Environmental predictability as a cause and consequence of animal movement

The impacts of environmental predictability on the ecology and evolution of animal movement have been the subject of vigorous speculation for several decades. Recently, the swell of new biologging technologies has further stimulated their investigation. This advancing research frontier, however, still lacks conceptual unification and has so far focused little on converse effects. Populations of moving animals have ubiquitous effects on processes such as nutrient cycling and seed dispersal and may therefore shape patterns of environmental predictability. Here, we synthesise the main strands of the literature on the feedbacks between environmental predictability and animal movement and discuss how they may react to anthropogenic disruption, leading to unexpected threats for wildlife and the environment

Communal and efficient movement routines can develop spontaneously through public information use

Animal aggregations occur in almost all taxa and can be strongly influential for consumer-resource dynamics and population health. Their adaptive value and underlying mechanisms are thus fundamental questions. Many animals use information about resource locations inadvertently broadcasted by other individuals through visual, acoustic, or olfactory cues. Such simple, involuntary information transfer is commonly employed in groups of social animals. However, it remains unknown whether public information use could have been the initial cause of social aggregations. Here, using agent-based modeling, in the absence of inclusive fitness benefits or direct conspecific attraction, we show that the use of ephemeral public information about resource locations can cause memory-based foragers to spontaneously and permanently aggregate into communal home ranges that take the form of movement circuits (also called traplines) along which individuals travel asynchronously. Even though experienced individuals only rely on their personal memory to inform their movement decisions, we find that the use of public information during the learning phase is very beneficial in the long term because the communal circuits are more efficient than those established by individuals that do not use public information. Our results reveal how simple, inadvertent information transfer between naïve, selfish foragers can cause the emergence of long-term aggregations, which are a prerequisite for the evolution of more complex social behaviors. They also suggest that individuals may not necessarily need to witness the entire sequences of actions performed by others to converge to the same behavioral routines

Spatial memory shapes density dependence in population dynamics

Most population dynamics studies assume that individuals use space uniformly, and thus mix well spatially. In numerous species, however, individuals do not move randomly, but use spatial memory to visit renewable resource patches repeatedly. To understand the extent to which memorybased foraging movement may affect density-dependent population dynamics through its impact on competition, we developed a spatially explicit, individual-based movement model where reproduction and death are functions of foraging efficiency. We compared the dynamics of populations of with- and without-memory individuals. We showed that memory-based movement leads to a higher population size at equilibrium, to a higher depletion of the environment, to a marked discrepancy between the global (i.e. measured at the population level) and local (i.e. measured at the individual level) intensities of competition, and to a nonlinear density dependence. These results call for a deeper investigation of the impact of individual movement strategies and cognitive abilities on population dynamics

Mobility and its sensitivity to fitness differences determine consumer–resource distributions

An animal's movement rate (mobility) and its ability to perceive fitness gradients (fitness sensitivity) determine how well it can exploit resources. Previous models have examined mobility and fitness sensitivity separately and found that mobility, modelled as random movement, prevents animals from staying in high-quality patches, leading to a departure from an ideal free distribution (IFD). However, empirical work shows that animals with higher mobility can more effectively collect environmental information and better sense patch quality, especially when the environment is frequently changed by human activities. Here, we model, for the first time, this positive correlation between mobility and fitness sensitivity and measure its consequences for the populations of a consumer and its resource. In the absence of consumer demography, mobility alone had no effect on system equilibria, but a positive correlation between mobility and fitness sensitivity could produce an IFD. In the presence of consumer demography, lower levels of mobility prevented the system from approaching an IFD due to the mixing of consumers between patches. However, when positively correlated with fitness sensitivity, high mobility led to an IFD. Our study demonstrates that the expected covariation of animal movement attributes can drive broadly theorized consumer–resource patterns across space and time and could underlie the role of consumers in driving spatial heterogeneity in resource abundance

Foraging efficiency in temporally predictable environments: is a long-term temporal memory really advantageous?

Cognitive abilities enabling animals that feed on ephemeral but yearly renewable resources to infer when resources are available may have been favoured by natural selection, but the magnitude of the benefits brought by these abilities remains poorly known. Using computer simulations, we compared the efficiencies of three main types of foragers with different abilities to process temporal information, in spatially and/or temporally homogeneous or heterogeneous environments. One was endowed with a sampling memory, which stores recent experience about the availability of the different food types. The other two were endowed with a chronological or associative memory, which stores long-term temporal information about absolute times of these availabilities or delays between them, respectively. To determine the range of possible efficiencies, we also simulated a forager without temporal cognition but which simply targeted the closest and possibly empty food sources, and a perfectly prescient forager, able to know at any time which food source was effectively providing food. The sampling, associative and chronological foragers were far more efficient than the forager without temporal cognition in temporally predictable environments, and interestingly, their efficiencies increased with the level of temporal heterogeneity. The use of a long-term temporal memory results in a foraging efficiency up to 1.16 times better (chronological memory) or 1.14 times worse (associative memory) than the use of a simple sampling memory. Our results thus show that, for everyday foraging, a long-term temporal memory did not provide a clear benefit over a simple short-term memory that keeps track of the current resource availability. Long-term temporal memories may therefore have emerged in contexts where short-term temporal cognition is useless, i.e. when the anticipation of future environmental changes is strongly needed

Recursion to food plants by free-ranging Bornean elephant

Plant recovery rates after herbivory are thought to be a key factor driving recursion by herbivores to sites and plants to optimise resource-use but have not been investigated as an explanation for recursion in large herbivores. We investigated the relationship between plant recovery and recursion by elephants (Elephas maximus borneensis) in the Lower Kinabatangan Wildlife Sanctuary, Sabah. We identified 182 recently eaten food plants, from 30 species, along 14 × 50 m transects and measured their recovery growth each month over nine months or until they were re-browsed by elephants. The monthly growth in leaf and branch or shoot length for each plant was used to calculate the time required (months) for each species to recover to its pre-eaten length. Elephant returned to all but two transects with 10 eaten plants, a further 26 plants died leaving 146 plants that could be re-eaten. Recursion occurred to 58% of all plants and 12 of the 30 species. Seventy-seven percent of the re-eaten plants were grasses. Recovery times to all plants varied from two to twenty months depending on the species. Recursion to all grasses coincided with plant recovery whereas recursion to most browsed plants occurred four to twelve months before they had recovered to their previous length. The small sample size of many browsed plants that received recursion and uneven plant species distribution across transects limits our ability to generalise for most browsed species but a prominent pattern in plant-scale recursion did emerge. Plant recovery time was a good predictor of time to recursion but varied as a function of growth form (grass, ginger, palm, liana and woody) and differences between sites. Time to plant recursion coincided with plant recovery time for the elephant’s preferred food, grasses, and perhaps also gingers, but not the other browsed species. Elephants are bulk feeders so it is likely that they time their returns to bulk feed on these grass species when quantities have recovered sufficiently to meet their intake requirements. The implications for habitat and elephant management are discussed

EDITORIAL: Stuck in motion? Reconnecting questions and tools in movement ecology

In this editorial for the Special Feature, I firstly briefly review the major milestones in tool development for movement ecology research, from the first mark–recapture techniques to the current techniques allowing users to collect high-frequency movement data and high-resolution environmental data, as well as the methods for statistical and mathematical analyses. I then briefly describe the methods covered in the Special Feature and conclude with a brief outlook on ongoing and future developments

Cognition in a changing world: Red-headed Gouldian finches enter spatially unfamiliar habitats more readily than do black-headed birds

Human activities are increasingly confronting animals with unfamiliar environmental conditions. For example, habitat change and loss often lead to habitat fragmentation, which can create barriers of unsuitable and unfamiliar habitat affecting animal movements and survival. When confronted with habitat changes, animals’ cognitive abilities play an important, but often neglected part in dealing with such change. Animals must decide whether to approach and investigate novel habitats (spatial neophilia) or whether to avoid them (spatial neophobia) due to potential danger. For species with strict habitat preferences, such as the Gouldian finch (Erythrura gouldiae), which is an open habitat specialist, understanding these novelty responses may be especially important for predicting responses to habitat changes. The Gouldian finch is a polymorphic species, with primarily red or black head colors, which are linked to differing behavioral phenotypes, including novelty reactions. Here we investigate responses to novel habitats (open, dense) in the Gouldian finch, manipulating the color composition of same-sex pairs. Two experiments, each consisting of novel open and novel dense habitat, tested birds in opposite head color combinations in the two experiments. We measured the number of approaches birds made (demonstrating conflict between approach and avoidance), and their entry latency to novel habitats. Gouldian finches showed more approach attempts (stronger approach-avoidance conflict) towards the dense as compared to the open habitat, confirming their open habitat preferences. Black-headed birds also hesitated longer to enter the dense habitat as compared to the open habitat, particularly in experiment 1, appearing less neophilic than red-headed birds, which showed similar entry latencies into both habitat types. This is surprising as black-headed birds were more neophilic in other contexts. Moreover, there was some indication that pairings including at least one black-headed bird had a stronger approach-avoidance conflict than pairings of pure red-headed birds. Results suggest that the black-headed birds use a cognitive strategy typical for residents, whereas red-headed birds use a cognitive strategy known for migrants/nomads, which may cognitively complement each other. However, as 70% of the population in the wild are black-headed, the spatial wariness we document could be widespread, which may negatively affect population persistence as habitats change

Drivers of site fidelity in ungulates

1. While the tendency to return to previously visited locations—termed ‘site fidelity’—is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals’ recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. 2. We compared inter‐year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance‐based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. 3. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren‐ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a ‘win‐stay, lose‐switch’ strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. 4. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter‐annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species‐specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. 5. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change