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.

As global heating and other anthropogenic influences alter tropical marine environments, it is unclear how marine bird populations will be impacted and whether their current roles in tropical marine ecosystems will change. Although marine birds roost and breed on tropical islands in large numbers, the direct trophic interactions between these birds and their prey across the tropics are poorly documented. We present a first framework for evaluating the dependence on and contributions of marine birds to tropical coral reef ecosystems and use it to examine the evidence for different kinds of interaction, focusing primarily on avian diets. We found 34 publications between 1967 and 2023 that presented a total of 111 data sets with enough detail for quantitative dietary analysis of tropical marine birds. Only two bird species out of 37 (5.4%) had diets of >50% coral reef fishes and only one, the Pacific Reef Egret, appeared to depend almost entirely on reef-based production. Marine birds are also prey for other marine organisms, but insufficient data are available for quantitative analysis. Evidence for indirect effects of birds in tropical marine environments is stronger than for direct dependence on coral reefs, particularly in relation to nutrient concentration and the fertilisation impacts of guano on corals. Dispersal of propagules (e.g. seeds, spores, invertebrate eggs) by bathing, drinking, resting or foraging birds is under-studied and poorly documented. Although the degradation of coral reefs appears unlikely to have a significant direct impact on food availability for most marine bird populations, indirect effects involving marine birds may be disrupted by global environmental change.

The Eurasian woodcock prefers habitats where its main prey, earthworms, can be found in higher densities. Although they are forest-dwelling birds, they regularly visit pastures and natural grasslands at night, where earthworm abundance is generally higher. However, there is little information on fine-scale habitat use in relation to variation in habitat characteristics and prey availability, particularly beyond the breeding season. In our study, we investigated if the nocturnal occurrence of woodcocks during migratory stopover periods differed between two neighbouring fields, or management units, with similar vegetation structure, and if within-field variation in the spatial patterns of woodcock sightings were associated with fine-scale earthworm densities and soil parameters. Specifically, we used GPS tracking data of two tagged woodcocks and direct observation data to study patterns of occurrence of birds in a mixed forest-pasture landscape in Hungary during pre- and post-breeding periods. We compared these patterns with fine-scale soil characteristics and earthworm abundance, acquired by field sampling. We found that the field with higher earthworm abundance was visited by woodcocks more frequently, and this correlation was similarly observed at the intra-field level. Our results demonstrate that woodcocks select foraging sites with higher earthworm densities at multiple spatial scales, both between fields (coarse scale), and within fields (fine-scale). Considering that woodcocks tended to return to the same field to forage at night, the strong associations between occupancy and resources provide a basis for developing habitat management strategies at the field level for conservation. As earthworm densities and soil parameters are good indicators of woodcock foraging habitat, measuring those variables, at least at a coarse scale, could aid in predicting important habitats for the species across the landscape.

Bumblebee activity typically decreases during rainfall, putting them under the threat of the increased frequency of precipitation due to climate change. A novel rain machine was used within a flight arena to observe the behavioural responses of bumblebees (Bombus terrestris) to simulated rain at both a colony and individual level. During rainfall, a greater proportion of workers left the arena than entered, the opposite of which was seen during dry periods, implying that they compensate for their lack of activity when conditions improve. The proportion of workers flying and foraging decreased while resting increased in rain. This pattern reversed during dry periods, providing further evidence for compensatory activity. The increase in resting behaviour during rain is thought to evade the high energetic costs of flying while wet without unnecessarily returning to the nest. This effect was not repeated in individual time budgets, measured with lone workers, suggesting that the presence of conspecifics accelerates the decision of their behavioural response, perhaps via local enhancement. Bumblebees probably use social cues to strategize their energetic expenditure during precipitation, allowing them to compensate for the reduced foraging activity during rainfall when conditions improve.

The activity of many animals follows recurrent patterns and foraging is one of the most important processes in their daily activity. Determining movement in the search for resources and understanding temporal and spatial patterns in foraging has therefore long been central in behavioural ecology. However, identifying and monitoring animal movements is often challenging. In this study we assess the use of camera traps to track a very specific and small-scale interactions focused on the foraging behaviour of Heliconiini butterflies. Data on floral visitation was recorded using marked individuals of three pollen-feeding species of Heliconius (H. erato, H. melpomene and H. sara), and two closely related, non-pollen feeding species (Dryas iulia and Dryadula phaetusa) in a large outdoor insectary. We demonstrate that camera traps efficiently capture individual flower visitation over multiple times and locations and use our experiments to describe some features of their spatial and temporal foraging patterns. Heliconiini butterflies showed higher activity in the morning with strong temporal niche overlap. Differences in foraging activity between males and females was observed with females foraging earlier than males, mirroring published field studies. Some flowers were more explored than others, which may be explained by butterflies foraging simultaneously affecting each other\'s flower choices. Feeding was grouped in short periods of intense visits to the same flower, which we refer to as feeding bouts. Heliconius also consistently visits the same flower, while non-Heliconius visited a greater number of flowers per day and their feeding bouts were shorter compared with Heliconius. This is consistent with Heliconius having more stable long-term spatial memory and foraging preferences than outgroup genera. More broadly, our study demonstrates that camera traps can provide a powerful tool to gather information about foraging behaviour in small insects such as butterflies. © 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

Social learning is a mechanism used by many species to efficiently gain information about their environment. Although many animals live in an environment where members of other species are present, little is known about interspecific social learning. Domesticated and urbanized species provide the opportunity to investigate whether nonhuman animals can learn from heterospecifics such as humans, who are integral parts of their social landscape. Although domestic dogs Canis familiaris have been intensively researched for their ability to learn from humans, most studies have focused on dogs living as pets. However, free-ranging dogs represent the majority of the world\'s dog population, they live alongside humans, scavenge on human refuse, and are subject to natural and sexual selection. Thus, free-ranging dogs with extensive exposure to humans and their artifacts provide the opportunity to investigate interspecific social learning in a naturalistic setting, where learning from humans might be a benefit for them. Here we tested individual free-ranging dogs in a between-subject design: Dogs in the control group could spontaneously choose between two novel and differently patterned food-delivering boxes. In the experimental group, instead, dogs could first observe an unfamiliar human approaching and eating from 1 of the 2 boxes. We provide the first evidence that free-ranging dogs match the choice of an unfamiliar human. These results show that at least simple forms of interspecific social learning might be involved in dogs\' success in living alongside humans in a complex urbanized environment.

The optimal foraging theory posits that animals aim to maximize energy intake while minimizing predation and handling costs during foraging. Most observed animal behaviour supports this theory, but occasional deviations provide insights into the ecological factors that shape foraging decisions. We tested prey-size preference using a two-choice test between different prey sizes in tokay geckos. We expected geckos to prefer larger prey and decision latencies to be longer when discrimination was more difficult and when small prey was offered. Geckos preferred larger prey when the size difference was large, although decision latency remained consistent. This aligns with prior research on sit-and-wait predators. Together with previous findings showing freezing behaviour after prey capture in tokay geckos, our findings suggest a strong influence of predator avoidance on foraging decisions opening up a new avenue for future research investigating the link between decision making and predator avoidance in tokay geckos.

Conservation havens free of invasive predators are increasingly relied upon for fauna conservation, although havened populations can lose anti-predator traits, likely making them less suitable for life \'beyond the fence\'. Sustaining low levels of mammalian predator pressure inside havens may prevent the loss of anti-predator traits from havened populations. We opportunistically compared behavioural and morphological anti-predator traits between four woylie (Bettongia penicillata ogilbyi) populations- one haven isolated from all mammalian predators, one haven containing a native mammalian predator (chuditch; Dasyurus geoffroii), and their respective non-havened counterparts (each containing both chuditch and invasive predators). Havened woylies existing without mammalian predators were smaller (shorter hindfeet, smaller body weight) and less reactive (consumed more food from fox-treated and control feeding stations, less agitated during human handling) than a non-havened reference population. However, in the haven containing chuditch, we found no difference in behaviour or morphology compared to the adjacent non-havened population. Across populations, anti-predator responses tended to appear stronger at sites with higher predator activity, suggestive of an adaptive response across a gradient of predation pressure. Our findings suggest that maintaining mammalian predation pressure in conservation havens could be effective for preventing or slowing the loss of anti-predator traits from these populations.

While various marine predators form associations, the most commonly studied are those between subsurface predators and seabirds, with gulls, shearwaters or terns frequently co-occurring with dolphins, billfish or tuna. However, the mechanisms underlying these associations remain poorly understood. Three hypotheses have been proposed to explain the prevalence of these associations: (1) subsurface predators herd prey to the surface and make prey accessible to birds, (2) subsurface predators damage prey close to the surface and thereby provide food scraps to birds, and (3) attacks of underwater predators lower the cohesion of prey groups and thereby their collective defences making the prey easier to be captured by birds. Using drone footage, we investigated the interaction between Indo-Pacific sailfish (Istiophorus platypterus) and terns (Onychoprion sp.) preying on schooling fish off the eastern coast of the Malaysian peninsula. Through spatio-temporal analysis of the hunting behaviour of the two predatory species and direct measures of prey cohesion we showed that terns attacked when school cohesion was low, and that this decrease in cohesion was frequently caused by sailfish attacks. Therefore, we propose that sailfish created a by-product benefit for the bird species, lending support to the hypothesis that lowering cohesion can facilitate associations between subsurface predators and seabirds.

Cognition helps wildlife exploit novel resources and environments. Raccoons (Procyon lotor) have successfully adapted to human presence, in part due to their cognitive abilities. However, interactions between humans and wildlife can create conflict. A better understanding of the raccoon\'s behavioral flexibility and learning ability could mitigate some conflicts. Our objective was to evaluate wild raccoons learning in contexts varying in terms of exposure to humans (recreational and preservation zoning within protected areas) and task difficulty. Learning can be evaluated over multiple exposures to a cognitive task. Across three years of experiment, we employed 2 food extraction tasks to gauge the change in problem-solving performance over trials. This assessment considered the success probability (the number of successful trials divided by the total number of trials) and the time taken to solve the puzzles. We also looked at the effects of 2 behavioral traits, exploratory diversity and persistence. We found strong evidence for learning over consecutive trials in terms of improved success probability. Improvement in terms of success probability and solving time was more pronounced with the initially easier task. We detected an increase in success probability over trials only in the recreation zones, and there was no evidence of an effect of behavioral traits. The improved performance attributed to learning was also maintained over consecutive years. We provide additional evidence that raccoons can learn how to solve a problem, resulting in a more effective solution in consecutive trials. Finally, we consider the management implications of dealing with raccoons accessing anthropogenic resources.