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.
2 December 2008 in Proceedings of the National Academy of Sciences
Increasing global concentrations of atmospheric CO 2 are predicted to decrease ocean pH, with potentially severe impacts on marine food webs, but empirical data documenting ocean pH over time are limited. In a high-resolution dataset spanning 8 years, pH at a north-temperate coastal site declined with increasing atmospheric CO 2 levels and varied substantially in response to biological processes and physical conditions that fluctuate over multiple time scales. Applying a method to link environmental change to species dynamics via multispecies Markov chain models reveals strong links between in situ benthic species dynamics and variation in ocean pH, with calcareous species generally performing more poorly than noncalcareous species in years with low pH. The models project the long-term consequences of these dynamic changes, which predict substantial shifts in the species dominating the habitat as a consequence of both direct effects of reduced calcification and indirect effects arising from the web of species interactions. Our results indicate that pH decline is proceeding at a more rapid rate than previously predicted in some areas, and that this decline has ecological consequences for near shore benthic ecosystems.
Patterns of resource selection by animal populations emerge as a result of the behavior of many individuals. Statistical models that describe these population‐level patterns of habitat use can miss important interactions between individual animals and characteristics of their local environment; however, identifying these interactions is difficult. One approach to this problem is to incorporate models of individual movement into resource selection models. To do this, we propose a model for step selection functions (SSF) that is composed of a resource‐independent movement kernel and a resource selection function (RSF). We show that standard case‐control logistic regression may be used to fit the SSF; however, the sampling scheme used to generate control points (i.e., the definition of availability) must be accommodated. We used three sampling schemes to analyze simulated movement data and found that ignoring sampling and the resource‐independent movement kernel yielded biased estimates of selection. The level of bias depended on the method used to generate control locations, the strength of selection, and the spatial scale of the resource map. Using empirical or parametric methods to sample control locations produced biased estimates under stronger selection; however, we show that the addition of a distance function to the analysis substantially reduced that bias. Assuming a uniform availability within a fixed buffer yielded strongly biased selection estimates that could be corrected by including the distance function but remained inefficient relative to the empirical and parametric sampling methods. As a case study, we used location data collected from elk in Yellowstone National Park, USA, to show that selection and bias may be temporally variable. Because under constant selection the amount of bias depends on the scale at which a resource is distributed in the landscape, we suggest that distance always be included as a covariate in SSF analyses. This approach to modeling resource selection is easily implemented using common statistical tools and promises to provide deeper insight into the movement ecology of animals.
27 July 2010 in Philosophical Transactions of the Royal Society B: Biological Sciences
Modern animal movement modelling derives from two traditions. Lagrangian models, based on random walk behaviour, are useful for multi-step trajectories of single animals. Continuous Eulerian models describe expected behaviour, averaged over stochastic realizations, and are usefully applied to ensembles of individuals. We illustrate three modern research arenas. (i) Models of home-range formation describe the process of an animal ‘settling down’, accomplished by including one or more focal points that attract the animal's movements. (ii) Memory-based models are used to predict how accumulated experience translates into biased movement choices, employing reinforced random walk behaviour, with previous visitation increasing or decreasing the probability of repetition. (iii) Lévy movement involves a step-length distribution that is over-dispersed, relative to standard probability distributions, and adaptive in exploring new environments or searching for rare targets. Each of these modelling arenas implies more detail in the movement pattern than general models of movement can accommodate, but realistic empiric evaluation of their predictions requires dense locational data, both in time and space, only available with modern GPS telemetry.
Explaining and predicting animal movement in heterogeneous landscapes remains challenging. This is in part because movement paths often include a series of short, localized displacements separated by longer‐distance forays. This multiphasic movement behavior reflects the complex response of an animal to present environmental conditions and to its internal behavioral state. This state is an autocorrelated process influenced by preceding behaviors and habitats visited. Movement patterns depending on the behavioral state of an animal represent the broad‐scale response of that animal to the environment. Quantifying how animals respond both to local conditions and to their internal state reveals how animals respond to spatial heterogeneity at different spatial scales. We used a state–space statistical approach to model the internal behavioral state and the proximate movement response of elk (Cervus elaphus) to available forage biomass, landscape composition, topography, and wolf (Canis lupus) density during summer in Yellowstone National Park, USA. We analyzed movement paths of 16 female elk fitted with global positioning system (GPS) radio collars that recorded locations at 5‐h intervals. Habitat variables were quantified within 175 m radii (one‐half of the median 5‐h displacement) centered on the beginning location of each interval. Stepwise model selection identified models that best explained the movement distances of each animal. The behavioral state changed very slowly for most animals (median autocorrelation r = 0.93), and all animals responded strongly to time of day (with more movement in the crepuscular hours). However, the spatial variables included in the best‐fitting models varied substantially among individual elk. These results suggest that strong patterns of habitat selection observed in other studies may result from frequent visits to preferred areas rather than a reduction of movement in those areas.