or adequately used by, ecologists. In this review, we examine the nature and usage of 56 geomorphometrics extracted from the ecological modelling literature over a period of 32 years (1979-2011). We show that, in studies of mobile predators, numerous topographic variables have largely been overlooked in favour of single basic metrics that do not, on their own, fully capture the complexity of continuous landscapes. Based on a simulation approach, we assess the redundancy and correlation structure of these metrics and demonstrate that a majority are highly collinear. We highlight a suite of 7-8 complementary metrics which best explain topographic patterns across a bathymetric grid of the west Australian seafloor, and contend that field and analytical protocols should prioritise variables of these types, particularly when the responses of predator populations to physical habitat features are of interest. We suggest that prominent structures such as canyons, seamounts or mountain chains can serve as useful proxies for predator hotspots, especially in remote locations where access to high-resolution biological data is often limited.