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Many predators ambush prey rather than pursue them or shift between foraging modes. Active predators typically encounter prey more frequently than ambush predators. I designed a simulation model to examine whether this always holds and how active and ambush predators fare in capturing mobile prey. Prey foraged for clumped resources using area-restricted search, shifting from directional movement before resource encounter to less directional movement afterward. While active predators succeeded more than ambush predators, the advantage of active predators diminished when ambush predators were positioned inside resource patches rather than outside. I investigated the impact of eight treatments and their interactions. For example, regeneration of prey resources increased the difference between ambush predators inside and outside patches, and uncertain prey capture by predators decreased this difference. Several interactions resulted in outcomes different from each factor in isolation. For instance, reducing the directionality level of active predators impacted moderately when applied alone, but when combined with resource regeneration it led to the worst success of active predators against ambush predators inside patches. Ambush predators may not always be inferior to active predators, and one should consider the key traits of the studied system to predict the relative success of these two foraging modes., (© 2024 The Author(s). Annals of the New York Academy of Sciences published by Wiley Periodicals LLC on behalf of The New York Academy of Sciences.)

Desert ant foragers are well known for their visual navigation abilities, relying on visual cues in the environment to find their way along routes back to the nest. If the inconspicuous nest entrance is missed, ants engage in a highly structured systematic search until it is discovered. Searching ants continue to be guided by visual cues surrounding the nest, from which they derive a location estimate. The precision level of this estimate depends on the information content of the nest panorama. This study examines whether search precision is also affected by the directional distribution of visual information. The systematic searching behavior of ants is examined under laboratory settings. Two different visual scenarios are compared - a balanced one where visual information is evenly distributed, and an unbalanced one where all visual information is located on one side of an experimental arena. The identity and number of visual objects is similar over both conditions. The ants search with comparable precision in both conditions. Even in the visually unbalanced condition, searches are characterized by balanced precision on both sides of the arena. This finding lends support to the idea that ants memorize the visual scenery at the nest as panoramic views from different locations. A searching ant is thus able to estimate its location with equal precision in all directions, leading to symmetrical search paths., (© 2023. The Author(s).)

Background: Studying animal movement in the context of the optimal foraging theory has led to the development of simple movement metrics for inferring feeding activity. Yet, the predictive capacity of these metrics in natural environments has been given little attention, raising serious questions of the validity of these metrics. The aim of this study is to test whether simple continuous movement metrics predict feeding intensity in a marine predator, the southern elephant seal (SES; Mirounga leonine), and investigate potential factors influencing the predictive capacity of these metrics., Methods: We equipped 21 female SES from the Kerguelen Archipelago with loggers and recorded their movements during post-breeding foraging trips at sea. From accelerometry, we estimated the number of prey encounter events (nPEE) and used it as a reference for feeding intensity. We also extracted several track- and dive-based movement metrics and evaluated how well they explain and predict the variance in nPEE. We conducted our analysis at two temporal scales (dive and day), with two dive profile resolutions (high at 1 Hz and low with five dive segments), and two types of models (linear models and regression trees)., Results: We found that none of the movement metrics predict nPEE with satisfactory power. The vertical transit rates (primarily the ascent rate) during dives had the best predictive performance among all metrics. Dive metrics performed better than track metrics and all metrics performed on average better at the scale of days than the scale of dives. However, the performance of the models at the scale of days showed higher variability among individuals suggesting distinct foraging tactics. Dive-based metrics performed better when computed from high-resolution dive profiles than low-resolution dive profiles. Finally, regression trees produced more accurate predictions than linear models., Conclusions: Our study reveals that simple movement metrics do not predict feeding activity in free-ranging marine predators. This could emerge from differences between individuals, temporal scales, and the data resolution used, among many other factors. We conclude that these simple metrics should be avoided or carefully tested a priori with the studied species and the ecological context to account for significant influencing factors., (© 2023. The Author(s).)

Given the patchiness and long-term predictability of marine resources, memory of high-quality foraging grounds is expected to provide fitness advantages for central place foragers. However, it remains challenging to characterize how marine predators integrate memory with recent prey encounters to adjust fine-scale movement and use of foraging patches. Here, we used two months of movement data from harbour seals ( Phoca vitulina ) to quantify the repeatability in foraging patches as a proxy for memory. We then integrated these data into analyses of fine-scale movement and underwater behaviour to test how both spatial memory and prey encounter rates influenced the seals' area-restricted search (ARS) behaviour. Specifically, we used one month's GPS data from 29 individuals to build spatial memory maps of searched areas and archived accelerometery data from a subset of five individuals to detect prey catch attempts, a proxy for prey encounters. Individuals were highly consistent in the areas they visited over two consecutive months. Hidden Markov models showed that both spatial memory and prey encounters increased the probability of seals initiating ARS. These results provide evidence that predators use memory to adjust their fine-scale movement, and this ability should be accounted for in movement models.

Background: Diving marine predators forage in a three-dimensional environment, adjusting their horizontal and vertical movement behaviour in response to environmental conditions and the spatial distribution of prey. Expectations regarding horizontal-vertical movements are derived from optimal foraging theories, however, inconsistent empirical findings across a range of taxa suggests these behavioural assumptions are not universally applicable., Methods: Here, we examined how changes in horizontal movement trajectories corresponded with diving behaviour and marine environmental conditions for a ubiquitous Southern Ocean predator, the Adélie penguin. Integrating extensive telemetry-based movement and environmental datasets for chick-rearing Adélie penguins at Béchervaise Island, we tested the relationships between horizontal move persistence (continuous scale indicating low ['resident'] to high ['directed'] movement autocorrelation), vertical dive effort and environmental variables., Results: Penguins dived continuously over the course of their foraging trips and lower horizontal move persistence corresponded with less intense foraging activity, likely indicative of resting behaviour. This challenges the traditional interpretation of horizontal-vertical movement relationships based on optimal foraging models, which assumes increased residency within an area translates to increased foraging activity. Movement was also influenced by different environmental conditions during the two stages of chick-rearing: guard and crèche. These differences highlight the strong seasonality of foraging habitat for chick-rearing Adélie penguins at Béchervaise Island., Conclusions: Our findings advance our understanding of the foraging behaviour for this marine predator and demonstrates the importance of integrating spatial location and behavioural data before inferring habitat use., (© 2021. The Author(s).)

Free-recall tasks suggest human memory foraging may follow a heavy-tailed distribution, such as a Lévy flight, patch foraging, or area-restricted search - walk procedures that are common in other activities of cognitive agents, such as food foraging in both animals and humans. To date, research merely equates memory foraging with hunting in the physical world based on similarities in statistical structure. The current work supports that memory foraging follows a heavy-tailed distribution by using categories with quantitative distances between each item: countries, which have physical distances, and animals, from which cognitive distances can be derived using a multidimensional scaling (MDS) procedure. Likewise, inter-item lag times follow a heavy-tailed distribution. The current work also demonstrates that inter-item distances and times are positively correlated, suggesting the organization of items in memory may be akin to the organization of a physical landscape. Finally, both studies show that participants' original, heavy-tailed lists of countries and animal names produce shorter overall distances traveled than random selection. Human memory foraging follows the same pattern as foraging in the natural world - perhaps because exposure to ecological settings informs our inner cognitive experience - leading to a processing and retrieval time benefit.

Food exploration is an essential survival behavior in organisms. To find food efficiently, many organisms use a foraging strategy called area-restricted search (ARS) wherein individuals first turn more frequently, restricting their search to one area, then turn less frequently, moving along a straight path to widen the search area. Previous research suggests that the nematode Caenorhabditis elegans shows ARS behavior by changing turn frequency, and that dopamine is a crucial determinant. However, the effects of dopamine on multiple behavioral parameters have remained unknown. Here, we evaluated turn (pirouette) frequency, moving velocity, and specific area occupancy (cell occupancy) over time by using a multiple-worms tracking system. In the control (mock) experiments, all parameters changed over time, but no changes were observed in experiments with dopamine pre-exposed and dopamine-deficient animals. In inverse reinforcement learning analysis, the value function for specific velocity was found to modulate over time in mock animals only. These results demonstrate that dopamine regulates ARS via changes not only to pirouette frequency change but also to velocity., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Understanding how animals forage is a central objective in ecology. Theory suggests that where food is uniformly distributed, Brownian movement ensures the maximum prey encounter rate, but when prey is patchy, the optimal strategy resembles a Lévy walk where area-restricted search (ARS) is interspersed with commuting between prey patches. Such movement appears ubiquitous in high trophic-level marine predators. Here, we report foraging and diving behaviour in a seabird with a high cost of flight, the Atlantic puffin ( Fratercula arctica), and report a clear lack of Brownian or Levy flight and associated ARS. Instead, puffins foraged using tides to transport them through their feeding grounds. Energetic models suggest the cost of foraging trips using the drift strategy is 28-46% less than flying between patches. We suggest such alternative movement strategies are habitat-specific, but likely to be far more widespread than currently thought.

We develop a method of analysis for testing the marginal value theorem (MVT) in natural settings that does not require an independent definition or mapping of patches. We draw on recent theoretical work on area-restricted search (ARS) that links turning-angle and step-size changes to geographically localized encounter-rates. These models allow us to estimate "giving-up times" using encounter-annotated GPS tracking data. Applied to a case study of Nahua mushroom foragers, these models identify distinct forms of intrapatch and interpatch search behavior, with intrapatch search transitioning to interpatch search after a predictable interval of time since the last encounter with a harvested mushroom. Our empirical estimate of giving-up time coincides with the theoretically optimal giving-up time derived under the MVT in the same environment. The MVT is currently underused in studies of human foraging and settlement patterns, due in large part to the difficulty of identifying discrete resource patches and quantifying their characteristics. Our methods mitigate the need to make such discrete maps of patches and thus have the potential to broaden the scope for empirical evaluations of the MVT and related theory in humans. Beyond studies of naturalistic foraging in humans and other animals, our approach has implications for optimization of search behavior in a range of applied fields where search dynamics must be adapted to shifting patterns of environmental heterogeneity affecting prey density and patchiness., Competing Interests: The authors declare no conflict of interest., (Copyright © 2019 the Author(s). Published by PNAS.)

Individuals do not have complete information about the environment and therefore they face a trade-off between gathering information (exploration) and gathering resources (exploitation). Studies have shown individual differences in components of this trade-off but how stable these strategies are in a population and the intrinsic drivers of these differences is not well understood. Top marine predators are expected to experience a particularly strong trade-off as many species have large foraging ranges and their prey often have a patchy distribution. This environment leads these species to exhibit pronounced exploration and exploitation phases but differences between individuals are poorly resolved. Personality differences are known to be important in foraging behaviour but also in the trade-off between exploration and exploitation. Here we test whether personality predicts an individual exploration-exploitation strategy using wide ranging wandering albatrosses (Diomedea exulans) as a model system. Using GPS tracking data from 276 wandering albatrosses, we extract foraging parameters indicative of exploration (searching) and exploitation (foraging) and show that foraging effort, time in patch and size of patch are strongly correlated, demonstrating these are indicative of an exploration-exploitation (EE) strategy. Furthermore, we show these are consistent within individuals and appear stable in the population, with no reproductive advantage. The searching and foraging behaviour of bolder birds placed them towards the exploration end of the trade-off, whereas shy birds showed greater exploitation. This result provides a mechanism through which individual foraging strategies may emerge. Age and sex affected components of the trade-off, but not the trade-off itself, suggesting these factors may drive behavioural compensation to maintain resource acquisition and this was supported by the evidence that there were no fitness consequence of any EE trait nor the trade-off itself. These results demonstrate a clear trade-off between information gathering and exploitation of prey patches, and reveals for the first time that boldness may drive these differences. This provides a mechanism through which widely reported links between personality and foraging may emerge., (© 2017 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

In the last thirty years, the emergence and progression of biologging technology has led to great advances in marine predator ecology. Large databases of location and dive observations from biologging devices have been compiled for an increasing number of diving predator species (such as pinnipeds, sea turtles, seabirds and cetaceans), enabling complex questions about animal activity budgets and habitat use to be addressed. Central to answering these questions is our ability to correctly identify and quantify the frequency of essential behaviours, such as foraging. Despite technological advances that have increased the quality and resolution of location and dive data, accurately interpreting behaviour from such data remains a challenge, and analytical methods are only beginning to unlock the full potential of existing datasets. This review evaluates both traditional and emerging methods and presents a starting platform of options for future studies of marine predator foraging ecology, particularly from location and two-dimensional (time-depth) dive data. We outline the different devices and data types available, discuss the limitations and advantages of commonly-used analytical techniques, and highlight key areas for future research. We focus our review on pinnipeds - one of the most studied taxa of marine predators - but offer insights that will be applicable to other air-breathing marine predator tracking studies. We highlight that traditionally-used methods for inferring foraging from location and dive data, such as first-passage time and dive shape analysis, have important caveats and limitations depending on the nature of the data and the research question. We suggest that more holistic statistical techniques, such as state-space models, which can synthesise multiple track, dive and environmental metrics whilst simultaneously accounting for measurement error, offer more robust alternatives. Finally, we identify a need for more research to elucidate the role of physical oceanography, device effects, study animal selection, and developmental stages in predator behaviour and data interpretation.

Foraging and foraging-like processes are found in spatial navigation, memory, visual search, and many other search functions in human cognition and behavior. Foraging is commonly theorized using either random or correlated movements based on Lévy walks, or a series of decisions to remain or leave proximal areas known as "patches". Neither class of model makes use of spatial memory, but search performance may be enhanced when information about searched and unsearched locations is encoded. A video game was developed to test the role of human spatial memory in a canonical foraging task. Analyses of search trajectories from over 2000 human players yielded evidence that foraging movements were inherently clustered, and that clustering was facilitated by spatial memory cues and influenced by memory for spatial locations of targets found. A simple foraging model is presented in which spatial memory is used to integrate aspects of Lévy-based and patch-based foraging theories to perform a kind of area-restricted search, and thereby enhance performance as search unfolds. Using only two free parameters, the model accounts for a variety of findings that individually support competing theories, but together they argue for the integration of spatial memory into theories of foraging., (Copyright © 2015 Elsevier B.V. All rights reserved.)

The benefits of spatial memory for foraging animals can be assessed on two distinct spatial scales: small-scale space (travel within patches) and large-scale space (travel between patches). While the patches themselves may be distributed at low density, within patches resources are likely densely distributed. We propose, therefore, that spatial memory for recalling the particular locations of previously visited feeding sites will be more advantageous during between-patch movement, where it may reduce the distances traveled by animals that possess this ability compared to those that must rely on random search. We address this hypothesis by employing descriptive statistics and spectral analyses to characterize the daily foraging routes of a band of wild hamadryas baboons in Filoha, Ethiopia. The baboons slept on two main cliffs--the Filoha cliff and the Wasaro cliff--and daily travel began and ended on a cliff; thus four daily travel routes exist: Filoha-Filoha, Filoha-Wasaro, Wasaro-Wasaro, Wasaro-Filoha. We use newly developed partial sum methods and distribution-fitting analyses to distinguish periods of area-restricted search from more extensive movements. The results indicate a single peak in travel activity in the Filoha-Filoha and Wasaro-Filoha routes, three peaks of travel activity in the Filoha-Wasaro routes, and two peaks in the Wasaro-Wasaro routes; and are consistent with on-the-ground observations of foraging and ranging behavior of the baboons. In each of the four daily travel routes the "tipping points" identified by the partial sum analyses indicate transitions between travel in small- versus large-scale space. The correspondence between the quantitative analyses and the field observations suggest great utility for using these types of analyses to examine primate travel patterns and especially in distinguishing between movement in small versus large-scale space. Only the distribution-fitting analyses are inconsistent with the field observations, which may be due to the scale at which these analyses were conducted., (© 2013 Wiley Periodicals, Inc.)