Integrating learning in movement using step-selection analyses

Abstract

1. Understanding how animals acquire and use information from the environment is critical for linking movement to population dynamics, species distributions and conservation. Advances in tracking technologies and growing interest in learning processes have opened opportunities to study behaviours such as habitat exploration in translocated animals or ontogeny of migration and dispersal movements. However, accessible statistical methods for studying these behavioural processes are lacking. 2. We present a learning-explicit step-selection analysis (SSA) that integrates movement, habitat selection, and learning into a single evolving process. At the start of the movement trajectory, the animal is assumed to have no knowledge of the landscape, i.e. its internal habitat quality map is initialized to a constant. As the animal moves, this map is updated dynamically: with each step, only the habitat within its perceptual range becomes known and contributes to future decisions. 3. Through simulations, we show that this method separates true habitat preferences from learning effects and reveals when large-scale behaviours, such as attraction to resources or avoidance of risks, emerge as knowledge accumulates. Critically, we demonstrate that ignoring learning, as in traditional SSA, can lead to biased estimators of habitat selection. 4. Finally, we apply our approach to a real-world GPS dataset of naive individuals, consisting of 10 juvenile red kites dispersing in Switzerland. Using leave-one-individual-out cross-validation, we show that learner SSFs consistently outperform traditional SSFs in predicting movement decisions. 5. We conclude by discussing methodological considerations and future directions for integrating learning into movement ecology. By explicitly modelling the formation of memory that integrates both spatial information and habitat quality, our approach advances process-based movement ecology while remaining compatible with standard SSA workflows, offering practical tools for both theoretical research and applied wildlife management.