Unveiling the roles of temporal periodicity, the spatial environment and behavioural modes in terrestrial animal movement.

BACKGROUND: Animal movement arises from complex interactions between animals and their heterogeneous environment. To better understand the movement process, it can be divided into behavioural, temporal and spatial components. Although methods exist to address those various components, it remains challenging to integrate them in a single movement analysis. METHODS: We present an analytic workflow that integrates the behavioural, temporal and spatial components of the movement process and their interactions, which also allows for the assessment of the relative importance of those components. We construct a daily cyclic covariate to represent temporally cyclic movement patterns, such as diel variation in activity, and combine the three components in a multi-modal Hidden Markov Model framework using existing methods and R functions. We compare the trends and statistical fits of models that include or exclude any of the behavioural, spatial and temporal components, and perform variance partitioning on the model predictions that included all components to assess their relative importance to the movement process, both in isolation and in interaction. RESULTS: We apply our workflow to a case study on the movements of plains zebra, blue wildebeest and eland antelope in a South African reserve. Behavioural modes impacted movement the most, followed by diel rhythms and then the spatial environment (viz. tree cover and terrain slope). Interactions between the components often explained more of the movement variation than the marginal effect of the spatial environment did on its own. Omitting components from the analysis led either to the inability to detect relationships between input and response variables, resulting in overgeneralisations when drawing conclusions about the movement process, or to detections of questionable relationships that appeared to be spurious. CONCLUSIONS: Our analytic workflow can be used to integrate the behavioural, temporal and spatial components of the movement process and quantify their relative contributions, thereby preventing incomplete or overly generic ecological interpretations. We demonstrate that understanding the drivers of animal movement, and ultimately the ecological phenomena that emerge from it, critically depends on considering the various components of the movement process, and especially the interactions between them.