Quantifying animal movement for caching foragers: the path identification index (PII) and cougars, Puma concolor

Kirsten E. Ironside, David J. Mattson, Tad Theimer, Brian Jansen, Brandon Holton, Terence Arundel, Michael Peters, Joseph O. Sexton, Thomas C. Edwards

Many studies of animal movement have focused on directed versus area-restricted movement, which rely on correlations between step-length and turn-angles and on stationarity through time to define behavioral states. Although these approaches might apply well to grazing in patchy landscapes, species that either feed for short periods on large, concentrated food sources or cache food exhibit movements that are difficult to model using the traditional metrics of turn-angle and step-length alone. We used GPS telemetry collected from a prey-caching predator, the cougar (Puma concolor, Linnaeus), to test whether combining metrics of site recursion, spatiotemporal clustering, speed, and turning into an index of movement using partial sums, improves the ability to identify caching behavior. The index was used to identify changes in movement characteristics over time and segment paths into behavioral classes. The identification of behaviors from the Path Identification Index (PII) was evaluated using field investigations of cougar activities at GPS locations. We tested for statistical stationarity across behaviors for use of topographic view-sheds. Changes in the frequency and duration of PII were useful for identifying seasonal activities such as migration, gestation, and denning. The comparison of field investigations of cougar activities to behavioral PII classes resulted in an overall classification accuracy of 81%. Changes in behaviors were reflected in cougars' use of topographic view-sheds, resulting in statistical nonstationarity over time, and revealed important aspects of hunting behavior. Incorporating metrics of site recursion and spatiotemporal clustering revealed the temporal structure in movements of a caching forager. The movement index PII, shows promise for identifying behaviors in species that frequently return to specific locations such as food caches, watering holes, or dens, and highlights the potential role memory and cognitive abilities play in determining animal movements.