Memory is critical to understanding animal movement but has proven challenging to study. Advances in animal tracking technology, theoretical movement models and cognitive sciences have facilitated research in each of these fields, but also created a need for synthetic examination of the linkages between memory and animal movement. Here, we draw together research from several disciplines to understand the relationship between animal memory and movement processes. First, we frame the problem in terms of the characteristics, costs and benefits of memory as outlined in psychology and neuroscience. Next, we provide an overview of the theories and conceptual frameworks that have emerged from behavioural ecology and animal cognition. Third, we turn to movement ecology and summarise recent, rapid developments in the types and quantities of available movement data, and in the statistical measures applicable to such data. Fourth, we discuss the advantages and interrelationships of diverse modelling approaches that have been used to explore the memory-movement interface. Finally, we outline key research challenges for the memory and movement communities, focusing on data needs and mathematical and computational challenges. We conclude with a roadmap for future work in this area, outlining axes along which focused research should yield rapid progress.

Analyses of animal movement data have primarily focused on understanding patterns of space use and the behavioural processes driving them. Here, we analyzed animal movement data to infer components of individual fitness, specifically parturition and neonate survival. We predicted that parturition and neonate loss events could be identified by sudden and marked changes in female movement patterns. Using GPS radio-telemetry data from female woodland caribou (Rangifer tarandus caribou), we developed and tested two novel movement-based methods for inferring parturition and neonate survival. The first method estimated movement thresholds indicative of parturition and neonate loss from population-level data then applied these thresholds in a moving-window analysis on individual time-series data. The second method used an individual-based approach that discriminated among three a priori models representing the movement patterns of non-parturient females, females with surviving offspring, and females losing offspring. The models assumed that step lengths (the distance between successive GPS locations) were exponentially distributed and that abrupt changes in the scale parameter of the exponential distribution were indicative of parturition and offspring loss. Both methods predicted parturition with near certainty (>97% accuracy) and produced appropriate predictions of parturition dates. Prediction of neonate survival was affected by data quality for both methods; however, when using high quality data (i.e., with few missing GPS locations), the individual-based method performed better, predicting neonate survival status with an accuracy rate of 87%. Understanding ungulate population dynamics often requires estimates of parturition and neonate survival rates. With GPS radio-collars increasingly being used in research and management of ungulates, our movement-based methods represent a viable approach for estimating rates of both parameters.

State-space models (SSMs) are increasingly used in ecology to model time-series such as animal movement paths and population dynamics. This type of hierarchical model is often structured to account for two levels of variability: biological stochasticity and measurement error. SSMs are flexible. They can model linear and nonlinear processes using a variety of statistical distributions. Recent ecological SSMs are often complex, with a large number of parameters to estimate. Through a simulation study, we show that even simple linear Gaussian SSMs can suffer from parameter- and state-estimation problems. We demonstrate that these problems occur primarily when measurement error is larger than biological stochasticity, the condition that often drives ecologists to use SSMs. Using an animal movement example, we show how these estimation problems can affect ecological inference. Biased parameter estimates of a SSM describing the movement of polar bears (Ursus maritimus) result in overestimating their energy expenditure. We suggest potential solutions, but show that it often remains difficult to estimate parameters. While SSMs are powerful tools, they can give misleading results and we urge ecologists to assess whether the parameters can be estimated accurately before drawing ecological conclusions from their results.

State-space models (SSMs) are an important modeling framework for analyzing ecological time series. These hierarchical models are commonly used to model population dynamics, animal movement, and capture-recapture data, and are now increasingly being used to model other ecological processes. SSMs are popular because they are flexible and they model the natural variation in ecological processes separately from observation error. Their flexibility allows 1 This article has been accepted for publication and undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as

Like many species, movement patterns of southern elephant seals (Mirounga leonina) are being influenced by long‐term environmental change. These seals migrate up to 4,000 km from their breeding colonies, foraging for months in a variety of Southern Ocean habitats. Understanding how movement patterns vary with environmental features and how these relationships differ among individuals employing different foraging strategies can provide insight into foraging performance at a population level. We apply new fast‐estimation tools to fit mixed effects within a random walk movement model, rapidly inferring among‐individual variability in southern elephant seal environment–movement relationships. We found that seals making foraging trips to the sea ice on or near the Antarctic continental shelf consistently reduced speed and directionality (move persistence) with increasing sea‐ice coverage but had variable responses to chlorophyll a concentration, whereas seals foraging in the open ocean reduced move persistence in regions where circumpolar deep water shoaled. Given future climate scenarios, open‐ocean foragers may encounter more productive habitat but sea‐ice foragers may see reduced habitat availability. Our approach is scalable to large telemetry data sets and allows flexible combinations of mixed effects to be evaluated via model selection, thereby illuminating the ecological context of animal movements that underlie habitat usage.